Handbuch der Zoologie

Handbook of Zoology
Band/Volume IV Arthropoda: Insecta

Niels P. Kristensen (ed.)

Lepidoptera, Moths and Butterflies
Volume 1: Evolution, Systematics, and Biogeography
Teilband/Part 35
Handbuch der Zoologe
Eine Naturgeschichte der Stmme des Tierreiches

Handbook of Zoology
A Natural History of the Phyla of the Animal Kingdom

Gegrndet von / Founded by Willy Kkenthal

Fortgefhrt von / Continued by M. Beier, M. Fischer, J.-G. Helmcke,
D. Starck, H. Wermuth

Band / Volume IV Arthropoda: Insecta Teilband / Part 35

Herausgeber / Editor Maximilian Fischer

W
DE

G Walter de Gruyter Berlin New York 1999

Niels P. Kristensen (ed.)

Lepidoptera, Moths and Butterflies

Volume 1:
Evolution, Systematics,
and Biogeography

W
DE

G Walter de Gruyter Berlin New York 1999

Autor und Herausgeber Schriftleiter und Herausgeber
Author and Scientific Editor Managing and Scientific Editor

Professor Dr. Niels P. Kristensen Hofrat Univ.-Doz.

Zoological Museum Mag. Dr. Maximilian Fischer
University of Copenhagen Direktor am
Universitetsparken 15 Naturhistorischen Museum Wien
DK-2100 Copenhagen Burgring 7
Denmark A-1014 Wien
Austria

Verlag / Publishers

Walter de Gruyter GmbH & Co. Walter de Gruyter, Inc.

Genthiner Strae 13 200 Saw Mill River Road
D-10785 Berlin Hawthorne, N. Y. 10532
Germany USA

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Handbook of Zoology

agr 2 3 - 1 4 3 6

Printed on acid-free paper which falls within the guide-

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Handbuch der Zoologie : eine Naturgeschichte der

Stmme des Tierreiches / gegr. von Willy Kkenthal.
Fortgef. von M. Beier ... Berlin ; New York : de
Gruyter
Teilw. mit Paralleli.: Handbook of zoology
Bd. 4. Arthropoda: Insecta / Hrsg. Maximilian Fischer
Teilbd. 35. Lepidoptera, moths and butterflies
Vol. 1. Evolution, systematics, and biogeography /
Niels P. Kristensen (ed.). - 1998
ISBN 3-11-015704-7
Copyright 1998 by Walter de Gruyter GmbH & Co.,
Berlin. All rights reserved, including those of translation
into foreign languages. No part of this book may be
reproduced in any form - by photoprint, microfilm or
any other means nor transmitted nor translated into a
machine language without written permission from the
publisher. - Printed in Germany.
Typesetting and Printing: Arthur Collignon GmbH, Ber-
lin. - Binding: Lderitz & Bauer G m b H , Berlin.
 Editor's preface
Thirty years have elapsed since the publication
of the first issues in this second edition of the
Insecta section of the Handbuch der Zoologie!
Handbook of Zoology. Not surprisingly there-
fore, editorial principles and standards are now
changing to ensure that the series can continue
to fulfill its intended role in the biological refer-
ence literature.
While treatments of the majority of the small
and mediumsized insect groups conventionally
ranked as 'orders' have been issued already, only
a single treatment of a 'megadiverse' order has
been completed, viz., Willi Hennig's 1973 volume
on the Dipera. By the 1980s it had become clear
that accounts of the larger insect groups of the
breadth and depth required in the Handbuch/
Handbook series were far beyond the capacity of
any single author or even of a small team of
authors. The present Lepidoptera volume, then,
is the first example of the genuine multi-author
treatment which will be adopted in forthcoming
volumes on the large insect orders. Moreover, it
is our aim that future volumes in the series focus-
sing on systematic entities will be published in
English.
Two Lepidoptera volumes are planned. The
present volume on systematics, evolution and
biogeography will be followed by one on mor-
phology and physiology. The latter is in an ad-
vanced stage of preparation. Plans for a third vol-
ume dealing with Lepidoptera ecology and behav-
iour were abandoned after consultation with a
number of potential editors: While ecological and
behavioural research on moths and butterflies has
long been vigorous, there is greater motivation
for reviewing findings emerging therefrom in
contexts other than one centred just on the order
Lepidoptera. Needless to say, a certain body of
information on ecology and behaviour is indeed
being provided in relevant places in both volume
1 and 2. Moreover, volume 1 was broadened to
accommodate a chapter on larval food relation-
ships, a topic which certainly can be appropri-
ately summarized in a Lepidoptera context, i.e.,
within a large-scale evolutionary framework.
Cross references between chapters in this two-
volume Handbook treatment of the Lepidoptera
is eased by means of emboldened volume-and-
chapter numbers; thus 1 - 8 and 28 refer to
chapter 8 in volume 1 and 2 respectively.
Name sequence of contributors to coauthored
chapters is alphabetical (except in the case of
chapter 120, where this principle was aban-
doned at an express request from within the
author team), hence it does not necessarily indi-
cate who was the principal contributor (if the
workload was indeed unevenly shared). How-
ever, in the case of some chapters where the
responsibility for different section has been
clearly divided between authors, this has been
explained.
As is usual in works of this kind an approxi-
mate total length of the volumes was agreed
from the outset. The subsequent page allocation
to individual chapters was evidently fraught with
problems. As far as the systematics chapters are
concerned a balance was sought between criteria
concerning species richness, diversity (there is
not necessarily a simple correlation of the two),
importance in a phylogenetic context, biological
phenomena of general interest, and knowledge
available. Unquestionably different traditions in
the study of different major lineages, as well as
contrasting maturity of their classifications, con-
tribute to any imbalance that may remain in the
treatment. For instance, while all tortricoid
moths are included currently in just a single fam-
ily, a globally applicable tribal taxonomy is now
available for this lineage; the detailed treatment
down to this level is likely to be welcomed by
users. By contrast, the preliminary state of the
higher classification of the much more species-
rich family Geometridae largely explains the rel-
atively more brief treatment of this taxon. Since
most authors probably considered alloted space
inadequate for a proper treatment of their sub-
jects), exesses were the rule rather than the ex-
ception! Some manuscripts were shortened by
the editor. But variation between authors in ob-
serving space limitations remains another source
of imbalance in chapter length.
Differences between individual chapters in
style and presentation have been left unharmo-
nized, as long as they are unlikely to create con-
fusion. For instance, the use of both Roman and
Arabic numerals to designate abdominal segment
numbers has been accepted. Similarly, examples
of both English and American spelling will be
found.
While works of this kind are intended primar-
ily to provide state-of-the-art reviews, the prepa-
ration of contributions for the present volume
prompted some authors to undertake a substan-
tial amount of original research. Hence the
content of some chapters is truly innovative.
Necessarily, subjective decisions had to be made
concerning what could appropriately be included
in the treatment published here. For instance, the
sizable original analyses of butterfly inter-rela-
tionships upon which chapter 1 16 is largely
based were found to exceed the scope of a Hand-
book chapter, and they were accordingly pub-
lished as a separate journal article (de Jong, R.,
Vane-Wright, R. I. & Ackery, R R.: Ent. scand.
VI
27: 6 5 - 1 0 1 , 1996). But it was decided that the
similarly innovative analysis of Gelechioid phy-
logeny, which forms the basis of 19, should be
included, with the primary material printed as
appendices.
While some contributions were completed just
before type-setting of the entire colume was initi-
ated, others had been completed considerably
earlier. However, most authors were able to take
the opportunity to make additions to their con-
tributions in 1997, so the volume as a whole will
appear reasonably updated by the time of its
publication.
Editor's preface
As an editor I am greatly indebted to the team
of authors for their commitment, faith and pa-
tience. I similarly thank the numerous specialists
who provided reviews of individual chapters. Fi-
nally I thank D r Maximilian Fischer (former sole
editor, Insecta section of the Handbook series)
as well as P D Dr Mario Noyer-Weidner and Ms
Christiane Bowinkelmann (Walter de Gruyter,
Biosciences Division) and Ms Ingeborg Klak
(Walter de Gruyter, Production Department) for
their collaboration on this enterprise.
Copenhagen, April 1998
Niels P. Kristensen
 Contents

Editor's preface V 12. The Tortricoidea.

Contributors IX Marianne Horak 199
1. Historical Introduction. 13. The Smaller Microlepidopteran-Grade
Niels P. Kristensen 1 Superfamilies.
John S. Dugdale, Niels P. Kristensen,
2. Phylogeny and Palaeontology.
Gaden S. Robinson & Malcolm J. Scoble 217
Niels P. Kristensen & Andrzej W. Skalski 1
14. The Pyraloidea.
3. Classification and Keys to Higher Taxa.
Eugene Munroe & M. Alma Solis . . . . 233
David J. Carter & Niels P. Kristensen. . 27
4. The Non-Glossatan Moths. 15. The Axioidea and Calliduloidea.
Niels P. Kristensen 41 Jol Minet 257
5. The Homoneurous Glossata. 16. The Butterflies: Hedyloidea,
Niels P. Kristensen 51 Hesperioidea and Papilionoidea.
Phillip R. Ackery, Rienk de Jong &
6. The Monotrysian Heteroneura. Richard I. Vane- Wright 263
Donald R. Davis 65
17. The Drepanoid/Geometroid
7. The Tineoidea and Gracillarioidea.
Assemblage.
Donald R. Davis & Gaden S. Robinson 91
Jol Minet & Malcolm J. Scoble 301
8. The Yponomeutoidea.
18. The Bombycoidea and their Relatives.
John S. Dugdale, Niels P. Kristensen,
Claude Lemaire & Jol Minet 321
Gaden S. Robinson & Malcolm J. Scoble 119
19. The Noctuoidea.
9. The Gelechioidea.
Ian J. Kitching & John E. Rawlins . . . 355
Ronald W. Hodges 131
10. The Zygaenoidea. 20. Evolution of Larval Food Preferences
Marc E. Epstein, Henk Geertsema, Clas in Lepidoptera.
M. Naumann & Gerhard M. Tarmann 159 Jerry Powell, Charles Mitter & Brian
Farrell 403
11. The Cossoid/Sesioid Assemblage.
Edward D. Edwards, Patricia Gentili, 21. Biogeography of the Lepidoptera.
Niels P. Kristensen, Marianne Horak & Jeremy D. Holloway <6 Ebbe S. Nielsen 423
Ebbe S. Nielsen 181 Index of Scientific Lepidoptera Names . . . 463
 Contributors
P. R. Ackery
Department of Entomology
The Natural History Museum
Cromwell Road
London SW7 5BD, England
email: pra@nhm.ac.uk
D. J. Carter
Department of Entomology
The Natural History Museum
Cromwell Road
London SW7 5BD, England
email: d.carter@nhm.ac.uk
Donald R. Davis
Department of Entomology
National Museum of Natural History
Smithsonian Institution
Washington, DC 20560, USA
email: Davis.Don@NMNH.SI.edu
John S. Dugdale
do Landcare Research
Private Bag 6
Nelson, New Zealand
email: DugdaleJ@landcare.cri.nz
E. D. Edwards
Entomology Division, CSIRO
PO Box 1700, Canberra City
ACT 2601, Australia
email: ted.edwards@ento.csriso.au
Marc E. Epstein
Department of Entomology
National Museum of Natural History
Smithsonian Institution
Washington, DC 20560, USA
email: epstein. marc@nmnh.si.edu
Brian Farrell
Museum of Comparative Zoology
Harvard University
Cambridge, MA 02138, USA
email: farrellb@oeb.harvard.edu
Patricia Gentili
Department of Entomology
National Museum of Natural History
Smithsonian Institution
Washington, DC 20560, USA
email: MNHEN087@SIVM.SI.EDU
Henk Geertsema
Department of Entomology and Nematology
University of Stellenbosch
Stellenbosch, 7600 South Africa
email: hge@maties.sun.ac.za
Ronald W. Hodges
Systematic Entomology Laboratory, USDA
Present address: 85253 Ridgetop Drive
Eugene, OR 97405-9535, USA
email: rwhodges@continet.com
J. D. Holloway
do Department of Entomology
The National History Museum
Cromwell Road
London SW7 5BD, England
email: jdh@nhm.ac.uk
Marianne Horak
Entomology Division, CSIRO
PO Box 1700, Canberra City
ACT 2601, Australia
email: marianne@ento.csiro.au
R. de Jong
Department of Entomology
Naturalis/Nationaal Naturhistorisch Museum
Postbus 9517
NL-2300 RA Leiden, Netherlands
email: jong@nnm.nl
Ian J. Kitching
Department of Entomology
The National History Museum
Cromwell Road
London SW7 5BD, England
email: ijk@nhm.ac.uk
Niels P. Kristensen
Department of Entomology
Zoological Museum
University of Copenhagen
DK-2100 Copenhagen 0 , Denmark
email: npkristensen@zmuc.ku.dk
Claude Lemaire
La Croix des Baux
F-84220 Gordes, France
Jol Minet
Musum National d'Histoire Naturelle
Entomologie
45, Rue de Buffon
F-75005 Paris, France
email: minet@mnhn.fr
Charles Mitter
Dept of Entomology
University of Maryland
College Park, Maryland 20742, USA
email: Charles_MITTER@umail.umd.edu

Eugene Munroe
3093 Barlow Crescent
R.R. 1 Dunrobin
Ontario, KOA ITO, Canada
Clas M. Naumann
Zoologisches Forschungsinstitut
und Museum Alexander Koenig
Adenauerallee 160
D-53113 Bonn 1, Germany
email: c.naumann@zfmk.uni-bonn.de
Ebbe S. Nielsen
Entomology Division, CSIRO
PO Box 1700, Canberra City
ACT 2601, Australia
email: ebben@ento.csiro.au
Jerry Powell
Department of Entomological Sciences
201 Wellmann Hall
University of California
Berkeley, California 947200-0001, USA
email: powellj@nature.berkeley.edu
John E. Rawlins
Department of Invertebrate Zoology
Carnegie Museum of Natural History
4400 Forbes Avenue
Pittsburgh, PA 15213, USA
email: rawlinsj@clpgh.org
Contributors
Gaden S. Robinson
Department of Entomology
The Natural History Museum
Cromwell Road
London SW7 5BD, England
email: gsr@nhm.ac.uk
Malcolm J. Scoble
Department of Entomology
The Natural History Museum
Cromwell Road
London SW7 5BD, England
email: mjs@nhm.ac.uk
Andrzej W. Skalski (deceased)
Pedagogical University in Czestochowa
Poland
M. Alma Solis
Systematic Entomology Laboratory, USDA
c/o National Museum of Natural History
Washington, DC 20560, USA
email: asolis@sel.barc.usda.gov
Gerhard M. Tarmann
Tiroler Landesmuseum
Museumstrasse 15
A-6020 Innsbruck, Austria
email: gtarmann@ping.at
R. I. Vane- Wright
The Natural History Museum
Department of Entomology
Cromwell Road
London SW7 5BD, England
email: riw@nhm.ac.uk
 1. Historical Introduction
Niels P. Kristensen
While the references in the following chapters of
these volumes by necessity will be focussed on
twentieth-century works, a brief outline is given
below of principal pre-1900 contributions to the
foundation of contemporary lepidopterology (as
covered in the present Handbook).
Systematics and Bionomics Before 1860
The Lepidoptera include some of the most con-
spicuous and attractive of all insects, and their
striking metamorphosis lends itself well to obser-
vation. Therefore, the book which demarcates
the beginning of the current naming and classifi-
cation of animals, C. Linnaeus' Systema Naturae
(tenth edition, 1758), could already refer to a
substantial body of published work on lepidop-
teran natural history. Principal references include
T. Mouffet's Insectorum sive Minimorum Ani-
malium Theatrum (published posthumously 1634
and including significant contributions by other
naturalists, notably the Swiss scientific pioneer
K. Gesner, 151665), J. Goedart's Metamorpho-
sis Naturalis (1662-67), Maria S. Merian's il-
lustrated accounts of European Lepidoptera as
well as her renowned Metamorphosis insectorum
Surinamensium (1705), J. Ray's Historia Insec-
torum (1710), J. Petiver's Papilionum Britanniae
icones (1717), . Wilkes' English Moths and But-
terflies (1747-60), A. J. Rsel von Rosenhof's
Die monatlich herausgegebene Insekten-Belusti-
gung 1 - 4 (1746-61), A. R. Reaumur's M-
moires pour servir l'histoire des insectes 16
(1734-42) and K. de Geer's similarly titled work
( 1 - 7 , 1752-78).
Linnaeus, Fabricius and Latreille
Linnaeus' system profoundly influenced later
classifications. His basic groups in the Lepidop-
tera Papilio (Eques, Danaus, Nymphalis, Plebe-
jus, Barbarus), Sphinx, and Phalaena (Bombyx,
Noctua, Geometra, Tortrix, Pyralis, Tinea, Alu-
cita) persisted in some general zoology text-
books for more than a century, although in the
meantime Lepidoptera classification had un-
dergone considerable refinement in specialists'
writings. Linnaeus' pupil J. C. Fabricius de-
scribed numerous lepidopteran species from
many parts of the world and subdivided several
Linnean groups into what were to become recog-
nized as valid genera; his latest and most com-
prehensive works dealing with Lepidoptera are
Entomologica Systematica (179294) and Sys-
tema Glossatorum (1807, incomplete). P. A. Lat-
reille similarly refined the system by dividing
Linnean supraspecific taxa.
Early works on European Lepidoptera
While Fabricius' descriptive works all had a
global scope, Linnaeus himself had devoted a
special publication to the treatment of his local
(national) fauna: Fauna Svecica (1746, first no-
menclatorically valid edition 1761), and this ex-
ample was soon followed by workers in many
European countries. Principal early 'faunas'
from Central Europe, with many original species
descriptions, include J. A. Scopoli's Entomologia
Carniolica (1763), J. N. C. M. Denis & I. Schif-
fermller Ankndigung eines systematischen Wer-
kes von den Schmetterlingen der Wienergegend
(1776), and F. von Schranck's Fauna Boica 13
(1798-1804); the lastmentioned work was the
first to expand the Linnean classificatory hierar-
chy for Lepidoptera (see Emmet 1991). Among
British works M. Harris' The Aurelian or Natural
History of English Insects, namely Moths and
Butterflies (1766) is renowned for its exquisite
coloured illustrations, but truly comprehensive
accounts of the British fauna only came with
A. H. Haworth's Lepidoptera Britannica (1803
28), followed by two works of a more modern
scope, J. F. Stephen's Illustrations of British En-
tomology: Haustellata (1827-35) and Curtis'
British Entomology (1823-40). Principal early
works on Lepidoptera in the Mediterranean sub-
region are by J. P. Rambur, including Catalogue
systmatique des Lpidoptres de l'Andalusie
(1858-66, 'Macrolepidoptera' only). Principal
pioneering contributions to the knowledge of the
lepidopteran fauna of Russia (including Siberia)
are the works of E. F. Eversmann, e. g. Fauna
Lepidopterologica Volgo-Uralensis (1844).
Several important works on the European
fauna were of a broader geographical scope.
Early examples were E. J. C. Esper's Die Schmet-
terlinge in Abbildungen nach der Natur 15
(177794 [second edition, with additions, by T.
von Charpentier 1829-39]) and M. B. Bork-
hausen's Naturgeschichte der Europischen
Schmetterlinge 1 - 5 (1788-94). Later followed
the influential ten-volume treatise Die Schmetter-
linge von Europa (1807-34), initiated by F. Och-
senheimer and completed by his collaborator F.
Treitschke. Its title notwithstanding the Histoire
Naturelle des Lpidoptres de France, initiated in
1820 by J. B. Godart and completed (in 11 vol-
2 Niels P. Kristensen
umes, with four supplements) in 1842 by P. A. J.
Duponchel also was Europe-wide in scope.
J. B. A. D. de Boisduval's Genera et Index Me-
thodicus Europaeorum Lepidopterorum (1840)
catalogued the European 'Macrolepidoptera',
with Guene's incomplete Europaeorum Micro-
lepidopterorum Index Methodicus (1845) as a
companion volume; these works were paralleled
by Duponchel's Catalogue mthodique des Lpi-
doptres d'Europe (1844-46). The artist J.
Hbner's volumes of outstanding coloured
plates, Sammlung europischer Schmetterlinge
1796[1838] and Geschichte europischer Schmet-
terlinge [1793][1842], deserve particular men-
tion; he also published illustrations of extra-
European Lepidoptera (cp. below) and intro-
duced a wealth of new taxon names. Hbner's
work was continued and completed by G. A. W.
Herrich-Schffer and the painter C. Geyer, The
former's six-volume supplement to Hbners
Sammlung, entitled Systematische Bearbeitung
der Schmetterlinge von Europa (1843 1856), is a
milestone in Lepidoptera systematics with a fam-
ily classification largely foreshadowing that
adopted today. Herrich-Schffer made extensive
use of characters provided by the wing venation
(preliminary studies on which had been made by
some earlier workers), but also took characters
of e. g. vestiture, palpi, and legs into account. J.
Lederer's Die Noctuiden Europas (1857) is one of
the first works in which characters in the male
genital apparatus are used (an even earlier use
was in Rambur's study of Andalusian skipper
butterflies). Of outstanding significance are P. C.
Zeller 's numerous works on micro-moths; his
1839 article (in Oken's Isis) Versuch einer natur-
gemssen Eintheilung der Schaben (meaning Lin-
nean 'Tinea', not cockroaches) was followed by
a suite of high-quality monographs of several
micro-moth groups, and eventually his writings
were extended to extra-European faunas. He fur-
ther contributed, as did H. Frey and J. W. Doug-
las, to the unique and lavishly illustrated 13-vol-
ume The Natural History of the Tineina (1855
73) published (with text in English, French, Ger-
man and Latin) by H. T. Stainton.
Early works on the world fauna
Expectedly, local/regional 'faunas' by resident
workers outside Europe were few in this period.
J. Abot & J. E. Smith's Natural History of Rarer
Lepidopterous Insects of Georgia 12 (1797) and
E. Blanchard's treatment (1852) of the Lepidop-
tera in C. Gay (ed.) Historia fisica y politica de
Chile are examples.
Already in the earliest phase of descriptive
taxonomy P. Cramer's De Uitlandsche Kapellen
in de drie warereld-deelen Asia, Africa en America
(177582, with posthumous supplements by C.
Stoll in 1782 and 1787-1790) provided substan-
tial pictorial documentation of extra-European
Lepidoptera; so did D. Drury's Illustrations of
Natural History 1 - 3 (1770-82). Hbner's
Sammlung exotischer Schmetterlinge (1806
[38]), completed by Geyer and supplemented by
Herrich-Schffer's Sammlung neuer oder wenig be-
kannter auereuropischer Schmetterlinge (1850
1869) time-wise overlapped the author's works
on the European fauna, and the scope of Her-
rich-Schaeffer's system was broadened by the in-
clusion of exotic taxa. The Spcies Gnral des
Lpidoptres (eight volumes, 1836-75) by Bois-
duval & A. Guene in 'Nouvelles Suites Buffon'
was planned as a complete treatise of the lepi-
dopteran species, but remained incomplete. E.
Doubleday & J. O. Westwood's The genera of di-
urnal Lepidoptera (1846-1852), illustrated by
W. C. Hewitson is the foundation for modern
butterfly classification, while F. Walker's List of
Specimens of Lepidopterous Insects in the Collec-
tion of the British Museum (35 parts, 18541866)
is the last synopsis of the World's moths to have
been completed.
Systematics and Bionomics 18601900
C. Darwin's On the Origin of Species is a turning
point in biological history. The principal contri-
bution of Lepidoptera research to the early shap-
ing of evolutionary biology came with the mim-
icry theory, first formulated in H. W. Bates' A
Naturalist on the Amazonas (1861) and later elab-
orated on by several lepidopterists including
A. R. Wallace and F. Mller. Towards the end of
the century K. Jordan's theoretical work {Novit.
Zool. 3, 1896) rooted in Lepidoptera taxonomy
laid an important foundation for the later preva-
lent species concept. Truly important progress
towards an evolutionary higher classification of
the order were made in the last decade of the
century, particularly through studies on imma-
ture stages by T. A. Chapman (e. g., Trans, ent.
Soc. London 1893, 1896) and on wing structure
by A. Spuler (Z. Wiss. Zool 53, 1892) as well as
by J. Comstock and J. Needham (articles in Am.
Nat., also Comstock's Manual for the Study of
Insects, 1895).
Inventory of the World fauna
Outstanding among the European 'fauna hand-
books' of the period was Die Schmetterlinge
Deutschlands und der Schweiz 1 - 2 (1863-77) by
H. von Heinemann & M. F. Wocke. The Catalog
der Lepidopteren Europas by O. Staudinger &
M. F. Wocke (1861) and its later editions in 1871
and 1901 (the lastmentioned, covering the entire
Palaearctic region, was published by H. Rebel)
have been invaluable tools. Major works on
central- and eastern-Palaearctic taxa by H. T.
Christoph, S. N.-Alpheraky and O. Staudinger
appeared in the series Mmoires sur les Lpidop-
Historical Introduction
teres edited by . M. Romanoff from 1884 on-
wards. Principal comprehensive works on Lepi-
doptera from E. and SE. Asia include J. H.
Leech's Butterflies from China, Japan and Corea
(1892-94), W. L. Distant's Rhopalocera Malay-
ana (188286), F. Moore's The Lepidoptera of
Ceylon 1 - 3 (1880-87) and G. F. Hampson's
four-volume treatment (1893-96) of the macro-
moths and pyraloids in the Fauna of British In-
dies series. Early comprehensive works on Afro-
tropical Lepidoptera were few. Suffice it to note
R. Trimen's contributions, culminating in the
monograph (with J. H. Bowker) S. African But-
terflies 1 - 3 (1887-89), and M. Saalmller's
Lepidopteren von Madagaskar 12 (188491),
completing ealier works by Boisduval and Ma-
bille. In the Nearctic J. G. Morris' Catalogue of
the Described Lepidoptera of North America
(1860) was followed by several monographs in-
cluding W. H. Edwards' Butterflies of North
America 1 - 3 (1868-97) and S. H. Scudder's
Butterflies of New England 1 - 3 (1888-89). A. S.
Packard's Monographs of the Geometrid Moths
or Phalaenidae of the United States (1876) and
Monograph of the Bombycine Moths of America
North of Mexico ... I, 1 Notodontidae (1895)
stand out for craftsmanship and attention to de-
tail. J. Smith monographed several noctuid
groups. Principal N. American micro-moth au-
thors were B. Clemens, whose papers on moths
of the lower grades were reprinted by Stainton
as Tineina of North America (1872), and V. T.
Chambers. The prodoxid/ Yucca symbiosis, one
of the most intriguing insect-plant relationships
known, was first discovered by C. V. Riley (Na-
ture 6, 1872). Outstanding in the New World lit-
erature is the regional monograph Biologia
Centrali-Americana edited by F. D. Godman &
O. Salvin. Lepidoptera parts include the editors'
account of butterflies, 1 - 3 (1879-1901) and H.
Druce's of macro-moths, 1 - 3 (1881-1900)
(while T. de Grey Walsingham's treatment of the
'Tineina, Pterophorina, Orneodina, Pyralidina
and Hepialina [part]' appeared later).
Of global coverage were E. L. Ragonot's
Monographie des Phycitinae et des Galleriinae (in
Romanoff's 'Mmoires' 7, 8; 1893-1901) as well
as the accounts of butterfly genera in O. Stau-
dinger's Exotische Tagfalter (188488) and the
companion volumes Die Familien und Gattungen
der Tagfalter by E. Schatz & J. Rber (1885-92).
W. F. Kirby's A synonymie catalogue of diurnal
Lepidoptera (1871), supplement 1877) was com-
plete for its time, the same author's Catalogue of
Lepidoptera Heterocera (1892) covers "Sphinges
and Bombyces". By the end of the 19th century
Lepidoptera systematics of a worldwide scope
was pursued by several workers whose most in-
fluential works date from the 20th century. Par-
ticularly outstanding were E. Meyrick (all-time
most prolific descriptive systematist of micro-
moths; his Lepidoptera classification presented
3
in A Handbook of British Lepidoptera, 1895,
takes the World fauna into account), G. F.
Hampson (author of monographs on pyraloid-
grade moths, but best known for the monumen-
tal Catalogue of the Lepidoptera Phalaenae in the
British Museum [treating a major part of the
Noctuoidea] which started appearing in 1898),
and the remarkable team of W. Rothschild and
K. Jordan, whose first joint monograph (on
Charaxes butterflies, Novit. Zool., 5 - 7 ) was pub-
lished 1898-1900.
Morphology and Physiology
Two of the most renowned early general works
on insect anatomy deal with larger moths: M.
Malpighi's Dissertatio epistolica de Bombyce
(1663) treating the silkworm moth Bombyx, and
P. Lyonet's remarkably accurate (with respect to
description of structure, not to interpretation of
function) Trait anatomique de la chenille qui
ronge le bois de saule (1762) treating a Cossus
larva. A comparative approach to insect mor-
phology was only developed in the late 18th cen-
tury, mainly by French workers. Particularly
outstanding was J.-C. Savigny's account of the
mouth parts in Mmoires sur les Animaux sans
Vertbres (1816).
Integumental structures
Lepidopteran vestiture was an attractive subject
for early microscopists. 19th century studies on
hair and scale structure (often focussing on scent
organs) include papers by, i. a., R. Maddox, R.
Schneider, C. Aurivillius, A. Spuler, V. Kellog,
and particularly numerous contributions by F.
Mller and E. Haase. F. G. Hopkins (Phil.
Trans. R. Soc., 186B, 1895) proved that excretory
products are stored in wing pigments in pierid
butterflies. There are numerous smaller 19th cen-
tury accounts of defensive glands, urticating
hairs and other integumental specializations of
caterpillars. Dyar (Ann. New York Acad. Sci., 8,
1894 and later papers) made a pioneering study
of the larval setal pattern, foreshadowing 20th
century taxonomic use of this system.
The head
The coilable proboscis, a unique specialization
present in most adult Lepidoptera, had been
noted by early naturalists. Lepidopteran mouth-
parts were treated in some detail in Savigny's
aforementioned work. E. Burgess described the
sucking pump in his succint, but influential, ac-
count of the anatomy of the monarch butterfly
(Danaus) (Anniv. Mem. Boston Soc., 1880). The
discovery that primitive biting mouthparts are
indeed retained in some Lepidoptera was re-
4 Niels P. Kristensen

ported by A. Walter (Jena Zschr. Naturw., 18 Viscera

((. F.), 11, 1885). Antennal structure was sur-
Important 19th century contributions include
veyed by D. Bodine {Trans. Amer. ent. Soc. 23,
numerous papers by E. Brandt (nerve system)
1896) for Lepidoptera in general, and by . Jor-
and N. Cholodkovsky (internal genitalia, mal-
dan (Novit. Zool. 5, 1898) for butterflies in par-
pighian tubules); the latter discovered (Z. wiss.
ticular. Micro-moth mouthparts were treated ex-
Zool. 42, 1885) that a non-ditrysian female geni-
tensively by K. Gen the (Zool. Jahrb. Syst. 10,
tal configuration does exist in some Lepidoptera.
1897), and there is a massive memoir on butterfly
W. Petersen's monumental Beitrge zur Morpho-
(labial) palps by E. Reuter (Act. Soc. Sei. Fenn.
logie der Lepidopteren (Mm. Acad. Sci. St. Pe-
22, 1896). O. vom Rath described the sensilla
tersb. 8 [9] 6, 1900) summarized these findings as
later named after him in 1888 (Z. wiss. Zool, 46).
well as original comparative data. Giant fibres
Compound eye structure was already examined
were discovered by A. Benedicenti (Arch. Ital.
by H. Grenacher in the 1870s, and the different
Biol., 24, 1895) in the nerve cord of Bombyx.
configurations found in moths and papilionoid
Early insights into the structure and function of
butterflies became classical examples of the 'su-
insect circulatory systems owe much to New-
perposition' and 'apposition' eye-types proposed
port's studies (Phil. Trans. Roy. Soc., 122, 1832;
by S. Exner in Die Physiologie der facettierten
127, 1837) on sphingid moths; the discovery of
Augen von Krebsen und Insekten (1891). Larval
antennal hearts was reported in Burgess' Danaus
stemmata structure was studied by Landois (.
memoir. The larval silk glands naturally
wiss. Zool, 16, 1866) and O. Pankrath (Ibid., 49,
attracted interest at an early date and were de-
1890). scribed by, e. g. F. Helm (Z. wiss. Zool., 26, 1876)
and G. Gilson (Cellule 6, 1890).

Thorax
Most early studies concern applications of wing
characters to systematics (see above). The tho-
racic trunk attracted little attention; the principal
early account of the musculature is by Amans
(Annal. Sci. nat. (Zool.), (6) 19, 1885). The
tympanic organ of noctuoids was first described
by A. Swinton (Ent. month. Mag., 14, 1877).
Abdomen
The early applications of genital characters to
Lepidoptera systematics were paralleled by in-
quiries into their structure and function; often
the approaches were combined. The presence of
two female genital openings characteristic of
most female Lepidoptera had been recognized
already by Malpighi, and the functional design
of this ditrysian female apparatus was examined
in more detail by C. T. von Siebold (Arch. Anat.
Physiol, wiss. Med., 1837) and H. Lacaze-Du-
thiers (Ann. Sci. Nat. (Zool.), (3), 19, 1853); the
former also (Z. wiss. Zool. 3, 1850) studied the
sphragis in Parnassius butterflies. Principal con-
tributors to the discussion of the morphological
interpretation of the genital structures in the two
sexes and to the development of the nomencla-
ture of these structure include F. B. White, au-
thor of a paper on male genitalia of European
butterflies (Trans. Linn. Soc. Lond. Zool. 2 (I),
1876), H. Gosse (Ibid 2 (II) 1883), W. Jackson
(Ibid. 2 (V), 1890) and A. Peytoreau, author of
Contributions l'tude de la morphologie de
l'armure gnitale des insectes (1895). J. Wood's
account of lepidopteran 'ovipositors', particu-
larly the piercing types (Ent. month. Mag 27,
1891) ist still useful.
Development
Worthwhile studies on lepidopteran embryogen-
esis started with A. Kowalewsky (Mm. Acad.
Sci. St. Ptersb., (7) 16, 1871). Egg diapause was
early discovered in the silkworm Bombyx, and its
dependence upon temperature regimes was
worked out in some detail by E. Duclaux (C. R.
Acad. Sci. Paris 69, 1869). H. Dyar discovered a
widely applicable 'rule' of geometrical pro-
gression of head capsule widths at moults (Psy-
che 5, 1890). M. Herold's Entwicklungsgeschichte
der Schmetterlinge, anatomisch und physiologisch
bearbeitet (1815) is a pioneer (albeit crude) ac-
count of metamorphic changes in the viscera of
a Pieris butterfly; it also demonstrated the pre-
hatching determination of sex, as well as the de-
velopment of male genital rudiments in a peri-
podial cavity. Principiai early studies on the mor-
phogenesis of wings, scales and hairs include
those of C. Semper (Z. wiss. Zool. 8, 1857) and
H. Landois (Ibid. 21, 1871). E. Verson made sev-
eral important observations on the structure and
metamorphosis of Bombyx, including the first
discovery of insect dermal glands (e. g. Zool.
Anz. 13., 1890) and with E. Bisson he described
the morphogenesis of the genitalia of both sexes
(Z. wiss. Zool., 61, 1896).
Information Sources
Principal sources for this outline are the chapters
by Tuxen, Lindroth, Richards and Wigglesworth
in Smith et al. (eds.) 1973, as well as the litera-
ture compilations in Kirby (1897) and Kuznet-
sov (1967).
Historical Introduction
References
Kirby, W. F. (1897): A Hand-book to the Order Lepi
doptera. IV. London, Allen & Co.
Kuznetsov, N. Y. (1967): Lepidoptera I. Introduction.
Fauna of Russia and Adjacent Countries. Jerusa
lem, Israel Program for Scientific Translations.
Lindroth, C. H. (1973): Systematics specializes. Be
tween Fabricius and Darwin: 1800-1859. Pp. 119-
154 in Smith, R. F., Mittler, T. E. & Smith, C. N.
(eds.) History of Entomology. Palo Alto, Annual
Reviews.
5
Richards, A.G. (1973): Anatomy and morphology.
Pp. 185-202 in Smith, R. F., Mittler, T. E. &
Smith, C. N. (eds.) History of Entomology. Palo
Alto, Annual Reviews.
Tuxen, S. L. (I 973): Entomology systematizes and de
scribes: 1700-1815. Pp. 95-118 in Smith, R. F.,
Mittler, T. E. & Smith, C. N. (eds.) History of Ento
mology. Palo Alto, Annual Reviews.
Wigglesworth, V. B. (I 973): The history of insect phys
iology. Pp. 203-228 in Smith, R. F., Mittler,
T. E. & Smith, C. N. (eds.) History of Entomology.
Palo Alto, Annual Reviews.
 2. Phylogeny and Palaeontology
Niels P. Kristensen & Andrzej W. Skalski*
The monophyly of the taxon Lepidoptera is
firmly established by an impressive suite of syna-
pomorphies of its constituent basal lineages. The
position of the group within the insect hierarchy
is similarly well established: It has a sistergroup
relationship to the Trichoptera (caddisflies), con-
stituting with the latter the high-rank taxon Am-
phiesmenoptera. The Amphiesmenoptera, in
turn, together with the Antliophora (Mecoptera
+ Siphonaptera + Diptera) constitute the 'pan-
orpoid' clade within the Endopterygota (see
Kristensen 1991, and references therein, for high-
level relationships).
This chapter first treats the ground plan and
intrinsic phylogeny of the Lepidoptera, as de-
duced from the character distribution in extant
taxa. Subsequently the palaeontological contri-
butions to the knowledge about Lepidoptera
evolution are surveyed. No known fossil Lepi-
doptera represent organisation types which
markedly differ from extant ones, and as in sev-
eral other insect groups, fossils have so far con-
tributed little, if anything, to the understanding
of phylogenetic interrelationships. But fossils, of
course, provide the only direct means for estab-
lishing the minimum ages of individual evolu-
tionary lineages.
Major ecological aspects of lepidopteran evo-
lution are dealt with in 120.
The Lepidopteran Ground Plan and
the Superorder Amphiesmenoptera
Detailed accounts of the lepidopteran ground
plan will be presented in the morphological
chapters in volume 2 of the present work. Kris-
tensen (1984) identified 27 probable lepidopteran
autapomorphies, and another 20 groundplan
traits were identified as probable synapomor-
phies with the Trichoptera, i. e., autapomorphies
of the Amphiesmenoptera. The great majority of
the characters in question are in the adult insect
(Fig. 2.1). Subsequent work has so far entailed
few modifications to these conclusions.
The lepidopteran groundplan autapomorphies
are briefly enumerated here; for more detailed
discussions see Kristensen (1984) and references
therein. Adult (numbers in brackets refer to Fig.
2.1): Median ocellus lost (1). Corporotentorium
with posteromedian process (2). Intercalary scler-
ite present laterally in membrane between anten-
* deceased 1996
nal scapus and pedicellus (3). Maxillary palp
with points of flexion between segments 1/2 and
3/4; segment 4 longest; intrinsic palp musculature
not comprising antagonistic pairs (4). Cranio-
stipital muscle present, slender and running close
to craniocardinal muscle (5). Postlabium an
arched sclerite with long piliform scales (6). Ter-
minal segment of labial palp with group of sen-
silla in depresseion ('vom Rath's organ') (7). Sali-
varium devoid of dorsolongitudinal muscle (8)
(note: this character needs additional scrutiny,
since recent observations on Trichoptera and
Mecoptera cast doubt on the primitive nature of
a muscle of this kind in the panorpoid orders; M.
Engvall & N. P. Kristensen unpublished). Labral
nerve and frontal ganglion connective separating
immediately at their origin on the tritocerebrum
(9). Nervus recurrens running inside aorta until
reaching retrocerebral complex (10). Laterocer-
vical sclerite with proprioceptive 'hair plate'
close to anterior apex (11). Prothoracic endoskel-
eton with prominent free arm arising from
bridge between sternum and lower posterior cor-
ner of pleuron (12). Mesothorax with 'tergopleu-
ral apdeme' issued from upper part of pleural
suture and accommodating insertion of a ter-
gopleural muscle (13). Metathorax with 'prescu-
tal arm' (14). Fore tibia with movable 'epiphysis'
on inner surface (15). Fore tibia with at most a
single spur (15) (Shields 1993; note: not included
in the 1984-enumeration this regressive trait ap-
parently represents yet another lepidopteran au-
tapomorphy, since 3 spurs can be ascribed to the
trichopteran ground plan). Wings with dense
covering of broad scales (16). Metathoracic spi-
racle with single, anteriorly situated, external lip
(17). Tergum I extensively desclerotized, with
concomitant loss of external layer of 'short' dor-
solongitudinal I/II muscles (18). Tergum I with
lateral lobes extending posteroventral to articu-
late with anterior corners of sternum II (19).
Male 'valve' (gonopod) primarily undivided (20).
Phallic protractor muscle originating inside valve
(21). Cerci lacking in both sexes (22). Abdominal
nerve cord with at most five ganglionic masses,
and unpaired connectives (23). Sperm of apyrene
variety (24). Larva: Pleurostome elongated, cran-
iocardinal articulation far behind mandibular
base. Maxillary palp with fewer than 5 segments.
Kristensen (1984), on the basis of Hessel's
(1969) information about neighbouring endop-
terygotes considered the dorsal curvature of the
adult's mesothoracic aorta to be another lepi-
dopteran groundplan autapomorphy. However,
recent comparative studies (Krenn & Pass 1995)
8 Niels P. Kristensen & Andrzej W. Skalski

Fig. 2.1. Hypothetical ancestral adult lepidopteran. Amphiesmenopteran and lepidopteran groundplan autapo-
morphies are indicated by Roman and Arabic numerals respectively (see text). Modified from Kristensen (1984).

indicate that the curved configuration is plesio-
morphic at the panorpoid level.
In the light of this suite of synapopmorphies
of the basal lepidopteran lineages inclusive of the
Micropterigoidea, earlier suggestions that the
latter should be assigned to a separate order, be-
cause they are "much more closely related to the
Trichoptera than to the Lepidoptera" (Hinton
1958) now have only historical interest. Two re-
gressive traits in larval structure (loss of lacinia,
marked weakening of corporotentorium) are
known to be shared by Trichoptera and non-
Phylogeny and Palaeontology
micropterigoid Lepidoptera, but on grounds of
parsimony, they can be construed only as homo-
plasies.
The following lepidopteran groundplan traits
are apparently autapomorphic of the superorder
Amphiesmenoptera. Adult (Roman numerals re-
fer to Fig. 2.1): Prelabium fused with hypophar-
ynx (I). Lower posterior corner of laterocervicale
produced towards the prosternum (II). Prono-
tum with paired setose 'warts' (III). Prothoracic
epistema with unique suture pattern (IV). Sec-
ondary furcal arms of pterothorax fused with
posterior margins of corresponding epimera (V).
Metathorax with setose, presumably propriocep-
tive, sclerite in wing base membrane behind/be-
low subalare (VI). Pretarsus above claw with
'pseudempodium' (strong seta on socket) (VII).
Wings with dense vestiture of setae (forerunners
of the lepidopteran scales) (VIII). Fore wing anal
veins looping up into double-Y formation (IX).
One ventral (tentorial) neck muscle originating
on fore coxa (X). Conical furcopleural muscle in
mesothorax with broad end on pleural ridge
(XI). Paired glands opening on sternum V (XII).
Male segment IX with tergum and sternum fused
into closed ring (XIII). Anterior margin of fe-
male segments VIII and IX with long rod-like
apodemes accomodating insertions of protractor
and retractor muscles of extensible oviscapt
(XIV) (note: recent work shows that the inter-
pretation of the female postabdomen in the low-
est Amphiesmenoptera is more problematical
than hitherto believed, and the apodemes in
question may not all be homologous, see 2 - 4 ) .
Ventral diaphragm muscles inserting on the
nerve cord (XV). Male sex homogametic (XVI).
Spermatozoa with outer accessory filaments
thickened, filled with proteinaceous and glyco-
gen-like material (XVIII). Chromosome number
unusually high (basic number 3031), chromo-
somes holocentric and oogenesis achiasmatic
(XVII). Larva: stemmata each with one crystal-
line cone cell transformed into primary pigment
cell. Prelabium and hypopharynx fused into
composite lobe with silk gland orifice on apex.
It is straightforward to assume that the mem-
bers of the amphiesmenopteran stem lineage had
larvae which were 'soil animals' and lived in wet
conditions, like those of most extant Lepidop-
tera-Micropterigidae. The step from here to the
truly aqatic lifestyle, which is autapotypic of im-
mature Trichoptera, is but a small one. In the
context of the recent debate over ancestral be-
haviour within the Trichoptera (see Weaver
1992 a, b; Wiggins 1992) it is relevant to note
that lepidopteran larvae which live in silken gal-
leries in the soil are not encountered until after
the eighth splitting event identifiable among ex-
tant taxa (viz., the one leading to the Exoporia
and Heteroneura). The out-group criterion
therefore lends no support to the assumption,
that ancestral caddisfly larvae lived in silken
tubes.
9
Lepidoptera Phylogeny above
Superfamily-Level
A total of 46 lepidopteran superfamilies are rec-
ognized in this work; they are all diagnosed in
the relevant systematics chapters (14 through
19). A tentative phylogeny of the superfamilies
is illustrated in Fig. 2.2; for superfamilies/super-
family assemblages with more than 1000 de-
scribed species the species number is indicated
by the width of the clade lines. It is immediately
obvious, that the most speciose lineages are cla-
distically quite subordinate. This particular phy-
logenetic pattern was ascribed considerable gen-
eral significance in Hennig's writings (e. g., 1953,
1966), and indeed the basic diversification
pattern within the Lepidoptera remains an unu-
sually instructive example of what has subse-
quently come to be known as a 'Hennigian
comb', with the first diverged extant lineages ex-
hibiting a step-by-step acquisition of the apo-
morphies which characterize the most subordi-
nate (and successful) groups.
While the current estimate of described lepi-
dopteran species is close to 150.000 the total
number of extant species may be some
3-500.000 (Kristensen in press). Undoubtedly
the proportional representation of individual su-
perfamilies will change markedly as descriptive
Lepidoptera taxonomy approaches completion.
Little growth is foreseeable in the butterfly and
bombycoid assemblages, whereas particularly
strong increases are expected for the Gelechi-
oidea (see 120) and the Pyraloidea (Shaffer in
Scoble 1992).
Basal Splitting Events. The present account is
based on Kristensen (1984) which may be con-
sulted for further details. The basal splitting
event among known extant Lepidoptera is here
presumed to have given rise to the Micropteri-
goidea (suborder Zeugloptera auct., see 13) on
one hand, and all other members of the order on
the other. Synapomorphies of the latter clade are
enumerated in 14. Another proposal deserving
serious attention, is that the Agathiphagoidea
are the sister group of all other Lepidoptera.
Shared derived traits of non-agathiphagoid Lepi-
doptera (note: the internal anatomy of Hetero-
bathmiidae remains largely unstudied) include:
Absence of M4 in both wing pairs; absence of
true spurs on the foretibia and (Shields 1993) ab-
sence of spur pair near mid-length of mesotibia;
testes follicles small, closely appressed; few (4 or
fewer) discrete abdominal ganglionic masses
(higher numbers in some glossatane interpreted
as autapomorphic neotenic traits, Nielsen &
Kristensen 1996); few (5 or fewer) ovarioles per
ovary (higher numbers in Glossata-Incurvari-
oidea interpreted as autapomorphic); absence of
the mesocoxal 'outer tergal remotor'; presence of
10 Niels P. Kristensen & Andrzej W. Skalski

Micropterigoidea
Agathiphagoidea
Heterobathmioidea
Eriocranioidea
Acanthopteroctetoidea
Lophocoronoidea
GLOSSATA *
Neopseustoidea
COELOLEPIDA"
Mnes/Hepialoidea
Nepticuloidea
MYOGLOSSATA
Incurvariodea
NEOLEPIDOPTERA 1 Palaephatoidea
HETERONEURAL- Tischerioidea
Simaethistoidea

Tineoidea

Gracillarioidea
Yponomeutoidea

DITRYSIA
Gelechioidea

Galacticoidea

Zyg/Coss/Sesioidea

Choreutoidea

Tortricoidea

Urodoidea
Schreckensteinioidea
Epermenioidea
Aluc/Pterophoroidea
Whalleyanoidea
Immoidea
APODITRYSIA
Copromorphoidea
Hyblaeoidea

Pyraloidea

Thyridoidea
Mim/Las/Bombycoidea
Axioidea
Calliduioidea
OBTECTOMERA Hedyloidea

Hesp/Papilionoidea

Drepanoidea

Geometroidea
MACROLEPIDOPTERA

Noctuoidea

10.000
species
described

Fig. 2.2. Phylogeny of extant lepidopteran superfamilies. Width of superfamily lines indicate approximate numbers
of described extant species (where these numbers are > 1000). Large sectors within the Ditrysia remain unresolved,
and all proposed groupings above superfamily level in the Ditrysia must be considered very tentative.
Phylogeny and Palaeontology
only one (not two) pairs of male accessory
glands. The interpretation of the two lastmen-
tioned traits as apomorphies at the basic lepidop-
teran level is debatable, however; particularly the
occurrence of the coxal muscle in agathiphagoids
(and, as far as known, nowhere else among pan-
orpoid insects !) could well be an autapomorphic
character reversal. The reason why the phylog-
eny here preferred has the Micropterigoidea
rather than the Agathiphagoidea as the first di-
verged lepidopteran lineage, is that the suite of
derived characters shared by non-micropteri-
goids in addition to a number of regressive char-
acters includes some striking neoformations
(metafurcal process, double-compartment sper-
mathecal duct, larval lateral head-capsule mus-
cle, hypertrophied and angularly bent pupal
mandible), while those shared by non-agathipha-
goids all are regressions/simplifications which are
more likely to be homoplasious. This issue is not
addressed in Shield's (1993) advocacy of the the-
ory of the Agathiphagoidea being the sister
group of all other Lepidoptera. Ivanov (1994)
has recently found support for the same theory
in characters in the wing base; these await eval-
uation.
The next splitting event is here believed to
have been between the Agathiphagoidea and the
Heterobathmioidea + Glossata. Derived traits
characterizing the latter entity include some very
characteristic traits of the larval head (adfrontal
and hypostomal ridges, interrupted hypostomal
bridge) in addition to the presence of sensilla au-
ricillica and loss of Sc-R crossvein in the adult,
and presence of coarse apical teeth on the mandi-
ble in the pupa. Alternatively the dichotomy
could be between the Agathiphagoidea + Heter-
obathmioidea and the Glossata, but the derived
traits of the former entity seem less impressive.
Most notable are the Y-configuration of the an-
terior tentorial arms and the posteriorly widened
postlabium in the adult, as well as the subapical
ventral tooth on the mandible in the pupa; other
similarities are in regressive traits (indistinct lat-
eral lobe from tergum I, loss of labral compres-
sors, intrinsic maxillary muscles, and dorsal
postcerebral pharynx dilators in the endopha-
gous larvae). The anastomosis pattern of the
mesepisternal para- and precoxal sutures is
somewhat different in the Agathiphagoidea and
Heterobathmoidea, and the suggested presence
(Kristensen & Nielsen 1979, 1983) in the Hetero-
bathmioidea of a phallotheca-type intromittent
organ, similar to that of the Agathiphagoidea,
has been proved wrong by a later reinvestigation.
Needless to say, an Agathiphagoidea + Hetero-
bathmioidea monophyly model also necessitates
that the unique agathiphagoid plesiomorphies
listed above should have been lost three times
independently (in the Micropterigoidea, Hetero-
bathmioidea and Glossata).
11
Basal Glossatan Phylogeny. The monophyly of
the Glossata is very strongly supported. Not only
do the glossa tan autapomorphies (listed in 15)
include some very remarkable neoformations/
modifications, but there is at present no conflict-
ing evidence at all (i. e., evidence that subordi-
nate glossatan taxa should have their closest rel-
atives outside the Glossata).
Six basal clades are recognized within the
Glossata: the Eriocraniidae, Acanthopterocteti-
dae, Lophocoronidae, Neopseustidae, Exoporia
(Mnesarchaeoidea + Hepialoidea) and Hetero-
neura (all other Glossata). A phylogenetic analy-
sis of these clades, based on 46 characters (Niel-
sen & Kristensen 1996) gave best support to the
arrangement illustrated in Fig. 2.2, viz., Eriocra-
niidae + (Acanthopteroctetidae + (Lophocoro-
nidae + (Neopseustidae + (Exoporia + Hetero-
neura)))). Apomorphies supporting each inter-
node are listed in 15. Names have been given
to three of the identified high-rank taxa: Coelo-
lepida (the non-eriocraniid Glossata), Myoglos-
sata (Neopseustidae + (Exoporia + Hetero-
neura)) and Neolepidoptera (Exopria + Hetero-
neura).
Basal Heteroneuran Phylogeny. The monophyly
of the Heteroneura is here accepted as a reason-
able working hypothesis supported by several
putative groundplan autapomorphies discussed
in more detail by Nielsen & Kristensen (1996):
Heteroneurous venation (hindwing Sc and R
fused beyond a short distance from wing base,
Rs unbranched). Male with composite frenulum.
Forewing jugum reduced, projecting little be-
yond general wing contour. Dorsal tentorial
arms in adult strongly reduced. Prothoracic dor-
sum with median sclerotization behind principal
paired 'warts', and precoxal bridge present. Mes-
osternum with suture-delimited 'mesoclidium',
and fusion of the latter with the prospina. Two
discrete bundles, with different posterior inser-
tion sites, of mesofurco-metafurcal muscles.
Metaprescutal arm and sternum I obliterated. In
embryogenesis 'fault-type' formation of amnion
and serosa developed, germ band immersed in
yolk, and temporary epithelium-like membrane
formed by peripheral yolk nuclei/vitellophages
absent. This evidence arguably outweighs the dif-
ficulties with heteroneuran monophyly empha-
sized by Dugdale (1974) and Minet (1984), viz.,
respectively, the somewhat similar female genital
systems (with separate copulatory orifice and
ovipore, and with ventral bursa copulatrix) in
Exoporia and Heteroneura-Ditrysia (here con-
sidered parallelisms), and the absence of muscu-
lated, crochet-bearing prolegs in Heteroneura-
Nepticuloidea (here considered a secondary re-
gression, linked to endophagy).
The basal splitting events within the Hetero-
neura remain problematical (see 16). A mono-
phyletic group comprising (Palaephatoidea +
12
Fig. 2.3. Davis' (1986) alternative proposals for the
basal phylogeny of the Heteroneura.
Tischerioidea) + Ditrysia (Fig. 2.2) could be sup-
ported by the presence of a medial ridge between
the 'ovipositor' lobes in the two firstmentioned
taxa and a derived female frenulum type (few,
closely approximated setae) in all (Nielsen 1985,
1989). Moreover, the eggs in Tischerioidea (pa-
laephatid egg ultrastructure remains unstudied)
and Ditrysia share a unique trabecular chorionic
layer (Fehrenbach 1995, 2 - 1 8 ) . Also, the mono-
phyly of all non-nepticuloid Heteroneura is now
being supported by data from female 'calling' be-
haviour (oviscapt extrusion) and 'long-chain' fe-
male sex pheromenes (Lfstedt & Kozlov 1995,
2 - 1 3 ) . Alternatively, Davis (1986) advocated the
phylogeny shown in Fig. 2.3, implying that all
non-ditrysian Heteroneura constitute a mono-
phyletic 'Monotrysia' the groundplan of which is
characterized by the presence of a 'pectinifer' (a
valve area bearing a comb of specialized stout
setae) and a derived metafurcal configuration. A
pectinifer, however, is absent in the Palaepha-
toidea and Tischerioidea, and the ground plan
of the 'monotrysian' metafurca is probably not
different from that of the Heteroneura (24).
The monophyly of the Nepticuloidea + Tischeri-
oidea was supported only by the flattened head
and reduction of larval legs (commonplace modi-
fications in miners) as well as a narrowing of the
hind wings and loss of M2 in both wings. Recent
evidence from 18S rDNA data support the mo-
nophyly of Tischerioidea + Ditrysia as well as of
a clade comprising the remaining monotrysian
Heteroneura. However, the nepticulid examined
comes out as subordinate in the Incurvarioidea,
which appears unacceptable on the basis of mor-
phology, and it is emphasized, that the above-
mentioned alternative phylogenies "are not
strongly contradicted by the 18S rDNA gene"
(Wiegmann 1994).
Ditrysian Phylogeny. The monophyly of the Di-
trysia is corroborated by very notable ground-
Niels P. Kristensen & Andrzej W. Skalski
plan autapomorphies including the specialized
female genital apparatus (separate copulatory
orifice and ovipore, and internal communication
between sperm-receiving and oviduct systems),
prominent internal apodemes on abdominal ster-
num II; moreover, the intrinsic proboscis muscu-
lature is organized as numerous short bands in-
stead of longitudinal fibres (Kristensen unpub-
lished).
While the clade Ditrysia perhaps comprises
close to 99% of the extant lepidopteran species,
it is structurally overall homogeneous, and not
surprisingly, therefore, its intrinsic phylogeny
has proved particularly difficult to unravel. The
demand that taxa should be monophyletic in the
strict (cladistic) sense has led to the establish-
ment, in recent decades, of an increasing number
of superfamilies comprising a single or very few
families. In the classification adopted here 33 di-
trysian superfamilies are recognized. The scheme
of their hypothesized interrelationships shown in
Fig. 2.2 is mainly based on Minet's seminal 1991
study, which should be consulted for further de-
tails.
A large assemblage of ditrysian superfamilies,
including the apparently first differentiated su-
perfamilies, share some conspicuous (but plesio-
morphic) traits with most non-ditrysians: They
are generally small moths with a distinct CuP (at
least in the hindwing; this vein, however, is re-
peatedly lost in subordinate clades, particularly
those characterized by pronounced wing narrow-
ing), and with larvae that are concealed feeders
and usually have the proleg crochets arranged in
a circle/ellipse (which may be more or less in-
terrupted). Together with the non-ditrysians the
members of this ditrysian grade are colloquially
referred to as microlepidopterans or micro-
moths ('micros').
The monophyly of the non-tineid Ditrysia is
only supported by a few regressive traits (Minet
1991): shortening of the maxillary palp which is
at most 4-segmented (the only known exception
being some Alucitoidea-Alucitidae in the Obtec-
tomera; a character reversal?) and has lost the
sharp bend between segments 3 and 4; absence
of projecting lateral bristles on the labial palp.
The significance of the lastmentioned character
is ambiguous; the bristles in question have tenta-
tively been considered a tineoid autapomorphy
by Robinson & Nielsen (1993, see also 1 - 7 ) , but
these authors, like Minet, noted their occurrence
also in a variety of non-ditrysian Heteroneura,
and hence admitted their possibly plesiomorphic
nature at the ditrysian level. Robinson & Nielsen
(1993) suggested that all Ditrysia apart from the
Tineoidea and Gracillarioidea constitute a mo-
nophylum, to whose ground plan they ascribed,
as an autapomorphy, the presence of spines on
the abdominal terga in the adult moths.
All Ditrysia, apart from the Tineoidea, Gracil-
liarioidea, Yponomeutoidea and Gelechioidea
Phylogeny and Palaeontology
are tentatively united in the taxon Apoditrysia
(Minet 1983), whose monophyly is supported by
an apomorphic sternum II structure. The apo-
demes are shortened, with enlarged bases, not
appearing as mere continuations of the 'venulae'
(longitudinal costae on the sternal plate). Pro-
duced anterolateral corners are another charac-
teristic specialization of sternum II in most
Apoditrysia, but it is uncertain whether they can
be ascribed to the ground plan of the clade. Rob-
inson & Nielsen (1993) considered the loss of ter-
gal spines an additional apoditrysian autapo-
morphy (as a character reversal), but tergal
spines are actually retained in some apoditrysian
lineages, including Zygaenoidea, Epermenioidea
and Alucitoidea (1 11, 113).
The monophyly of the Gelechioidea + Apodi-
trysia has been suggested on the basis of their
shared reduction of the basal apdeme of the
male valve and the dorsal displacement of the
origin of the valve 'protractor' (Robinson &
Nielsen 1993; see also 24): Additional support
for this clade has recently come from the discov-
ery of putative synapomorphies in its members'
proboscis structure: two (not one) rows of sen-
silla styloconica per galea, and presence of
strongly sclerotized transverse ribbons (Ram-
mert 1994); however, considerable homoplasy in
these traits render their significance questionable
(Minet, pers. comm.). The close approximation
of setae LI and 2 on larval segments IVIII may
be yet another groundplan autapomorphy of this
clade; the wide spacing of these setae in many
advanced Apoditrysia would, then, have be con-
sidered character reversals. In the light of the evi-
dence for an alternative placing of the Gelechi-
oidea (as the sister group of the Yponomeu-
toidea, see below) the question has here been
left open.
The classification of the Apoditrysia is partic-
ularly problematical. Fig. 2.2 reflects Minet's
view that a large assemblage of apoditrysian su-
perfamilies constitute a monophylum 'Obtec-
tomera'. When this taxon was first proposed
(Minet 1986) its monophyly was supported only
by the immobility of pupal abdominal segments
I - I V ; related to this immobility the pupal abdo-
men has lost the dorsal transverse rows of spines
(which were acquired by ancestral Neolepidop-
tera) and are not protruded from the pupal shel-
ter prior to eclosion. This character complex is
subject to considerable homoplasy. In the non-
apoditrysian grade it occurs also in the Ypono-
meutoidea and Gelechioidea; for this reason the
monophyly of an entity comprising these two su-
perfamilies has been suggested by Minet (1991),
who recently (1996) has found this monophyly
supported by details in labial palp structure.
Among the Apoditrysia the 'obtectomeran-type'
pupa also occurs in the Epermenioidea and Alu-
citoidea; hence these two superfamilies were ini-
tially included in the Obtectomera. However,
13
subsequently they were excluded from this taxon
when it was redefined (Minet 1991) by the addi-
tion of another groundplan autapomorphy, viz.,
the differentiation of a dorsal lobe (sometimes
merely appearing as a group of bristle-like pro-
cesses) on the adult's pretarsal pulvillus.
Eleven superfamilies of non-obtectomeran
Apoditrysia are presently recognized; the Tortri-
coidea are by far the largest of these, and the
whole assemblage may conveniently be referred
to as the 'tortricoid grade'. Interrelationships
above the superfamily level are largely unclari-
fied within this grade. The Sesioidea (which as
now delimited include the Brachodidae, Sesiidae
and Castniidae) and Cossoidea may be each oth-
er's closest relatives (1 11), and it is just possible
that the two together are the sister group of the
Zygaenoidea. A possible synapomorphy accord-
ing to Minet (1991) is the position of the sensil-
lum Pb on the larval head capsule, which in these
three superfamilies lies distinctly dorsad from,
rather than below, a line between the P2 setae;
admittedly, however, the polarization of this
character is difficult, because both states occur
within the Tineoidea. According to Ehnbom
(1948) the ocellus structure in Zygaenidae and
Sesiidae was unique among the Lepidoptera ex-
amined by him because of the presence of an ex-
tra cell layer between the cornea and the retina
(he had not correctly identified the vitreous body
subsequently described from Eriocranioidea; see
15, 212). A reinvestigation, including an ex-
amination of ocellus structure in the Brachodi-
dae and Castniidae, is now an obvious priority.
The proposed monophyly of the Alucitoidea
+ Pterophoroidea (Minet 1991) is based on their
shared specialized configuration of the meso-pre-
coxal suture, and the resting posture with fore-
wings markedly held out from the body.
The Alucitoidea have previously been in-
cluded, together with the Epermenioidea, in the
Copromorphoidea. Specializations shared by
these superfamilies include, in addition to the pu-
pal type (see above), a process on the larval post-
labium (Heppner 1987) and a bisetose protho-
racic L group (the significance of the trisetose L
in Isonomeutis, hitherto placed in 'Copromorphi-
dae' is ambiguous: see 113): Common (1970)
mentioned some further characteristics of the
larval spiracle VIII of copromorphoids and aluc-
itoids (larger size and more dorsal position than
the preceding spiracles), but these are actually of
widespread occurrence (Minet 1983) and ques-
tionably apomorphic at the apoditrysian level.
The arrangement in Fig. 2.2 reflects Minet's
rejection (1991) of the monophyly of the Copro-
morphoidea . lai., since only the Copromor-
phoidea s. str., but not the Epermenioidea and
Alucitoidea, possess the lobed pulvillus attrib-
uted to the obtectomeran groundplan. But the
relatively immobile, non-spinose and non-pro-
truded pupa of Epermenioidea and Alucitoidea
14
may represent a genuine synapomorphy with the
Obtectomera. If the Alucitoidea + Pteropho-
roidea are indeed monophyletic, this would evi-
dently require that the pupal mobility and spi-
nosity in pterophorids are character reversals/
neoformations. In the case of the spinosity this
has already been claimed by Kuznetzov & Stekol-
nikov (1979) and Scott (1986). Minet (1991)
noted that the Epermenioidea share with the
Alucitoidea and Pterophoroidea the apparently
apomorphic trait of having groups of lamellar
scales in the forewing fringe. But obviously the
acceptance of a monophyletic Epermenioidea +
(Alucitoidea + Pterophoroidea) will require ad
hoc explanations for the specializations (in im-
matures) shared by Epermenioidea and Aluci-
toidea but not Pterophoroidea.
Witihin the Obtectomera there is an assem-
blage of superfamilies, which have retained the
'microlepidopteran' characteristics. The Pyra-
loidea are by far the largest of these, and the
whole assemblage, including also the Copromor-
phoidea, Immoidea, Hyblaeoidea, Thyridoidea
and Whalleyanoidea, may conveniently be re-
ferred to as the 'pyraloid grade'; indeed some of
these taxa have previously been formally in-
cluded in the Pyraloidea. However, as empha-
sized by Minet (1983), there is no evidence that
the superfamily, if more broadly delimited, is
monophyletic.
The Simaethistoidea, with 'pyraloid' facies as
adults (immatures are unknown), are at present
entirely unplaced within the Ditrysia.
The remaining Ditrysia, which include some
very speciose superfamilies, are mostly medium-
sized to large, frequently rather broad-winged in-
sects with predominantly exophagous larvae.
Most members of this assemblage have long been
known to share a few structural specializations
rarely encountered among 'microlepidopterans',
such as the complete loss of CuP, and the ar-
rangement of larval proleg crochets in a 'mesose-
ries'. In the tree in Fig. 2.2 these superfamilies
constitute a monophylum Macrolepidoptera
supported by at least the apparently unique elon-
gation of the 1st axillary sclerite in the forewing
base (Minet 1991). The Macrolepidoptera com-
prise the mimallonoid/lasiocampoid/bombycoid
assemblage, the Axioidea, Calliduloidea, the but-
terflies (i. e., the hedyloid/hesperioid/papilionoid
assemblage), the Drepanoidea, Geometroidea
(including Uraniidae) and Noctuoidea. Scott
(1986, Scott & Wright 1990; see also Scott 1989)
listed a suite of additional putative autapomor-
phies of an almost identically delimited taxon
Macrolepidoptera (Mimallonidae excluded).
These apomorphies, however, either occur else-
where among apoditrysians, or the plesiomor-
phic states are actually retained in scattered
macrolepidopterans, so that the apomorphic
state cannot be attributed to the macrolepidop-
teran ground plan unless ad hoc hypotheses of
Niels P. Kristensen & Andrzej W. Skalski
character reversals are invoked (Brock 1988, see
also Robbins 1987).
As far as the butterflies are concerned Minet
(1991) suggested that they constitute a subordi-
nate rather than a basal macrolepidopteran lin-
eage, inasmuch as he proposed the monophyly
of a large entity comprising the Axioidea, Calli-
duloidea, butterflies (s. lat., including the Hedy-
loidea) and Drepanoidea + Geometroidea.
Probable synapomorphies of these taxa include:
concealment of pupal labial palps; reduction of
the metapostnotal 'fenestrae laterales' (see 24);
internal laminae of secondary metafurcal arms
widened, their mesal edges adjacent and parallel,
or posteriorly convergent. It was emphasized,
however, that at least the first two of these traits
are pronouncedly homoplasious, occurring also
in subordinate members of other macrolepidop-
teran lineages. No hypothesis on the precise posi-
tion of the butterflies s. lat. (or s. str. for that
matter) can at present be considered at all well
founded in spite of considerable recent effort to
clarify this matter on the basis of morphological
as well as molecular data; see 117 as well as
Weller & Pashley (1995), de Jong et al. (1996)
and Scoble (1996) for further details.
It has been a central theme in recent discus-
sions about ditrysian phylogeny whether a taxon
'Macrolepidoptera' inclusive of the butterflies is
indeed a monophylum (Scott 1986, 1989; Scott &
Wright 1990; Brock 1988, 1990 a, b). In a recent
phylogeny reconstruction by Rammert (1994)
neither the Obtectomera nor the Macrolepidop-
tera came out as monophyla. Instead the basal
dichotomy within the Apoditrysia was found to
lie between to major lineages, one being Cos-
soidea + (Zygaenoidea + (Hyblaeoidea + ((Eper-
menioidea + Thyridoidea) + (Sesioidea + 'but-
terfly assemblage')))), the other (Schreckensteini-
oidea + Tortricoidea) + ((Copromorphoidea +
(Alucitoidea + Pterophoroidea)) + Choreu-
toidea + (Pyraloidea + 'macromoths')). How-
ever, Rammert's results must be considered in-
conclusive, since (1) the analysis was focussed on
characters in the abdominal base and omitted
most other characters currently discussed in
apoditrysian phylogeny, and (2) the scorings of
many of the character states for individual super-
families/superfamily groups were highly debat-
able (Minet, pers. comm.).
The abovementioned 'butterfly assemblage' in-
cludes the Papilionoidea, Hesperioidea, Hedy-
loidea, Calliduloidea and Axioidea. The 'macro-
moths' include the Mimallonoidea, Lasiocam-
poidea, Bombycoidea, Drepanoidea, Geome-
troidea and Noctuoidea.
Any Apoditrysia cladogram based on current
morphological knowledge will contain so much
homoplasy 'noise' that its value for making bio-
logical inferences is very limited. It remains to
be seen to what extent future morphological and
molecular investigations can remedy this situa-
Phylogeny and Palaeontology
tion. The field of comparative morphology still
contains vast, and largely untapped sources of
information, e. g. several aspects of proboscis
structure, myology, respiratory organs, internal
genitalia, larval mouthparts: also, a detailed re-
assessment of some already available morpho-
logical information (particularly the wealth of
data on male genitalia musculature emerging
from the research programme of Kuznetznov,
Stekolnikov and coworkers) from a cladistic
viewpoint is challenging. The molecular ap-
proach is still in a very initial phase (Weller et al.
1994, Weller & Pashley 1995, review by Regier
et al. 1995), but many more contributions can be
expected in the foreseeable future. It seems likely
that 'total-evidence' studies integrating morpho-
logical and molecular data hold the greatest
promise for genuine progress in Lepidoptera
phylogeny.
Fossil Lepidoptera
The fossil record of Lepidoptera is meagre com-
pared with that of the other major endopterygote
insect orders; this can be at least partly attrib-
uted to the delicateness of their bodies and
wings. The record is also a short one, and unam-
biguous lepidopteran fossils of pre-Tertiary age
were not recorded until 1970. It is estimated that
only some 600700 specimens of fossil Lepidop-
tera are presently known, and of these about 500
are preserved in resins. Most of the remaining
are from fine-grained sediments: clay, sand, silt,
mud, shale, limestone, marl, asphalt and vol-
canic ash/dust, and there is even an old record
of a moth larva in a rock crystal (Kawall 1876).
Most lepidopteran fossils are adults, but also
fossil larvae, larval cases and pupae (Rebel 1934,
Kusnezov 1941; MacKay 1969; Nel & Nel 1985),
and even a moth egg (Gall & Tiffney 1983) have
been recorded. Mines in fossil leaves have with
more or less certainty been attributed to Lepi-
doptera (Opler 1973, 1974, 1982; Hickey &
Hodges 1975; Skalski 1979 a, 1990 a; Kuroko
1987; Kozlov 1988; Davis 1994; Labandeira et al.
1994). Lepidopteran scales have been identified
in fossil resins (Khne, Kubig & Schlter 1973;
Schlter 1974; Skalski 1976; Whalley 1978), and
together with cuticular remnants in alimentary
canal of fossil bats (Richter & Storch 1980).
A succinct account of fossil Lepidoptera de-
scribed up to 1983 is given by Carpenter (1992).
Other recent reviews include Skalski (1979 b,
1990 a), Whalley (1986) and Kozlov (1988), the
lastmentioned with a list of alleged Lepidoptera
fossils which have subsequently been transferred
to other orders. Lepidoptera from fossil resins
have long attracted particular attention; they
were monographed by Kusnezov (1941) and
Skalski (1976), and summarily reviewed in more
15
general accounts by Larsson (1978) and Poinar
(1992). General problems in the classification
and taxonomy of fossil Lepidoptera were dis-
cussed by Skalski (1976, 1977; at that time still
from a non-cladistic viewpoint).
Fossil Sites. The Eocene/Oligocene (ca. 55
54 myr) Baltic amber, of which the main deposit
is located on the southeast shore of the Baltic
Sea, is the richest source of fossil Lepidoptera.
Another rich, and now intensively investigated,
source is the mostly younger Dominican amber
(ca. 4015 myr, Poinar 1992). Cretaceous amber
from Lebanon, N. Siberia (Tajmyr) and N.
America have so far yielded only few Lepidop-
tera which, however, are of particular interest be-
cause of their age.
The rocky sediments best known for lepidop-
teran remains are probably the Oligocene lacus-
trine beds at Florissant, Colorado, USA (Scud-
der 1889; Brown 1976, Tindale 1985, Emmel
et al. 1992). Other rich formations are sandy-clay
continental deposits near Stavropol, N. Cauca-
sus in Russia (Danilewskij & Martynova 1962;
Nekrutenko 1965; Kozlov 1988; Skalski 1988),
calcareous marls of the gypsum quarries at Aix-
en-Provence in France (Theobald 1937 a, 1937 b;
Leestmans 1983), and argillaceous limestone of
the Isle of Wight in England (Jarzembowski
1980); all of these are similarly from the Tertiary.
Further important strata for fossil Lepidoptera
are the Tertiary Gabbro in Italy (Rebel 1898),
Radoboj in Croatia (Scudder 1875), Rott (Scud-
der 1875, Burgeff 1951) and Randecker Maars
(Zeuner 1942) in Germany, Mo-clay in Denmark
(Larsson 1975), Creste (Nel & Descimon 1984)
and Dauphin (Henrotay 1986) in France, as well
as some Mid-Late Jurassic/Early Cretaceous for-
mations in Middle Asia (Skalski 1979 b, 1984;
Rasnitsyn 1983; Kozlov 1988, 1989).
The main sites for fossil Lepidoptera were
mapped by Schepdeal (1974) and Skalski (1976,
1977, 1979 b, 1979 c, 1979 d). These localities are
all in the Northern Hemisphere. It is only quite
recently that Pre-Quarternary Lepidoptera have
been reported from the Southern Hemisphere
(Martins-Neto & Vulcano 1989; Rozefelds
1988 a, b).
Systematics of Fossil Lepidoptera
Numerous lepidopteran families (in the classifi-
cation adopted here) have been identified (Skal-
ski 1990 b) in fossil resins, most of which in the
Baltic amber. The majority of the recorded fami-
lies belong to the microlepidopteran grade. Fos-
sil Lepidoptera recorded from rocky sediments
include several families that have not been found
in resins (Skalski 1990 b).
Lepidoptera inclusions in fossil resins are fre-
quently well preseved, and when lying in a fa-
vourable position they may be examined in some
16
detail. In Lepidoptera fossils from rock beds
usually only the wings lend themselves to closer
examination; the venation and/or a wing pattern
composed of light and dark elements are fre-
quently discernible (Figs. 2.7 8), but some early
interpretations have been marred by artifacts.
An instructive example is the widely reproduced
reconstruction (Rebel 1898) of the Tertiary papi-
lionid butterfly Doritites bosniaskii, in which
grooves on the rock surface near the wing mar-
gins appear as traces of a parnassiine-like wing
pattern.
A recurrent problem in the identification of
fossil Lepidoptera (as indeed of most fossil or-
ganisms) is that they so rarely display the auta-
pomorphies that are diagnostic of taxa recog-
nized in our classification. A very considerable
proportion of the family assignments of fossil
Lepidoptera proposed in previous literature are
based on superficial phenetic similarity and these
interpretations can therefore be characterized at
best as tentative, at worst as misleading. Several
of these family assignments have now been
rightly replaced by transfers to the incertae sedis
category by Carpenter (1992) and de Jong (in
press), but it is almost certain that several addi-
tional transfers of this kind still remain to be
made for fossil moths.
About one-third of the known lepidopteran
fossils have been described and named. Named
fossils have generally received more attention in
the literature than those (sometimes more impor-
tant ones) which have been described, but not
named. Mesozoic to Middle Tertiary Lepidop-
tera fossils have usually been assigned to extant
families but to extinct genera and species. Plio-
cene and Pleistocene fossils are recognized as re-
cent species.
Overview of fossil Lepidoptera
The oldest known evidence of the Lepidoptera is
from the Lower Jurassic (Whalley 1985). Triassic
fossil insects such as Eoses trissica (Tindale
1945), Eocorona iani (Tindale 1980), Mesoses op-
tata and M. magna (Riek 1976), and Curvicubitus
triassicus (Hong 1984) have been referred to the
Lepidoptera on inadequate evidence; their sys-
tematic status is discussed by Hennig (1969),
Skalski (1979 b), Sukatsheva (1982), Willmann
(1984, 1989), Whalley (1986) and Kozlov (1988).
Also some younger Mesozoic fossils have been
included erroneously in the Lepidoptera; the
most widely cited are probably the Jurassic
Palaeontinidae (Scudder 1875; Handlirsch
19061908), which were eventually transferred
to be Hemiptera. On the other hand some of the
Mesozoic fossils currently placed in the Trichop-
tera may well be stem-lineage Lepidoptera or
stem-lineage Amphiesmenoptera; the majority of
these fossils (Microptysmatidae, Cladochoristi-
dae, Dasyneuridae, Liassophilidae, Necrotaulii-
Niels P. Kristensen & Andrzej W. Skalski
dae etc.) do not exhibit any apomorphies to per-
mit their unambiguous assignment to the
Trichoptera.
Evidence of Basal Splits. Interpretations of early
lepidopteran diversification are inextricably
linked to interpretations of their host associa-
tions; see chapter 120 for a fuller discussion.
The bulk of the Mesozoic Lepidoptera so far de-
scribed are believed to belong to the non-glos-
satan grade, but it is admittedly only in excep-
tional cases (Parasabatinca) that head structures
are so well conserved that there is direct evidence
for this assumption. Two fossils, Protolepis cu-
prealata (Fig. 2.5 A) and Karataunia lapidaria
from the Upper Jurassic of Kazakhstan, have
been assigned to the Glossata-Ditrysia (Kozlov
1989), and while the evidence for this assignment
have proved spurious upon reexamination, the
presence of a coiled proboscis in Protolepis is still
believed to be correct (M. Kozlov pers. comm.).
If the existence of the Glossata in the Jurassic
can really be confirmed, then this would have
important consequences for the debate over
multiple origins of early lepidopteran angio-
sperm-feeding, since the earliest current records
of angiosperms are from the Lower Cretaceous
(Crane et al. 1995). In any case the attributing of
mines (of gallery type, with no frass observable)
in Upper Jurassic pteridospermophyte leaves
from Queensland to the Heteroneura-Nepticuli-
dae (Rozefelds 1988 b) cannot easily be accepted
on the basis of present knowledge. The earliest
convincing evidence for the existence of the Di-
trysia are leaf mines from the N. American Da-
kota Flora (97 myr BP) which can with great
certainty be referred to the Gracillariidae-Phyl-
locnistinae (Labandeira et al. 1994); obviously,
therefore, the origin of the Glossata must have
been considerably earlier. It would be unsurpris-
ing if future fossil discoveries reveal that several
splitting events even within the higher Ditrysia
had taken place by the end of the Cretaceous.
Unplaced Homoneurous Moths. The oldest
known fossil which can with great certainty be
referred to the Lepidoptera is the small scaly-
winged Archeolepis mane (Whalley 1985) from
the Lower Jurassic (Lias) of Dorset, England. It
was originally placed in a family of its own
(Archaeolepidae). Four moths from Upper Ju-
rassic to Lower Cretaceous aleuronitic tuftes in
Central Asia, described in the genera Eolepiop-
terix, Undopterix, Daiopterix and Palaeolepidop-
terix (Skalski 1979 b, 1984; Rasnitsyn 1983; Koz-
lov 1989) were similarly assigned to a family, and
even suborder, of their own (Eolepidopterigina-
Eolepidopterigidae), but there is no convincing
support for the monophyly of the latter. These
fossils are all female specimens in which extensi-
ble oviscapts and/or one or two pairs of long
oviscapt apophyses have been identified; hence
Phylogeny and Palaeontology
Fig. 2.4. Wing scales of Archeolepis mane, the oldest
lepidopteran so far identified with reasonable certainty
(photograph courtesy of dr. P. Whalley).
the female postabdomen conforms to the puta-
tive amphiesmenopteran ground plan. Palaeosa-
batinca and Auliepterix from the same deposits
as well as Gracilepterix from a Lower Cretaceous
limestone in Brazil (Martins-Neto & Vulcano
1989) were described in the Micropterigidae
(Kozlov 1988, 1989), but no apomorphies sup-
port this placement. Two moths described by
Martins-Neto & Vulcano (1989) from the same
formation as Gracilepterix were assigned to Un-
dopterix and Parasabatinca respectively (also
these genera were both considered by the authors
to be micropterigids), but again there are no reli-
able criteria for a family placement of the fossils
in question. The same is true for Cockerell's
(1919) Micropteryx pervetus from Burmese (Mio-
cene?) amber. The assignment of Electrocrania
immensipalpia, a Baltic amber moth with ex-
traordinarily large maxillary palps, to the Eri-
ocraniidae (Kusnezov 1941) is also not sup-
ported by convincing synapomorphies. It does
have unpaired mesotibial spurs, but the mesotib-
ial spur complement has been reduced indepen-
dently in several primitive moth lineages. Kozlov
(1988) synonymized Electrocrania with Micro-
pterix (Micropterigidae), but this family assign-
ment is contradicted by the presence of a meso-
tibial spur. It is unknown whether Electrocrania
was indeed homoneurous (or glossatan, for that
sake).
Micropterigoidea. Parasabatinca from lower Cre-
taceous amber of Lebanon (Whalley 1978) does
exhibit some apomorphies (shortened labial
palps, pronouncedly moniliform antennae)
which may permit its placement in the socalled
Sabatinca group of genera in the Micropterigidae
(Kristensen & Nielsen 1979). In the Tertiary the
family is represented by the extant genus Micro-
pterix, recorded from the Baltic amber and marls
of the Isle of Wight, and Sabatinca-group taxa
17
ae
Fig. 2.5. Mesozoic basal Lepidoptera. A. Protolepis cu-
prealata. The appendage indicated by arrow has been
interpreted as the proboscis, but it cannot be ruled out
that it is a maxillary palp (which in extant basal Glos-
sata is a much more conspicuous formation than the
proboscis). B. Lophocoronid-like moth from Siberia,
forewing venation and male postabdomen. A after
Kozlov (1989).
from the Baltic and Saxonian amber (Jarzem-
bowski 1980; Skalski 1989, 1995, in press b).
'Eriocraniid-Grade' Families. Fossil evidence of
Eriocraniidae comes in the form of Tertiary leaf-
mines (Opler 1973; Jarzembowski 1980).
An interesting moth (Fig. 2.5 B) in Upper Cre-
taceous amber from Siberia (Skalski 1979 a)
shares two apomorphies with the Lophocoroni-
dae, viz., a postapical Rs4 and a bilobed valve
with a strong apical seta on the lower lobe (in
extant lophocoronids this seta is, however, short
and blunt); the very long phallus has, beyond
mid-length, a dorsal process which is without
counterpart in extant homoneurous moths. The
moth is now being described in the Lophocoroni-
dae (in a subfamily of its own, Skalski in press
c), but its assignment to this family, the extant
range of which is restricted to Australia, must be
considered very tentative indeed; the support for
this assignment is considered inadequate by Niel-
sen & Kristensen (1996).
Exoporia. The surprise record of Mnesarchaei-
dae, presently restricted to New Zealand, in Up-
18
per Cretaceous Siberian Amber (Zherihin & Su-
katsheva 1973) is inadequately substantiated; the
strong family autapomorphies are not in the
wing venation.
Hepialoids have been recorded from Palaeo-
gene Isle of Wight limestone (Robinson 1977,
Jarzembowski 1980) and mid-Miocene continen-
tal deposits in China (Zhag 1989).
Unplaced Heteroneura. Stem-lineage Hetero-
neura were undoubtedly superficially similar to
Incurvarioidea in many aspects, and records of
this superfamily from the Cretaceous (Whalley
1978, Skalski 1979) must be considered uncer-
tain, since the pertinent autapomorphies (16)
cannot be observed in these specimens.
A larva from Canadian Cretaceous amber (the
first pre-Tertiary lepidopteran fossil to be discov-
ered, MacKay 1970) has been assigned to the Ti-
neoidea, but while it certainly belongs in the
Glossata (it has a distinct spinneret), it cannot
be placed with any greater precision. About 30%
of all lepidopteran inclusions in the Baltic amber
have been referred to the Tineidae. However,
since the monophyly of this family remains de-
batable, and that of the superfamily Tineoidea
is supported by characters which are difficult to
observe (17), many of these fossils may well
belong to basal grades in other ditrysian, or just
heteroneurous, lineages. Tineoid-grade moths
are also numerous in Dominican amber, and
have been found in Mexican amber as well. A
few have been found in Tertiary rock beds (Rebel
1934; Kusnezov 1941; Skalski 1974, 1977; Jar-
zembowski 1980; Kozlov 1987). Prolyonetia from
the Baltic amber was assigned to a family of its
own (Kusnezov 1941), close to the Lyonetiidae.
Its venation is less modified than in extant Lyo-
netiidae, and it has no eye-cap. Its fully devel-
oped maxillary palp precludes a systematic as-
signment 'above' the tineoid grade.
Nepticuloidea. While the above-mentioned al-
leged presence of nepticulids in the Jurassic is
questionable, several leaf mines in the mid-Creta-
ceous Dakota Flora exhibit so detailed similari-
ties with those made by extant nepticulids in
Stigmella and Ectoedemia that their assignment
to these genera seems justified (Labandeira et al.
1994). A minute undescribed moth from Cana-
dian Upper Cretaceous amber (Mutuura & Kris-
tensen, unpublished) may also be nepticulid. The
family is represented by two adults in the Baltic
amber (one referred to Ectoedemia), and one in
African copal (referred to the Recent genus Aca-
lyptris (= Niepeltia)), and many leaf-mines from
the Tertiary have been assigned to it (Skalski
1976, 1979 a, 1990 a, 1992; Kozlov 1988, Roze-
felds 1988 a).
Incurvarioidea are recorded mainly from the Bal-
tic amber. Two adelid species have been de-
Niels P. Kristensen & Andrzej W. Skalski
scribed in the recent genus Adela. See also 'Un-
placed Heteroneura'.
Tineoidea. See 'Unplaced Heteroneura'. All de-
scribed tineoids are placed in fossil genera, ex-
cept that species in the Dominican amber have
been referred to the extant tineid genera Choro-
pleca and Opogona (Skalski unpublished).
Psychidae are known mainly from cases
('bags') in the Baltic amber; a few compression
fossils of adults have also been reported (Kozlov
1988). Acrolophidae are recorded from Domini-
can amber (Skalski in press d). Bucculatricidae
are recorded from the Late Cretaceous to Oligo-
cene on the base of leaf-mines (Opler 1973, 1982;
Strauss 1977; Crane & Jarzembowski 1980; Koz-
lov 1988).
Gracillarioidea. Leaf mines exhibiting detailed
similarities with those of extant Gracillariidae-
Phyllocnistinae are present in the mid-Creta-
ceous D a k o t a Flora (Labandeira et al. 1994),
and somewhat similar mines have been identified
in (similarly N. American) Eocene deposits
(Hickey & Hodges 1975). Adult Gracillaridae are
recorded from Baltic and Dominican amber.
Yponomeutoidea. The families Yponomeutidae,
Plutellidae, Heliodinidae are recorded from Bal-
tic amber (see also 'Unplaced Heteroneura'). An
(undescribed) heliodinid species is represented by
more than 70 specimens, and is a dominant
taxon in this resin (Skalski 1976).
Gelechioidea. About 30% of the Baltic amber
Lepidoptera have been referred to the Oecopho-
ridae (almost as many as to the 'Tineidae'), but
since the Oecophoridae as previously delimited
are paraphyletic, many of these fossils presuma-
bly belong in the stem-lineages of other gelechi-
oid families. Oecophorid-like moths are also nu-
merous in the Dominican amber. Among 20 de-
scribed species 19 are placed in fossil genera and
one is refered to the recent genus Schiffermuel-
leria.
A few Gelechiidae and Elachistidae-Elachisti-
nae are recorded from the Baltic amber, while
records of Cosmopterigidae have come from the
Mexican amber only (Rebel 1935/1936; Kus-
nezov 1941; Skalski 1973 a, 1976, 1977, 1979 c;
Kozlov 1988).
Zygaenoidea. Zygaenids are known from the Mi-
ocene of Germany (Reiss 1936; Burgeff 1951;
Naumann 1987) and Spain (Fernandez-Rubio
et al. 1991). The systematic position of an Oligo-
cene moth from Aix-en-provence, determined as
a Zygaena species (Leestmans 1983), remains un-
clear. Fossil zygaenids are discussed by Faille
(1994).
Cossoidea. Cossidae are recorded from the Flo-
rissant (Cockerell 1926); the alleged cossid Xy-
Phylogeny and Palaeontology
leutites miocenicus from Russian Miocene (Koz-
hanchikov 1957; Martynova 1960) was transfer-
red to the Noctuidae by Kozlov (1988).
Sesioidea. Castniidae are recorded from the Flo-
rissant on the basis of a well preserved fore wing
(Tindale 1985).
Tortricoidea. Tortricidae are recorded from fossil
resins (Baltic, Mexican and Dominican amber;
African copal; Skalski 1973 b, 1976, 1992 b; Koz-
lov 1988; Poinar & Brown 1993) and the Floris-
sant (Cockerell 1909).
Pterophoroidea. The Pterophoridae are known
from one specimen, well preserved in the Oligo-
cene shales near Aix-en-Provence and described
in the extant genus Pterophorus (Bigot, Nel &
Nel 1986).
Pyraloid-Grade Families. A copromorphid is re-
corded from English Palaeogene marls (Jarzem-
bowski 1980). Thyrididae are recorded from the
Baltic amber (Skalski 1992 a), but the Eocene
Hexeritis from Colorado, described in this fam-
ily almost certainly does not belong here (Whal-
ley 1971). A few Pyraloidea are recorded from
the Baltic amber (Kusnezov 1941), and wing
fragments in Palaeocene to Oligocene sediments
from England and France are likewise referred
to this superfamily (Jarzembowski 1980).
Bombycoidea. The German Pliocene Sphingid-
ites, known from remains of a larva with a poste-
rior horn, is presumably a sphingid (Kernbach
1967).
Geometroidea. Geometridae are recorded from
the Florissant (Cockerell 1922). Wing fragments
from Paleocene to Oligocene English sediments,
and a larva from a German Pliocene deposit are
also referred to this family (Jarzembowski 1980,
Kernbach 1967). So are specimens in African co-
pal (Evers 1907, Skalski unpublished).
Hesperioidea + Papilionoidea. The fossil repre-
sentatives of butterflies are unusually well under-
stood, due to de Jong's (in press) critical review
of previous accounts. Butterfly fossils are partic-
ularly abundant in the Florissant formation
(Scudder 1889; Brown 1976; Tindal 1985, Emmel
el al. ); in other localities usually only single spec-
imens have been found. The oldest named but-
terflies are Praepapilio colorado, P. gracilis (Papi-
lionidae) and Riodinella nympha (Lycaenidae)
from the Eocene Green River Shale of Colorado
(Durden & Rose 1978), but an apparent hesperi-
oid (de Jong 1. c.) butterfly (Fig. 2.6) has recently
been identified from the Upper Paleocene of Fur,
Denmark. Interestingly, the Praepapilio species
exhibit a plesiomorphic trait (trifid, not quadri-
fid hind margin of forewing discal cell) not re-
19
Fig. 2.6. An apparent hesperioid butterfly from the
Upper Paleocene, Fur, Denmark; wings folded below
body. If correctly identified this is the oldest butterfly
fossil so far known.
tained in extant members of the family. About
20 fossil species have been assigned to the
Nymphalidae, including 2 species tentatively re-
ferred to the Libytheinae and 3 to the Satyrinae;
in several cases an 'open cell' in the hindwing
suggests assignments to the Nymphalinae. All
other butterfly families have very few known fos-
sil representatives: Lycaenidae s. lat. 1, Pieridae
3 (family assignments tentative, except in the
case of a Pleistocene/post-Pleistocene African co-
pal specimen indistinguishable from an extant
Belenois species), Papilionidae 5 and Hesperiidae
1. Many fossil butterflies have the wing pattern
well preserved, e. g. the nymphalids Neorinops
sepulta, Prodryas persephone (Fig. 2.7), Apanth-
esis leuce and 'Agais' karaganica. Most butter-
flies from the Eocene to Miocene are placed in
extinct genera, but they are in some cases phenet-
ically similar to, and probably closely related to,
extant taxa. For instance, in the Papilionidae-
Parnassiinae the Miocene Doritites from Italy is
probably the sister group of Bhutanitis + Luehd-
orfia, and the Oligocene Thaites from France is
close to the most recent common ancestor of
Parnassius + Hypermnestra (de Jong in press).
On the other hand some previously reported
cases of particularly close relationships between
fossil and recent taxa (e. g. between the Oligo-
cene 'Vanessa' amerindica and the extant Vanessa
indica, and between the Miocene 'Vanessa' kara-
ganica and the extant Agais urticae) were shown
by de Jong to be founded on inadequate evi-
dence. Reports of extant forms of younger (Plio-
cene and Pleistocene) age are unsurprising
(Zeuner 1942, 1943; Branscheid 1968, 1969; Fuji-
yama 1968, 1983; Skalski 1976). While earlier
claims of findings of butterflies in the Baltic am-
ber have not been confirmed, they have been
found in Dominican amber (Poinar 1992) and
African copal.
Noctuoidea. Fossil Notodontidae, Noctuidae and
Arctiidae are recorded from the Eocene to the
20
Fig. 2.7. Prodryas persephone (probably N y m p h a l i d a e - N y m p h a l i n a e ) , a f a m o u s butterfly fossil f r o m the Floris-
sant (Oligocene).
Fig. 2.8. Stauropolia nekrulenkoi (Arctiidae?); a m o t h
fossil with unusually well preserved wing p a t t e r n .
Pleistocene (Schille 1914; Thobald 1937 a; Kern-
bach 1967; Leestmans 1983; Kozlov 1988; Skal-
ski 1988). The assignment of a Cretaceous lepi-
dopteran egg to the Noctuidae is uncertain (see
below). Most taxa have been identified on the
basis of single wings, and family assignments are
mostly weakly founded. Stauropolia nekrutenkoi,
described as an arctiid, is notable for its well pre-
served wing pattern (Fig. 2.8).
Niels P. Kristensen & A n d r z e j W. Skalski
Biogeographic and palaeoecological aspects
Fossil Lepidoptera add little to the k n o w n geo-
graphical ranges of individual lepidopteran
clades. The Baltic amber m o t h f a u n a seems to be
overall similar to that of the present-day Holarc-
tic, and Lepidoptera in Mexican and Dominican
ambers resemble recent taxa in the Neotropical
region.
Exceptions do occur. Micropterigids referred
to the Sabatinca group of genera occurred in the
Cretaceous in Lebanon and in the Eocene in the
'Baltic amber area'; these moths are now absent
f r o m the Western Palaearctic, but it must be em-
phasized that the monophyly of this genus group
is not established beyond d o u b t (14). Castnii-
dae, recorded f r o m Florissant, are now absent
f r o m N . America, but occur in the Neotropics.
And while the Tertiary parnassiine papilionids
Doritites amd Thaites were West Palaearctic,
their presumed extant sister taxa are East Pa-
laearctic.
Phoretic/ectoparasitic relationships between
Arachnida and Lepidoptera have (not unexpect-
edly) existed since the Eocene, as evidenced by a
Baltic amber gelechiid m o t h carrying a phoretic
pseudoscorpion (Skalski 1994). Poinar et al.
(1991) recorded parasitic erythraeid mites f r o m
Phylogeny and Palaeontology
a tineoid and a gracillarioid moth in Dominican
amber.
A lepidopteran egg in an Upper Cretaceous
deposit has aroused particular interest, because
it was referred to the Noctuidae (Gall & Tiffney
1983). It has been commonplace to think of bat
prdation as the chief selective agent promoting
the evolution of tympanic organs in nocturnal
Lepidoptera, and while tympanate moth lineages
were therefore not expected to predate the bats
in the fossil record, the latter group are actually
so far not known before the Eocene. Two caveats
are in place. Firstly, the bat lineage may, of
course, be much older than documented by its
fossil record. And secondly, no diagnostic auta-
pomorphies in the egg structure of the higher
lepidopteran lineages, including the Noctuidae
(or Noctuoidea), have actually been identified
(Hinton 1981); the family assignment, based on
phenetic similarity, therefore must be considered
very tentative at best.
As noted above, evidence for considerable an-
tiquity of some lepidopteran host relationships is
provided by leaf mines, closely resembling those
of extant 'microlepidopteran' taxa, in leaves as-
signed to extant plant families (Cretaceous) or
genera (Tertiary) (Hickey & Hodges 1975; Opler
1973, 1974, 1982; Labandeira et al. 1994). See
1-20.
Acknowledgments
We are indebted to Dr. M. J. Scoble for critical
comments of a draft of this chapter, and to Drs.
M. V. Kozlov and J. Minet for valuable informa-
tion. Mr. G. Brovad and Mr. S. Langemark pro-
vided much appreciated assistance with illustra-
tions.
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 3. Classification and Keys to Higher Taxa
David J. Carter & Niels P. Rristensen
(D. J. Carter: Key to families, larvae. . P. Kristensen: Classification; Key to superfamilies, adults)
Classification
The systematics chapters in this Handbook are all
written on the basis of 'Hennigian' phylogenetic
systematics or cladistics. Therefore, taxa which
are identified as likely poly- or paraphyletic (i. e.,
which do not include all descendants of the
members' last common ancestor) have been re-
jected, or they are accepted merely as prelimi-
nary groupings, pending further analysis.
Phylogenetic systematists are not, however, in
agreement about how even a fully resolved phy-
logeny is best transformed into a written classifi-
cation. Indeed it has been argued that 'classifica-
tion' is an inappropriate term in this context:
phylogentic systematists are concerned not with
'classifying', but with 'systematizing', because
monophyla/clades are individuals forming parts
of a 'system', rather than elements of definable
classes (Griffiths 1974). Adherents of this view
claim that the Linnean categories have outlived
their usefulness (see e. g. Ax 1984/1987; Will-
mann 1987, 1989; de Queiroz & Gauthier 1992),
and that the optimal written presentation of the
phylogenetic subordination is an indented (or
numerically coded) list, in which all recognized
clades are named but not given categorial ranks.
The principal alternative approach has been
termed the 'annotated Linnean classification'.
Here the Linnean categories are retained (even
an expanded array of them), but a suite of con-
ventions, most importantly that of 'sequencing',
may be adopted to permit an exact representa-
tion of the cladistic information while prolifera-
tion of taxon names as well as changes in previ-
ous systems are minimized (for more detailed ac-
counts see e. g. Wiley 1981, Forey 1992).
It seems immediately obvious that the two
kinds of approach each has its merits and prob-
lems, and the Lepidoptera system adopted here
embodies elements of both. It is accepted that
the notion of categorial ranks, rooted in pre-evo-
lutionary systematics, may represent an unneces-
sary impediment to efficient systematization.
Formal categories have therefore been largely
abandoned in the present system: no lepidop-
teran 'suborders', 'infraorders' etc. are recog-
nized here. Use of the redundant names Zeug-
loptera, Aglossata and Heterobathmiina (for the
superfamilies Micropterigoidea, Agathiphago-
idea and Heterobathmioidea respectively) is dis-
continued altogether, and named high-rank
monophyla (Glossata, Ditrysia, etc.) are simply
designated as 'clades'.
The important (suprageneric) exceptions to
this principle are the categories at subfamily/
family/superfamily level (for which the Interna-
tional Code of Zoological Nomenclature pre-
scribes specific suffixes). The system adopted re-
flects the firm belief that the family category has
come to play such an important role in the lan-
guage by which biologists communicate (thus it
is pivotal in many catalogue projects) that its
abandonment now, for whatever reason, would
be counterproductive. Hence every recent genus
should be assigned to a family (a view with
which 'pure systematizers' would not agree). A
similar case may be made for superfamilies. Ar-
guably the superfamily category has for some
time been so extensively used in Lepidoptera sys-
tematics (the same may not be true in all other
major insect groups) that it is desirable that every
family is assigned to a superfamily. For this
reason new, and admittedly redundant, super-
families are here created for the three families
which were left outside the superfamilies in pre-
vious Lepidoptera systems. Needless to say, it is
disturbing that no less than 27 of the 46 super-
families here recognized are redundant, compris-
ing only a single family. Extensive amalgam-
ations of such superfamilies with others are
highly desirable, to the extent future research can
disclose convincing synapomorphies to support
them. A subfamily classification has been
adopted in 48 of the 121 lepidopteran families
here recognized; subfamilies have very variable
status in the literature, but in some cases they
are entities of widespread and longstanding rec-
ognition.
Table 1 presents a synoptic classification of the
lepidopteran superfamilies recognized in this
Handbook; phylogenetic systems of individual
superfamilies have been worked out at very dif-
ferent levels of refinement, as discussed in chap-
ters 14 through 1 - 1 9 . The table adopts the se-
quencing, plesion, incertae sedis and sedis muta-
bilis conventions of the 'annotated Linnean clas-
sification' rather than a consistent subordina-
tion. Names have been applied only to a selec-
tion of the more inclusive clades recognized; the
choice at least partly reflects an assessment of
the firmness of the support for the monophyly
of the clades in question and/or the conspicuous-
ness of their autapomorphies. Naturally these
decisions are open to debate. Thus the hypothe-
sized clade comprising all non-micropterigoid
Lepidoptera has been left un-named, though its
28 David J. Carter & Niels P. Kristensen

Table 1. Synoptic phylogenetic classification of lepidopteran superfamilies recognized in the present work: The
sequencing convention applies unless taxa are stated to be incertae sedis or sedis mutabilis. Figures in brackets
after superfamily names indicate numbers of families recognized.
Plesions Archeolepis, Eolepidopterix, Undopterix, Dai- 22. COSSOIDEA (2)
opterix, Palaeolepidopterix, Auliepterix, Gracilepterix: 23. TORTRICOIDEA (1)
Lepidoptera incertae sedis* 24. CHOREUTOIDEA (1)
1. MICROPTERIGOIDEA (1) 25. URODOIDEA (1)
2. AGATHIPHAGOIDEA (1) 26. SCHRECKENSTEINIOIDEA (1)
3. HETEROBATHMIOIDEA (1) 27. EPERMENIOIDEA (1)
Superfamilies 4 - 4 6 : clade GLOSSATA 28. ALUCITOIDEA (2)
4. ERIOCRANIOIDEA (1) 29. PTEROPHOROIDEA (1)
Superfamilies 5-46: clade COELOLEPIDA Superfamilies 30-46: clade OBTECTOMERA
5. ACANTHOPTEROCTETOIDEA (1) Superfamilies 30, 31, 32, 33, 34, 35, 36-46: sedis
6. LOPHOCORONOIDEA (1) mutabilis
Superfamilies 7 - 4 6 : clade MYOGLOSSATA 30. WHALLEYANOIDEA superfam. nov. (1)
7. NEOPSEUSTOIDEA (1) 31. IMMOIDEA (1)
Superfamilies 8 - 4 6 : clade NEOLEPIDOPTERA 32. COPROMORPHOIDEA (2)
Superfamilies 8 - 9 : clade EXOPORIA 33. HYBLAEOIDEA (1)
8. MNESARCHAEOIDEA (1) 34. THYRIDOIDEA (1)
9. HEPIALOIDEA (5) 35. PYRALOIDEA (2)
Superfamilies 10-46: clade HETERONEURA Superfamilies 36-46: clade MACROLEPIDOP-
10. NEPTICULOIDEA (2) TERA
11. INCURVARIOIDEA (6) Superfamilies 3638, 39-45, 46: sedis mutabilis
Superfamilies 12, 13, 14-46: sedis mutabilis Superfamilies 36, 37, 38: sedis mutabilis
12. PALAEPHATOIDEA (1) 36. MIMALLONOIDEA (1)
13. TISCHERIOIDEA (1) 37. LASIOCAMPOIDEA (2)
Superfamilies 14-46: clade DITRYSIA 38. BOMBYCOIDEA (9)
Superfamily 14: Ditrysia incertae sedis Superfamilies 39, 40, 41 - 4 3 , 44-45: sedis muta-
14. SIMAETHISTOIDEA superfam. nov. (1) bilis
15. TINEOIDEA (5) 39. AXIOIDEA (1)
16. GRACILLARIOIDEA (4) 40. CALLIDULOIDEA (1)
17. YPONOMEUTOIDEA (8) 41. HEDYLOIDEA (1)
18. GELECHIOIDEA (15) 42. HESPERIOIDEA (1)
Superfamilies 19-46: clade APODITRYS1A 43. PAPILIONOIDEA (4)
Superfamilies 19, 20, 21-23, 24, 25, 26, 27, 2 8 - 44. DREPANOIDEA (2)
29, 30, 31-46: sedis mutabilis 45. GEOM ETROIDEA (3)
19. GALACT1COIDEA superfam. nov. (1) 46. NOCTUOIDEA (8)
20. YG A EN 01 DEA (11)
21. SESIOIDEA (3) * See 1 - 2 .

monophyly may be better corroborated than
that of, e. g., the Obtectomera.
It must be emphasized that the choice of clas-
sificatory approach ultimately depends on judge-
ments which are by necessity subjective. For ex-
ample, the systematizers' attitude to subordina-
tion/naming is dependent on the clearly sub-
jective belief that the (potentially very numerous)
new names needed for all recognized clades
above the generic level do not "represent a com-
plication worth mentioning" (Willmann 1987:
228); the refusal of consistently naming such
clades reflects the equally subjective view that
the opposite is true. Presumably, however, there
will be broad agreement that the principal con-
cern of phylogenetic systematists should be the
cladograms, their support and the biological in-
ferences that can be drawn from them, rather
than the written classifications/systems per se.
Key to superfamilies, adults
Modified from Nielsen & Common 1991. All
keys to higher taxa must be used with reserva-
tion, and the results should be checked against
descriptions. This key will not work for brachyp-
terous/wingless forms, for which reference
should be made to the reviews by Heppner
(1991) and, particularly, Sattler (1991).
1 Hindwing with Rs unbranched; fore-
wing with jugal lobe not markedly pro-
duced Heteroneura, 11
Hindwing Rs with 3 or 4 branches; fore-
wing almost always with jugal lobe
markedly produced 2
2 (1) Maxillary galeae forming a proboscis,
usually spirally coiled in repose, some-
times secondarily reduced or absent;
mandibles often strongly reduced, artic-
ulations with head capsule undeveloped
Glossata, 5
Maxillary galeae unmodified, not form-
ing proboscis; mandibles large; articula-
tions with head capsule well developed
3
3 (2) M4 present; ocelli absent; tibial spurs 1-
4-4; larger, caddisfly-like moths; fore-
Classification and Keys to Higher Taxa
wing pattern mottled brownish, non-iri-
descent; Australia, SW Pacific
Agathiphagoidea (1 - 4 )
M4 absent; ocelli present; tibial spurs 0-
0-4; smaller moths (wingspan at most 13(12)
ca. 16 mm., usually much smaller; fore-
wings often iridescent 4 prominent, blunt spines often forming
4 (3) Forewing Sc forked; pterostigma ab-
sent; hindwing without produced jugal
lobe; all regions
Micropterigoidea (14)
Forewing Sc simple; pterostigma pre-
sent; hindwing with distinct jugal lobe
(Fig. 4.1)); temperate S. America . . . .
Heterobathmioidea (14)
5(2) Mediumsized to very large, robust 14(13)
moths; proboscis and maxillary palps
strongly reduced
Hepialoidea (part, 15)
Mediumsized to small moths, wingspan
not exceeding 27 mm, often much
smaller; proboscis and maxillary palps
often well developed 6 Tischerioidea (16)
6(5) Forewing Rs4 postapical 7 15(11)
Forewing Rs4 apical or preapical . . . 9
7(6) Forewing Rs3 postapical; larger, more 16(15)
broad-winged moths (wingspan at least
ca. 12 mm, usually much larger)
Hepialoidea (part, 15)
Forewing Rs3 preapical; small, narrow-
winged moths 8 Labial palps porrect, beak-like or as-
8 (7) Maxillary palps well developed, 5-seg-
mented, with marked bends between
segments 1/2 and 3/4; Australia
Lophocoronoidea (15)
Maxillary palps very small, at most 3- 17(16)
segmented, porrect; New Zealand . . . .
Mnesarchaeoidea (15)
9 (6) Meso-tibiae with paired apical spurs;
larger, often markedly broad-winged
moths (wingspan 1327 mm) SE. Asia,
temperate S. America
Neopseustoidea (15) 18(15)
Mesotibiae with unpaired apical spurs;
small (wingspan at most ca. 16 mm usu-
ally markedly smaller) narrow-winged
moths 10 19(18)
10(9) Ocelli present, M l not stalked with Rs
veins Eriocranioidea (15)
Ocelli absent, M1 stalked with Rs . . .
Acanthopteroctetoidea (15)
11(1) Sternum II with paired anterior apo- 20(19)
demes; wing membranes devoid of
microtrichia; female genital system with
copulatory orifice (on VIII) separate
from more posterior ovipore
Ditrysia 15
Sternum II without anterior apodemes;
wing membranes usually with more or
less extensive microtrichiation; female 21 (18)
with single genital orifice 12
12(11) Antennal scape with eye-cap; hind tib-
29
iae with prominent spines
Nepticuloidea ( 1 - 6 )
Antennal scape without eye-cap; hind
tibiae without spines 13
Proboscis scaled; labial palp with lateral
bristles on segment 2; valvae with
comb (pectinifer); ovipositor piercing,
without medial sensory ridge
Incurvarioidea (16)
Proboscis scaled or unsealed; labial palp
without lateral bristles; $ valvae with-
out prominent, blunt spines; ovipositor
non-piercing, with medial sensory ridge
14
Proboscis unsealed; broad-winged
micro-moths, wingspan up to 30 mm.
Southern S. America, Australia, S.
Africa Palaephatoidea (16)
Proboscis scaled at base; narrow-
winged micro-moths, wingspan less
than 12 mm; Holarctic, tropical
Proboscis scaled 16
Proboscis unsealed or absent 18
Labial palps recurved, apical segment
may exceed vertex, usually tapering;
chaetosemata absent; sternum II of 'tin-
eid type' (see couplet 40); tympanal or-
gans absent Gelechioidea {1-9)
cending; chaetosemata present or ab-
sent; sternum II of 'tortricoid type';
tympanal organs present at base of ab-
domen, or absent 17
Labial palps ascending; fore wing with
CuP present near termen; tympanal or-
gans absent . . . Choreutoidea (1 13)
Labial palps porrect, beak-like or as-
cending; CuP usually absent; tympanal
organs present at base of abdomen . . .
Pyraloidea (1 14)
Antennae gradually or abruptly
clubbed, tip some times hooked . . . 19
Antennae filiform or tapered, often cili-
ate, pectinate or plumose 21
Hind wing with frenulum; ocelli pre-
sent, large; chaetosemata absent
Sesioidea (part, 111)
Hind wing usually without frenulum;
ocelli absent; chaetosemata present . 20
Antennae widely separated at base; with
subapical thickening; fore wing with all
veins from discal cell arising separately
Hesperioidea (1 16)
Antennae approximated at base, with
apical thickening; fore wing usually
with at least some peripheral veins
stalked Papilionoidea (1 16)
Tympanal organs present in metathorax
or ventrally in base of abdomen . . . 22
Tympanal organs absent 26
30
22 (21) Tympanal organs in metathorax, coun-
ter-tympanal cavities usually at base of
abdomen Noctuoidea (119)
Tympanal organs in abdomen . . . . 23
23 (22) Chaetosemata absent 24
Chaetosemata present
Geometroidea (1 17)
24 (23) Fore wing broad, often falcate
Drepanoidea (1 17)
Fore wing narrow, not falcate . . . . 25
25 (24) Fore wing reddish brown, with silvery
marks Cossoidea (part, 1 - 1 1 )
Fore wing grey . Tineoidea (part, 1 7)
26 (21) Wings more or less deeply cleft into two
or more plumes 27
Wings not cleft 28
27 (26) Hind wing divided into 3 plumes . . . .
Pterophoroidea (part, 1 13)
Hind wing divided into 6 - 7 plumes . .
Alucitoidea (part, 1 - 1 3 )
28 (26) CuP absent as a tubular vein in fore
wing 29
CuP well developed, at least near mar-
gin, as a tubular vein in fore wing . 40
29 (28) Abdominal segment VII with pocket-
like concavities associated with spira-
cles; medium-sized moths with silvery
markings on forewing (Figs. 15.2 A, B);
sub-tropical W. Palaearctic
Axioidea (1 15)
Abdominal segment VII without such
concavities 30
30 (29) Broad-winged micro-moths (wingspan
< 20 mm) with prominent ocelli; head
vestiture or very narrow scales with
deeply scalloped apices; Australasian . .
Simaethistoidea (1 13)
Not as above 31
31 (30) Head vestiture of piliform scales, if
mixed with lamellar scales (Mimallo-
noidea, part) frenulum small or absent;
robust, medium-sized to very large
moths . 'Bombycoid assemblage' (1 18)
Head vestiture of lamellar scales; frenu-
lum well developed; small to medium-
sized moths 32
32 (31) Hind wing CuA with pecten; fore wings
often with raised scales
Copromorphoidea (part, 1 13)
Hind wing CuA without pecten; fore
wing without raised scales 33
33 (32) Abdominal terga with anterior band of
spines; sternum II with V-shaped sclero-
tization . . . . Alucitoidea (part, 1 - 1 3 )
Abdominal terga without spines; ster-
num II without V-shaped sclerotization
34
34 (33) Labial palps beak-like; S hind coxa with
apical process and tibia with hair-pen-
cil; fore wing with 2 A sinuous
Hyblaeoidea (1 13)
Labial palps not beak-like; hind tibia
David J. Carter & Niels P. Kristensen
without apical lobe; 2 A in fore wing
not sinuous 35
35 (34) Robust medium-sized and relatively
narrow-winged moths with aposematic
colours . . . . Thyridoidea (part, 1 13)
Facies different 36
36(35) Pronouncedly narrow-winged (forewing
w/1 ratio 1/4 or less) and delicate moths
37
Wings broader, forewing w/1 ratio 1/3 or
more 38
37 (36) Small moths, wingspan < ca. 15 mm;
wings entire; body and legs not unusu-
ally long . Schreckeristeinioidea (113)
Mostly larger moths, wingspan often
considerably exceeding 15 mm (always
if wings entire); wings often cleft; body
and legs unusually long
Pterophoroidea (part) (1 13)
38 (36) Abdominal tergum I with large post-
spiracular expansion laterad from 'ter-
gal rim' (Fig. 13.3 H); medium-sized
moths, often with cryptic 'netted' wing
pattern . . . . Thyridoidea (part, 1 13)
Abdominal tergum I without such ex-
pansions 39
39 (38) Strong spines present on all tarsomeres;
facies as Fig. 13.41; Madagascar only .
Whalleyanoidea (113)
Strong spines present absent from api-
cal foreleg tarsomere; facies variable;
Madagascar, Asia Calliduloidea (1 15)
40 (28) Sternum II of 'tineid' type: anterolateral
corners not produced, apodemes long
and slender; usually small delicate spe-
cies, usually without raised scales on
fore wing 41
Sternum II of 'tortricoid' type: antero-
lateral corners usually distinctly pro-
duced, apodemes with broad bases, usu-
ally short and not appearing as continu-
ations of sternal ridges, ('venulae'); usu-
ally robust, small to very large species;
sometimes with raised scales on fore
wing 42
41 (40) Abdominal segment VIII of $ with
prominent lobes; fore wing with Rs4
often to termen and M usually absent
from discal cell Yponomeutoidea ( 1 - 8 )
Abdominal segment VIII of $ without
lobes; fore wing with Rs4 often to costa;
M usually present in discal cell
Tineoidea (part, 17)
42 (40) Fore wing with raised scales on surface
or along dorsum 43
Fore wing without raised scales . . . 44
43 (42) Fore wing with raised scales along dor-
sum; wing narrow, almost parallel-
sided Epermenioidea (13)
Raised scales on fore wing not restricted
to dorsum; wing relatively narrow, rect-
angular Copromorphoidea (part, 113)
Classification and Keys to Higher Taxa
44 (42) Chaetosemata present; ocelli absent;
fore wing without chorda and M stem
in cell Immoidea ( 1 - 1 3 )
Chaetosemata present or absent; ocelli
present or absent; fore wing often with
both chorda and M in cell 45
45 (44) Lower part of frons with ascending
scales; ocelli and chaetosemata present;
ovipositor lobes leaf-like
Tortricoidea (1 12)
Lower part of frons without ascending
scales; ocelli and chaetosemata rarely
both present; ovipositor lobes not leaf-
like 46
46 (45) M tubular and forked in cell 47
M absent in cell, or M not present in
cell as a complete, forked tubular vein
48
47 (46) Chaetosemata absent; ocelli rarely pre-
sent Cossoidea (1-11)
Chaetosemata and ocelli present . . . .
Zygaenoidea (part, 1 10)
48 (46) Patagia large, extending ventrad be-
yond anteroventral corres of pronotum
Sesioidea (part, 1 11)
Patagia relatively smaller, not extending
beyond anteroventral corres of prono-
tum 49
49(48) Ocelli present
Zygaenoidea (part, 1 - 1 0 )
Ocelli present 50
50 (49) Forewing CuP distinct throughout . . .
Zygaenoidea (part, 1 10)
Forewing CuP distinct only at margin
and (sometimes) at base. Some smaller
groups of apoditrysian moths with
poorly understood affinities will key out
here; they are currently placed in the su-
perfamilies . . . . Galacticoidea (1 13),
Urodoidea (1 13) and Zygaenoidea
(1-10).
Key to families: Larvae
Identification of lepidopterous larvae often
proves to be difficult, largely due to the fact that
the larvae of relatively few species have been de-
scribed in detail and comprehensive reference
collections of immature stages are scarce. This is
because Lepidoptera classification is tradition-
ally based on studies of imagines. However,
modern studies are increasingly taking into ac-
count the immatures stages, which provide many
vital clues to systematic and phylogenetic rela-
tionships. Identification of immatures is also ex-
tremely important to the economic entomologist
as the larva stage is almost invariably the one
causing damage to crops and other commodities.
The present key has evolved by the refinement
and expansion of previous more restricted keys
31
to larvae (Carter, 1984; Carter in Hollo way,
Bradley & Carter, 1987) to include as many fami-
lies as possible. It is based largely on final instar
larvae alhough earlier instars are taken into ac-
count where practically possible. It should be
borne in mind that larval instars may differ very
considerably. Many species with very hairy final
instar larvae have first instar larvae that bear pri-
mary setae only.
The larvae of many Lepidoptera families are
very poorly known and key characters for some
are based on only a single species. Some families
have been omitted due to lack of sufficient infor-
mation to provide key characters for immature
stages.
In the preparation of this key, ease of use has
been considered of high importance. Stehr (1987)
provides a much more detailed and precise key
to families of Nearctic Lepidoptera but omits
many families that only occur in other parts of
the World. In order to cover all families in the
present key without making it too long and diffi-
cult to use, it has been simplified by keeping to
fairly short couplets and omitting reference to
exceptions within families, excepting major sub-
families and other important groups. The key
should therefore be used with caution and identi-
fications checked against good family descrip-
tions.
Traditionally, the most important larval char-
acteristics used for identification are relative se-
tal positions. Unfortunately, study of larval
chaetotaxy has been hampered by a proliferation
of different systems of setal nomenclature. The
system adopted here for body setae is that of
Hinton (1946), while nomenclature of cranial se-
tae is based on that of Heinrich (1916) modified
by Gerasimov (1935) and Stehr (1987).
Not only are setae sometimes very difficult to
see, but asymmetry is fairly common with aber-
rant setal positions that may cause problems in
use of keys. When problems occur in use of keys,
it is always worth checking the setae on both
sides of the body.
Larval characters are not illustrated within the
key but the figures of general larval features (Fig.
3.1,2) should act as a simple glossary of terms
used.
1 Case-bearing 2
Not case-bearing 6
2(1) Pro thorax with L pinaculum fused
with thoracic shield 3
Prothorax with L pinaculum separate
or not developed 4
3 (2) Meso and metathorax with SV unise-
tose. Small larvae in flat, elongate-
oval cases of plant fragments on the
ground Adelidae (16)
Meso and metathorax with SV bise-
tose Psychidae (17)
32 David J. Carter & Niels P. Kristensen

vertical
triangle

Anterodorsal view of head-

setal nomenclature and general features

palp

Lateral view of head

Fig. 3.1. Head of generalized lepidopteran (glossatali) larva, diagram.

Classification and Keys to Higher Taxa 33

abdominal segments

D1 02

anal comb

J
JJ1* crochets 91h+ 10th
prothorax
mesothorax abdominal segment abdominal segments

simple
seta

planta
anal comb

chalaza scoli thoracic leg

uniserial jniserial uniserial multiserial

lili fff H
uniordinal biordinal triordinal uniordinal
homoideous
-mirheteroideous

HEAD
!# l?
MESON
^
transverse circle mesopenellipse lateropenellipse mesoseries pseudocircle
bands
Arrangement of crochets

Fig. 3.2. Taxonomically useful structures on generalized neolepidopteran larva; diagram.

4 (2) Abdominal prolegs with crochets in - Anal prolegs without crochets; pro-
uniserial, transverse bands. Case of thorax only with dorsal shield
plant material 5 Incurvariidae (16)
Abdominal prolegs with crochets in 6(1) Antennae longer than head, the latter
circle or penellipse. Case of animal or- strongly retracted into the thorax.
igin (eg hair). . . Tineidae (part, 17) Strongly tapered prolegs sometimes
5 (4) Anal prolegs with crochets; thoracic present on abdominal segments 18.
segments with well developed dorsal Very small, slug-like larvae with flat-
plates tened, often scale-like body setae . . .
. ColeophoridaeColeophorinae (19) Micropterigidae (14)
34
- Antennae much shorter than head.
Strongly tapered prolegs not present
on abdominal segments 1 - 8
7 (6) Head without adfrontal sutures or ec-
dysial lines. Thoracic legs and prolegs
absent. Body setae minute and diffi-
cult to detect. Larvae feeding within
seeds of kauri pine (Agathis)
Agalhiphagidae (14)
Head with adfrontal sutures and ecdy-
sial lines
8 (7) Thoracic legs absent or very weakly
developed. Prolegs with or without
crochets
- Thoracic legs present
9 (8) Numerous secondary setae present . .
Cecidosidae ( 1 - 6 )
Secondary setae absent
10 (9) Head with adfrontal suture passing
behind antenna . Eriocraniidae (15)
Head with adfrontal suture in front of
antenna
11 (10) Head with frons triangular, upper
edge not or hardly included in forma-
tion of hind edge of head capsule . 12
Head with frons parallel-sided or
shaped as an inverted trapezium . 14
12(11) Anal shield with a dorsal, hooked
prong Douglasiidae
(Tinagma, overwintering form) (17)
Anal shield without a dorsal pro-
jection
13 (12) Dorsal plates or calli present on tho-
racic and/or abdominal segments . . .
Heliozelidae (16)
Dorsal plates or calli absent, larva
usually pink or red when preserved . .
Prodoxidae (16)
14 (11) Prolegs with crochets, at least on anal
prolegs Tischeriidae (16)
Prolegs without crochets
15(14) Head with frons rectangular. Weakly
developed, stump-like, unsegmented
legs present on meso- and metathorax
Nepticulidae (16)
Head with frons shaped as an inverted
trapezium; thoracic legs absent . . 16
16(15) Head with frontal bridge connecting
hind margins of adfrontal ridges . . .
Gracillariidae (part, 1 - 7 )
Head without frontal bridge connect-
ing hind margins of adfrontal ridges
(Mandible sometimes with large dorsal
conical projection) Long, very slender
larvae Opostegidae (1 6)
17 (8) Larvae with thoracic legs but without
crochet-bearing prolegs or with only a
few crochets (less than 5 on each pro-
leg)
Larvae with thoracic legs and prolegs,
the latter usually with crochets . . 24
David J. Carter & Niels P. Kristensen
18 (17) Larvae slug-like, often with scoli.
Head strongly retracted into thorax
7 Limacodidae (1 10)
Larvae subcylindrical, without sec-
ondary setae. Head not retracted into
thorax 19
19(18) Small larvae, less than 3 cm long,
without patches of spicules on dorsal
surface 20
- Large larvae, more than 5 cm long
8 with prominent dorsal patches of spic-
ules Castniidae (part, 111)
20 (19) Meso and metathorax and abdominal
9 segments 18 each with a pair of dor-
17 sal calli. Head with spinneret not de-
veloped . . . Heterobathmiidae (14)
- Dorsal paired calli absent from thorax
10 and abdomen. Head with spinneret de-
veloped 21
21 (20) Abdominal segments 1 - 8 with LI di-
rectly caudad to spiracle
Acanthopteroctetidae ( 1 - 5 )
11
- Abdominal segments 1 - 8 with LI not
directly caudad to spiracle 22
22 (21) Pro thorax with two L setae and with
spiracle on sclerotised plate
Glyphipterigidae (18)
- Prothorax with three L setae. Spiracle
not on sclerotised plate 23
23 (22) A few rudimentary crochets present
on abdominal prolegs; abdominal se-
tae short . . . Gelechiidae (part, 1 - 9 )
13
- Abdominal crochets absent; abdomi-
nal setae long . . Douglasiidae (17)
24(17) Crochet-bearing prolegs present on
abdominal segments 2 - 7 . Slug-like
larvae with deciduous, jelly-like tuber-
cles Dalceridae (1 10)
- Abdominal segments 2 and 7 without
crochet-bearing prolegs. Body without
jelly-like tubercles 25
15 25 (24) Crochets of abdominal prolegs ar-
ranged in medial longitudinal bands
or curves (mesoseries), sometimes with
rudimentary lateroseries, forming a
pseudocircle or broken pseudocircle
82
- Crochets not in a mesoseries . . . . 26
26(25) Abdominal segment 10 with crochets
in a complete or almost complete
ellipse, with large biordinal crochets
anteriorly and smaller uniordinal or
biordinal crochets posteriorly
Mimallonidae (1 18)
- Abdominal segment 10 with crochets
not as above 27
27 (26) Body with secondary setae, sometimes
only in SV region 28
- Body without secondary setae . . . 33
18 28 (27) Head large and rounded with many
secondary setae, sometimes on chala-
zae. Prothorax narrowed to form dis-
Classification and Keys to Higher Taxa
tinct neck. Anal comb sometimes pre-
sent Hesperiidae (1 16)
Head not large and rounded; cranial 40 (39)
setae not on chalazae. Prothorax not
narrowed to form a "neck" 29 -
29 (28) Prolegs with crochets in a mesopenel-
lipse that may be mistaken for a meso-
series Pterophoridae {1-13)
Crochets otherwise arranged . . . . 30
30 (29) Secondary setae on Verrucae; setae
with minute lateral branches
Lecithoceridae {1-9)
Secondary setae not on Verrucae; setae
without lateral branches 31
31 (30) Abdominal prolegs with crochets in
uniordinal circle. Tarsi of thoracic legs
with flattened setae
. . ElachistidaeAgonoxeninae (19) 43 (40)
Abdominal prolegs with crochets not
uniordinal. Tarsi of thoracic legs with-
out flattened setae 32
32(31) Abdominal prolegs with crochets
biordinal. Secondary setae not con-
fined to SV group. Thoracic segments
with SV trisetose
. . . XyloryctidaeScythridinae (19)
Abdominal prolegs with crochets tri-
ordinal to multiordinal. Secondary se-
tae confined to SV group or to anal
prolegs only. Meso- and metathorax
with SV unisetose
. . . XylorytidaeXyloryctinae (19)
33 (27) Abdominal seg 6 without prolegs . . .
Gracillariidae (part, 17)
Abdominal seg 6 with prolegs . . . 34
34 (33) Body globular. Head reduced to small
triangular capsule. Stemmata grouped
on a raised tubercle. Larva parasitic
on Hemiptera . . Epipyropidae (1 - 1 0 )
Body not globular. Head normal.
Stemmata not on tubercle 35 -
35 (34) Abdominal spiracles on pinacula or
small sclerotized conical protuber-
ances 36
Abdominal spiracles not on pinacula
or protuberances 39
36 (35) Spiracles on raised or conical protu-
berances 37 48 (43)
Spiracles not raised 38
37 (36) Prothorax with L group trisetose;
some dorsal setae spatulate or bifur-
cate . . . . Schreckensteiniidae (1 13)
Prothorax with L group bisetose; dor-
sal setae normally developed
Epermeniidae (1 13)
38 (36) Pinacula surrounding spiracles ex-
tended dorsocranially. Prolegs with
few crochets or without crochets . . .
Ypsolophidae Ochsenheimeriinae (1 8)
Pinacula surounding spiracles not ex-
tended dorsocranially. Prolegs with
crochets in a circle
Acrolepiidae (18)
35
39 (35) Prothorax with L group trisetose . . 40
Prothorax with L group bisetose . . 70
Prothorax with L pinaculum fused
with thoracic shield 41
Prothorax with L pinaculum separate
or not developed 43
41 (40) Prolegs with crochets in a uniserial cir-
cle, small larvae 42
Prolegs with crochets in a biserial or
multiserial circle. Larvae medium to
large Hepialidae (15)
42(41) Abdominal segments 1 - 6 with LI and
L2 on a common pinaculum
Galacticidae (1-13)
- Abdominal segments with LI and L2
remote, not on a common pinaculum
Tineidae (part, 17)
Abdominal prolegs with crochets in
uniserial transverse bands 44
Abdominal prolegs with crochets in a
circle or penellipse 48
44(43) Abdominal prolegs usually with one
transverse band of crochets, occasion-
ally with two bands. Anal shield with
six setae. Small larvae in fruit and
buds
. . . ProdoxidaeLamproniinae (16)
(early instar, leaf-mining larvae of In-
curvariidae may also key out here)
- Abdominal prolegs each with two
bands of crochets. Anal shield with
eight setae 45
45 (44) Abdominal segments with LI and L2
always remote from each other . . . .
Bucculatricidae (17)
Abdominal segments with LI and L2
close together 46
46 (45) Anal prolegs with crochets divided
into two groups
Gelechiidae (part, 19)
Anal crochets in undivided bands . 47
47 (46) Dorsal surface of body with raised
patches of spicules. Large larvae (c
7 cm) Castniidae (part, 1 - 1 1 )
- Dorsal surface of body without raised
patches of spicules. Small to medium
sized larvae Sesiidae (111)
Small larvae mining in grasses
. . . ElachistidaeElachistinae (19)
- Not mining in grasses 49
49 (48) Abdominal segment 3 with LI and L2
remote, or in very small larvae L2
missing 50
Abdominal segment 3 with LI and L2
close, often on same pinaculum . . 54
50 (49) Crochets in a uniserial circle or penel-
lipse 51
- Crochets in a multiserial circle or a
penellipse surrounded by a circle of
small crochets 53
51 (50) Prolegs long, almost as long as tho-
racic legs Plutellidae &
Ypsolophidae Ypsolophinae (18)
36
Prolegs considerably shorter than tho-
racic legs
52 (51) Abdominal segments with SD2 absent
. . . . YponomeutidaeArgyresthiinae
(part, 1 - 8 )
Abdominal segments with SD2 pre-
sent Tineidae (part, 17)
53 (50) Abdominal prolegs with crochets in a
multiserial circle
Yponomeutidae- Yponomeutinae ( 1 - 8 )
Abdominal prolegs with crochets in a
penellipse surrounded by a circle of
small crochets Plutellidae (part, 18)
54 (49) Abdominal segment 9 with setae D2
closer together than setae D1 of abdo-
minai segment 8, often on a common
pinaculum
Abdominal segment 9 with setae D2 at
least same distance apart as setae D1
of abdominal segment 8
55 (54) Abdominal prolegs slender and pencil-
like, almost as long as thoracic legs. .
Abdominal prolegs short, distinctly
shorter than thoracic legs
56 (55) Abdominal prolegs each with a sclero-
tised collar . . Gelechiidae (part, 19)
Abdominal prolegs without sclerotised
collars Choreutidae {1-13)
57 (55) Prothorax with spiracle dorsad of L
pinaculum
. ColeophoridaeBlastobasinae (19)
Prothorax with spiracle posterior to L
pinaculum
58 (57) Metathoracic legs swollen and club-
shaped Chimabachidae (19)
Metathoracic legs not swollen . . . 59
59 (58) Head rugulose. Each abdominal seg-
ment with all four D setae on a large
sclerotised plate
. . . . CossidaeMetarbelinae (1 11)
Head smooth, D setae not on large
plates
60 (59) Head wedge-shaped. Large larvae. . .
CossidaeZeuzerinae (1 11)
Head not wedge-shaped
61 (60) Prolegs of abdominal segments 36
with crochets in a multiserial penel-
lipse Palaephatidae (16)
Prolegs with crochets otherwise ar-
ranged
62(61) Prolegs each with a surrounding collar
and/or with a sclerotised, central foot
patch Gelechiidae (part, 19)
Prolegs without sclerotised collar or
foot patches
63 (62) Abdominal segment 8 with SD usu-
ally distinctly anterior to spiracle; anal
comb sometimes present
Tortricidae (1 12)
Abdominal segment 8 with SD ante-
rodorsad of spiracle. Anal comb ab-
sent . . . . Oecophoridae (part, 19)
David J. Carter & Niels P. Kristensen
64 (54) Head with LI further from A3 than
52 A3 is from A2
Gelechiidae!Oecophoridae (part, 19)
Head with LI nearer to A3 than A3 is
to A2 or equidistant 65
65 (64) Metathorax with distance between
coxae twice as great as their breadth
Cosmopterigidae (19)
Metathorax with distance between
coxae less than twice their breadth 66
66 (65) Abdominal segments with D1 and D2
close together . . . Heliodinidae { 1 - 8 )
Abdominal segments with D1 and D2
remote 67
67 (66) Head with frons reaching only half
way to vertical triangle 68
55 - Head with frons reaching at least 2/3
distance to vertical triangle 69
68 (67) Abdominal segment 9 with LI and L2
64 on separate pinacula
Cossidae Cossinae (1 11)
_ Abdominal segment 9 with LI and L2
56 on a common pinaculum
. . Elachistidae-Stenomatinae (1-9)
57 59 Prolegs with crochets usually biserial.
Anal segment, excluding anal shield,
with more than nine setae on each
side . . . . Oecophoridae {part, 1 - 9 )
_ Crochets uniserial. Anal segment, ex-
cluding anal shield, with not more
than nine setae on each side . Ypono-
meutidaeArgyresthiinae (part, 1 - 8 )
Abdominal segments with LI and L2
70 (39)
58 remote (LI distinctly posterior to spi-
racle) . . . Tineidae-Scardiinae (17)
Abdominal segments with LI and L2
close together, often on a common pi-
naculum 71
71 (70) Crochets of abdominal prolegs in two
transverse bands 72
Crochetsof abdominal prolegs in a cir-
60 cle or penellipse 73
72 (71) Head with frons reaching 3 or more
to vertical triangle Coleo-
61 phoridae- Momphinae (part, 19)
~~ Head with frons reaching less than Vi
way to vertical triangle
Brachodidae (1 11)
73 (71) Meso- and metathorax with SV group
62 bisetose 74
~ Meso- and metathorax with SV group
unisetose 75
74(73) Meso- and metathorax with L2 absent
Thyrididae (1 13)
63 - Meso- and metathorax with L2 pre-
sent Pyraloidea (part, 1 14)
75 (73) SV group above prolegs with four
macroscopic setae 76
SV group, on or above prolegs, with
three macroscopic setae 78
76 (75) Abdominal segments 1 6 with LI and
L2 on a common pinaculum . . . . 77
Classification and Keys to Higher Taxa
Abdominal segments 16 with LI and
L2 more widely separated, not on a
common pinaculum
ColeophoridaeMomphinae
(part, 1 - 9 )
77 (76) Anal prolegs with 2 rows of crochets;
abdominal segment 9 with D1 absent
or minute. . Copromorphidae {113)
Anal prolegs with 1 row of crochets;
abdominal segment 9 with D1 nor-
mally developed. Carposinidae (113)
78 (75) Prothorax with SD absent (ie protho-
racic shield with only 10 setae); lateral
leaf-like gland present on abdominal
segment 1 . . . . Callidulidae {1-15)
Pro thorax with SD present (pro tho-
racic shield with 12 setae); no lateral
gland on abdominal segment 1 . . 79
79 (78) SV setae situated low-down on abdo-
minai prolegs. Metathorax with SD
approximately equal in length to SD2.
Abdominal prolegs with crochets tri-
ordinal Hyblaeidae (1 13)
SV setae situated at base of abdominal
prolegs. Metathorax with SD con-
siderably longer than SD2 80
80 (79) Crochets of prolegs biordinal or trior-
dinal (exceptionally uniordinal) . . . .
Pyraloidea (part, 1 14)
Crochets of prolegs uniordinal. . . 81
81 (80) Abdominal prolegs long and slender;
crochets in a mesal penellipse
Tineodidae (1 13)
Abdominal prolegs short; crochets in
a circle Alucitidae (1 - 1 3 )
82 (25) Abdominal prolegs long, often as long
as thoracic legs; with crochets in
strongly curved mesoseries, closely re-
sembling a penellipse
Uraniidae (1 17)
Abdominal prolegs not long, or if so,
crochets no arranged in a strongly
curved mesoseries resembling a penel-
lipse 83
83 (82) Thoracic legs very short, with adhesive
pads on tarsi; body with broad lateral
flanges. Small head usually retracted
into thorax. . . Cyclotornidae {1 10)
Thoracic legs normally developed . 8 4
84 (83) Well developed prolegs present on ab-
dominal segments 6 and 10 only;
sometimes with small or rudimentary
prolegs on abdominal segments 4 and
5; anal prolegs strongly developed . .
Geometridae (1-17)
Well developed prolegs present, at
least on abdominal segments 5 and 6
85
85 (84) Abdominal prolegs with an additional
lateroseries of crochets, forming a
pseudocircle or broken pseudocircle .
86
37
Abdominal prolegs with crochets in
simple mesoseries 88
86 (85) Anal prolegs reduced to small lobes
without crochets. Anal shield extended
to a single point or process. Secondary
setae in SV region only
Drepanidae (part, 1 17)
Anal prolegs normally developed.
Anal shield not extended to a single
point or process 87
87 (86) Head with dorsal protuberances. Anal
shield bifurcate extending to form two
long processes. Anal comb present . .
Hedylidae( 1 - 1 6 )
- Head without protuberances. Anal
shield not modifidied. Anal comb ab-
sent. Dorsal eversible gland present on
prothorax . Papilionidae (part, 1 - 1 6 )
88 (85) Larvae without numerous secondary
setae, except in some cases in the SV
region and on the prolegs 89
Larvae with many secondary setae (in
some cases, secondary setae are micro-
scopic) 98
89 (88) Prothorax with L group trisetose; pro-
legs long and slender 90
- Prothorax with L group bisetose . 91
90 (89) Abdominal prolegs medially con-
stricted Urodidae (1 13)
- Abdominal prolegs not medially con-
stricted Immidae (1 13)
91 (89) Abdominal prolegs with secondary se-
tae 92
Abdominal prolegs without secondary
setae 94
92 (91) Setae on raised pinacula. Head with
epicranial index more than 0.8; stipital
lobe absent
NoctuidaeDilobinae (1 19)
Setae not on raised pinacula. Head
with epicranial index less than 0.6;
stipital lobe present 93
93 (92) Abdominal segment 1 with only 1 MD
seta; thorax swollen
Doidae-Doa (1-19)
- Abdominal segment 1 with 2 MD se-
tae or thorax not swollen
Notodontidae (part, 1 19)
94 (91) Thorax and abdomen with crown-like
circlets of spines scattered over the
dorsal and lateral surfaces
Heterogynidae (1 10)
- Thorax and abdomen without circlets
of spines 95
95(94) Abdominal segments 1 - 8 with two
extra L setae, one caudad of spiracle,
one distinctly ventrocaudad of spira-
cle . DrepanidaeThyatirinae ( 1 17)
Abdominal segments without extra L
setae caudad of spiracle 96
96 (95) Meso- and metathorax with SV group
bisetose 97
38
- Meso- and metathorax with SV group
unisetose Noctuidae (part,
(including Aganainae; 1 - 1 9 )
97 (96) Body covered with minute spinules.
D2 and LI arising from elongate tu-
bercles or conical projections espe-
cially on meso- and metathorax.
Crochets homoideous
NycteolidaeEarias (1 19)
- D1 and D2 not arising from elongate
tubercles. Prolegs with ends of plantae
drawn out. Crochets usually heteroi-
deous Arctiidae (part, 1 - 1 9 )
98 (88) Additional rudimentary prolegs, with-
out crochets, on abdominal segments
2 and 7 99
- Abdominal segments 2 and 7 without
prolegs 100
99 (98) Prothorax and abdominal segments
18 with peg-like, glandular protu-
berance adjacent to spiracle
Megalopygidae (1 10)
- Prothorax and abdominal segments
without peg-like protuberances adja-
cent to spiracle
Somabrachyidae (1 10)
100(98) Abdominal segments 6 and 7 with
mid-dorsal, eversible glands, or, if ab-
dominal segment 6 is without a gland,
then abdominal segment 8 with a
large, mid-dorsal hair pencil. Body
usually with series of dorsal hair tufts
or pencils . . . . Lymantriidae ( 1 - 1 9 )
- Abdominal segments 6 and 7 without
mid-dorsal glands 101
101 (100) Prolegs with crochets heteroideous, or
ends of plantae drawn out beyond ex-
tent of crochets 102
- Prolegs with crochets homoideous;
ends of plantae not drawn out beyond
the extent of the crochets 103
102(101) Mesothorax with only one verruca
above the spiracle
. . . . ArctiidaeCtenuchinae { 119)
- Mesothorax with more than one ver-
ruca above the spiracle
Arctiidae (part, 1 19)
103 (101) Abdominal prolegs with a row of soft
basal plates below the crochets; head
retracted into thorax
Zygaenidae (110)
- Abdominal prolegs without soft basal
plates below crochets 104
104 (103) Prolegs with crochets uniordinal 105
Prolegs with crochets biordinal or tri-
ordinal Ill
105 (104) Anal shield produced to form a pair of
anal projections. Head with stemma 3
enlarged
. . . . NymphalidaeSatyrinae (1 16)
- Anal shield not forked 106
106(105) Spiracles small and circular. Prolegs
David J. Carter & Niels P. Kristensen
long and thin, with enlarged plantae
107
Spiracles large and usually elliptical.
Prolegs short 108
107 (106) All setae on distinct Verrucae; protho-
racic L group with more than 3 setae
NycteolidaeNolinae (1 19)
Only some setae on Verrucae, others
scattered; prothorax with L group tri-
setose Pterophoridae (1 13)
108 (106) Prolegs with crochets biserial and with
3 or less SV setae
Oenosandridae (1 19)
Prolegs with crochets uniserial and
with more than 3 SV setae . . . . 109
109 (108) Some setae arranged on Verrucae or
flat, verruca-like plates. Head with
stipi tal lobes absent 110
Setae not grouped on Verrucae or sim-
ilar plates. Head with stipital lobes
present . . Notodontidae {part, 119)
110(109) Abdominal segment 8 with spiracle at
least 1.5 the height of that in abdo-
minal segment 7. Head with mandibles
lacking secondary setae
Noctuidae (part, 1 - 1 9 )
Abdominal segment 8 with spiracle
less than 1.5 (usually about 1.25 x)
the height of that on abdominal seg-
ment 7. Head with mandibles bearing
secondary setae. Larvae usually living
communally
. . . Notodontidae Thaumetopoeinae
(1-19)
111 (104) Secondary setae very variable in
length (some at least 5 length of oth-
ers), not arising from raised Verrucae
or scoli 112
Secondary setae fairly equal in length
or larvae with distinct warts or spines
114
112(111) Pointed anal lobe present between
anal prolegs; body sometimes with lat-
eral or subventral lappets
Lasiocampidae (1 18)
Anal lobe absent; lappets not present
113
113 (112) Mid-dorsal tapered hair tufts or pen-
cils present
. . BombycidaeApatelodinae (1 18)
Mid-dorsal hair tufts or pencils absent
Lemoniiidae (1 18)
114(111) Secondary setae minute, many flat-
tened and scale-like. Dorsal paired fil-
aments on meso and metathorax and
abdominal segment 9 and mid-dorsal
filament on abdominal segment 8 are
lost in the final instar
Brahmaeidae (118)
Secondary setae if minute not scale-
like. Body without deciduous fila-
ments 115
Classification and Keys to Higher Taxa
115 (114) Abdominal segment 8 with a mid-dor-
sal horn, tubercle or spine . . . . 116
- Abdominal segment 8 without a mid-
dorsal horn or spine 122
116(115) Body with spines or large protuber-
ances or expansions on thorax and
elsewhere 117
- Body without spines or projections on
thorax and rarely elsewhere, except on
abdominal segment 8 120
117 (116) Head angular or covered with spines.
Abdominal prolegs with crochets usu-
ally triordinal
Nymphalidae (part, 1 16)
- Head rounded and smooth. Abdomi-
nal prolegs with crochets biordinal . .
118
118(117) Dorsal and subdorsal primary setae
clubbed or reduced to stout cones;
eyespots surround spiracles
Carthaeidae (penultimate instar, 1 18)
- Primary setae not so modified or lat-
eral circumspiracular eyespots absent
119
119(118) Metathorax expended laterally, with
dorsal eyespots; Horn on abdominal
segment 8 with bifid tip
. . . SaturniidaeOxyteninae (1 18)
- Metathorax not expanded laterally or
without dorsal eyespots; Uorn/tuber-
cle on abdominal segment 8 without
bifid tip . . . Saturniidae (part, 1 18)
120 (116) Abdominal segments each divided into
six to eight annuii. Prolegs not widely
separated . . Sphingidae (part, 1 - 1 8 )
Abdominal segments indistinctly di-
vided into two or three annuii. Prolegs
often widely separated 121
121 (120) Abdominal prolegs very widely sepa-
rated and splayed. Thoracic segments
often enlarged
Bombycidae (part, 1 18)
- Abdominal prolegs not widely sepa-
rated or splayed. Thoracic segments
taper towards head
Endromidae (1 18)
122 (115) Abdominal prolegs with crochets in a
mesoseries divided by a central gap, or
reduced centrally, with a pad-like,
central protuberance. Head usually re-
tracted into thorax
Lycaenidae (1 16)
123 (122) Metathorax dorsally expanded into
forward-pointing conical peak; anal
shield produced into a cone-like back-
ward-pointing structure
. Saturniidae Cercophaninae (1 18)
- Metathorax and anal shield not modi-
fied in this way 124
124(123) Prothorax with a dorsal eversible
gland situated in a groove or pit . . .
Papilionidae (part, 1 16)
39
Prothorax without a dorsal eversible
gland 125
125 (124) Head and body without protuber-
ances 126
Head and/or body with spines, scoli or
other conspicuous protuberances 129
126(125) Primary setae reduced to small stout
cones; abdominal spiracles surrounded
by eyespots
. . . . Carthaeidae (final instar, 1 18)
Primary setae not reduced to cones;
circumspiracular eyespots absent 127
127 (126) Some setae grouped on Verrucae; setae
sometimes plumose
. . . . AntheiidaelEupterotidae (1 18)
Verrucae absent; setae short, not plu-
mose 128
128 (127) Head and body densely covered with
short hairs, often on small warts; abd
seg 8 without mid-dorsal patch; anal
comb sometimes present
Pieridae (1-16)
Body with minute secondary setae, ap-
pearing completely smooth. Abdomi-
nal segment 8 with a sclerotised dorsal
patch . . . . Sphingidae (part, 1 - 1 8 )
129(125) Head with protuberances
Nymphalidae (part, 1 16)
Head without protuberances
Saturniidae (part, 118)
Acknowledgments
NPK thanks Dr. N. M. Andersen and Mr. O.
Karsholt for critical perusal of the introductory
section. The abandonment of sub- and infraor-
ders in Lepidoptera systematics was laid before
of a number of leading Lepidoptera systematists,
several, but not all of whom sympathized with
the step; all expressions of views were informa-
tive and are appreciated.
References
Ax, P. (1984): Das Phylogenetische System. Stuttgart,
New York G. Fischer [English edition 1987 Thephy-
logentic system. Chichester, New York J. Wiley]
Carter, D . J . (1984): Pest Lepidoptera of Europe with
special references to the British Isles. - Series Ento-
mologica 31. W. Junk, The Hague,
de Queiroz, K. & Gauthier, J. 1992. Phylogenetic tax-
onomy. - A. Rev. Ecol. Syst. 23: 4 4 9 - 4 8 0 .
Forey, P. L. (1992): Formal classification. Pp. 1 6 0 - 1 6 9
in Forey P. L., Humphries, C. J., Kitching, I. J.,
Scotland, R. W., Siebert, D. J. & Williams, D. M.
Cladistics. A practical Course on Systematics. The
Systematics Association Publication N o 10. Ox-
ford, Clarendon Press.
Gerasimov, A. M. (1935): Zur Frage der Homodyna-
mie der Borsten von Schmetterlingsraupen. Zool.
Anz. 112: 1 1 7 - 1 9 4 .
Griffiths, G. C. D. (1974): On the foundations of bio-
logical systematics. Acta Biotheor. 23: 8 5 - 1 3 1 .
40
Heinrich, C. (1916): On the taxonomic value of some
larval characters in the Lepidoptera. - Proc. ent.
Soc. Wash. 18: 154-164.
Heppner, J. B. (1991): Brachyptery and aptery in Lepi-
doptera. Tropical Lepidoptera 2: 1140.
Hinton, H. E. (1946): On the homology and nomencla-
ture of the setae of lepidopterous larvae, with some
notes on the phylogeny of the Lepidoptera. -
Trans. R. ent. Soc. Lond. 97: 1 - 3 7 .
Holloway, J. D Bradley, J. D. & Carter, D. J. (1987):
Lepidoptera. CIE Guides to Insects of Importance
to Man 1. CAB International Institute of Ento-
mology & British Museum Natural History,
London.
David J. Carter & Niels P. Kristensen
Sattler, . (1991): A review of wing reduction in Lepi-
doptera. - Bulletin of the British Museum (Natural
History) (Entomology) 60: 243-288.
Stehr, F. (1987): Order Lepidoptera. Pp. 288-340 in
Stehr, F. (ed.) Immature Insects. Kendall/Hunt, Du-
buque.
Wiley, E. O. (1981): Phylogenetics. The Theory and
Practice of Phylogenetic Systematics. New York
etc., J. Wiley.
Willmann, R. (1987): Phylogenetic systematics, classi-
fication and the plesion concept. - Verh. naturwiss.
Ver. Hamburg (NF) 29: 221-233.
(1989): Palaeontology and the systematization of
natural taxa. - Abh. naturwiss. Ver. Hamburg (NF)
28: 267-291.
 4. The Non-Glossatan Moths
Niels P. Kristensen
Three lepidopteran clades are primitively devoid
of the coilable proboscis and other apomorphic
traits which characterize the huge monophylum
Glossata. These clades are the Micropterigoidea
Micropterigidae, Agathiphagoidea - Agathipha-
gidae and Heterobathmoidea-Heterobathmii-
dae; their phylogeny is discussed in 12 and
their formal ranking in 13. Inferences about
the ground plan of lepidopteran structure and
function are by necessity based largely on condi-
tions in the non-glossatan clades, and their mor-
phology is therefore treated in some detail in vol-
ume 2.
The adult moths in the non-glossatan grade
share a suite of plesiomorphic traits. Heads with
'rough' vestiture of hair-scales. Dorsal tentorial
arms variably developed, but never large. La-
brum large, movable, with frontal retractors.
Mandibles have well developed articulations
with the head capsule and strong muscles. Maxil-
lae generalized, with long, folded 5-segmented
palps, distinct, sclerotized laciniae and non-elon-
gate galae. Labial palps with vom Rath's organ
well developed. Hypopharynx broad, with varia-
bly armed infrabuccal cavity. Laterocervicale
with 'hair plate' close to the apex. Pronotal plate
not subdivided, with at least one pair of seta-
bearing 'warts'. First thoracic spiracle of 'primi-
tive type' (see 2 - 8 ) . Wing-'coupling' jugo-fre-
nate (but there is presumably no functional
coupling, judging from the photographic obser-
vations on the overall similar Glossata-Eriocran-
iidae by Grodnitsky & Kozlov 1985). Wings held
tectiform in repose, set with microtrichia ('acu-
lei'); wing surface scales (not marginal 'fringes')
'primitive type' exclusively, with 'herringbone'
pattern (see 2 - 2 ) . Sternum I a distinct, discrete
plate with produced corners. In male genitalia
valve consistently devoid of movable medial
(morphologically distal) appendage. Female with
single genital opening, common oviduct commu-
nicating with genital chamber ventrad from
bursa copulatrix. Cephalic CNS with free trito-
cerebral commissure. All three suborders in
ground plan with well-developed sternum V
gland (at least in male) and with hindwing Rs
four-branched; secondary reductions in these
traits occur in Micropterigidae. The larvae in this
grade are more or less distinctly prognathous,
without a spinneret on the apex of the prelabio-
hypopharyngeal lobe; larval structure otherwise
very diverse. The decticous, exarate pupae have
the cephalic setae very prominent; clypeus dis-
tinctly demarcated, setose.
MICROPTERIGOIDEA
Micropterigidae. Probable autapomorphies: An-
tennae with 'ascoid' sensilla (branched basiconic
sensilla, Fig. 4.1 E; paralleled in Nepticuloidea-
Opostegidae). Labrum extensively desclerotized.
Strong mandibles asymmetrical, incisor cusps on
left mandible only present apically. Labial palps
greatly shortened. Spurs 0-0-4 (absence of meso-
tibial spurs paralleled in the Heterobathmi-
oidea). Prothorax unusually small, with anterior
notopleural articulation and with unsclerotized
areas in notum. Spiracle VIII nonfunctional.
Male with sternal sclerotization on VIII strongly
reduced, represented by small paired plates or
altogether absent. Phallus with close-set radial
folds around gonopore. Female postabdominal
segments without apophyses. 6 malpighian tu-
bules grouped into two bundles (of 3), each dis-
charging through a common duct (paralleled in
the Neolepidoptera). Brain usually with distinct
'deutocerebral loop' (paralleled in Glossata-
Acanthopteroctetoidea and Lophocoronoidea).
Larval head with antennae unusually long (seg-
ment 2 particularly elongate). Anterior tentorial
pits close to antennal base. Larval trunk approx-
imately hexagonal in cross section. Trunk integu-
ment with cuticle specializations unique among
arthropods: exocuticle high, with liquid-filled
chambers (each corresponding to one epidermal
cell) in a honeycomb-like pattern, exo-and endo-
cuticle separated by fluid-filled space, con-
tinuous with 'honeycomb' chambers via pores in
lower exocuticular layer; cuticle overlaid by
sticky pellicle to which foreign bodies often ad-
here. Thoracic legs short, with at most 4 seg-
ments, subterminal one presumably a tibiotarsus.
It is surprising, that a Micropterix species has
been found to have eupyrene sperm exclusively,
while an Epimartyria like other Lepidoptera in-
vestigated has both eupyrene and apyrene sperm.
However, the monophyly of the Micropterigidae
is strongly supported by the numerous synapo-
morphies of Micropterix and other micropterigid
genera, including Epimartyria.
Small moths (wingspan up to ca. 15 mm, usu-
ally ca. 10 mm or less). Head with headcapsule
only moderately extending above upper margin
of compound eyes. Ocelli usually prominent,
very rarely lost. Antennae moniliform to fili-
form, scaling variable, without typical sensilla
auricillica (probably a plesiomorphy); antennae
and palps with a diversity of little understood
sensory (and glandular?) organs (Le Cerf 1926,
Faucheux 1992). In cibarium epipharynx usually
with complex asymmetrical armature of sclerites
42
and microtrichial brushes (which serve to clean
the mandibles, Hannemann 1956) and hypophar-
ynx with 'triturating basket', i. e. infrabuccal
cavity set with strong, transversely aligned spines
(similar cibarial specializations present in Heter-
obathmiidae, parallelism or symplesiomorphy?).
Labial palps usually 2-segmented (3 segments re-
tained in a few 'Sabatinca', only a single present
in Neomicropteryx). Metathoracic furcal stem
without anterior process (apparently a uniquely
primitive condition in the Lepidoptera, absence
of this formation elsewhere in the order interpre-
ted as secondary loss). Protibial epiphysis lost in
a few genera. Wings narrow, but outline variable,
apex sometimes pointed, sometimes obtusely
rounded. Forewing pattern often iridescent, very
variable (Figs. 4.3 AC), in some genera promi-
nent transverse fasciae are commonplace. Fore-
wing (Fig. 4.1 A) with Sc forked (except in an
undescribed dwarfish New Caledonian taxon),
crossvein Sc-R usually present, R forked or sim-
ple, position of Rs4 relative to apex variable, A
veins sometimes in typical 'double-Y' configura-
tion, or A 2 and A 3 strongly reduced; hindwing
with configuration of Sc-R region variable, posi-
tion of Rs4 variable as in fore wing, in Squami-
cornia only three Rs branches present (Fig.
4.1 B). Sternum V gland orifice in some genera
on sculptured process which may, or may not, be
set with hair scales, in other genera simple, or
gland lost. Segment IX usually a completely scle-
rotized ring, but dorsalmost part sometimes
membranous, its posterodorsal margin in Micro-
pterix with a variable complement of processes
(Fig. 4.1 C). Phallus not of phallotheca-type; in
Epimartyria (Fig. 4.1 D) with accessory ventral
branch. Segment X variable, presumably primi-
tively with paired tergal lobes. Female abdomen
(Fig. 4.1 F) with ten distinct segments and termi-
nal cloaca; segment IX sclerotization variable, a
complete ring or reduced/absent on dorsum; seg-
ment X sclerotization a pair of lateral setose
plates; segment XI probably represented only by
circumcloacal membrane. Terminal abdominal
segments in repose telescoped into segment VIII,
extensible through blood pressure. Bursa copu-
latrix in some genera with highly characteristic
'signa' sclerotizations (Fig. 4.1 F). Ductus sper-
mathecae, as far as known, without double-com-
partment structure (a unique plesiomorphy
within the Lepidoptera). Nature of formation
termed 'circumcloacal chamber' (Dugdale 1974)
awaiting reinvestigation. Gut without stomo-
daeal crop and with long midgut commencing
in thorax, few rectal papillae (3 in Micropterix).
Nerve cord with 4 or fewer (2 in some genera)
abdominal ganglia; abdominal nerve sheath dor-
sally with enlarged cells. Tracheal system in
Micropterix without transverse commissures in
abdominal segments, but this condition is not
general in the family. Testes follicles minute, tes-
tes of the two sides not united; a single pair of
Niels P. Kristensen
male accessory glands. Each ovary with 4 or 5
ovarioles.
Chorion in the Micropterix egg highly special-
ized, with clubbed protuberances formed within
one hour after oviposition (see 218, 219); egg
in Neomicropteryx covered with hygroscopic, ge-
latinous mass. Larvae (Fig. 4.2 A) strikingly dif-
ferent from other lepidopteran caterpillars. Head
in repose completely retractible into thorax (an-
tennal length surely related to this extraordinary
retractability). Antennae 3-segmented. A median
seta on frontoclypeus between antennae proba-
bly homologous with a campaniform sensillum
in caddisfly larvae, but without obvious homo-
logues in other Lepidoptera. 5 or 6 stemmata,
contiguous or at least close-set. Headcapsule
without adfrontal ridges and with short hyposto-
mal bridge (plesiomorphies shared with Aglos-
sata), tentorial arms and corporotentorium
stout, labrum with both medial and lateral re-
tractors retained, maxilla with both galea and
lacinia distinct (unique plesiomorphies within
Lepidoptera). Trunk setae simple (some Saba-
tinca) or thickened, truncate or davate; Hinton's
(1958) claim of the distribution of primary setae
being "entirely different" in zeuglopteran and
other lepidopteran larvae has not been con-
firmed (Davis 1987), but the ground plan of the
former still remains to be worked out. Abdomen
usually without appendages, but Micropterix and
a species group within 'Sabatinca' with pointed,
non-musculated and non-crochet-bearing pro-
legs on IVIII. Abdominal end in Micropterix
with a threelobed 'sucker', with which the larva
may attach itself to the substrate. First thoracic
and all eight abdominal spiracles functional in
some genera; Hinton's report of a functional spi-
racle on metathorax may refer to a topographi-
cally forwards shifted spiracle I (Davis 1987).
Pupa (Fig. 4.2 B) with mandibles not hypertro-
phied.
Adults usually diurnal, but nocturnal activity
is reported for some species. They are mostly
pollen eaters. Micropterix species feed in the
flowers of a great variety of dicot and monocot
plants, often in trees. In contrast, the 'Sabatinca-
group' genera have almost exclusively been ob-
served on sedges; particularly interesting excep-
tions are the feeding of some New Caledonian
taxa on pollen of Winteraceae and fern spores,
the latter is also a food source of Malagasy taxa.
The larvae may be broadly characterized as soil
animals with high moisture demands. Most
known larvae in the 'Sabatinca-group' feed on
folise liverworts, but the taxon with distinct
prolegs has been found in rotten logs (feeding
on fungus hyphae?), and Micropterix larvae in
addition to detritus/fungus hyphae eat green an-
giosperm leaves. Pupation takes place in an oval
cocoon (Fig. 4.2 B), which may be a pure-silk
formation (Sabatinca, Epimartyria, Neomicro-
pteryx) or may have soil particles incorporated
The Non-Glossatan Moths
into the outer wall (Micropterix); at emergence it
is cut open in the anterior end, which at least
in Micropterix is less dense than other parts. In
temperate zones adult micropterigids are pre-
dominantly spring/early summer insects.
121 described species, but about half again as
many are recognized and still unnamed. The
family has been provisionally classified into the
'Micropterix group' (Micropterix only, 67 species,
Palaearctic, most speciose in Mediterranean sub-
region; Fig. 4.3 C) and the 'Sabatinca group'
comprising all other genera; the principal puta-
tive autapomorphy of the latter is the complete
absence of discrete male venter VIII sclerotiza-
tions, but its monophyly remains questionable.
One apparently monophyletic subgroup com-
prises 4 genera with an amphipacific N. Hemi-
sphere distribution: Taiwan/China/Japan (where
Neomicropteryx has been studied in some detail)
- N.America (Epimartyria only; Fig. 4.3B).
Otherwise the group is mainly SW Pacific: Sa-
batinca s. lat. occurring in E. Australia, New
Zealand and particularly New Caledonia, which
has a 'swarm' of about 50 species, of which only
three are so far named. There is one small genus
(Agrionympha) in S. Africa, one monotypic in
austral S. America, and a recently discovered,
still un-named taxon in Madagascar; apart from
the monotypic Squamicornia from montane
Equador the absence of records of micropterigids
from the continental tropics is conspicuous.
Principal references on taxonomy/biogeogra-
phy include Issiki (1931; with extensive discus-
sion of previous works on systematics and integ-
umental structure), Whalley (1978), Kristensen &
Nielsen (1979, 1982), Kaltenbach & Speidel
(1982), Minet (1985), Gibbs (1983, 1989), Kozlov
(1988-90). The morphological literature on
adults is relatively extensive; because of the long-
recognized phylogenetic key position of the
micropterigids they (usually at least a Micro-
pterix species) are considered in many compara-
tive accounts of lepidopteran structures, see
vol. 2. Studies devoted to this family deal with
the adult head (Le Cerf 1926, Tillyard 1923 a,
Hannemann 1956, Chauvin & Faucheux 1981,
Faucheux 1992), wings (Tillyard 1919, Philpott
1923), pregenital abdomen (Kristensen 1984 c),
genitalia (Philpott 1934, 1924, Hannemann 1957,
Kristensen 1984 a), brain (Buxton 1917), tracheal
system (Kristensen 1984 b), spermatogenesis
(Friedlnder 1983, Sonnenschein & Huser 1990,
Hamon & Chauvin 1992), ovary, egg structure
and embryogenesis (Chauvin & Chauvin 1980,
Ando & Kobayashi 1978, Kobayashi 1994, Ko-
bayashi & Ando 1981, 1982). The extraordinary
larval structure has so far been treated only in a
preliminary way (Tillyard 1923 b, Mutuura 1956,
Hinton 1958, Lorenz 1961, Yasuda 1962, Davis
1987, Kozlov 1991, Kristensen 1990). Principal
works on natural history and behaviour include
43
Pirngruber (1944), Lorenz (1961), Carter & Dug-
dale (1982), Tuskes & Smith (1984) and Pellmyr
et al. (1990).
Clade Agathiphagoidea + Heterobathmioidea +
Glossata
Probable autapomorphies of the ground plan of
this clade (Kristensen 1984 f) include: Loss of
paraglossae. Presence of anterior process (ac-
commodating origin of sternal trochanter de-
pressors) on metafurcal stem. Ductus sperma-
thecae with thickwalled and thinwalled compart-
ment. Larval head with tentorium delicate, me-
dian labral retractor lost, only single maxillary
endite present, cranial flexor of dististipes lost,
unique dorsoventral muscle laterally spanning
foramen magnum. Pupa (Fig. 4.2 D) with mandi-
ble hypertrophied, angularly bent (this special-
ization lost again in higher Glossata).
In the currently preferred phylogeny of this
clade the Heterobathmioidea and the Glossata
constitute a monophylum; derived characters
shared by the Agathiphagoidea and Heterobath-
mioidea are therefore considered homoplasious.
AGATHIPHAGOIDEA
Agathiphagidae. Probable autapomorphies: Ocelli
absent. Antennae with bifurcate sensilla. Ster-
num V gland in male opening on long, smooth
process; gland structure uniquely specialized, se-
cretory section a long and twined tube leading
into large thinwalled reservoir; gland absent in
female. Male segment IX ring greatly indented in
dorsal and ventral midline. Genital chamber in
roof with distinctive complement of sclerotiza-
tions, including median 'spiny plate'. Larva with
but two stemmata (without corneal lenses) pre-
sent on each side. Antenna reduced, one-seg-
mented. Endophagous larva with trunk unpig-
mented and with greatly reduced setae, thoracic
legs and abdominal appendages absent.
Additional agathiphagid apomorphies are
shared with the Heterobathmioidea, but must be
homoplasious autapomorphies of both taxa, if
the Heterobathmioidea are indeed the sister
group of the Glossata (12): Anterior tentorial
arms fused into Y-shaped formation. Postlabium
proximally widenend. Mesepisternal pre- and
paracoxal sutures anastomosing below anapleu-
ral cleft. Tergum I with lateral process indistinct
and hind corners fused with tergum II. In larval
head labral compressors, intrinsic maxillary
muscles and dorsal postcerebral pharynx dilators
lost. Pupal mandible with strong subapical tooth
on lower surface.
Medium-sized micro-moths, wingspan up to
ca. 27 mm. Facies singularly caddisfly-like, with
the antennae stretched forward in repose (other
primitive moths have them upward, in a V-like
position) and the forewing pattern non-iridescent
44 Niels P. Kristensen

Fig. 4.1. Micropterigidae: A - F ; Agathiphagidae: GI; Heterobathmiidae: J - L . A, Hypomartyria venation; ,

Squamicornia hindwing venation; C, Micropterix male genitalia lateral; D, Epimartyria phallus lateral; E, Micro-
pterix antennal flagellum segments with ascoids; F, Sabatinca female genitalia, lateral; G, Agathiphaga venation;
H, same, male postabdomen dorsal; I, same, female postabdomen ventral; J, Heterobathmia hindwing, note jugal
lobe (arrow); K, same, male genitalia lateral, left valve removed, note dorsal anus (arrow); L, same, female
genitalia lateral and (lower right) abdominal apex ventral, note separate slitlike gonopore (large arrow) and anus
(small arrow), ag: accessory gland; be: bursa copulatrix; cr: [severed] phallocrypt; dl: dorsum X lobe; re: rectum;
sp: spermatheca; tl: terminal (segment XI) lobes; numbers (Roman and Arabic) denote segments. A, after
Kristensen & Nielsen (1983); C after Kozlov (1989); D after Kristensen (1984a); E after le Cerf (1926); F after
Minet (1985); G after Nielsen & Common (1991); H after Kristensen (1984e); I after Common (1973); J - L original
(B. R u b a * del.)
The Non-Glossatan Moths
Fig. 4.2. Micropterigidae: A B; Heterobathmiidae: C; Agathiphagidae: D. A, Sabatinca larva; B, Micropterix
pupa in cocoon; C, Heterobathmia larva dorsal (above) and ventral (below); D, Agathiphaga pupa. A, C original
(B. Rubaek del.), after Lorenz (1961), D, after Nielsen & Common (1991).
Fig. 4.3. Micropterigidae: - C ; Agathiphagidae: D;
Heterobathmiidae: E. A, Sabatinca lucilia-, B, Epimar-
tyria pardella; C, Micropterix aureatella\ D, Agathi-
phaga vitiensis; E, Heterobathmiapseuderiocrania. Uni-
form magnification, scale (milimetres) in D.
and cryptic, brownish mottled (Fig. 4.3 D).
Headcapsule extending far above compound
eyes, with high internal crest medially above an-
tennae. Antennae filiform. Dorsal tentorial arms
short. Mandibles large, but without incisivus
cusps, post-ecdysial function unknown; adductor
muscle huge, origin covering large parts of dorsal
headcapsule, including median crest. Basal
maxillary piece with costa between cardo and ba-
sistipes; galea a large membranous pad with nu-
merous sensilla basiconica. Cibarium with simple
epipharynx; armature in infrabuccal pouch slen-
der, scattered microtrichia (not strong, aligned
spines as in 'triturating baskets' of Micropteri-
goidea and Heterobathmioidea; another family
autapomorphy?). Legs with spur formula 1-4-4
45
antenna claws
I labial palp
(protibial spur and midlength spurs on mestoti-
bia unique plesiomorphies within Lepidoptera).
Forewing (Fig. 4.1 G) Sc and (in one of the two
extant species) R forked, Rs4 to apex. Both wing
pairs with M four-branched (unique plesiomor-
phy within Lepidoptera - if not an autapomor-
phic character reversal!). In mesothorax outer
tergal coxa-remotor present (unique condition
within panorpoid endopterygotes; remarkable
plesiomorphy or, probably most likely, autapo-
morphic character reversal). In male genitalia
(Fig. 4.1 H) segment IX a synscleritous ring, val-
vae long, simple; segment X with pair of broad
tergal lobes; a rather prominent, medially in-
dented 'terminal lobe' above eversible perianal
area possibly pertaining to XI; phallus of phallo-
theca-type, with well-developed eversible aedea-
gus. Each testis comprising four large saccular
follicles with narrow apex (unique condition in
Lepidoptera, most probably a plesiomorphy);
seminal vesicle close to testis, questionably ho-
mologous with vesicles in other Lepidoptera; two
pairs of long (one particularly long, twined), tu-
bular accessory glands (similarly unusual lepi-
dopteran condition, interpretation questionable).
Female segment VIII (Fig. 4.11) posteriorly nar-
rowing, tergal and ventral plate separate, each
with pair of long apophyses on anterior margin;
segments I X - X I partly retractible into VIII,
forming narrow, weekly sclerotized probe with
Y-shaped sclerotization in dorsal wall, arms of
' continuing into long internal apophyses; clo-
acal opening subterminal. Bursa copulatrix
small, without signa. Colleterial glands very
large sacs. Very numerous (40 +) ovarioles per
ovary (isolated case among homoneurous Lepi-
doptera, probably a plesiomorphy). Gut with
large, thinwalled stomodaeal crop extending into
46
Fig. 4.4. Agathiphaga vitiensis, larva (length ca. 6 mm.)
in Agathis seed.
abdominal base; midgut short; six malpighian tu-
bules each discharging separately into gut; nine
rectal papillae. Nerve cord with five abdominal
ganglia (discrete ganglia I I - V , plus composite
terminal ganglion); sheath thickened dorsally,
but thickened mass apparently mainly cellular,
rather t h a n composed of extracellular material.
Egg undescribed. Larva (Fig. 4.3 F) endopha-
gous (seed miner). Head capsule weekly mela-
nized; adfrontal ridges and distinct ecdysial lines
lacking; complete hypostomal bridge present;
short tentorial dorsal arms present. Maxilla
without discrete cardo. Labial glands very long,
thick tubes.
Adults are nocturnal and have been encoun-
tered in nature on but very few occasions; it is
u n k n o w n whether they feed at all. The eggs are
believed to be inserted below the cone scales of
the kauri pines (Agathis, Araucariaceae) in the
seeds of which the larvae subsequently feed. The
excrements are incorporated into the lining of
the larval cell. U p o n maturity the larva coats the
cell wall with a (labial gland?) secretion, produc-
ing a smooth, hard layer. Larvae probably al-
ways enter a diapause, which seems to be termi-
nated only under high moisture regimes, and
which under laboratory conditions is very diffi-
cult to terminate anyway. Before eclosion the
pharate adult grinds an emergence whole by
means of the hypertrophied pupal mandibles. In-
festation of pine seeds may locally attain pest
levels.
The single genus Agathiphaga comprises two
closely related species, occurring in Australia
(Queensland) and Fiji/Solomon Islands/New
Hebrides/New Caledonia respectively.
The original description of Agathiphaga by
Dumbleton (1952) contained much information
on biology and integumental structure; subse-
quent biological observations are reviewed by
Robinson & Tuck (1976) and C o m m o n (1990).
Detailed structural accounts deal with the anten-
Niels P. Kristensen
nal sensilla (Faucheux 1990), male genitalia
(Kristensen 1984 e) and larval head (Kristensen
1984 d); further observations on agathiphagid
a n a t o m y are reported in Kristensen (1984 f) and
in vol. 2 of the present work.
HETEROBATHMIOIDEA
Heterobathmiidae. Probable autapomorphies:
Absence of costa between cardo and basistipes.
Proximal prelabial sclerite tapering inwards. Lat-
erocervicalia with anteromedial corners rounded,
not produced toward presternum. Pro-katepi-
sternum narrow, almost parallel-sided. Free arm
on profurcal bridge bent backwards. Setose plate
behind metasubalare syncleritous with epimeron.
Spurs 0-0-4 (loss of mesotibial spurs paralleled
in Micropterigoidea). Forewing Sc and R simple.
Forewing scales on obverse surface with smooth
rounded plates on inter-ridge areas. Male seg-
ment VIII desclerotized in middle (parallelism
with Micropterigoidea); segment IX (Fig. 4.1 K)
with short, stout (sometimes markedly bifid)
ventromedian process; two pairs of variably
shaped processes arising f r o m posterolateral cor-
ners of dorsal plate; base of phallus with posteri-
orly directed apdeme; postgenital complex with
variable armature of spines.
If the phylogenetic hypothesis preferred here
(see above, and chapter 2) is correct, suites of
specializations shared with the Micropterigoidea
(shortened labial palps, lack of mesotibial spurs,
medial desclerotization of male venter VIII, and
perhaps cibarial armature) and with the Agathi-
phagoidea (listed under the latter above) are ho-
moplasious autapomorphies of the Heterobath-
mioidea as well as the other taxa.
Small m o t h s (Fig. 4.3 E), wingspan usually ca.
10 mm. Headcapsule extending markedly above
c o m p o u n d eyes. Ocelli present. Antennae with
sensilla auricillica (probable s y n a p o m o r p h y with
Glossata). Mandibles with strong incisivus cusps.
Wings narrow (forewing index < 0.3) but not
acutely pointed. Forewings iridescent purple,
with silvery-white scales forming more or less ex-
tensive suffusion and often distinct spots at least
in tornai area; superficial similarity with Glos-
sata-Eriocraniidae sometimes striking. Forewing
Sc and R unforked, crossvein Sc-R absent (but
weak basal crossvein S c - R + R S sometimes pre-
sent); base of 1A sometimes appearing forked;
pterostigma distinct in forewing, small in hind-
wing. Hindwing base (Fig. 4.1 J) with distinct,
rounded jugal lobe (apparently a unique plesio-
m o r p h y in the Lepidoptera). Sternum V gland
present in b o t h sexes, orifice on sculptured pro-
cess. Male segment IX (Fig. 4.1 K) usually com-
pletely divided into dorsal and ventral plate, but
in Hetorobathmia valvifer an anteriorly complete
sclerotized ring; phallus not of phallotheca-type
(contrary to statement in original description),
ejaculatory duct partly melanized. In female
The Non-Glossatan Moths
postabdomen (Fig. 4.1 L) segment VIII with sim-
ple tergum and functional spiracle; large subgeni-
tal plate on venter VIII covering base of mem-
branous ventral side of terminal segments. Seg-
ment IX bilaterally flattened, high tergal plate
with variably developed process arising above
anterolateral corner; in the best known species,
Heterobathmia pseuderiocrania, this process is
very short indeed, in H. valvifer it is a genuine,
albeit rather short, apophysis. Segment X proba-
bly represented by small paired, unmelanized
and setose lobes which are immovably fused with
tergum IX; genital chamber and rectum opening
separately in shallow median groove of membra-
nous ventral surface of terminal segments. Duc-
tus bursae opening on prominent, thickwalled
papilla in genital chamber; intima in ductus and
proximal corpus bursae with delicate spines and
ridges; no true signa.
The species H. valvifer is unique in the Lepi-
doptera in having a pair of long, flattened ap-
pendages arising from the ventral VIII/IX
boundary region, their bases being laterally very
close to the anterolateral corners of tergum IX.
The only musculature associated with these ap-
pendages is the ventralmost fibre bundle of a
broad muscle extending from the subgenital
plate to the membranous body wall adjacent to
the anterolateral margin of tergum IX; this bun-
dle inserts immediately in front of the appendage
base. The morphological interpretation of these
appendages is highly problematical; if they are in
any way derivable from an element in the gener-
alized insect ovipositor, they can only be consid-
ered an autapomorphic character reversal. The
female postabdomen of H. valvifer is also notable
for the presence of ribbonshaped longitudinal
sclerites flanking the median groove on the mem-
branous ventral surface of the terminal segments.
Gut without stomodaeal crop; long midgut
commencing in thorax; 6 malpighian tubules
opening separately into gut. Ventral diaphragm
with well developed segmental alae. Other as-
pects of visceral anatomy still remain to be
worked out.
Egg undescribed, covered by a jellylike sub-
stance. Leafmining larva (Fig. 4.2 C) progna-
thous. Headcapsule with prominent adfrontal
and hypostomal ridges, hypostomal lobes medi-
ally separated by distinct membranous area
(noteworthy probable synapomorphies with
Glossata). Antennae short, 3-segmented. 7 stem-
mata (this high number a unique plesiomorphy
within the Lepidoptera) present on each side. La-
brum with even epipharyngeal surface sclero-
tized. Mandible with medial tuft of barbed hairs.
Prothorax with well developed notai and sternal
sclerites, trunk otherwise membranous and un-
pigmented. Thoracic legs well developed, with
large, subdivided trochanters (unique plesiomor-
phy within the Lepidoptera). Eversible calli pre-
sent on dorsum and venter of all segments from
47
mesothorax to VIII, venter I excepted. Terminal
segment a trilobed formation, with subanal lobes
forming paired 'anal feet' (without crochets).
Pupa with apical teeth on hypertrophied mandi-
bles markedly asymmetrical; right mandible with
prominent tooth on lower edge, one-third from
apex.
All known heterobathmiids are associated
with deciduous Nothofagus. The adults are very
early spring insects and only fly actively in sun-
shine. They visit Nothofagus flowers and are be-
lieved to be pollen feeders. Eggs are deposited on
the lower surface of the host leaf. The full-depth
mine starts as a narrow gallery, but after the first
moult a blotch mine is formed, often obliterating
the gallery. Frass is initially deposited at the egg
site, which thereby becomes very conspicuous.
As leaf miners heterobathmiid larvae are unusual
in being able to leave one leaf and start mining in
another. Feeding is rapidly completed. Pupation
takes place in the earth, in a silken cocoon cov-
ered with soil particles.
The single genus Heterobathmia comprises at
least 9 species, of which 3 have been named so
far. They occur with deciduous Nothofagus in
austral. S. America.
Principal references: Kristensen & Nielsen
(1979, 1983, in press), Nielsen (1985).
Acknowledgements
Communication of unpublished information on
Malagasy Micropterigidae from D. C. Lees in
gratefully acknowledged.
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Addendum
A SEM account of the antennal sensilla of Mi-
cop ter ix calthella was given by Faucheux (1997)
and the larval integument of the same species
was described by Hamon & Chauvin (1995). A
S. African micropterigid taxon phenetically sim-
ilar to generalized SW Pacific Sabatinca has been
discovered (Kristensen unpublished).
Upton (1997) reported on Agathiphaga larvae
staying alive after 12 years in diapause.
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nae of Micropterix calthella L. (Lepidoptera,
Micropterigidae). - Acta zool. 78: 1 - 8 .
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and its differentiation in Micropterix calthella L.
Lepidoptera: Micropterigidae): anatomy and ultra-
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- The Entomologist 116: 142-143.
 5. The Homoneurous Glossata
Niels P. Kristensen
As outlined in 1 - 2 the monophyly of the clade
Glossata is very strongly corroborated, while the
monophyly of the clade Heteroneura, compris-
ing the vast majority of the glossatan superfami-
lies, and characterized i. a. by the 'heteroneu-
rous' wings, is a reasonably supported working
hypothesis. Within the 'homoneurous' grade in
the Glossata five extant lineages are currently
recognized, four of them with but one included
family. In the presently preferred cladogram
(12, Fig. 2.2) the six extant basal glossatan lin-
eages are seen to have arisen in a single sequence
of five splitting events.
Clade GLOSSATA
Primarily characterized by notable specializa-
tions in the adults' mouthparts: mandibles non-
functional after pupal/adult ecdysis, without
clearly defined articulations with head capsule
(and often strongly reduced); maxilla without
distinct, sclerotized lacinia, galeae elongate,
forming a proboscis, which is coiled in repose
and primitively has the walls of the food canal
lined with discrete cuticular plates; in maxillary
palp muscle between segments 2/3 lost; prelabio-
hypopharyngeal lobe narrowed, hypopharynx
without infrabuccal pouch; on head capsule epi-
stomal suture reduced, not accompanied by cu-
ticular thickening (regression surely related to re-
duction of mandibular function). Dorsal tentor-
ial arms considerably larger than in non-glos-
satans, but apparently reduced again in ground
plan of the Heteroneura. Trend towards complex
fusion of adult brain and suboesophageal gan-
glion recognizable already in ground plan
through incorporation of tritocerebral commis-
sure in suboesophageal mass. Larva with spin-
neret on apex of prelabio-hypopharyngeal lobe.
Pupal clypeus non-setose, not distinctly demar-
cated.
All lineages in the homoneurous grade have
(in their ground plans) retained microtrichiated
('aculeate') wing surfaces and distinct jugal lobes
on the forewings. Members of the first differenti-
ated lineages (Eriocraniidae, Acanthopterocteti-
dae, Lophocoronidae) are all small moths (wing-
span nearly always less than 15 mm). Like the
non glossatans they have a rough cephalic vesti-
ture of hair-scales, long (5-segmented) and folded
maxillary palps, and they are relatively narrow-
winged. The antennae are simple and filiform.
The proboscis is devoid of an intrinsic muscula-
ture. The females in these families have a charac-
teristic 'Eriocrania-type' piercing oviscapt (Fig.
4.1 C, see also 24): the sclerotized abdominal
apex is flattened, pointed and has serrate edges,
forming an 'oviscapt saw'; a subterminal mem-
branous area below the saw bears a pair of elon-
gate, near-tubular sclerotizations ('ventral rods'
in 2 - 4 , 'tertiary apophyses' of Davis 1978), at
the posterior apex of which the cloacal orifice is
located. The oviscapt saw is in repose completely
retracted into the preceding 'oviscapt cone', a
more or less extensively synscleritous ring which
tapers posteriorly. The oviscapt saw and oviscapt
cone both have long paired apophyses invagi-
nated from their anterior borders. The segmental
assignment of the elements in the 'Eriocrania-
type' oviscapt is debatable (24); since the three
families in which this oviscapt occurs apparently
do not form a monophylum, it is arguably a glos-
satan groundplan trait.
The name Dacnonypha (introduced by Hin-
ton, 1946) has been used for a high-rank taxon
centred on the Eriocraniidae. However since the
Eriocraniidae do not appear to form a monophy-
lum together with any other subset of the Glos-
sata (Nielsen & Kristensen 1996), the category
Dacnonypha is redundant and here abandoned.
ERIOCRANIOIDEA
Eriocraniidae. Probable autapomorphies: tibial
spur formula 0-1-4 (unpaired mesotibial spur
paralleled in the Acanthopteroctetidae). Valvae
greatly shortened and phallus with accessory
ventral branch (Fig. 5.1 E, F). Female with com-
plex 'vaginal sclerite' (sensu Davis 1978) in poste-
rior part (behind spermathecal orifice) of genital
chamber encircling egg passage (Fig. 5.1 D).
Ventral diaphragm musculature exceedingly re-
duced, a single fibre pair present per segment (re-
duction parallelled in Acanthopteroctetidae).
Male accessory gland with swollen and thin-
walled (non-secretory?) apical portion, adhering
to counterpart on other side. Larval head Fig.
5.2 B) with dorsal ecdysial line running laterad
from antennal base, and only a single pair of
stemmata. Legs and prolegs absent.
Small moths, wingspan usually ca. 10 mm, al-
ways below 15. Head capsule (Fig. 5.1 A) extend-
ing far above compound eyes. Ocelli present. La-
brum large, movable, with frontal retractors.
Epiphysis present or (in most taxa) absent. Fore-
wing (Fig. 5.1 B) often with distinct pterostigma;
forking of Sc and R retained in some taxa; pres-
ence of only one Rs 1 + 2 branch (also in hind-
wing) characteristic of the dominant Palaearctic
taxon Eriocrania. Veins A in 'double-Y' configu-
52 Niels P. Kristensen

Fig. 5.1 Eriocraniidae: AF; Acanthopteroctetidae: GI; Lophocoronidae: J - K ; Neopseustidae: L. A, Dys-

eriocrania head; , Dyseriocrania venation; CD, Eriocrania female genitalia, ventral (C), 'vaginal sclerite' lateral
(D); EF, Eriocrania male genitalia, ventral (E), phallus (F); G, Acarithopteroctetes, venation; H - I , Acanthopter-
octetes, male genitalia, phallus (H), armature lateral (I); J, Lophocorona venation; , Lophocorona male genitalia
lateral; L, Neopeustis venation; M, Synempora female genitalia lateral; N, Neopeustis male genitalia ventral. BAS:
Sclerotized plate on base of anal cone; IS: intercalary sclerite; J: median plate ('juxta') below phallus; ME: mandi-
ble; Oc: ocellus; SI: seta-bearing process on valve. A - I after Davis (1978); J - K after Common (1973); L, after
Davis (1975); M after Davis & Nielsen (1980).
The Homoneurous Glossata
ration. Hindwing with several distinct 'frenu-
lum'-bristles, wing-coupling apparently non-
functional (Grodnitsky & Kozlov 1985). Wing-
surface scales of 'primitive-type', without 'her-
ringbone crests'. Central hindwing scales of vari-
able shape, sometimes hairlike. Forewings irides-
cent. Orifice of sternum V gland on sculptured
process. Sternum IV in females nearly always
with unpigmented and un-microtrichiated 'win-
dow' on areas to which gland reservoir adhere;
similar 'windows' present in males of some taxa;
sternum V glands altogether lost in some male
Eriocrania. Male segment IX (Fig. 5.1 F) a com-
plete synscleritous ring, with paired lateroventral
apophyses from its anterior margin accomodat-
ing the origins of phallic retractors; segment X
with paired tergal lobes. Gut with short stomo-
daeum widened posteriorly, but much narrower
than midgut; midgut large, extending from meso-
thorax well into abdomen; 6 malpighian tubules
discharging separately into gut; 3 - 6 rectal papil-
lae. 3 - 4 abdominal ganglia. Testes small, not
united. 4 ovarioles per ovary; colleterial gland
relatively small, lobed or simple.
Egg oval, with thin, smooth chorion. Larva
(Fig. 5.2A) permanently leafmining, progna-
thous, head partially retracted within prothorax.
Mandible with or without bundle of barbed bris-
tles on inner base. Melanized plates present on
prothoracic dorsum and venter, anal shield usu-
ally poorly defined; trunk with variable comple-
ment of dorsal and ventral eversible calli. Pupa
with hypertrophied, angularly bent mandibles.
Adults are usually diurnal, although nocturnal
flight is of regular occurrence in at least one Dys-
eriocrania. They are not flower-visitors, but have
been observed drinking water, and sucking sap
from injured leaf buds. A sexual pheromone
function of the sternum V gland has recently
been proved (Kozlov et al. 1996). Eggs are in-
serted in the host leaves, in pockets cut by the
female's oviscapt saw. The host range is almost
restricted to the Fagales (Quercus and Betula be-
ing the principal host genera), but a few taxa are
known to utilize hosts belonging to Salicales
(120) and Rosales (Holodiscus, Sorbus). Mines
are fulldepth blotch mines, which may or may
not start as a distinct gallery. Frass deposition
usually highly characteristic, in thread-like pieces
(sometimes very long and twined, Fig. 5.2 C).
Larval growth is rapidly completed; full-grown
larvae drop to the ground and bury into the soil,
where the cocoon (covered by soil particles on
the outer surface) is constructed. Most eriocrani-
ids are early spring insects, the adult activity
period coinciding with the leafing of the host,
and the larvae completing their feeding stage
while the leaves are still young. A few eriocraniid
species occasionally occur in such large numbers
that they attain pest levels, particularly at high
latitudes.
53
24 species are named. The family is restricted
to the Holarctic. 6 genera are currently recog-
nized, but the family is structurally and biolo-
gically overall homogeneous, and it is debatable
whether this degree of generic splitting is justi-
fied. Dyseriocrania, Eriocraniella and Eriocrania
(Fig. 5.3 A) are the principal genera.
Davis (1978; principal reference), Wood (1890,
1891), Kristensen (1968), Birket-Smith & Kris-
tensen (1974), Kobayashi & Ando 1987; a review
of Old World eriocraniids by Karsholt, Kris-
tensen & Kozlov is in preparation. Like the
Micropterigidae the Eriocraniidae are treated in
many comparative accounts of lepidopteran
structure; see 24. A report by Koricheva (1994)
provides an entry into the ecological literature
on eriocraniids, and Kozlov et al. (1996) give ref-
erences to works on their pheromone biology.
Clade COELOLEPIDA
The clade Coelolepida (Nielsen & Kristensen
1996), comprising all non-eriocraniid Glossata,
is characterized primarily by the presence of
'normal-type scales (22) on the wing surfaces,
and of the 'normal type' (28) of first thoracic
spiracle in the adult. Ocelli are absent in all ho-
moneurous Coelolepida as well as in nearly all
non-ditrysian Heteroneura. Ocelli probably dis-
appeared in ancestral Coelolepida, and their re-
appearance in heteroneuran lineages may be seen
as apomorphic character reversals.
Three basal lineages are recognized within the
Coelolepida: The Acanthopteroctetoidea, the
Lophocoronoidea and the high-rank clade My-
oglossata.
ACANTHOPTEROCTETOIDEA
Acanthopteroctetidae. Probable autapomorphies:
apices of dorsal tentorial arms fused with head
capsule. Epiphysis absent. Tibial spurs 0-1-4 (see
Eriocraniidae). Labial palp greatly shortened.
Wings (Fig. 5.1 G) unusually narrow. M l
'stalked' with Rs in both wing pairs; forewing
with only two A veins, both extending to wing
margin; hindwing M-Cu crossvein vestigial,
hence 'cell open'. Sternum II with anterior rim
separate from main plate. Sternum V gland ab-
sent. Valve with membranous bulge on inner sur-
face; prominent, posteriorly produced and
ventrally serrate transtilla present (Fig. 5.11);
phallus (Fig. 5.1 H) with apparently eversible
membranous vesica set with cornuti (a similar
phallic configuration is commonplace in Hetero-
neura; this may be a parallelism).
Moths (Fig. 5.3 B) small, a single species with
ca. 15 mm wingspan, others smaller. Head ex-
tending far above compound eyes. Labrum nar-
row, movable, with frontal retractors. Labial
palp 3-, 2- or 1-segmented. Forewing pattern
quite variable, iridescent in some species. Fore-
54 Niels P. Kristensen

Fig. 5.2. Eriocraniidae: - C ; Acanthopteroctetidae: D; Neopseustidae: E; Mnesarchaeidae: F, Hepialidae: G - H .

A, Dyseriocrania larva; , same, head; C, Eriocrania mine in Betula; D, Acanthopteroctetes larva; E, Apoplania
moth in resting posture; F, Mnesarchaea pupa; G, Hepialus larva; H, Fraus pupa. A after Davis (1987); C after
Hering (Biology of Leaf Miners, Junk 1951); D after Davis & Frack (1987); F drawn (. Rubk) after sketch by
G. W. Gibbs; G after Wagner (1987); H after Nielsen & Kristensen (1989).

wing Sc simple; R forked or simple. Jugal lobe
completely reduced in Acanthopteroctetes unifas-
cia and Catapterix; the former with two (stout)
frenulum bristles, the latter, remarkably, with
only a single large bristle recalling that of typical
male Heteroneura. Male segment IX (Fig. 5.11)
a synscleritous ring, anteroventral margin more
or less strongly concave; paired tergum X lobes
small and narrow. Female genital chamber in
some taxa with prominent sclerotization (extend-
ing into wall of bursa copulatrix) in front of
spermathecal orifice (hence apparently nonho-
mologous with eriocraniid 'vaginal sclerite').
Few data on soft anatomy available, all pertain-
ing to Acanthopteroctetes unfascia: brain with
'deutocerebral loop', and optic lobes pressed
against suboesophageal ganglion (as in Lopho-
coronidae); 3 abdominal ganglia; ventral dia-
phragm very reduced; dorsal nerve-cord sheath
not thickened. Gut without stomodaeal crop,
midgut large, extending from prothorax into ab-
domen; only 2 malpighian tubules present.
Immature stages and biology known only for
Acanthopteroctetes unifascia, which is a leafminer
in Ceanothus (Rhamnaceae). Egg undescribed,
inserted in upper epidermis. Larva (Fig. 5.2 D)
prognathous, head partially concealed within
prothorax; 6 stemmata, one of which with re-
duced lens; spinneret very prominent. Prothorax
with melanized tergal and sternal plates. Tho-
racic legs present, with indistinct coxae. Prolegs
present only on X, no crochets. Thoracic and
first seven abdominal segments with paired dor-
sal and ventral eversible calli (in thorax arising
on coxal membranes). Mine a full-depth blotch
mine in the leaf; larva feeding in summer/autumn
and again in spring, leaving mine before pupa-
tion, which takes place in a cocoon on the
ground. Pupa with hypertrophied, angularly
bent mandibles.
Two genera are currently recognized, Acan-
thopteroctetes with 4 species in western N. Amer-
ica and the monotypic Catapterix from Crimea.
Sinev & Zagulajev (1988) described the latter in
a family of its own, but clearly recognized its
closest affinities to be with Acanthopteroctetes;
indeed it is likely, that it is actually cladistically
subordinate within the latter. There is an unde-
scribed taxon from montane Peru.
Davis (1978; principal reference), Davis
(1987), Sinev & Zagulajev (1988), Sinev (1988),
Zagulajev (1990), Nielsen & Kristensen (1996).
Clade LOPHOCORONOIDEA +
IVI YOG LOSS ATA
In the currently best supported hypothesis of
coelolepidan phylogeny (Nielsen & Kristensen
1996) the Lophocoronoidea are the sister group
of the Myoglossata. Derived characters shared
by the two include: frontal labrum retractors
lost. Pronotum with two sclerotizations, the an-
terior articulating with propleuron (parallelled in
Micropterigoidea). Vein R always unforked.
The Homoneurous Glossata
Mesosternum markedly produced anteromedi-
ally. Thickened dorsal sheath of ventral nerve
cord with copious connective tissue.
LOPHOCORONOI DEA
Lophocoronidae. Probable autapomorphies: cra-
nium with weakly developed dorsal condyles at
antennal base (in addition to commonplace
ventral ones, apparent parallelism with Exopo-
ria). Mandibles completely devoid or muscula-
ture (parallelled in many Neolepidoptera).
Epiphysis absent. Rs4 postapical (Fig. 5.1 J).
Male segment IX ring (Fig. 5.1 K) with anterior
margin nearly straight; valve deeply cleft, lower
lobe with stout apical sensillum and 23 long
basal setae; phallus in apical half with promi-
nent, near-tubular infolding from outer wall, and
tergum X lobes small, set with characteristic con-
ical sensilla. Stomodaeum with expanded crop
followed by narrow, tubular section. Only four
malpighian tubules present, each opening into
gut. Brain with 'deutocerebral loop', and optic
lobes pressed against suboesophageal ganglion
(similarities with Acanthopteroctetidae, presum-
ably parallelisms). Abdominal ganglion II
discrete, so 5 ganglia are discernible (autapomor-
phic character reversal?).
Small moths, wingspan 10-15 mm. Head cap-
sule not markedly extending above compound
eyes. (Fronto)clypeolabral boundary indistinct.
Mandibles medium-sized, simple lobes. Spurs 0-
2-4. Forewing pattern quite diverse in the few
known species: unicolorourous fuscous, cryptic
('bird-dropping' type, Fig. 5) or with bold black-
and-white fasciae. Forewing (Fig. 5.1 J) Sc
forked; A veins in 'double-Y' pattern; 'coupling'
jugofrenate. Sternum V gland present in both
sexes, opening on sculptured process. Male seg-
ment IX (Fig. 5.1 K) a synscleritous ring. Female
genital chamber and bursa without sclerotiza-
tions. Large, delicate stomodaeal crop extending
into abdominal base; malpighian tubules each
opening separately into gut. Ventral diaphragm
with well developed segmental alae.
Early stages unknown. Since the female has a
piercing oviscapt type, the larva is supposed to
be an internal feeder, and since the adult moth is
completely devoid of mandibular muscles (there
are also no apodemes for such muscles), the
pupa must be adecticous. The moths are noctur-
nal and mainly autumn fliers. Nearly all lopho-
coronid sites are in eucalypt forest, with growth
of Melaleuca and usually Atriplex. Females have
been encountered much more rarely than males.
The single genus Lophocorona (6 species, Fig.
5.3 C) occurs in the southern parts of West and
South Australia, with outliers in the ACT and
N. S. Wales.
Common (1973: benchmark paper with origi-
nal description), Nielsen & Kristensen (1996
comprehensive account).
55
Clade MYOGLOSSATA
All Glossata-Coelolepida other than the Acan-
thopteroctetidae and Lophocoronidae probably
constitute a monophylum characterized primar-
ily by the presence of an intrinsic musculature in
the proboscis. Other putative myoglossatan au-
tapomorphies include the somewhat enlarged
median mesonotal wing process, as well as the
upwards bend of acrotergite I and concomitant
insertion of the indirect hind wing depressor on
its anterior-facing surface (Nielsen & Kristensen
1996).
NEOPSEUSTOIDEA
Neopseustidae. Probable autapomorphies: facial
scales restricted to paired lateral, usually promi-
nently swollen, patches. Antennae submonili-
form/subserrate, usually (always?) longer than
forewing; scape with distinct condyle fitting pedi-
cellar base; flagellar scales longitudinally aligned.
Proboscis scaled, with a unique double-tube
structure, the inner wall of each galea being
strongly concave and forming by itself a func-
tionally closed tube. Well-developed labial palp
without distinctly concave vom Rath's organ.
Prominent apodemal plate invaginated from up-
per base of propecoxal bridge. Mesofurcal arms
fused into transverse bridge. Male sternum VII
with medial spinose process; 'socius' setae with
prominently swollen base.
Intriguing similarities with the Heteroneura in
thoracic structure include presence of distinct
proprecoxal bridge, transverse suture delimiting
'mesoclidium', and fusion of the latter with the
prospinasternum.
Medium-sized moths, wingspan ca. 15
27 mm. Head capsule not markedly extending
above compound eyes. Labrum large and dis-
tinctly demarcated. Mandibular lobes with well
developed musculature. Maxillary palp similarly
primitive, long and 5-segmented. Epiphysis
much reduced in S. American taxa. Tibial spurs
0-2-4. Wings in Nematocentropus and Synempora
with rather commonplace proportions and dense
scaling, in Neopseustis (Fig. 5.1 L) and, in partic-
ular, Apoplania wings unusually (for homo-
neurous moths) broad and thinly scaled (scales
narrow and widely spaced). Rs (of both wing
pairs) 4-branched only in Nematocentropus, in
other genera Rsl + 2 represented by single vein.
Rs4 to apex. Forewing, except in Nematocentro-
pus, with 'anal pocket', formed by wing mem-
brane behind curved base of 1 A covering base
of Cu. Forewing A veins forming 'single Y \
Frenulum bristles undifferentiated. Female usu-
ally with hyaline 'fenestrae' on IV, but sternum
V glands have not been identified. Genital seg-
ments in both sexes highly modified and inade-
quately understood. Male (Fig. 5.1 N) segment
IX ring with variably developed lateroventral
56
Fig. 5.3. Eriocraniidae: A; Acanthopteroctetidae: ; Lophocoronidae: C; Neopseustidae: E; Mnesarchaeidae: D;
Palaeosetidae: F - G ; Anomosetidae: H; Prototheoridae: I; Neotheoridae: J. A, Eriocrania sparrmannella; B, Acan-
thopleroctetes bifascia; C, Lophocorona pediasia; D, Mnesarchaea acuta; E: Neopseustis meyricki; F: Palaeoses
scholastica; G, Osrhoes coronta; , Anomoses hylecoetes; I, Prototheora petrosema; J. Neotheora chiloides. Scale:
A - , 5 mm; FJ, 10mm.
apophyses from anterior margin; base of valvae
usually (always?) fused with IX ring. 'Anellus'
with armature of processes. Phallus simple and
tubular in Nematocentropus, highly modified in
other genera; paired processes ('parameres') in
Neopseustis are probably phallic derivatives.
Postgenital complex with paired tergum X lobes;
variably shaped 'tegumenal lobes' usually pre-
sent between IX ring and tergum X, sometimes
apparently synscleritous with the former; anus
flanked by mostly conspicuous patches of setae
with raised sockets. Testes not united in midline.
Female (this sex poorly known in Nematocentro-
pus) postabdomen (Fig. 5.1 M) with deep pocket
invaginated behind sternum VII. Tergum VIII
with short, blunt 'apophyses anteriores' from an-
terolateral corners. 'Apophyses posteriores' aris-
ing from minute terminal sclerotization, which
usually bears a pair of distinct lateral teeth and
which may have a rasping/piercing function. The
presence of an apparently distinct segment IX,
with more or less divided tergal plate, between
the apophysis-bearing VIII and the terminal unit
on which the posterior apophyses are born, is
morphologically intriguing (see 2-4). Alimen-
tary canal without stomodaeal crop and with
large midgut; 6 malpighian tubules, each opening
separately into gut. Ventral nerve cord with
paired connectives between suboesophageal and
prothoracic ganglia and between the 4 abdomi-
nal ganglia; autapomorphic neotenic trait?
Nerve cord sheath not thickened dorsally;
ventral diaphragm fibres completely absent.
Niels P. Kristensen
(Data on soft anatomy available only for Sy-
nempora).
The immature stages are unknown, but since
mandibular muscles are strong in the adult, the
pupa is believed to be decticous (and surely exa-
rate). Adult activity has been recorded by day as
well as by night; feeding has not been observed.
While Synempora show the tectiform resting pos-
ture which is commonplace among primitive
moths, the broad-winged Apoplania hold the
wings flat over the abdomen (one overlapping
the other), and in complete rest the antennae are
hidden below the wings (Fig. 5.2 E).
This small and little-known family has a pecu-
liarly disjunct distribution. Nematocentropus
(= Archepiolus, 2 species), the sister group of the
remaining taxa, occurs in Assam, Burma and
Szechuan, Neopseustis (5 species, Fig. 5.3 E)
ranges from Assam to Taiwan, while Synempora
(monobasic) and Apoplania (3 species) occur in
austral. S. America.
Davis (1975; principal reference), Davis &
Nielsen 1980, 1985), Kristensen & Nielsen
(1981), Nielsen & Kristensen (1996), Kristensen
(in preparation).
Clade NEOLEPIDOPTERA
All Myoglossata apart from the Neopseustidae
probably constitute a monophylum, Neolepidop-
tera, characterized by adecticous, obtect pupae.
In the lower neolepidopteran grades the pupae
have backwards-pointing spines on the dorsum
The Homoneurous Glossata
of several abdominal segments; at eclosion this
spinosity aids the movement of the pharate adult
out of the (usually silk-lined) pupal shelter.
Correlated with their loss of function, even in
the pharate stage, the adult mandibles (which in
several neolepidopterans are retained as quite
distinct lobes) have at most a vestigial muscula-
ture. The complement of crochet-bearing prolegs
on abdominal segments IIIVI and X in the
larva is another notable apomorphy ascribed to
the ground plan of the Neolepidoptera. Addi-
tional probable neolepidopteran autapomorphies
include the lateral displacement of the free pro-
furcal arm, the concealment of the upper part
of the mesopleural suture below a mesepimeral
bulge, the disassociation of the aorta from the
metathoracic dorsum, the general absence of
sternum V gland, the arrangement (non-aliform)
of ventral diaphragm fibres in II, the presence of
a spacious stomodaeal crop; the arrangement (in
2 groups of 3) of the malpighian tubules, the
coiling of the spermathecal duct. Some of these
traits are paralleled in one or more non-neolepi-
dopteran clades; caveats concerning their phylo-
genetic interpretation are discussed by Nielsen &
Kristensen (1996).
All homoneurous Neolepidoptera belong to
the clade Exoporia.
Clade EXOPORIA
The monophyly of this clade is now well substan-
tiated (summary in Nielsen & Kristensen 1996),
primarily by the unique configuration of the fe-
male genital apparatus (detailed account in
24): the ostium bursae (copulatory orifice) is
separate from the ovipore and located ventrad
from the latter (as it is in the Ditrysia), but there
is (in the exoporian ground plan) no internal
communication between the two parts of the
genital system. Sperm deposited in the bursa
must be transported to the spermatheca via the
ovipore, the body wall below the latter being
typically grooved one way or another. Accessory
(colleterial) glands lost. No apodemal apophyses
present. Male genitalia also distinctive: phallus
membranous and devoid of protractor and re-
tractor muscles; segment X with transverse (su-
pra- and subrectal) muscles very strongly devel-
oped. Maxillary palp always small, hidden in the
vestiture and at most 3-segmented. Wing scales
mostly with strong 'secondary ridges' (2-1).
High number (5 or 6) of abdominal ganglia in
adult probably an autapomorphic character re-
versal. Egg shell structure and embryogenesis
distinctive (2-18, 2-19); eggs turn black after
being deposited. The 'epicranial notch' of the
larval head is unusually small, and there are
several potential autapomorphies in larval chae-
totaxy, including e. g., the absence of the thoracic
MXD1 and the approximation of the meso-/
metathoracic MD1 to the two MSD setae; pro-
57
thoracic L-pinaculum fused with notai shield in
second and later instars in known hepialoids and
some, but not all mnesarchaeoids (Nielsen &
Kristensen 1989). Apomorphies paralleled in Lo-
phocoronoidea include the presence of (some-
times strongly developed) dorsal cranial condyles
at the antennal base, and the distinctly postapi-
cal Rs4.
Homologies of exoporian postabdominal
structures (in both sexes) with those of other
Lepidoptera have been difficult to establish. In
males the prominent dorsal sclerotization pre-
viously identified as the tegumen most probably
pertains to segment X (Nielsen & Kristensen
1989), and it is now referred to as the 'pseudoteg-
umen' (or 'pseudoteguminal plates', since it is
usually a distinctly paired formation); conspicu-
ous processes from the pseudotegumen and the
tergal area just above it occur in several exopo-
rian lineages. In females the main morphological
problem concerns the segmental assignment of
the sclerite, which forms the lower lip of the cop-
ulatory orifice.
Female pheromone activity has recently been
recorded from the hindwing of a hepialid
(Kuenen et al. 1994) and also from mnesarchaeid
wings (Kozlov, pers. comm.). The nature and ex-
act source of exoporian female pheromones re-
main to be identified.
Larvae of ancestral exoporians were probably
primarily 'soil animals' with generalized feeding
habits similar to those of extant mnesarchaeids.
A checklist of the World Exoporia (by E. S.
Nielsen, G. S. Robinson & D. L. Wagner) is
forthcoming.
MNESARCHAEOIDEA
Mnesarchaeidae. Mnesarchaeids are by far the
smallest exoporian moths, and the only ones
with functional mouthparts in the adults. They
were long grouped with the 'eriocranioid-grade'
families, their close relations to the hepialoids
only being revealed in the 1970s. Probable auta-
pomorphies: wings narrow, lanceolate. Rsl + 2
in both wing pairs represented by one vein only
(Fig. 5.4 A). Wing coupling achieved by band of
long piliform scales on lower side of forewing
dorsum entangling with correspond band of pili-
form scales on hindwing costa. In male genitalia
anterodorsal corner of valve produced and fused
with the lateral part of the pseudoteguminal
plate (which in turn is indistinctly separated
from the counterpart of the hepialoid trulleum).
Elongate and apically bifid 'dorsal plate' on fe-
male postabdominal apex comprising fused terga
of VIII and following segments; anterolateral
corner of dorsal plate more or less extensively
fused with elongate subgenital plate (the latter
forms the lower lip of the copulatory orifice and
presumably at least partly pertains to segment
VIII; Fig. 5.4 C). Roof of antrum with paired,
58 Niels P. Kristensen

Fig. 5.4. Mnesarchaeidae: - C ; Palaeosetidae: IJ; Prototheoridae: D - ; Hepialidae: F - H . A, Mnesarchaea,

venation; , same, head; C, same, female genitalia; D, Prototheora, venation; E, same, maxilla; F, Afrotheora male
genitalia lateral; G, Phymatopus hind leg (scales omitted); H, Aoraia female genitalia ventral; I, Osrhoes venation;
J, same, female genitalia lateral, an: antevaginal lamella; cs: closed margins of antevaginal lamella; db: ductus
bursae; dc: expanded chamber on ductus seminalis; dp: dorsal plate; ds: ductus seminalis; ip: 'intermediate plate'
(lateral part of pseudoteguminal plate); ju: in A jugal lobe, in F juxta; mp: maxillary palp; te: pseudoteguminal
plate; s: sternum; su: subanal plate; t: tergum; tp: process on pseudoteguminal plate; va: valve; vi: vinculum;
Roman numerals denote segment numbers. - B after Dugdale (1988, Fauna of New Zealand 14); C redrawn
after Dugdale (1974); D - after Davis (1995); F after Nielsen & Scoble (1986); G redrawn after Bourgogne (in
Trait de Zoologie 10, 1, 1951); H after Dugdale (1994), I - J after Kristensen & Nielsen (1994).

posteriorly converging grooves (probably sperm
tracts). Pupa (Fig. 5.2 F) on mesothorax and ab-
dominal segments I - V I I I with mid-dorsal
spines only.
Antennae filiform, fully scaled, in one species
group with 3-5-branched trichoid sensilla. La-
brum very small (in anterior view practically hid-
den by bulging frontoclypeus). Maxillary palp 3-
segmented, or second and third segments more
or less completely fused; proboscis (Fig. 5.4 B)
well developed, coilable. Labial palp 3-seg-
mented; apical segment with sensilla cluster at
most slightly depressed. Protibial epiphysis vari-
ably developed, occasionally absent. Spurs 0-2-4.
Forewing in one species group with Ml stalked
with stem of Rs3 + 4; only one A vein present.
Sc branching and presence of inter-M crossvein
have occasionally been observed, but not as a
stable condition in any species.
Egg relatively very large, with smooth
chorion. Larval facies characteristic, with very
long spinneret, anteriorly tapering thorax, and
slender prolegs with biserial crochets. Pupa with
cephalic cocoon-cutter.
Adults mainly diurnal, but have occasionally
been attracted to light. Males fly frequently
around low vegetation, while females are rarely
encountered by day; adults not flower-frequent-
ing. Eggs are attached singly to moss or liverwort
leaves. Larvae live in silken sheets and galleries
spun in liverwort/moss periphyton on moist soil,
logs etc., feeding on leaves and rhizoids, fern
sporangia, fungal hyphae and other detritus. Pu-
pation in an oval, debris-covered cocoon.
The Homoneurous Glossata
14 mnesarchaeid species are known (half of
them still unnamed), all included in the genus
Mnesarchaea (Fig. 5.3 D). The family is endemic
to New Zealand.
Gibbs (1979); comprehensive treatment by
Gibbs & Kristensen in progress.
HEPIALOIDEA.
The Hepialoidea are the first differentiated lepi-
dopteran lineage in which the number of extant
species runs into several hundreds; they are also
the first differentiated lineage which includes
truly large and robust taxa (indeed some of the
largest moths belong here). A long-recognized
hepialoid autapomorphy is the strong regression
of the proboscis, which is at most as long as the
head capsule (usually much shorter, or com-
pletely absent), never truly coilable and probably
always non-functional; the food canal walls are
without discrete plates. Other autapomorphies
include: intercalary sclerite between scapus and
pedicellus elongate, partly lowered into pocket in
intersegmental membrane (primitive, i. e., small
and superficial, sclerite in some palaeosetids now
interpreted as autapomorphic character rever-
sal). Posterior Rs fork backwards displaced, Rs3
reaching termen, not costa (Figs. 5.4 D, I). Meta-
furca without anterior process, sterno-trochan-
teral muscle originating directly from (anteriorly
tilted and convex) furcal stem. Male genitalia
with characteristically hinged 'juxta + trulleum'
sclerites (at least partly homologous with 'me-
dian plate' in lepidopteran ground plan, see
24) below phallocrypt entrance (Fig. 5.4 F).
Antennae structurally diverse; a marked short-
ening is commonplace, but can at most be an
'underlying synapomorphy' of the members of
the superfamily. The same is probably true of the
sensillum-bearing 'nipples' on the vertex and
pronotum found in a variety of hepialoids. While
the maxillary palp in the Prototheoridae (Fig.
5.4 E), and hence in the hepialoid ground plan,
is now known (Davis 1996) to be as well devel-
oped as in the Mnesarchaeidae, it is strongly re-
duced (with indistinct segmentation) or com-
pletely lost in other hepialoids. Hairs on hind-
wing front margin soft, with no structural differ-
entiation between 'frenular' and 'costal' hairs
(sensu Braun 1924, see 2-4). Pro tibial epiphysis
and tibial spurs variably developed. Abdominal
venter I usually (but according to Brner 1938
not always) membranous. Female postabdomen
(Figs. 5.4 H, J) with tergum VIII separate from
'dorsal plate', the latter posteriorly more or less
deeply indented (in Neotheoridae apparently
completely divided, but confirmation of this is
desirable). Distinct sternum VII sometimes pre-
sent, but ventral surface often extensively unscle-
rotized behind VI. The sclerotized 'antevaginal
lamella' anteriorly bordering the ostium bursae
probably belongs to IX; however, it cannot be
59
ruled out that it pertains to VIII (as the mnes-
archaeid subgenital plate apparently does) or is
a composite formation. Occasionally there is a
distinct sclerotization between sternum VII and
the antevaginal lamella. Primitively narrow and
paired 'subanal plates' (pertaining to X or IX?)
flank the grooved (or otherwise modified) 'inter-
genital field' between ostium bursae and ovipore;
their anterior/lateral corners may be closely ap-
proximated to, or fused with, the anterolateral
corners of the dorsal plate.
Male scent organs on wings and hindlegs have
evolved independently on a number of occasions
in hepialoids. It is noteworthy that male lekking,
an unusual behavioral trait in the Lepidoptera,
may also be evolved on more than one occasion
within the superfamily: in Palaeosetidae-Ogj-
gioses (Davis et al. 1995) and at least once within
the Hepialidae s. str. (Wagner & Rosovsky 1991).
Or will the lekking 'palaeosetids' actually
prove to be cladistcally subordinate within the
latter?
Larvae of most of the basal hepialoid lineages
are unknown, so diagnostic traits of the super-
family remain unidentified. Known pupae (Fig.
5.2 H) with dorsal abdominal spine rows basi-
cally similar to those of the lower Heteroneura;
this presumably represents the neolepidopteran
ground plan condition.
The Hepialoidea are currently classified into
five families, of which three are monogeneric and
the remaining two without unambiguous auta-
pomorphies. A total of 12 terminal taxa (species
and assumedly monophyletic supraspecific as-
semblages) can at present be recognized within
the superfamily, and a re-analysis of their inter-
relationships, based on an extensive character
set, is much needed. While Scoble's (1992) inclu-
sion of all 12 in a single family may have much
to recommend it, the time-honoured family cate-
gories are retained here, pending results from an
analysis of the said kind.
Family key
1 Inter-M crossvein (between M3 and M2 or
stem of M1 + M2) present, at least in fore-
wing (Fig. 5.4 D) 2
Inter-M crossvein absent (Fig. 5.41) . . . .
Palaeosetidae
2(1) Metatibia almost always without spurs,
very rarely 1 or 2 p r e s e n t . . . . Hepialidae
Metatibia with 4 spurs 3
3 (2) Entire wing surfaces with microtrichia
('aculei'). All wing scales of 'normal type'
(hollow, with internal trabeculae and per-
forated upper lamella). Forewing with anal
loop. S. African Prototheoridae
Wings microtrichiated at base only. Wings
with lower layer of 'primitive type' wing
scales (devoid of perforations and trabecu-
60
lated lumen). Forewing without anal loop
4 rect and densely scaled. Wings entirely micro-
4 (3) Labial palps long and snout-like, pro-
jecting far beyond antennal pedicellus; api-
cal segment longer than two basal seg-
ments together. Protibial epiphysis present.
Neotropical Neotheoridae
Labial palps shorter, not projecting beyond
antennal pedicellus; middle segment equal-
ling basal and distal segments together.
Protibial epiphysis absent. Australian . . .
Anomosetidae
Palaeosetidae. It is debatable whether the diag-
nostic lack of an inter-M crossvein (Fig. 5.41) is
apomorphic or not. Moths unusually small (wing
span no more than ca. 12.5 mm in some Ogy-
gioses), slender-bodied and broad-winged for
hepialoids, Palaeoses itself least pronouncedly
so. Maxillary appendages completely reduced (as
in most Hepialidae).
Protibial epiphysis absent. Spurs 0-1-2
(Osrhoes), 0-1-1 (Genustes), otherwise 0-0-0.
Male metatibia in Osrhoes, Genustes and (Tai-
wanese) Ogygioses thickened, in two latter gen-
era with hair-pencil (surely a scent apparatus) in
a groove. Male Ogygioses with hair pencil in
folded expansion of hindwing anal area. 'Primi-
tive type' wing scales absent from the genera so
far examined for this trait (Osrhoes, Ogygioses).
Osrhoes is notable for its unusually reduced
dorsal tentorial arms and, in particular, for its
unique female genital apparatus (Fig. 5.4J): a
long, narrow 'ductus seminalis' (presumably a
functional analogue to the similarly termed duct
in Ditrysia) extends backwards from the ductus
bursae to an opening adjacent to the ovipore.
Information on biology and immatures avail-
able only for Ogygioses: egg with prominent tu-
bercles on chorion. 1st instar larva (only instar
known) with only 3 stemmata. Slender prolegs
with crochets in a uniordinal circle. Adult moths
diurnal, males exhibiting lekking behaviour.
Four genera with a total of 8 species are in-
cluded in the Palaeosetidae: Osrhoes (Colombia,
monobasic; Fig. 5.3 G), Palaeoses (Queensland,
2 species, one named; Fig. 5.3 F), Genustes (As-
sam, monobasic) and Ogygioses (Taiwan, Thai-
land, 4 species).
Issiki & Stringer (1932), Common (1990),
Kristensen & Nielsen (1994), Davis et al. (1995),
Heppner et al. (1995).
Prototheoridae. Probable autapomorphies: geni-
talia with unique apparent copulation locking
device, comprising a median 'conjugal process'
on the female antevaginal lamella and a corre-
sponding male 'conjugal pouch', the roof of
which is formed by the arched ventral trulleum
wall. 'Primitive type' wing scales absent.
Maxilla (Fig. 5.4 E) with 3-segmented palp
and distinct proboscis vestige; medial surface of
Niels P. Kristensen
each galea densely spined. Labial palp long, por-
trichiated; Rs3 + 4 'stalked' in both wings; fore-
wing anal loop present (Fig. 5.4 D). Protibial
epiphysis well developed. Spurs 0-2-4. Hind tibia
unusually long, more than twice femoral length.
Eggs and larvae undescribed; a pupa has been
incidentally found "in moss"; it had no kind of
silken enclosure. The moths are nocturnal. Fe-
male specimens are rarely observed, their flight
activity is likely to differ markedly from that of
males.
The nine described species are now all in-
cluded in the genus Prototheora (Fig. 5.31), re-
stricted to Southernmost Africa (Cape Floristic
Region and Afromontane forests, northwards
to Natal).
Davis (1996; exhaustive account of available
knowledge.)
Neotheoridae. Monobasic. Neotheora chiloides
(Fig. 5.3 J) is known from a single, incomplete
female specimen. It is unique among exoporians
in having several (9) round and crested signa in
the bursa copulatrix, and the antevaginal lamella
preceded by an asymmetrical sclerite with two
posterior processes.
Maxillary palp short, with indistinct segmen-
tation; proboscis vestige distinct, medial galea
surface more distinctly concave than in other
hepialoids examined, somewhat spinose. Labial
palp long, porrect and densely scaled. Protibial
epiphysis present; metatibia just above twice
femoral length, with 4 spurs. Wings devoid of
microtrichia (forewing base excepted), with ter-
men slightly convex, apices produced; vestiture
with lower layer of 'primitive type' scales:
Rs3 + 4 'sessile'; forewing without anal loop. Ap-
parent tergum VIII narrow, anterolateral corners
produced. 'Dorsal plate' probably completely
cleft.
Brazil (Matto Grosso).
Kristensen (1978 b.)
Anomosetidae. Monobasic. Anomoses hylecoetes
(Fig. 5.3 H) is distinctive by the enormously de-
veloped male trulleum and a 'chambered' roof
of the female genital sinus hidden by the large
antevaginal lamella.
Maxillary palp short, with indistinct segmen-
tation; proboscis vestige distinct, medial galea
surface densely spinose. Labial palp not conspic-
uously long. Protibial epiphysis absent; hind
tibia only about 1.5 times femoral length; spurs
0-2-4. Wings devoid of microtrichia (bases ex-
cepted); vestiture with lower layer of 'primitive
type' scales; Rs3 + 4 variable, 'sessile or stalked;
forewing without anal loop; male hind wing with
hair-pencil concealed in folded posterior margin.
Immature stages unknown. The adult moth is
nocturnal. An Australian rainforest species,
The Homoneurous Glossata 61

known from southern Queensland and northern

New South Wales.
Kristensen (1978 a), Common (1990).

Hepialidae, 'ghost' or 'swift' moths. As currently

circumscribed the Hepialidae are a large, but
poorly characterized, assemblage of hepialoids
with the inter-M crossvein present and the meta-
tibial spur complement reduced (2 or 1 in Gazor-
yctra and Bipectilus, 0 in all others).
Antennae in some taxa dentate, or bi- or tri-
pectinate, in others simple, occasionally very
small and unsealed. Male compound eyes some-
times enlarged conspicuously (the exact distribu-
tion of this trait deserves investigation; is it re-
stricted to taxa with 'reversed-calling' mating be-
haviour?). Epiphysis variable, sometimes com-
pletely reduced; hind legs usually smaller than
fore and mid legs. Venational specializations,
such as unforked forewing Se, 'sessile' Rs4/Rs5,
and 'pectinate' Rs branches, probably evolved on
more occasions. Wing patterns diverse, sexual di-
morphism often pronounced; female brachyptery
occurs in some high-altitude Pharmacis and Aer-
ala (Sattler 1991). In male genitalia connections
between trulleum and pseudotegumen variable
and apparently of taxonomic importance. Fe-
male 'intergenital lobes' (strengthened by sub-
anal plates) flanking groove between ovipore
and ostium bursae, in some taxa fused medially,
thereby forming an internal tubular sperm pas-
sage; this variability and its phylogenetic implica-
tions deserving further scrutiny.
Larvae with development of melanized pina-
cula variable between taxa, sometimes also
markedly variable between instars of the same
species. Proleg crochets uni-, bi- or multiserial,
the latter arrangement (with crochet size increas-
ing towards centre) probably representing the
ground plan condition in mature larve.
Adults crepuscular/nocturnal; mass emer-
gences stimulated by rainfall being of regular oc-
currence in some taxa. Mating behaviour diverse
(see below). Egg numbers may be remarkably
high, > 30000 in larger species; in some taxa
they are broadcast by the females in flight. Many
larvae tunnel in soil, or construct silken galleries
among litter, and feed on roots (or, at night, on
leaves adjacent to retreat) of pteridophytes, gym-
nosperms and angiosperms; others are root/stem/
branch borers, some feeding exclusively on callus
tissue. Fungivory is probably widespread, the
transition from fungivory to phytophagy during
larval ontogeny being recorded in a number of
cases.
Four genera of medium-sized hepialids, re-
ferred to by convenience term 'primitive Hepiali-
dae', are devoid of specializations shared by
other members of the family. Fraus (Australian,
25 structurally diverse species) include some spe-
cies with almost conventional 'micro-moth fa-
cies' (slender bodies, relatively long antennae)
Fig. 5.5. Hepialidae: A, Hepialus humuli, male (the
'ghost moth' par excellence, the name refers to the ag-
gregations of pendulating white moths seen at dusk);
B, Sthenopis argenleomaculatus; C, Zelotypia stacyi.
Scale 50 mm.
and the only hepialids 5. lat. with sizable probos-
cis vestiges; known larvae grass-feeders. Afro-
theora and Antihepialus (8 and 4 named species,
respectively) are Afrotropical; their immatures
undescribed. Gazoryctra (Holarctic, 13 species)
have meso- and metatibial spurs; like the Hepia-
lidae s. str., but unlike Fraus their pupae have
two dorsal spine rows (instead of a single) on
abdominal segments IIIVII.
Remaining hepialid genera, 'Hepialidae s. str. '
probably constituting a monophylum, as indi-
cated by specialized male genitalia (trulleum
fused with pseudoteguminal plates), reduction of
pseudempodial seta, and presence of a ventral
spine row on pupal segment VII. Medium-sized
to very large, robust moths. Pretarsal arolium
and pulvilli sometimes greatly reduced. Ca. 500
named species, currently in 50 + genera; cosmo-
politan. Previous subfamily classifications con-
sidered untenable (Nielsen & Robinson 1983).
An assemblage characterized by androconial
scales on the male metatibia may prove mono-
phyletic or at least have a monophyletic core;
well-known genera include Oncopera, Hepialus
(Fig. 5.5 A), Zenophassus, Sthenopis (Fig. 5.5 B,
Nearctic, 4 species), Phymatopus (with metatibia
club-like, and metatarsi lost, Fig. 5.5), and the
probably closely related Aenetus (with many
large and colourful species) and Endoclita whose
late-instar larvae are wood-borers, sometimes
62
pests. While female pheromone-calling m a y still
occur in s o m e Oncopera, calling is 'reversed' else-
where, with male territoriality and 'pendulating'
lek flights being d o c u m e n t e d in s o m e taxa.
A m o n g the largest and/or m o s t striking hepialids
(indeed m o t h s ) are the Australian Zelotypia
stacyi (Fig. 5.5 C; w i n g s p a n u p t o 25 cm), the S.
African Leto venus, and the N e o t r o p i c a l Tricho-
phassus giganteus, all with w o o d - b o r i n g larvae.
While the t w o first-mentioned taxa probably are
closely related to the EndoclitalAenetus lineage
the latter appears isolated. Important pasture
pests occur in, e. g., Gorgopis, Dalaca, Oncopera,
Oxycanus and Wiseana. Other m a j o r and/or
wellstudied genera include, e. g., Bipectilus, Eu-
dalaca, Callipielus, Phassus, Aepytus, Aoraia, Tri-
odia, Korscheltellus, Pharmacis, Thitarodes, Tri-
ctena and Abantiades.
Hepialids are treated in m a n y regional hand-
b o o k s o n 'macro-moths', including v o l s 2, 6, 10,
14 o f Die Groschmetterlinge der ErdelThe
Macrolepidoptera of the World/Les Macrolpi-
doptres du Globe (A. Seitz ed.). Recent w o r k s
providing references t o the v o l u m i n o u s previous
literature o n the m o r p h o l o g y , b i o l o g y and taxon-
o m y o f the family include R o b i n s o n & Nielsen
(1983), G r e h a n (1989), Mallet (1984), Wagner
(1987, 1988), Wagner et al. (1989), Nielsen &
Scoble (1986), Nielsen (1988), Nielsen & Kris-
tensen (1989: principal account o f a n a t o m y ) ,
C o m m o n (1990), Wagner & R o s o v s k y (1991),
Leonard et al. (1992) and D u g d a l e (1994: exten-
sive treatment centred o n rich N e w Zealand
fauna).
Acknowledgments
I a m indebted to Drs. D . R . D a v i s , G . W. Gibbs,
M . V. K o z l o v and E. S. Nielsen for i n f o r m a t i o n
and illustrations and t o Drs. G . S. R o b i n s o n and
M . J. Scoble for c o m m e n t s o n parts o f the m a n u -
script.
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 6. The Monotrysian Heteroneura
Donald R. Davis
Including more than 99% of the extant species of
Lepidoptera (Nielsen 1985, Heppner 1991), the
infraorder Heteroneura overwhelmingly consti-
tutes the most successful clade within the order.
The monotrysian Heteroneura, a questionably
monophyletic assemblage of four distantly re-
lated superfamilies, are believed to represent the
basal group of the infraorder, as indicated by the
retention of a primitive monotrysian repro-
ductive system (Figs. 6.8 G, LN, Dugdale
1974) and more generalized distribution of wing
microtrichia. Understanding the phylogeny of
this archaic group is important because one of
its components probably is the sister group of
most of the remaining Lepidoptera (Nielsen
1985 b). Major synapomorphies shared with the
ditrysian Heteroneura include the reduction of
the hindwing Rs system (i. e., heteroneurous ve-
nation), the frenulum-retinaculum wing cou-
pling, and loss of the first abdominal sternum
(Fig. 6.7, Kristensen 1984). Monophyly of the
monotrysian Heteroneura (or Monotrysia) is in-
adequately supported, with only two possible sy-
napomorphies proposed (Davis 1986): (1) pres-
ence of spine combs (pectinifers, Figs. 6.8 B, D)
on the male valva (absent in Tischerioidea and
Palaephatoidea), and (2) metafurcasternum with
two pairs of dorsal tendons and without second-
ary arm lamellae (Figs. 6.6 I -U). Unfortunately,
the metafurcasterna of most ditrysian genera re-
main too poorly known for adequate evaluation,
and it cannot be ruled out that the monotrysian
configuration represents the groundplan state in
the Heteroneura. The subdivision of sternum 2
into a smaller anterior sclerite (S2a) and a much
larger caudal region (S2b, see Fig. 6.7) may con-
stitute another synapomorphy of the monotry-
sian Heteroneura, although considerable varia-
tion in this condition has developed within the
non-incurvarioid Monotrysia (Figs. 6.7 N - R ,
Kristensen and Nielsen 1980, Davis 1986, 1989).
The four superfamilies comprising the mono-
trysian Heteroneura have been partitioned into
two subgroups (Davis 1986): (1) Incurvariina
represented only by Incurvarioidea, charac-
terized by an extensible, piercing ovipositor with
its associated tendons for inserting eggs into
plant tissue (Figs. 6.8 G - K ) , and an elongate
cloaca terminating on or near the apex of the
tenth abdominal segment (AIO, Fig. 6.8 G); and
(2) Nepticulina - including Nepticuloidea,
Tischerioidea, and Palaephatoidea, all with
greatly shortened, nonpiercing ovipositors and a
separate ovipore opening on A8A9 (Figs. 6.8
L - N ) . Acceptance of the piercing ovipositor as
a feature of the glossatan ground plan (Nielsen
1985 b, Davis 1986), supports that Incurvari-
oidea be considered the most basal of the mono-
trysian superfamilies. Preliminary studies involv-
ing the 18S rDNA gene tentatively support this
view with the tendancy to position prodoxine
genera basal to all Heteroneura (Wiegmann
1994). In contrast, the basis for considering the
vestigial jugum in Nepticulidae as indication of
this taxon's basal position (Hennig 1953, Minet
1984) has been overemphasized (Davis 1989). Al-
though sequence data from 18S rDNA suggest
the Incurvarioidea to be paraphyletic, our cur-
rent knowledge based on morphology strongly
supports its monophyly (Nielsen and Davis
1985). Resolution of phylogenetic relationships
at this level utilizing the above, very conservative
gene was often shown to be inconclusive. Data
from genes with higher levels of sequence varia-
tion are needed to test hypotheses on heteroneu-
ran relationships (Wiegmann 1994). Hypothe-
sized morphological relationships between Nep-
ticuloidea and Tischerioidea have been estab-
lished only on the basis of reduction characters
in the adult (loss of M2) and larva (depressed
body, stemmatal reduction) - the latter often as-
sociated with a leaf-mining habit (Davis 1986,
1989). Palaephatoidea and the highly apomor-
phic Tischerioidea are suprisingly linked by two
morphological synapomorphies the presence
of a median ovipositor lobe (Figs. 6.9 GH) and
enlarged, closely set female frenular setae (Davis
1989, Nielsen 1989). The tineid-like Palaepha-
toidea had been suggested as a possible sister
group to the ditrysian Heteroneura (Davis 1986),
but Wiegmann's (1994) molecular studies more
strongly supported the monophyly of Tischeri-
oidea + Ditrysia, as proposed by Nielsen
(1985 b). See also 1 - 2 .
Heteroneura - Monotrysia
Key to the Superfamilies and Families
1 Female with elongate piercing ovipositor
(Figs. 6.8 G - K ) . Male genitalia usually with
elongate, sagittate, or deeply furcate juxta
(Figs. 6.8 C, F, Incurvarioidea) 2
Ovipositor short, non-piercing. Juxta absent
or variably shaped, never sagittate nor
deeply furcate 7
2 Hindwing with discal cell open (M-CuA
crossvein absent, (Figs. 6.5 A B)
Heliozelidae
66 Donald R. Davis

Fig. 6.1. Incurvarioidea. AC, Heliozelidae: A, Heliozela sp., Australia (4.2 mm); , Antispila nysaefoliella, USA
(3.2 mm); C, Hoplophanes chlorochrysa, Australia (6.1 mm); D - , Adelidae, D, Ceromitia iolampra, Australia
(7 mm); E, Nematopogon metalicus, Germany (8 mm); F, Nemophora amatella, Japan (11 mm); G, Nemophora
aurisparsella, Indonesia (7.5 mm); H, Adela trigrapha, USA (6.5 mm); IO, Prodoxidae: I, Lampronia russatella,
USA (6.5 mm); J, Greya punctiferella, USA (6.6 mm); K, Agavenema barberella, USA (6 mm); L, Prodoxus
y-inversus, USA (6 mm); M, Prodoxus quinquepunctellus, USA (11 mm); N, Tegeticula yuccasella, USA (12 mm);
O, Parategeticula pollenifera, USA (13.5 mm); PQ, Cecidosidae: P, Cecidoses eremita, Brazil (11.8 mm);
Q, Dicranoses congregateli, Argentina (3.2 mm); R, Incurvariidae: R, Incurvarla masculella, Germany (6 mm);
S, Perthida sp., Australia (5.8 mm); T, Paraclemensia acerifoliella, USA (5 mm). (Forewing length in parentheses).

Hindwing with discall cell closed 3 tate, either broadly angulate or deeply fur-
3 Antenna usually longer than forewing . . . . cate. Mature larva casebearing 5
Adelidae Ovipositor compressed (Fig. 6.8 K). Male
Antenna shorter than forewing 4 juxta usually slender, sagittate. Larva en-
4 Female with apex of ovipositor depressed dophagous, never casebearing 6
(Figs. 6.8 HI) Male with juxta not sagit- 5 Male pectinifer consisting of a single large
The Monotrysian Heteroneura 67

Fig. 6.2. A, Crinopterygidae: Crinopteryx familiella, France (4.5 mm); BD, Nepticulidae: B, Peclinivalva sp.,
Australia (2.7 mm); C, Ectoedemia sericopeza, Germany (3.9 mm); Stigmella malella, Germany (2.2 mm); E - G ,
Opostegidae: E, Notiopostega atrata, Chile (8.3 mm); F, Opostega spatulella, Europe (4 mm); Opostegoides gephy-
raea, Australia (3.5 mm); Tischeriidae: H, Tischeria zelleriella, USA (4 mm); I - L , Palaephatidae: I, Sesommata
leuroptera, Chile (5.6 mm); Palaephatus falsus, Chile (11 mm); H, Azaleodes micronipha, Australia (10 mm); L,
Ptyssoptera sp., Australia (7.7 mm). (Forewing length in parentheses).

spine. Larva leaving case to mine host leaf;
A36 with biserial crochets Crinopterygidae
Male pectinifer multispined or absent. Late
instar larvae usually exophytic; A 3 - 6 with
uniserial crochets Incurvariidae
6 Adult with haustellum usually reduced, less
than first labial palpal segment in length, or
absent; maxillary palpy short, 3-segmented
to absent. Frenulum-retinaculum absent.
Larva cecidogenic Cecidosidae
Adult with haustellum developed; maxillary
palpi elongate, 4-5-segmented. Frenulum-ret-
inaculum present. Larva usually boring in
fruits, leaves or stems, rarely cecidogenic . .
Prodoxidae
7 Discal cell open in forewing; antennal scape
usually greatly broadened, covering eye
(Nepticuloidea, Figs. 6.3 J - K ) 8 length of labial palpi. Prosternum reduced
Discal cell closed in forewing; scape unmodi-
fied, never forming eyecap 9 cal cell usually absent in both wings or reduced
8 Forewing with branched or stalked, often ar-
cuate veins (Fig. 6.5 I) Nepticulidae
Forewing with all veins separate and rela-
tively straight (Fig. 6.5 J) . . . . Opostegidae
9 Hindwings slender; width 0.23 the length or
less; third antennal segment 2 . 5 - 3 X length
of first (Fig. 6.3 M, Tischerioidea)
Tischeriidae
- Hindwings broader, width 0.33 the length or
more; third antennal segment less than
length of first (Palaephatoidea)
Palaephatidae
NEPTICULOIDEA.
Includes the smallest Lepidoptera; length of
forewing 1.5-8.3 mm. Vertex of head rough,
with erect piliform scales. Ocelli and chaetosem-
ata absent. Antennae with scape typically ex-
panded to form large eyecap (Figs. 6.3 J - K ) ; fla-
gellum, with specialized, often branched sensilla
(Figs. 6.9 DE). Maxillary palpi 5-segmented,
folded. Haustellum naked, short, 0.75 1.5 the
(Davis 1986). Wings with reduced venation; dis-
in forewing; hindwing with (Nepticulidae) or
without (Opostegidae) frenulum; a dense row
of elongate subcostal pseudofrenular setae (Fig.
6.9 C) usually present in hindwing of both sexes;
microtrichia scattered over all wing surfaces in
Nepticulidae, or reduced in Opostegidae. Legs
68
with tibial spur pattern of 0-2-4; epiphysis ab-
sent; hindtibia with prominent spinose scales.
Abdomen with S2a separate in Nepticulidae (Fig.
6.7 N), fused to lateral ridges (venulae) in Opos-
tegidae (Fig. 6.7 P); tergosternal connection usu-
ally absent, preserved in some Nepticulidae
(Nieukerken 1986). Ovipositor extremely short,
non-piercing, largely enclosed ventrally by en-
larged S7 (Fig. 6.8 L). Larva apodal leaf or stem
miners, with a single pair of stemmata. Pupa
with all coxae exposed and tergal spines on
A 2 - 8 ; pupa partially extruded from oval, lentic-
ular cocoon (Fig. 6.11 E) prior to eclosion.
Nepticulidae (Fig. 6.2 BD). Very small moths,
with forewings 1.54.6 mm in length. Head (Fig.
6.3 K): Vertex rough; lower portion of frons and
clypeus naked; occipital scales either relatively
broad or piliform. Ocelli absent. Eyes relatively
small, interocular index 0.671.0; cornea naked.
Antennae 0.40.6 the length of forewing, rarely
up to 1.0; scape usually greatly enlarged, cover-
ing dorsum of eye, reduced in some genera; pec-
ten present or absent; flagellum filiform, possess-
ing paired, variably branched sensilla vesicu-
loclada (Fig. 6.9 E, Nieurkerken and Dop 1987)
per segment and a single row of slender, biden-
tate scales encircling each segment. Pilifers and
mandibles absent. Haustellum short, usually less
than, rarely up to 3 X, the length of labial palpi
(Scoble 1983). Maxillary palpi long, 5-seg-
mented. Labial palpi 3-, rarely 2-segmented,
drooping; lateral bristles present or absent. Tho-
rax: Moderately depressed (Davis 1989). Meta-
furca (Fig. 6.6 Q) with primary apophyses free,
erect, elongate. Forewings (Fig. 6.5 I) slender to
lanceolate; index 0.22-0.25; venation reduced,
usually faint, without true crossveins; forewing
with Sc extremely short; M anastromosing with
R and sometimes basally with Cu; R with 1 - 4
branches terminating on costa; M with 12
branches; Cu reduced, unbranched; 1A + 2A
usually fused, rarely with anal fork; $ retinacu-
lum consisting of several stiff curled scales from
costal fold as well as subdorsal scales in some
species. Hindwing lanceolate with 3 6 major
veins and no closed cells; index 0,160.31; cilia
24X wing width; sex scaling frequently present
in S frenulum a single stout seta, sometimes
accompanied by a series of stout, subcostal pseu-
dofrenular setae; frenulum composed only of
subcostal pseudofrenular setae. Pretarsi with un-
guitractor plate composed of rows of divided
scutes. Abdomen: S2 (Fig. 6.7 N) divided into a
short, broad, anterior fragment and a much
larger posterior sclerite; a pair of small tubercu-
late plates present in pleura; wart- like processes
absent. S4 of $ with a pair of small, circular fe-
nestra; $ sometimes with sex scaling mid-dor-
sally on A68, or with paired lateral tufts on
A48; corythrogyne absent. Male genitalia. Un-
cus usually distinct and either acute or bilobed,
Donald R. Davis
sometimes completely fused to tegumen or ab-
sent. Tegumen a narrow to broad, dorsal band.
Vinculum usually well developed, elongate,
either U-shaped or like an inverted U (with lat-
eral arms), less commonly V-shaped or truncate.
Gnathos a well developed U-to -shaped bridge
with or without a median caudal process, rarely
absent. Anellus membranous. Juxta sometimes
present as a lightly sclerotized, broad plate. Val-
vae relatively large and broad, often tapering to
a narrow, distal process or apex, occasionally
with slender lobes arising from costal or mesal
surfaces; an elongate, sessile pectinifer present in
many species of Pectinivalva (Fig. 6.8 D).
Transtilla either present or absent, usually well
developed. Aedoeagus a relatively broad, elon-
gate tube usually heavily armed with numerous
spinose internally as well as various spinose scle-
rotizations externally in the surrounding dia-
phragma; base of vesica almost always enclosed
by a thickened collar, or cathrema (Scoble 1983,
Nieukierken 1986, Johansson et al. 1990). Female
genitalia: Ovipositor short, with two pairs of
apophyses; posterior pair usually longer and
more slender, terminating on T9, sometimes in
paired, setose pads (papillae anales); anterior
apophyses usually shorter and stouter, some-
times indistinct, arising from T8 which continues
as a short band across tergum. Oviporus located
terminally, between S8 and 9. Vagina short, usu-
ally enlarging anteriorly into thick walled vesti-
bulum; latter with or without spicules. Sperma-
theca with a lateral lagena. Corpus bursae usu-
ally gradually enlarged, rarely reduced, with or
without accessory (appendix) bursae; signa pre-
sent or absent, variable, usually with a large pair
of oval, reticulated signa in Trifurculini (Johans-
son et al. 1990), or with numerous spicules scat-
tered over inner walls.
Egg: Largely undescribed due to cuplike cov-
ering secreted by over egg (Johansson et al.
1990); egg very flat, laid singly and cemented to
epidermis of host. Larva (Fig. 6.10 LM):
Small, usually less than 5 mm, sometimes up to
9 mm long; body cylindrical to slightly de-
pressed, apodal, with translucent cuticle ranging
in color from yellow to green or whitish. Epicra-
nial notch deep, extending ca. 0.5 the length of
head; lobes of vertex well developed, extending
deep into T l ; frontoclypeus triangular to rectan-
gular, generally outlined by strong internal
ridges. One pair of stemmata. Antenna usually
reduced to one segment, 3-segmented in Pectini-
valva (Johansson et al. 1990). T 2 - 3 with paired
ventral ambulatory calli, sometimes with dorsal
calli. Chaetotaxy reduced; D2 absent on A l - 9 .
Paired ventral calli usually present on Al8,
sometimes lost on A8; dorsal calli present in
some Ectoedemia\ AIO with a slender pair of
sclerotized anal rods. Larvae are miners through
all instars, usually in leaves, less commonly in
bark of woody twigs, in fruit, or in galls. Leaf
The Monotrysian Heteroneura
mines are typically slender and serpentine, with
a relatively broad median frass trail (Figs.
6.12 - C ) , occasionally widening to form oval
blotch mines in some species. More than 30 plant
families reported as hosts, with Betulaceae, Fa-
gaceae, Rhamnaceae, Rosaceae, and Salicaceae
particularly favored in the Holartic region
(Nieukerken 1986). Larvae usually with 4 instars,
less commonly 5 - 8 (Johansson et al. 1990). The
number of generations can vary according to
species, with some obligatorily univoltine and
others undergoing as many as 3 generations ac-
cording to climatic conditions. Pupa: Vertex
smoothly rounded. Antennal scape usually
greatly enlarged. Wings extending to A9. A28
with one or more rows or small, tergal spines.
Al8 apparently moveable in both sexes
(Mosner 1916). Cremaster usually consisting of
a small pair of dorsal and/or ventral spines. Pu-
pation rarely inside mine, usually occurring out-
side, either attached to some plant surface or in
ground detritus beneath host, within an oval,
lenticular cocoon of densely woven brown to yel-
lowish silk.
Currently 11 genera and nearly 800 species
have been described from every major region ex-
cept Antarctica (Braun 1917, Borkowski 1969,
1972, Wilkinson and Scoble 1979, Newton and
Wilkinson 1982, Puplesis 1984 a, b, 1985, 1994,
Kemperman and Wilkinson 1985, Nieukerken
1985, 1990, Donner and Wilkinson 1989, Hepp-
ner 1991, Hoare et al. 1997). Serpentine mines,
allegedly caused by Nepticulidae (but see 1 - 2 ) ,
have been reported on fossilized leaves of one of
the most primitive seed plants, Pachypteris (Pte-
ridospermophyta) from the upper Jurassic/lower
Cretaceous of north Queensland, Australia
(Rozefields 1988). Two subfamilies are recog-
nized (Scoble 1983).
Pectinivalvinae. With pectinifer on S valva.
One genus, Pectinivalva, restricted to Australia.
Nepticulinae. With pectinifer lost, subdorsal
retinaculum present on forewing. Acalyptris, Are-
ticulata, Bohemannia, Ectoedemia, Enteucha,
Parafomoria, Simplimorpha, Stigmella, and Tri-
furcula (often treated as a subgenus of Ectoede-
mia), and Varius.
Opostegidae (Figs. 6.2 EG). Extremely to mod-
erately small moths, with forewings 1.8 8.3 mm
in length. Head (Fig. 6.3 J): Vestiture variable;
vertex typically rough with erect piliform scales
possessing minutely bidentate apices; most of
frons and cranium caudad to vertex covered with
broad, lamellar scales with 35 dentate apices;
lower frons naked. Ocelli absent. Eyes small to
large; interocular index 0 . 5 - 1 . 2 ; cornea naked.
Antennae 0.70.9 the length of forewing; scape
usually greatly enlarged; pecten absent; flagellum
filiform, typically with 3 ascoid sensilla per seg-
ment (Fig. 6.9 D) and one row of slender biden-
tate scales. Pilifers and mandibles absent. Haus-
69
tellum short, 0 . 7 5 - 1 . 5 the length of labial palpi.
Maxillary palpi long, geniculate, 5-segmented.
Labial palpi 3-segmented, moderately short,
slightly drooping; lateral bristles absent. Thorax:
Depressed, relatively broad. Metafurca (Fig. 6.6
RS) variable, usually with furcal apophyses
free; apophyses connected to secondary arms in
Pseudopostega. Forewings (Fig. 6.5 J) lanceolate,
index 0.20.23; venation extremely reduced, typ-
ically only 4 major veins present, without sec-
ondary branches or crossveins; microtrichia gen-
erally restricted to base of ventral forewing sur-
face. Hindwing index 0.140.17; a uniform
series of 418 subcostal pseudofrenular setae
(Fig. 6.9 C) arising from base of hindwing in
both sexes to couple with a subdorsal retinacu-
lum consisting of a row of stiff scales on un-
derside of Cu in forewing. Hindtibiae with rough
spinose scales; pretarsus with unguitractor plate
composed of several rows of undivided scutes.
Abdomen. S2 (Fig. 6.7 O) large, weakly sclero-
tized, usually with a large, central, partially or
completely divided hyaline area. Coremata and
corethrogyne absent. Male genitalia: Uncus re-
duced, typically a slender bridge between a pair
of widely separated setose lobes (socii?), rarely a
medium connate lobe (in Eosopostega). Tegumen
reduced to a narrow dorsal ring. Vinculum usu-
ally a narrow, rounded ventral ring, anterior
margin deeply concave in Opostegoides and Eo-
sopostega. Gnathos fused, variable, usually a
well developed arch. Transtilla typically absent,
rarely present as a slender, medium bridge. Juxta
either absent, or, if present, a broad menbranous
plate. Valvae variable in outline, with a distinct
costal apophysis extending inward from base but
articulated in Pseudopostega, apex of valva (cu-
cullus) an ovoid, pectinated lobe joined to basal
half of valva by a slender pedicel. Aedoeagus
usually absent, present in Notiopostega, Opos-
tega, Paralopostega, and Eosopostega. Female
genitalia: Ovipositor (Fig. 6.8 L) non-piercing,
short, barely extending beyond A7. Dorsal,
paired, anal papillae usually present and setiger-
ous, sometimes single, absent in Opostegoides.
Anterior apophyses usually absent; posterior
apophyses always present, either short or elon-
gate. Oviporus located terminally due to extreme
reduction of A8, 9 + 1 0 . Ductus usually mem-
branous. Corpus bursae usually a large membra-
nous sac, without a well defined signum but
often with inner walls variously covered with
minute spicules.
Egg: Cylindrical, elongate-oval, up to 1.1 mm
long. Chorion smooth and transparent, whitish
to yellow. Laid singly on epidermal surface of
plant host. Larva (Figs. 6.10 N - O ) : Body whit-
ish in color, extremely slender, 8 - 2 5 mm long,
and apodal, with paired ventral callosities on
T23. Head (Fig. 6.10 O) depressed, triangular,
with shallow epicranial notch and poorly defined
sutures. One pair of stemmata. Mandible with a
70 Donald R. Davis

Fig. 6.3. Head morphology. A, Heliozelidae: Heliozela aesella; B - C , Adelidae, Adela reaumurella: B, C, V;
D - , Prodoxidae: D, Tegeticulayuccasella, <j> maxilla (H = haustellum; MP = maxillary palpus; MT = maxillary
tentacle; ST = stipes); E, Lampronia luzella; FG, Cecidosidae: F, Oliera argentinana; G, Dicranoses congregateli,
(IS = intercalary sclerite); H, Incurvariidae: Incurvarla masculella\ I, Crinopterygidae: Crinopteryx familiella; J,
Opostegidae: Pseudopostega bistrigulella\ K, Nepticulidae: Ectoedemia phleophaga; L, Palaephatidae: Palaephatus
falsus, (Md = mandible, Oc = ocellus); M, Tischeriidae: Tischeria zelleriella (All scales = 0.5 mm).

single large setal brush. Cranium with a promi- metamorphic with spinneret not fully formed un-
nent pair of apophyses extending caudad into T1 til last (6th) instar. Larvae known to be leaf min-
and 3 pairs of prominent, internal, longitudinal ers in Pelea (Rutaceae) and cambium miners in
ridges (Fig. 6.10 N). Development slightly hyper- Betulaceae, Fagaceae, and Saxifragaceae. Pupa:
The Monotrysian Heteroneura
Simular to that of Nepticulidae. Cuticle rela-
tively smooth except for concentrations of min-
ute tergal spines on A28. Antennal scape usu-
ally extremely broad. Wings, antennae, and fore-
legs usually extending to or slightly surpassing
abdomen. All coxae flat and exposed. Pupation
occurring in a tough, silken, lenticular cocoon
(Fig. 6.11 F) outside of mine, usually in leaf lit-
ter. Family recently revised (Davis 1989) wherein
6 genera were recognized. Two subfamilies.
Opostegoidinae. Aedoeagus present. Notiopos-
tega (1 sp.), Eosopostega (1 sp.), Opostegoides
(7 sp.), Paralopostega (6 sp., all in Hawaii).
Oposteginae. Aedoeagus absent. Widely dis-
tributed in all major regions except Antarctica.
Opostega (58 sp.), and Pseudopostega (29 sp.).
INCURVARIOIDEA.
Small to very small moths; length of forewings
1.716 mm. Head usually with rough, piliform
scales, smooth and broadly scaled in most Helio-
zelidae. Chaetosemata and ocelli absent. Anten-
nae filiform, rarely pectinate in some <3 Incurv-
arla, scape without eyecap; pecten usually pre-
sent. Haustellum usually moderate in length,
typically elongate in Adelidae, to greatly reduced
or absent in most Cecidosidae; usually unsealed,
basally scaled in Adelidae and a few Prodoxidae.
Maxillary palpi variable, usually 5-segmented,
less commonly reduced or absent. Labial palpi
usually 3-segmented, with apical sensory organ
of vom Rath on segment III and erect bristles
on II; palpi rarely 2-segmented or absent. Wings
usually with complete heteroneurous venation
and distinct discal cell in forewing; venation
more reduced in Heliozelidae and Dicranoses
(Cecidosidae); microtrichia generally scattered
over all wing surfaces, reduced in some genera;
male frenulum usually consisting of a single large
bristle that inserts into a triangular, subcostal
lobe (retinaculum) under base of Sc in forewing;
frenulum absent in Cecidosidae; $ frenulum usu-
ally indistinct, consisting of 2 or more small, cos-
tal bristles along base of hindwing. Legs usually
with tibial spur pattern of 0-2-4, epiphysis usu-
ally present, absent in at least one genus in al-
most every family. Abdomen with second ster-
num (Figs. 6.7 A, C, G, H, J, L) divided trans-
versely into a mostly membranous anterior third
(S2a) and a more sclerotized, posterior two-
thirds with slender, anterolateral processes and a
bilateral pair of minute wart-like protuberances
(Fig. 6.7 A); usually with two rows of tuberculate
plates present in pleura, one near lateral margin
of sterna and one in a posterodorsal position to
spiracle (Nielsen 1980); anterolateral processes of
S2b articulating with first sternum by means of
slender tergosternal connection posterior (Fig.
6.7 A) to Al spiracle (Kyrki 1983). ? with A8
reduced and largely enclosed by enlarged S7; T7
(Fig. 6.7 E) less than half the size of S7; corethro-
71
gyne absent; without coremata. Male genitalia
typically with 1 - 3 rows of comb-like spines
(pectinifers) on valva (Figs. 6.8 AB); pectinifers
absent in Incurvariidae, Adelinae, or reduced in
Crinopterygidae and some Prodoxidae; juxta
usually elongate, either sagittate (Fig. 6.8 C) or
deeply furcate (Fig. 6.8 F). Female with elongate,
extensible, ovipositor specialized for piercing
plant tissue (Figs. 6.8 GK); 2 or more pairs of
threadlike retractor apodemes attached to vari-
ous sites along the ovipositor and associated in-
ternal organs (Fig. 6.8 G); cloaca well developed,
elongate; cloacal opening terminating near apex
of ovipositor (AIO). Larva with crochets con-
sisting of relatively simple spines typically ar-
ranged in 1 or more transversed rows Fig. 6.9 K);
crochets frequently lost within some families.
Pupa with 1 or more rows of abdominal tergal
spines; partially extruded from larval/pupal case
prior to eclosion.
Monophyly of the Incurvarioidea has been de-
termined by the presence of (1) an extensible,
piercing ovipositor, similarly developed in all
members of the clade; (2) two or more pairs of
retractor apodemes; (3) the enclosure of the $
eighth abdominal segment largely inside greatly
enlarged seventh sternum; and (4) sagittate juxta
in <5. Cladistic analysis of the six incurvarioid
families + a hypothetical outgroup was per-
formed using the computer program Hennig86
(Farris 1988). A morphological data set of 15
characters produced two parsimonius clado-
grams 36 steps long with consistency and reten-
tion indices of 69 and 54 respectively. One tree
was identical to a manually derived cladogram
previously proposed (Nielsen and Davis 1985).
Successive weighting selected the second tree that
differed only in the position of Prodoxidae +
Cecidosidae being inverted basal to Crinoptery-
gidae + Incurvariidae.
Heliozelidae (Figs. 6.1 A - C ) . Generally very
small moths with forewings 1.77.0 mm. Head
(Fig. 6.3 A): Vertex typically smooth with broad,
laminate, iridescent scales directed down over
similarly smooth frons; head rough with pilform
scales in one undescribed South American genus
(Nielsen 1985). Eyes small, interocular index
0.6-0.7; cornea with sparse, scattered inter-
facetal microsetae. Antennae 0.40.6 the length
of forewing; scape smooth, entirely covered with
usually iridescent scales; pecten absent; flagellum
filiform, fully scaled to apex but more sparsely
scaled ventrally; a single ring of scales per seg-
ment. Pilifers present or absent. Mandibles ves-
tigial. Haustellum moderate to long, 1.5-2.0 the
length of labial palpi, unsealed, occasionally with
base concealed by elongate, lamellar scales from
labrum. Maxillary palpi reduced in length, not
geniculate, usually 3 - 5 short segments, reduced
to 1 minute segment in Coptodisca. Labial palpi
moderately short, 3-segmented, drooping. Tho-
72
Fig. 6.4. 9 Tegeticula yuccasella (complex) collecting pollen from anther of Yucca elata.
rax: Lateral angles of laterocervical sclerites
broad, triangular (Fig. 6.6 A). Metafurca (Fig.
6.6 I) with dorsal apophyses moderate to well de-
veloped, arising perpendicular from mesal la-
mella free from secondary arms of metafurca-
sternum. Forewings (Fig. 6.5 AB) broadly lan-
ceolate, index 3.4-3.6; microtrichia mostly re-
stricted to subhumeral and subanal areas; scales
often lustrous; radius with 3 or 4 branches; Rs4
stalked to Ml and terminating on costa near
apex; discal cell either open or closed; accessory
cell absent; M with 1 or 2 branches; CuA with
12 branches; 1A + 2A without basal fork; <?
retinaculum a triangular lobe from costa; $ reti-
naculum consisting of 34 stout, curved spines
from a small swelling at base of Sc. Hindwings
more slender, lanceolate, index 0.20.28; M-
CuA crossvein absent, discal cell open; M with 1
or 2 branches; 1A + 2A usually fused, rarely di-
vided near margin; frenulum a single large bristle
in male with 46 smaller costal setae and with
usually 2 small bristles in . Legs with tibial spur
pattern of 0-2-4; epiphysis usually present, ab-
sent in Coptodisca. Abdomen. Anterior third of
S2 with broad U-shaped caudal rim (Fig. 6.7 A);
S7 of ? with caudal margin rounded (Fig. 6.7 B).
Male genitalia: Uncus indistinct, shallowly bi-
Donald R. Davis
lobed. Vinculum well developed with an elon-
gate, broad, V-shaped saccus. Valvae with a sin-
gle pair of pectinifers, each bearing about 520
spines. Juxta slender, sagittate. Aedoeagus an
elongate tube usually equalling length of saccus;
cornuti present. Female genitalia: Ovipositor
with depressed, moderately broad apex terminat-
ing in 45 minute cusps. Spermatheca with lat-
eral lagena (Razowski and Wojtusiak 1978).
Ductus bursae sometimes with small spines; cor-
pus bursae entirely membranous, without signa.
Egg\ Poorly known; inserted singly into host
tissue. Larva (Figs. 6.10 AB): Body white to
pale yellow or green, usually with brownish to
black pronotum and anal plate, sometimes with
a series of nearly contiguous, mid-dorsal dark
plates down length of body; 46 mm in length.
Head mostly prognathous, strongly depressed
with 2 - 5 pairs of stemmata. Legs usually absent,
well developed, 5-segmented in some Heliozela
(Davis 1987). Coxal plates, if present, separated.
Paired ventral and dorsal ambulatory calli some-
times present on T1 and 2. Prolegs either absent
or nearly so; crochets usually absent, present on
A3 6 in some Heliozela and consisting of
multiserial rows of simple, enlarged spines re-
sembling those of Adelidae except planta some-
The Monotrysian Heteroneura
times not separated mesally (Fig. 6.9 K);
crochets always absent on AIO. Some Antispila
and Coptodisca with fused, ventral ambulatory
calli on A3 6; paired dorsal and ventral calli
also present on A8 in Coptodisca, but only a sin-
gle mid-dorsal callus on A8 in Antispila. Most
first instar larvae begin a short, serpentine mine
that is abruptly enlarged into a small, full depth
blotch with the frass retained inside the mine. All
instars mine except the last, which constructs a
flat, oval case by cutting sections from the upper
and lower epidermis of the mine and joining
them with silk. The lenticular shape of the case
provides the origin of this family's common
name, 'shield bearers'. In most instances the lar-
vae release a silk strand to lower themselves usu-
ally to the ground to pupate amongst leaf litter.
Sometimes the cases are attached to the bark of
the host plant. Abandoned mines with small oval
holes are characteristic of the leaf damage caused
by these insects (Fig. 6.11 A). Some Heliozela
mine twigs, petioles, leaf ribs, or initiate galls
(Needham et al. 1928, McGufin and Neunzig
1985). At least 13 families of mostly woody trees,
shrubs, and vines have been reported as hosts
with the Betulaceae, Cornaceae, Fagaceae, and
Vitaceae particularly favored. Pupa: Very small,
length 2 - 3 mm. Cuticle thin, transparent. Head
smoothly round; antenna short, less than half the
length of wing; scape cylindrical. Abdomen with
dense patches of minute, scattered dorsal spines;
segments A 2 - 8 movable in A 2 - 7 in $
(Mosher 1916). Spiracles either raised or flush.
Pupation inside larval case with pupa partially
extruded at eclosion. Most species are probably
univoltine, less commonly bivoltine.
Because of their minute size and diurnal habit,
Heliozelidae are seldom collected. The neotropi-
cal fauna is especially poorly known. Approxi-
mately 12 genera and over 100 species (Heppner
1991) have been reported from nearly every ma-
jor region except Antarctica and New Zealand.
Most of the species are included within the gen-
era Heliozela, Antispila, and Coptodisca. The
principle autapomorphy for the family is the loss
of the M-CuA crossvein in the hindwing.
Adelidae (Figs. 6.1 D - ) . Small moths with
forewing length 3.5-12 mm. Head (Figs. 6.3
BC): Vertex rough, densely covered with pili-
form scales; frons usually rough, occasionally
covered by smooth, broad scales. Eyes small to
large, interocular index 0.5-1.7; interfacetal
microsetae scattered; eyes dimorphic in most
Adela and Nemophora with that of enlarged,
more approximate at vertex, and usually with
facets in upper 2 of eye enlarged. Antennae usu-
ally longer than forewing in both sexes, up to
3X forewing and arising nearly contiguous in
antennae shorter in Cauchas, 0.51.2 the length
of forewing in both sexes; scape sometimes
slightly swollen in 3; pecten usually present, ab-
73
sent in Cauchas; flagellum filiform, usually fully
scaled to apex with two annuii of appressed
scales per segment, more thinly scaled ventrally;
basal 0.20.6 of antenna roughly scaled in some
Adela; $ of some Adela and all Nemophora with
variable number of mediodorsal spines (Fig.
6.9 A) along base of flagellum (Nielsen 1980).
Pilifers present. Mandible vestigial. Haustellum
usually elongate, 1.52.5x the length of labial
palpus; reduced, about equal to labial palpus in
some African Ceromitia; basal '-'/ scaled.
Maxillary palpi 2-5-segmented. Labial palpi 3-
segmented, variable in length, usually upturned;
segments II and III usually rough and clothed
with long piliform scales. Thorax: Laterocervical
sclerites with slender, elongate, lateral processes
(Fig. 6.6 B). Metafurca (Fig. 6.6 J) with dorsal
apophyses well developed, arising perpendicular
from mesal lamella free from secondary arms of
metafurcasternum. Forewings (Fig. 6.5 C) slen-
der, index 0.30.37; microtrichia usually pre-
sent, reduced in some genera; radius with 5
branches, Rs2 and 3 either separate, connate or
stalked; Rs4 usually to costa in most genera, ter-
minating on termen below apex in Nematopogon\
accessory cell present; base of M faint, rarely
forked within cell; + 2A with short basal
fork; S retinaculum a long costal fold extending
under base of Sc. Hindwings similar to forewings
in width, index usually 0.35; Ml and 2 either sep-
arate or stalked; + 2A with short basal fork;
i frenulum a single stout bristle, usually accom-
panied by several smaller setae along costal mar-
gin; $ frenulum consisting of 3 - 4 smaller bristles
in a row along base of costa. Legs with tibial
spur pattern of 0-2-4; epiphysis present. Abdo-
men: Anterior third of S2 with a slender, U-
shaped caudal rim (Fig. 6.7 C); S7 of $ triangular
to subrectangular (Fig. 6.7 C), caudal margin
rounded to slightly emarginate. Male genitalia:
Uncus reduced, often bilobed; tegumen a moder-
ate to narrow, dorsal band. Vinculum well devel-
oped with an elongate, broad to slender, V- to Y-
shaped saccus. Valvae with 0 - 3 pairs of either
sessile or pedunculate pectinifers; pectinifers ab-
sent in Adela, Cauchas, Nemophora, and some
subgenera of Ceromitia. Juxta slender, sagittate.
Aedoeagus an elongate tube; cornuti usually pre-
sent, apex sometimes also armed with large, ex-
ogenous spines. Female genitalia (Fig. 6.8 G):
Ovipositor with a compressed, acute apex usu-
ally minutely serrated along ventral keel. Sper-
matheca without lateral lagena. Ductus and cor-
pus bursae membranous, without spines or
signa.
Egg: Poorly known; inserted singly into plant
tissue. Larva: Body slightly depressed, white to
green with darkly pigmented plates and head;
712 mm in length. Head mostly prognathous
with 6 pairs of stemmata; AF2 absent; adfrontal
sclerite elongate, extending to deeply incised epi-
cranial notch. Thorax with prespiracular sclerite
74 Donald R. Davis

Fig. 6.5. Wing venation. A - , Heliozelidae: A, Heliozela aesella; B, Coptodisca splendoriferella; C, Adelidae: Adela
trigrapha; D, Prodoxidae: Lampronia luzella; E - F , Cecidosidae: E, Cecidoses eremita (AC = accessory cell); F,
Dicranoses congregateli; G, Incurvariidae: Incurvarla masculella; H, Crinopterygidae: Crinopteryx familiella; I,
Nepticulidae: Ectoedemia phleophaga; J, Opostegidae: Opostega salaciella; K, Palaephatidae: Palaephatus falsus
(AnP = androconial pocket); L, Tischeriidae: Tischeria quercitella.

fused to pronotum; spiracle usually free from
prespiracular sclerite. Coxal plates fused, at least
on Tl. Legs well developed; tarsi without en-
larged, squamiform seta. Prolegs greatly reduced
on A 3 - 6 , absent on AIO; crochets arranged in
several more or less definite transverse rows that
gradually decrease in size from center. Largely
because of the secretive and sometimes omnivo-
rous feeding habits of the larva, surprisingly little
is known about the biology of most species in
this relatively well known family of moths. Eggs
are inserted into plant tissue in which the larva
may or may not feed (Heath and Pelham-Clinton
1976, Nielsen 1985). According to Chrtien
(1894), during late spring and early summer the
eggs of Nematopogon metaxella are inserted into
any convenient herbaceous plant. Upon hatch-
ing, the larva immediately drops to the ground
where it constructs a flattened, oval case from
soil particles and eventually dead leaves (Chrt-
ien 1894). The larva feeds on both living and
dead plants and does not complete its develop-
ment until the following spring. Kuroko (1961)
reports a somewhat different life history for
Nemophora raddei that may more accurately re-
flect the univoltine norm for the family. In this
species the eggs are inserted into the ovaries of
Salix sieboldiana in spring. The first instar larva
The Monotrysian Heteroneura
feeds on the ovules as well as the ovary wall.
After moulting, it constructs a small, oval case
and descends on a silken thread to the ground
where it prefers to feed on dead leaves of the
host Salix and Castanea crenata. The mature
larva (6th instar) pupates near the end of Octo-
ber, with the adult emerging the following
spring. The eggs of most Adela are inserted into
the flower ovary of their host wherein the first
instar larvae feed on the developing seeds. From
the second instar on, the larvae become caseb-
earers and feed on the lower or fallen leaves of
their host (Heath & Pelham-Clinton 1976). First
instar larvae of some Adelidae may mine leaves
(Common 1990). Over 20 families of angio-
sperms and one gymnosperm (Pinaceae) have
been reported as hosts (Kppers 1980). Pupa:
Length 5 - 7 mm. Vertex smoothly rounded. An-
tennae usually surpassing abdomen, then loosely
encircling caudal end of abdomen 1 - 4 times
(Figs. 6.10 C - D ) . Abdomen with 1 - 2 rows of
dorsal spines on most segments; segments 3 - 7
movable (Common 1990). Pupation inside larval
case with the pupal exuvium partially extruded.
Adelidae is a widely distributed family of
nearly 300 species that occur on all continents
except Antartica and are absent from New
Zealand (Heppner 1991, Janse 1945, Meyrick
1912 a, b). Autapomorphies for the family in-
clude the elongate antenna (particularly in the
<$), slender lateral processes of the lateral cervical
sclerites, and possibly the high number of ovari-
oles per ovary (10-20, Nielsen 1980, Nielsen and
Davis 1985). Two subfamilies have been pro-
posed (Kppers 1980):
Adelinae (maxillary palpi 2-3-segmented; $
valvae without pectinifers), including Adela,
Cauchas, and Nemophora.
Nematopogoninae (maxillary palpi usually 4-
5-segmented; $ valvae usually with pectinifers),
including Ceromitia and Nematopogon. Recent
mtDNA sequence data also links the problem-
atic North American genus Tridentaforma to this
family (Brown et al. 1994). The lateral cervical
sclerites of Tridentaforma (Fig. 6.6 C), however,
differ from the typical adelid form (Fig. 6.6 B).
In contrast to most Nematopogoninae which
tend to be drab in color and crepuscular or noc-
turnal, the adults of Adelinae are often metallic
and predominantly diurnal. The males of many
species of Adela and Nemophora swarm, usually
near the species' host plant or oviposition site.
The enlarged compound eyes in these males are
an adaptation for swarming (McAlpine and
Munroe 1968, Downes 1969). The development
of specialized, spinose setae and scales near the
base of the antennae of swarming males may be
further adaptations associated with this courship
behavior (Nielsen 1980), possibly for use in
sound production or visual signaling (Bland
1977).
75
Prodoxidae (Figs. 6.1 I - O ) . Small moths, with
forewings 4 to 16 mm in length. Head (Fig.
6.3 E): Vestiture usually rough, with dense pili-
form scales; frons and vertex covered with ap-
pressed, lamellar scales in Prodoxoides, with
erect tufts of slender, bidentate scales arising
more caudad on occiput. Eyes small to large; in-
terocular index 0.71.5; cornea either naked or
with sparse, scattered interfacetal microsetae.
Antennae 0.330.6 the length of forewing; scape
usually smooth, with pecten present or absent;
flagellum filiform, mostly naked except for dense
sensilla chaetica overall and a single row of nar-
row, dorsal scales per segment over basal seg-
ments which gradually disappear out to basal
third. Pilifers usually well developed, reduced in
Tetragma. Mandibles vestigial but relatively
prominent. Haustellum usually unsealed, basally
scaled in Tridentaforma and Greya pectinifera,
12X length of labial palpi. Maxillary palpi
elongate, 0.5 1.2 the length of haustellum, 3 to
5-segmented; an elongate, unsegmented, coiled
tentacle arising from intersegmental membrane
of basal segment of most Tegeticula and Para-
tegeticula (Fig. 6.3 D; Davis 1967). Labial palpi
relatively short, either porrect or upturned, usu-
ally 3-segmented, 2-segmented in Parategeticula;
lateral bristles present. Thorax: Lateral angles of
lateral cervical sclerites triangular, not produced
(Fig. 6.6 C - ) . Metafurca with apophyses either
free (Fig. 6.6 M) or joined as a bridge to second-
ary arms (Fig. 6.6 N). Forewings (Fig. 6.5 D)
slender; index 0.24-0.32; microtrichia usually
scattered over all surfaces, absent from dorsal
base of forewing in Tetragma, Rs4 terminating
on costa; Rs3 and 4 stalked in Tetragma, Acces-
sory and intercalary cells present; 1A + 2A with
basal fork; S retinaculum a triangular lobe in
costal membrane anterior to and tightly curling
under base of Sc. Hindwings ca. same width as
forewings, index 0.33-0.44; intercalary cell pre-
sent; $ frenulum a single stout seta; $ usually
with 23 sometimes indistinct frenular setae;
frenulum lacking in both sexes of Parategeticula,
sex scales typically absent from wings, a slender
hair pencil contained in hindwing anal fold of
Tetragma. Legs with tibial spur pattern of 0-2-4;
epiphysis usually present, absent in Parategeti-
cula. Abdomen: Caudal rim of S2a U to W-
shaped (Fig. 6.7 G); S7 of $ with caudal margin
rounded. Male genitalia (Figs. 6.8 AC): Uncus
variously fused to tegumen, usually with 2,
sometimes 1 terminal lobes. Tegumen a narrow
dorsal band. Vinculum usually well developed,
either V- or Y-shaped; saccus often well devel-
oped, total length 1 - 2 . 5 X the length of valva.
Anellus usually membranous. Valvae often with
a pectinifer (Fig. 6.8 B), variably developed,
sometimes reduced to a few short, contiguous
spines, or absent. Juxta a well defined, sagittate
sclerite. Aedoeagus an elongated tube, usually
without cornuti. Female genitalia: Ovipositor
76 Donald R. Davis

Fig. 6.6. Thoracic morphology. AH, Lateral cervical sclerites, Incurvarioidea. A, Heliozelidae: Heliozela aesella;
B, Adelidae: Adela trigrapha. (LP = lateral process, PS = proprioreceptor setae); C - E, Prodoxidae: C, Tridenta-
forma fuscoleuca; D, Lampronia russatella; E, Greya punctiferella; F, Cecidosidae: Cecidoses eremita, G, Incurvarii-
dae: Incurvarla masculella; H, Crinopterygidae: Crinopteryx familiella; I U, Metathoracic furcasterna, lateral
view. I, Heliozelidae: Heliozela aesella; J, Adelidae: Adela trigrapha; K - L, Prodoxidae: K, Lampronia russatella;
L, Prodoxus quinquepunctellus; M - N , Cecidosidae: M, Cecidoses eremita; , Dicranoses congregatala; O, Incurv-
ariidae: Incurvarla masculella; , Crinopterygidae: Crinopteryx familiella; Q, Nepticulidae: Ectoedemia phleophaga;
R - S , Opostegidae: R, Pseudopostega bistigulella, (APF = anteromedial process of furcasternum, FA = furcal
apophysis, M L = mesal lamella); S, caudal view of Fig. R, (SAF = secondary arm of furcasternum); T, Palaephati-
dae: Palaephatus falsus; U, Tischeriidae: Tischeria malifoliella (All scales = 0.5 mm).
The Monotrysian Heteroneura
elongate; apex (Figs. 6.8 JK) usually com-
pressed with dorsal and ventral margins often
raised and serrated, or smooth. Spermatheca
without lateral lagena. Corpus bursae usually
with paired, stellate signa which are occasionally
reduced or lost.
Egg: Poorly known; white, shape highly vari-
able, pedicellate in Tegeticula, ca. 2 mm long and
0.1 mm in diameter (Riley 1892) to oval, 0.3
0.5 mm long and 0.20.3 in diameter (Davis
et al., 1992). Chorion smooth with reduced
micropylar reticulation. Larva (Fig. 6.10 K):
Body white, green, or reddish in color, cylindri-
cal to fusiform, 6 - 2 2 mm in length. Head light
to dark brown, with frontoclypeus extending
0.5-0.66 the distance to epicranial notch. Usu-
ally 6 pairs of stemmata, reduced to 3 pairs in
apodal forms (Davis 1987). Legs usually present,
absent in Agavenema and Prodoxus; pretarsus
often with a large squamiform seta from lateral
base of claw. L series either bi- or trisetose on
T l . Prolegs reduced or absent; crochets often ab-
sent, at least in early instars, with a single
transverse row on A 3 - 6 in Lampronia and Mes-
epiola or as biserial circle in Greya; crochets al-
ways absent on AIO. Larva endophagous, boring
inside plant shoots, leaves, buds, flower recepta-
cles, fruits, or seeds. Typically there is a single
generation per year, with the larva overwinter-
ing. In synchrony with the irregular flowering
habit of their host, last instar larvae of some Pro-
doxinae are capable of prolonged diapause,
which in Prodoxus y-inversus may exceed 20
years (Powell 1992). Pupa: Vertex usually with
a prominent frontal beak, but smooth in Greya.
Wings extending to A 5 - 7 . A 2 - 7 movable in
both sexes (Mosher 1916); A 2 - 8 usually with a
single anterior row of tergal spines. Cremaster
usually consisting of a pair of dorsal spines or
tubercules on AIO, sometimes accompanied by
paired ventral spines. Pupation usually occuring
in silken cocoon either externally and attached
to host, or within larval gallery or underground,
with the adults of most species emerging in
spring to early summer.
Prodoxidae are almost entirely restricted to
the Holarctic region and are most diversified in
the Nearctic Region, with a single monotypic ge-
nus present in southern South America (Nielsen
and Davis 1985). Dugdale (1988) has reported
the discovery of an unidentified, questionable
prodoxid larva from cortical mines on Weinman-
nia sp. (Cunoniaceae) in New Zealand. The
paired, typically stellate signa are a major auta-
pomorphy for the family. Two subfamilies are
recognized, consisting of a total of 12 genera and
nearly 80 species.
Lamproniinae. Adults with metafurcal apoph-
yses free. Larvae with thoracic legs and crochets;
borers in seeds, fruits, or stems of Rosaceae, Sax-
ifragaceae, and possibly Myrtaceae, or in stem
galls of Betulaceae (larval biology unknown for
77
Prodoxoides and Tridentaforma). A typical life
cycle for a member of this subfamily normally
involves two feeding phases interrupted by a lar-
val diapause. Eggs are inserted singly into young
fruit in late spring to early summer, with the
early instar larvae first feeding on the developing
seeds. In mid to late summer, the larva (probably
third instar) leaves the fruit and spins a haberna-
culum in some crevice lower on the host or in
the soil where it overwinters. Early next spring
the larva moves back up the stem and bores into
developing buds or shoots, often entering and
damaging several. Pupation normally occurs in
the shoot with the adult emerging soon af-
terwards in late spring. Lampronia (28 sp.),
Tetragma (1 sp.), Prodoxoides (1 sp.), and Triden-
taforma (1 sp.). Relationships of Tridentaforma
remain problematical, both from the standpoint
of morphological and molecular data. Morpho-
logically, the genus possesses characters shared
with both Adelidae (basally scaled haustellum)
and Incurvariidae valva with rows of flattened
spines). Molecular data reported to date present
divergent results. Work with mtDNA (Brown
et al. 1994) associates the genus with Adela;
whereas, analyses involving 18S rDNA places
Tridentaform either at the base of the Hetero-
neura or near Cecidosidae (Wiegmann 1994).
Prodoxinae. Adults with metafurcal apophyses
joined to secondary furcal arms (Fig. 6.6 N).
Larvae either with legs and crochets or apodal.
All genera with the exception of Greya (17 sp.)
are restricted to the plant family Agavaceae. The
larval biology of those Greya studied appears to
resemble that of most Lamproniinae, with the
early instars boring in the seeds, fruits, or pedun-
cles of Saxifragaceae and Umbelliferae and then
exiting to the ground where they eventually over-
winter. Larvae complete their feeding the
following spring on the flower buds or foliage of
their host (Davis et al. 1992). Among the Agava-
ceae feeders, an unusual pollination mutualism
has evolved between two genera and their plant
hosts. Most females of Tegeticula (3 sp.) and Par-
ategeticula (1 sp.) purposefully pollinate their
host Yucca while visiting the inflorescenses for
oviposition without nectaring. This insures the
development of fruit upon which their larvae will
feed (Riley 1892; Powell and Mackie 1966; Davis
1967; Powell 1984, Pellmyr and Thompson
1992). Among all insects, only the females of
these two genera have developed specialized
maxillary tentacles (Fig. 6.3 D) which are used
to collect pollen from the anthers of their hosts
(Fig. 6.4). The females carry the pollen in a com-
pact mass under the head and force portions of
their load down the stigmatic tube of the yucca
during visits to the flower. Tegeticula oviposits in
the ovary, their larvae later consuming only a
portion of the seed crop. Parategeticula oviposits
in sterile tissue, usually in the flower scape or
outer perianth, with the larvae migrating to the
78 Donald R. Davis

Fig. 6.7. Abdominal morphology. A - R , sterna 2 (S2), 7 (S7) and tergum 7 (T7). - B , Heliozelidae, Heliozela
desella: A, S2; B, S7; C - , Adelidae, Adela trigrapha: C, S2; D, S7; E, T7; FG, Prodoxidae, Lampronia russa-
tella: F, S7; G, S2; H - I , Cecidosidae, Cecidoses eremita-, H, S2; I, S7; J - K , Crinopterygidae, Crinopteryx fami-
liella: J, S2; K., S7; LM, Incurvariidae, Incurvarla masculella: L, S2; M. S7; N, Nepticulidae: Ectoedemia phleo-
phaga, S2; O, Opostegidae: Notiopostega atraa, S2; PQ, Palaephatidae, Palaephatus falsus: P, S2; Q, Lateral
view TI2 and S2; R, Tischeriidae: Tischeria malifoliella, S2. (S = sternum, Spe = spiracle, = tergum, T P =
tuberculate piate, TSC = TI posterior ventral process of tergal sternal connection, Vn = venula, W P = wart-like
protuberance; all scales = 0.5 mm).
The Monotrysian Heteroneura
fruit to feed in a cyst within the fruit (Powell
1984). The interrelationships between Tegeticula
and their host yuccas are complex, involving sev-
eral sibling species including sympatric nonpolli-
nators, or "cheaters", with vestigial maxillary
tentacles (Davis 1967, in press; Miles 1983, Addi-
cott et al. 1990; Powell 1992). Females of the bo-
gus yucca moths, Prodoxus (10 sp.), also do not
transport pollen, but their larvae often depend
upon the development of inflorescenses and,
hence, the efforts of Tegeticula and Parategeti-
cula pollinators. Larvae of Prodoxus bore in the
flower stalks, sterile fruit tissue, or leaves of
Yucca (Wagner and Powell 1988), and those of
Agavenema (2 sp.) in the flower stalks of Agave.
Larvae of Mesepiola (1 sp.) bore in the seeds of
Nolina and Dasylirion (Frack 1982). All prodox-
ine larvae feeding on Agavaceae complete their
feeding in one season and are strictly endopha-
gous. The subfamily is largely restricted to the
Nearctic Region, with three species of Greya oc-
curring in eastern Russia and Japan (Kozlov
1996).
Cecidosidae (Figs. 6.1 PQ). Small moths, with
forewings 3.513 mm in length. Head: (Figs. 6.3
F - G ) Vestiture moderately smooth, with elon-
gate, slender, lamellar scales mostly curved for-
ward to smooth frons; all body scales piliform in
Dicranoses. Eyes large, interocular index 0.7
1.1. Antennae short, 0.5-0.6 length of forewing;
scape smooth except for dense pecten; flagellum
filiform, apparently 2 scale rows per segment but
actually with slender scales scattered only over
dorsal half; ventral half with elongate sensilla ca.
1.5x the length of flagellomere. Labrum greatly
reduced. Pilifere absent. All mouthparts absent
in Dicranoses (Fig. 6.3 G). Mandible reduced to
minute, sclerotized stubs or absent. Galea (haus-
tellum) usually greatly reduced to minute setose
lobes or absent (Figs. 6.3 G, 6.9 B), but moder-
ately well developed in Scyrotis. Maxillary palpi
0-3-segmented, reduced to a minute, unseg-
mented lobe from vestigial galea (Fig. 6.9 B) in
Cecidoses; labial palpi 3-segmented or absent,
without apical pit or lateral bristles. Thorax: An-
terior arms of laterocervical sclerites long and
slender (Fig. 6.6 F). Metafurca with slender,
elongate dorsal apophyses free from secondary
arms (Figs. 6.6 KL). Forewings (Figs. 6.5
EF) broad to slender, index 0.300.32; micro-
trichia usually present, reduced in some genera;
radius usually with 5 free branches, 4 in Scyrotis;
M 3-branched; accessory cell present; wing vena-
tion extremely reduced in Dicranoses (Fig. 6.5 F),
with most major veins indistinct; retinaculum ab-
sent. Hindwing nearly equal to forewing in
width, more lanceolate in Dicranoses; frenular
bristles absent in both sexes; wing coupling con-
sisting of long frenular scales arising in two to
five irregular rows near base of costa. Legs with
spurs 0-2-4 or 0-2-2; epiphysis present in most
79
genera, absent in Dicranoses. Abdomen: S2a with
broad, U-shaped caudal rim (Fig. 6.7 H); ter-
gosternal connection absent. S7 of with caudal
margin rounded (Fig. 6.7 I). Male genitalia. Un-
cus usually shallowly bilobed, deeply bilobed in
Dicranoses. Socii large. Valva slender, long, with
an elongated, sessil pectinifer along ventral mar-
gin at about half length of valva or with a promi-
nent swollen setose lobe along ventral margin.
Vinculum Y-shaped. Aedoeagus simple, slender
and tubular; cornuti absent. Juxta elongate, slen-
der, usually with caudal 0.2-0.3 deeply divided
and partially encircling aedoeagus. Female geni-
talia. Apex of ovipositor compressed, often sagit-
tate, ventral ridge with or without minute serra-
tions. Apophyses short, less than A7 in length.
Spermatheca without lateral lagena; caudal part
of ductus spermatheca not coiled. Vestibulum
without sclerotized structures; ductus and corpus
bursae membranous, signa absent.
Egg: Oval, up to 293 in length; chorion
relatively hard, up to 23 thick, brownish-yel-
low to dark brown; micropyle subapical, finely
reticulate, with reticulations enlarging toward
periphery and gradually dissipating. (Known
only for Cecidoses, Wille 1926). Larva: Body of
varying shades of greenish-yellow; length up to
11 mm. Head prognathous, with most structures
reduced, lightly sclerotized, light brown; stem-
mata absent. Thoracic legs reduced to circular,
minute, unsegmented tubercules. Prolegs and
crochets absent. Body setae often reduced in
length, sometimes with numerous secondary se-
tae. The larvae of all species are univoltine and
cecidogenic, developing within galls on Rhus and
Schinus (Anacardiaceae). Galls (pyxidiocecidea)
of the South American Cecidoses and Eucec-
idoses develop as hard, thick-walled, sessile
spheres (Figs. 6.11 GH) up to 18 mm in dia-
meter on the woody, terminal branches of Schi-
nus (Brthes 1916, Curtis 1835, Wille 1926, Mani
1964). Pyxidial galls are characterized by the
presence of a cap-like covering, or operculum
the formation of which is sometimes unclear. In
Cecidoses, operculum formation is largely inde-
pendent of larval development. Past a certain
developmental stage, the operculum forms even
after the young larva dies or is surgically removed
(Wille 1926). Prior to emergence, the pharate
adult easily forces open the operculum and exits.
Oliera and Dicranoses form colonies of much
smaller, elliptical to spindle-shaped galls en-
closed within swollen stems of Schinus (Brthes
1916, Keiffer & Jrgensen 1910). Stems contain-
ing Dicranoses eventually rupture, exposing the
galls inside (Fig. 6.11 I). The larvae of the South
African Scyrotis form similar galls on Rhus, ex-
cept their galls eventually dehisce and fall to the
ground. While on the ground and probably as a
means of avoiding excessive heat from the sun,
the very active pupa is capable of propelling the
gall for distances of up to 30 cm (Meyrick 1909,
80 Donald R. Davis

Fig. 6.8. Genital morphology. AF, $ genitalia: AC, Prodoxidae, Lampronia capitello: A, ventral view; B, Valva,
mesal view; C, juxta, ventral view; D, Nepticulidae: Pectinivalva commoni, <$ valva (from Scoble 1983); E, Crinopt-
erygidae: Crinopteryx familiella, juxta (from Petersen 1978); F, Incurvariidae: Incurvarla vetulella, S juxta; G ,
9 genitalia: G, Adelidae: Ceromitia fasciolata, lateral view; - K , apices of ovipositor: H, Crinopterygidae: Cri-
nopteryx familiella, ventral view; I, Incurvariidae: Perthida glyphopa, ventral view (from Common 1969); JK,
Prodoxidae: Prodoxus quinquepunctellus: J, lateral view; K, ventral view; L, Opostegidae: Pseudopostega bistrigu-
lella, lateral view; M, Palaephatidae: Palaephatus falsus, lateral view; N, Tischeriidae: Tischeria citrinipennella,
lateral view (AA = anterior apophysis, A C = afferent (transport) canal, A P = anal papilla (AIO), APA = ap-
deme of posterior apophysis, As = anus, AV apdeme of vestibulum, BC = bursa copulatrix, C G = colleterial
gland, CI = cloaca, CIA cloacal apdeme, CIO = cloacal opening, Cm = colliculum, C O = common oviduct,
EC = efferent (fertilization) canal, Os = oviporus, PA posterior apophysis, Pe = pectinifer, R - rectum, R A
rectal apdeme, R R = rectal reservoir, S = sternum, SP = spermathecal papilla, = tergum, U = utriculus of
spermatheca, V = vinculum, Va = vagina, Vs = vesicle, Vt = vestibulum). (All scales = 0.5 mm).

1917). B e c a u s e o f t h i s u n u s u a l h a b i t , t h e galls A 8 o r b e y o n d . Tergal s p i n e s o n A 2 - 8 , e i t h e r in a

of this species h a v e b e e n r e f e r r e d t o locally as
" j u m p i n g b e a n s " . J u m p i n g galls h a v e a l s o b e e n
r e p o r t e d f r o m o t h e r insect o r d e r s ( M a n i 1964).
Wille (1926) r e p o r t s 7 l a r v a l i n s t a r s f o r Cec-
idoses. Pupa: M a x i m u m l e n g t h 11 m m . H e a d
w i t h c o c o o n c u t t e r v a r i a b l e , r e d u c e d t o a small
a c u t e s p i n e o r e n l a r g e d as a t r u n c a t e , t r a n s v e r s e d
ridge o r a 3 - 5 - l o b e d p r o c e s s . A n t e n n a e m o d e r -
ately l o n g , a s l o n g a s w i n g s . W i n g s e x t e n d i n g t o
single r o w o r 2 - 3 i r r e g u l a r r o w s ; s e g m e n t s 2 - 8
m o v a b l e in $ , A 2 7 in . P u p a t i o n i n s i d e gall.
T h e C e c i d o s i d a e ( i n c l u d i n g R i d i a s c h i n i d a e ) is
a small f a m i l y (7 species) of exclusively gall
f o r m i n g m o t h s restricted t o the southern H e m i -
s p h e r e . F o u r g e n e r a ( C e c i d o s e s 1 sp., Dicra-
noses (= Ridiaschinia) 2 sp., Eucecidoses
1 sp., a n d Oliera 1 sp.) h a v e b e e n d e s c r i b e d
f r o m southern South America and two genera
The Monotrysian Heteroneura
(Ptisanora - 1 sp., Scyrothis - 1 sp.) f r o m South
Africa. Except for two gall-producing species of
Prodoxidae (Lampronia fuscatella and possibly
Prodoxus y-inversus), the Cecidosidae are the
only Incurvarioidea with this habit. In addition
to their cecidogenic biology, other m a j o r autapo-
morphies of this family include: (1) host restric-
tion to Anacardiaceae, (2) loss of the retinacu-
lum-frenulum wing coupling in male, and (3) loss
of the tergal-sternal connection between T1 and
S2 of the abdomen (Fig. 6.7 H). Together with
the Prodoxinae, Cecidosidae are the only incurv-
arioids to have adapted to arid and semiarid en-
vironments. Early reports associated Cecidosi-
dae with either Tortricidae or Tineidae (Curtis
1835, Brthes 1916). Biezanko (1961, actually
Forbes in litt. ) and later Becker ( 1977) were the
first to note the family's true affinities with the
Incurvarioidea.
Incurvariidae (Figs. 6.1 R - T ) . Small m o t h s with
forewing length 3 . 5 - 9 m m . Head (Fig. 6.3 H):
Usually entirely rough with piliform scales; irons
smooth in Paraclemensia, with broader, laminate
scales. Eyes moderately reduced; interocular in-
dex 0.61.0; cornea without interfacetal micro-
setae. Antennae 0.40.8 the length of forewing;
scape smoothly scaled; pecten present; flagellum
usually filiform, bipectinnate in $ Incurvarla, an-
nuii usually indistinct with scales scattered over
flagellomere. Pilifers moderate to well developed.
Mandibles vestigial. Haustellum unsealed, short,
ca. Vi the length of labial palpus. Maxillary palpi
5-segmented, geniculate. Labial palpi 3-seg-
mented, relatively short, drooping, with erect
bristles f r o m apex of second segment. Thorax'.
Ventral half of lateral cervical sclerites broad,
with lateral angles only slightly produced (Fig.
6.6 G). Metafurca (Fig. 6.6 O) similar to Adeli-
dae, with well developed dorsal apophyses aris-
ing perpendicular f r o m anteromedial process,
free f r o m secondary arms. Forewings (Fig.
6.5 G) slender, index 0 . 2 4 - 3 . 3 ; microtrichia usu-
ally present on all wing surfaces. Radius with
45 branches, Rs4 to costa above apex; acces-
sory cell present; M with 23 branches; base of
M faint or absent within cell; 1A + 2A with
short basal fork; $ retinaculum either a long cos-
tal fold or a short flap extending under base of
Sc. Hindwings nearly as b r o a d as forewings; M
with 3 branches; base of M usually preserved and
forked; 1A + 2A with short basal fork; frenulum
a single stout bristle in $ accompanied by several
smaller, costal setae; female with usually an un-
differentiated, basal series of much smaller costal
setae. Legs with tibial spur pattern of 0-2-4;
epiphyses either present or absent. Abdomen.
Caudal rim of S2a broadly U-shaped (Fig.
6.7 L); caudal margin of S7 truncate (Fig.
6.7 M). Male genitalia. Uncus usually indistinct
and shallowly bilobed, consisting of an elongate
rodlike, caudal appendage in Alloclemensia. Vin-
81
culum well developed, 1 - 2 X the length of val-
vae, U - to V-shaped. Valvae without a distinct
pectinifer but with flattened spines often ar-
ranged in 1 - 3 groups. Juxta (Fig. 6.8 F) with
1 - 2 pairs of usually elongate caudal arms par-
tially enclosing aedoeagus. Aedoeagus tubular,
usually with 1 or more large cornuti. Female gen-
italia: Ovipositor with a depressed, acute apex
usually bearing minute, lateral serrations (Fig.
68 I). Spermatheca without lateral lagena. Duc-
tus bursae often with internal sclerotizations;
signa absent.
Egg: Poorly known; inserted singly into plant
tissue. Larva: Body slightly depressed, variable
in color f r o m white, pale yellow to gray;
5 - 9 m m in length. Head mostly prognathous
with 6 pairs of stemmata; A F 2 absent; adfrontal
sclerite elongate, extending to deeply incised epi-
cranial notch. Thorax with prespiracular sclerite
bearing spiracle usually separate f r o m prono-
tum. Sternal plates reduced, coxae separate. Legs
well developed; tarsi sometimes with an enlarged,
squamiform seta adjacent to claw (Davis 1987).
Prolegs greatly reduced; crochets present on
A36 and arranged in a uniserial, transverse
band; AIO usually without crochets but present
in bi- to triserial rows in Excurvaria, Paraclemen-
sia, and Vespina. Eggs are frequently inserted
into the underside of the host leaf where the first
instar develops an irregular blotch mine (Dziur-
zynski 1958). O n completion of the mining stage,
most larvae cut through the upper and lower epi-
dermal layers of the mine and construct an oval,
lenticular case (Figs. 6.11 B E) of the two sec-
tions by sewing the edges together. At this stage,
leaf damage becomes most obvious as nearly cir-
cular holes begin to appear on the host, some-
times accompanied by circular patches of skele-
tonized leaf (Figs. 6.11 B, D, Jensen 1932). Drag-
ging the case, the larva continues to feed exter-
nally on its host (Alloclemensia, Basileura, Para-
clemensia, Vespina, Davis 1972, Nielsen 1981,
Parra & Ibarra 1994 a, Ross 1958) or descends
often to the ground to feed on dead or partially
withered leaves (Incurvarla). As the larva devel-
ops, it enlarges the case by cutting oval sections
f r o m leaves and adding these to the top and bot-
tom of the smaller, older case (Fig. 6.11 C). A
few genera (e. g., Perthida, Protaephagus) con-
tinue to mine throughout their larval develop-
ment and do not abandon the mine to construct
a case until ready to p u p a t e ( C o m m o n 1969,
Scoble 1980). Hosts include members of the
Aceraceae, Betulaceae, Caprifoliaceae, Corna-
ceae, Ericaceae, Fagaceae, Myrtaceae, Pro-
teaceae, and Rosaceae ( C o m m o n 1990, H e a t h
and Pelham-Clinton 1976, Scoble 1980) Pupa:
Length 410 mm. Vertex smooth. Haustellum
short, usually extending to end of labial palpi.
Wings extending to A 5 - 7 movable in $, A 2 - 6
in ? ( C o m m o n 1990). A 2 - 8 with scattered tergal
spines (Jensen 1932), reduced to 1 - 2 rows in
82 Donald R. Davis

Fig. 6.9. Adults and immatures, morphological details. A, Adelidae: Adela caerulella, r? antennal segment 10 with
dorsal spine (46 ); , Cecidosidae: Cecidoses cremila, head, ventral view (H = haustellum, L = labrum, Md =
mandible, M P = maxillary palpus, (100 ); C - D , Opostegidae, Pseudopostega bistrigulella: C, hindwing pseudo-
frenular setae (150 ); D, flagellomere, sensilla ascoidea (23 ); E, Nepticulidae: Stigmella juglandiella, flagel-
lomere, sensillum vesiculocladum (10 ); FG, Tischeriidae, Tischeria malifoliella: F, flagellomere, recurved
sensilla trichodea (17 ); G, dorsal view of 9 ovipositor (arrow = peg spines of A 10, 50 ); H - J , Palaephati-
dae: H, Metaphatus cirrhus, median sensory ridge, AIO, of 9 ovipositor (20 ); IJ, Melaphatus ochraceons,
larva: I, pretarsus of T1 with large squamiform seta (15 ); J, proleg, A4, crochets with mesal plate = MeP,
arrow = anterior (60 ); , Heliozelidae: Heliozela aesella, larval crochets, A3 (108 ); L, Opostegidae; Paralo-
postega callosa, spinose mandibular seta (MS) of larva, lateral view (25 ). (Scale lengths shown in parentheses;
bar scale for all photographs as in Fig. A).

s o m e g e n e r a ( R o s s 1958). P u p a t i o n o c c u r s w i t h i n t r a l i a n ( e n d e m i c Perthida a n d several o t h e r u n -

t h e l a r v a l case, either a t t a c h e d t o t h e h o s t o r o n d e s c r i b e d g e n e r a a n d species, Nielsen a n d C o m -
t h e g r o u n d in leaf litter. All species a r e believed m o n 1991) a n d P a l a e a r c t i c (Alloclemensia, Ex-
to be univoltine. curvaria, Incurvarla, Paraclemensia, Procacitas,
T h e I n c u r v a r i i d a e is a small, c o s m o p o l i t a n a n d Subclemensia, K o z l o v 1987, K u p r i j a n o v
f a m i l y o f a p p r o x i m a t e l y 100 d e s c r i b e d species 1992, 1993, Nielsen 1981) regions. It is p o o r l y
a n d 11 g e n e r a . It is b e s t r e p r e s e n t e d in t h e A u s - r e p r e s e n t e d in A f r i c a (Protaephagus capensis
The Monotrysian Heteroneura
Scoble 1980), South America (Basileura, Sima-
cauda Nielsen and Davis 1981, Parra and Ibarra
1994 a), and North America (endemic Vespina
quercivora, Davis 1972) and one species each of
Alloclemensia and Paraclemensia (Davis 1983,
Nielsen 1981, 1982). One autapomorphy of the
family is the presence of large, noncontiguous
spines on the $ valvae, in lieu of a pectinifer. The
depressed apex of the ovipositor and trunctate
caudal margin of A7 (?), synapomorphies shared
with its nearest relative, Crinopterygidae, consti-
tute other diagnostic features.
Crinopterygidae (Fig. 6.2 A). Small moths with
forewing length 3 - 3 . 5 mm. Head (Fig. 6.3 I):
Entirely rough, with piliform scales. Eyes moder-
ately reduced; interocular index ca. 1.0. Antenna
0.6 the length of forewing, smoothly scaled; pec-
ten present; flagellum filiform; annuii indistinct,
with probably scattered slender scales. Labrum
and pilifers reduced. Mandibles minute, ca.
the length of maxillary palpus I. Haustellum un-
sealed, short, ca. 0.5 the width of head. Maxillary
palpus 4-5-segmented, geniculate; apical segment
minute, sometimes lost. Labial palpi 3-seg-
mented, with erect bristles from second segment.
Thorax. Lateral projections of lateral cervical
sclerites broadly triangular (Fig. 6.6 H). Meta-
furca (Fig. 6.6 P) with anterior margin of ante-
romedial process (APM) broad, flared; dorsal
apophyses moderately short, free of secondary
arms. Forewings (Fig. 6.5 H) slender, index 0.27.
Radius with 5 separate branches; M with 3 sepa-
rate branches; base of M simple within cell of
both wings; accessory cell and chorda present;
+ 2A completely fused, without basal fork;
S retinaculum a relatively elongate, triangular,
costal fold extending under base of Sc. Hind-
wings nearly as broad as forewings, index 0.25;
M with 3 separate branches; S frenulum a single
large bristle accompanied by several smaller cos-
tal setae; $ with a basal series of much smaller,
undifferentiated costal setae. Legs with tibial
spur pattern of 0-2-4; epiphysis present. Abdo-
men: Caudal rim of S2a W-shaped (Fig. 6.7 J);
caudal margin of S7 truncate (Fig. 6.7 K).
Male genitalia: Uncus indistinct, with a broad,
shallow, bilobed caudal margin. Vinculum
broadly V-shaped. Valvae with a single large sub-
apical spine. Juxta broadly angulate (Fig. 6.8 E).
Aedoeagus short and relatively stout, with nu-
merous short cornuti (Petersen 1978). Female
genitalia: Ovipositor similar to some Incurvarii-
dae, with a depressed, laterally lobed apex (Fig.
6.8 H). Bursa copulatrix completely membra-
nous, signa absent.
Egg: Poorly known; inserted singly into plant
tissue. Larva: Morphology poorly known. Body
cylindrical, ca. 3.44.0 mm in length. Head
mostly prognathous with 6 pairs of stemmata.
Thoracic legs well developed, coxae separate.
Prolegs reduced; crochets present on A 3 - 6 and
83
arranged in two uniserial rows; crochets absent
on AIO. According to Nolcken (1882), the larva
constructs an elongate (3.54.0 mm long, 1.5
2.0 mm wide), approximately cylindrical case
(Fig. 6.10 E - G ) from the leaf epidermis of its
host, Cistus (Cistaceae). Prior to feeding, the
larva attaches its case to the underside of a leaf
and then enters through a small opening to mine
the leaf in a manner similar to that of Coleo-
phoridae. Pupa: Maximum length ca. 3.5 mm
A28 with a large patch of minute, scattered,
tergal spines (Petersen 1978).
Crinopterygidae is represented by a single
species, Crinopteryx familiella, from southern
France and Sicily. It has only been reared from
Cistus, a genus of evergreen shrubs found most
abundant in the Mediterranean region. On the
basis of the depressed ovipositor, Crinopteryx
shows greatest affinity to Incurvariidae. How-
ever, significant autapomorphies (e. g., coleo-
phorid-like larval biology, elongate-cylindrical
larval case, angulate juxta, and a single large
spine on the $ valva), exclude the genus from all
other incurvarioid families (Nielsen and Davis
1985).
PALAEPHATOIDEA
Palaephatidae (Figs. 6.2 IL). Small moths with
forewings 3.8-16 mm in length. Head (Fig.
6.3 L): Vestiture rough to semi-rough, consisting
of long piliform scales with acute to bidentate
apices. Ocelli usually absent, present in Sesom-
mata. Eyes small to large, interocular index 0.6
1.2; cornea naked. Antennae 0.50.9 the length
of forewing; pecten usually present, absent in
subgenus Palaephatus; flagellum filiform, each
segment with two rows of slender, bidentate
scales dorsally and a narrow strip ventrally
which is bare except for dense sensilla chaetica.
Pilifers usually well developed, reduced in Palae-
phatus. Mandibles vestigial, varying in size from
minute to moderately large. Haustellum short to
moderate in length, seldom exceeding length of
maxillary palpi. Maxillary palpi usually long, 5-
segmented, reduced to 4 short segments in Apo-
phatus. Labial palpi 3-segmented, curved slightly
upwards; second segment with bristles usually
arising apically. Thorax: Metafurca (Fig. 6.6 T)
with apophyses free, usually projecting dorso-
caudally between secondary arms. Forewings
(Fig. 6.5 K) slender to moderately broad, index
0.27-0.36; Radius 5-branched, Rs4 terminating
on costa; accessory and intercalary cells present;
1A + 2A with basal fork; microtrichia distrib-
uted over all surfaces of both wings, restricted
on dorsal surfaces of forewing of Sesommata and
Metaphatus to apical fifth; retinaculum arising
from costal membrane, usually forming a trian-
gular lobe curling under base of Se, or an elon-
gate fold in Sesommata. Hindwings usually as
broad as forewings, index 0.330.49; S frenulum
84 Donald R. Davis

Fig. 6.10. Immature stages. - B , Heliozelidae, Coptodisca arbutiella, larva: A, dorsal; , ventral; C - D , Adelidae,
Nemophora raddei, pupa (from Kuroko 1961): C, ventral; D. lateral; E - G , Crinopterygidae, Crinopteryx familiella,
larval case (from Petersen 1978): E, dorsal; F, anterior end, lateral (SH = stellate leaf hair); G, cross section; H - J ,
Tischeriidae, Tischeria malifoliella, larva: H, dorsal; I, Setae D l - 2 of A3-6; J, ventral; K, Prodoxidae: Prodoxus
quinquepunctellus, Larva, lateral; LM, Nepticulidae, Ectoedemia phleophaga, larva: L, Head, dorsal; M, lateral
(AR = anal rod); N - O , Opostegidae, Opostegoides scioterma, larva: N, Head, dorsal, (AfR = adfrontal ridge,
CA = cephalic apophysis, LR = lateral ridge, HR = hypostomal ridge); O, lateral; P, Palaephatidae: Metaphatus
ochraceus, larva, lateral. (All scales = 1 mm).

a single stout seta; $ frenulum with 24 frenular of $ often with folds or pockets containing spe-
setae; intercalary cell usually present, often ab- cialized sex scales (Fig. 6.5 K). Legs with tibial
sent and with discal cell sometimes open in those spur pattern of 0-2-4; epiphysis usually present,
S with modified venation; fore- and hindwings absent in Apophatus. Abdomen: 2S with anterior
The Monotrysian Heteroneura 85

Fig. 6.11. Immature stages. A, Heliozelidae: Antispila nysaefoliella, old mine damage on Nyssa sylvatica, holes are
areas cut out by larvae for pupal cases; E, Incurvariidae: B - C , Paraclemensia acerifoliella: B, Feeding injury
on Acer saccharum, larval cases indicated by arrows; C, Larva with larval case opened to show successively larger
cut leaf sections. D, Vespina quereivora on Quercus agri/olia, larval cases (see arrows) and leaf damage; E, Incurv-
arla masculella larval/pupal case with pupal exuvium, length 13 mm; F, Opostegidae; Notiopostega atraa, cocoon,
length 6 mm; G - I , Cecidosidae: G, Cecidoses eremita, galls on Schinuspolygamus, showing exit hole and detached
operculum (arrow), diameter of largest gall 17 mm; H, Eucecidoses minutanus, open galls on Schinus sp.; I,
Dicranoses congregateli, stem galls on Schinus sp.

margin truncate to excavate (V-shaped), with or
without a smaller, anterior sclerite (Fig. 6.7 P,
S2a); tergosternal connection well developed
(Fig. 6.7 Q). Coremata usually absent, sometimes
with pleural sex scales on pleuron of A23; cory-
throgyne absent. Male genitalia: Uncus usually
fused to tegumen, with rounded to bilobed apex.
Tegumen a narrow dorsal band. Vinculum usu-
ally reduced, broadly triangular. Gnathos usu-
ally present, absent in Metaphatus and Plesiopha-
tus. Juxta usually absent, reduced in Sesommata
and Plesiophatus. Anellus usually membranous,
sclerotized in Sesommata and Metaphatus. Val-
vae variable in form, from elongate and slender
to broadly rounded, often with lobes or pro-
cesses from distal third. Aedoeagus highly vari-
able, usually as long as genital capsule, with vari-
ably developed exogenous spines or lobes and
cornuti. Female genitalia (Fig. 6.8 M): Ovipositor
short and barely extending beyond A8, nearly
truncate and non-piercing, with a slightly raised
median sensory ridge (Fig. 6.9 H). Two pairs of
apophyses with anterior pair from A8 and more
slender. S8 short and broad; caudal margin
either divided or entire and often with paired
spine patches. Oviporus between S8 and 9 + 10.
86
Vagina relatively broad. Ductus bursae often
with thickened, internally spined walls, gradually
enlarging to elongate corpus bursae; internal
walls of corpus bursae usually with dense spic-
ules; signum present or absent.
Egg\ Unknown. Larva (Fig. 6.10 P): Small, ca.
712 mm in length. Body cylindrical, pale green
to white, with dark brown head, pronotal and
anal plates; setal pinacula extremely small,
slightly pigmented. Frontoclypeus extending ca.
0.50.6 the distance to epicranial notch; 6 stem-
mata present in a closely set anterodorsal ellipse.
Lateral series on T1 trisetose. Legs well devel-
oped; pretarsus with large squamiform seta from
lateral base of claw (Fig. 6.9 I); coxae widely sep-
arated. Prolegs well developed on A36 and
AIO; crochets on A 3 - 6 biserial with outer series
interrupted along mesal side by an elongate,
crescentic brownish plate (Fig. 6.9 J). Anal plate
with 3 pairs elongate setae; crochets of AIO with
ca. 10 hooks in a curved row. Larvae feed on
various species of Proteaceae (e. g., Embothrium,
Lomatia in South America and Persoonia in Aus-
tralia), at first mining in leaves and later living
in shelters formed by webbing together adjacent
leaves (Nielsen and Common 1991, Parra and
Ibarra - Vidal 1994 b). Pupa: Head with vertex
smooth. Wings extending almost to A5. Abdo-
men with a single, anterior row of small spines
on A 2 - 9 ; A 2 - 8 movable in . Cremaster con-
sisting of a pair of large, dorsal spines on AIO.
Pupation inside a loose shelter of plant material
spun together. Pupa partially extruded from
shelter prior to emergence.
The family currently includes ca. 60 species
equally divided between southern South America
(Davis 1986) and Australia (Nielsen 1987; Com-
mon 1990). Five genera (Sesommata, 6 sp.; Meta-
phatus, 6 sp.; Plesiophatus, 1 sp.; Palaephatus,
13 sp.; and Apophatus, 2 sp.) have been described
from South America and two (Azaleodes, 4 sp.
and Ptyssoptera, 25 sp.) from Australia. Autapo-
morphies for the Palaephatidae include the ter-
minal, median ridge covered with campaniform
sensilla of the ovipositor (Fig. 6.9 H) and possi-
bly the mesal plate on the abdominal prolegs of
the larva (Fig. 6.9 J). The squamiform seta lat-
erad to the pretarsal claw on the larva (Fig. 6.9 I)
is an unusual feature also shared with some In-
curvariidae and Prodoxidae. Unlike most mono-
trysian Heteroneura, adult male Palaephatidae
have also developed a diverse assortment of sex
scale pockets, most of them located in various
regions of the wings (Fig. 6.5 K).
TISCHERIOIDEA
Tischeriidae (Fig. 6.2 H). Small moths with fore-
wings 2.75 mm length. Head (Fig. 6.3 M): Ves-
titure of vertex generally rough with slender to
moderately broad scales directed mostly for-
ward; frons smooth. Ocelli absent. Eyes large; in-
Donald R. Davis
terocular index ca. 2.2; cornea naked; dorsal rim
slightly concave beneath antenna. Tentorial pits
situated high on frons and touching rim of eye.
Antennae 0.6-0.7 the length of forewing; scape
mostly smooth, with a prominent tuft of slender
pecten projecting over eye; flagellum filiform,
with 2 or more rows of slender scales per seg-
ment over dorsal half; venter naked except for
dense sensilla chaetica which are 23X longer in
$ than 9; sensilla trichodea with strongly re-
curved bases (Fig. 6.9 F, Davis 1986); flagellom-
ere 1 (third antennal segment) 2.5 3 X longer
than 2. Pilifers well developed. Mandibles re-
duced to a pair of minute lobes. Haustellum
moderately long, over 2.5X the length of labial
palpus; basal half scaled; distal half naked, scle-
rotized, covered with overlapping plates. Maxil-
lary palpi reduced to either 1 or 3 short seg-
ments. Labial palpi moderately short, 3-seg-
mented, drooping, smooth scaled. Thorax: Meta-
furca (Fig. 6.6 U) with apophyses connected as a
bridge to secondary arms. Forewings (Fig. 6.5 L)
lanceolate; index 0.17-0.28; microtrichia re-
stricted to basal area of both wings; forewings
usually with all 5 branches of R terminating on
costa; Rs2 sometimes absent; M 2-branched,
base of M present, usually faint within cell; CuA
unbranched; CuP faint. 1A + 2A usually with-
out basal fork; retinaculum a broad lobe aris-
ing in costal membrane immediately anterior to
and tightly coiling under base of Sc. Hindwings
usually narrower than forewings, sometimes
broader in $ than in ; index 0.15-0.23; M and
Cu usually unbranched; discal cell open; $ frenu-
lum a single stout seta; with 2 closely set frenu-
lar setae. Legs with tibial spur pattern of 0-2-
4; epiphysis present. Abdomen: 2S divided into
3 sclerites with caudal most sternite possessing
elongate lateral arms extending anteriorly (Fig.
6.7 R); postspiracular tergosternal connection
present but weak. 2 pairs of tuberculate plates
present, one on lateral margin of 2S, other near
2T; plates evident on most of caudal segments;
wart-like protuberances apparently absent. Core-
mata and corethrogyne absent. Male genitalia:
Uncus strongly bifurcate, often fused to short or
moderately long tegumen. Vinculum variable, a
very slender to broad ventral ring, U-V- or Y-
shaped, saccus present or absent. Gnathos ab-
sent. Juxta sometimes present. Anellus weakly
sclerotized. Transtilla present, sometimes sepa-
rated. Valvae usually simple, broad to slender.
Aedoeagus usually extremely slender, with a
strongly bifurcate apex with gonopore at or near
base of fork. Female genitalia (Fig. 6.8 N): Ovi-
positor short, non-extensible, and non-piercing.
T10 comprised of a bilateral pair of lobes bear-
ing short, stout "peg setae" and a median, termi-
nal, smooth ridge without campaniform sensilla
(Fig. 6.9 G). T9 represented by a bilateral pair of
subdorsal, usually smaller, flatter and finely se-
tose lobes which articulate with a moderately
The Monotrysian Heteroneura
Fig. 6.12. Leaf mines. AC, Nepticulidae: A, Trifurcula (Glaucolepsis) saccharella on Acer saccharum; B, Ectoede-
mia platanella on Platanus occidentella; C, Stigmella crataegifoliella on Crataegus sp.; D, Tischeriidae: Tischeria
quercitella on Quercus sp., note thickened nidus (arrow).
elongate pair of posterior apophyses. A8 a nar-
row sclerotized ring which is constricted laterally
where T8 is weakly joined to the greatly attenu-
ated anterior internal arms (i. e., apophyses) of
S8; in some species the median portion of S8 is
extended slightly anteriorly (the patibulum of
Braun 1972) before dividing to form the apophy-
ses. S9 extremely reduced to a minute ventral
band from which arise a usually short, slender,
lateral and more ventral pair of apophyses as
well as a broader and more dorsal, internal pair.
Both pairs of apophyses were collectively re-
ferred to as the prela by Braun (1972). Oviporus
situated in membrane of vestigial S9. Vestibulum
often enlarged with thickened walls. Bursa copu-
latrix membranous, without signum but often
with minute, spinulose sclerotizations.
Egg. Poorly known. Elliptic to broadly oval in
shape, laid singly and cemented to leaf surface.
Larva (Figs. 6.1 H - J ) : Small, not exceeding
6 mm. Body slightly flattened, pale green to yel-
low, with brownish head, pronotal and anal
plates. Head flattened; frontoclypeus and ecdy-
sial lines converge at epicranial notch; 6 stem-
mata, mostly horizontal. Thoracic legs greatly
reduced to absent, either with 2 vestigial seg-
ments (Emmet 1976) or with ambulatory calli;
latter also present on dorsum of T l - 3 . Prolegs
rudimentary with crochets in multiserial bands
or incomplete ellipses. D1 and 2 of A36 closely
approximate (Fig. 6.1 I), widely separated else-
where. Larvae are leaf-miners throughout the
87
larval stages and actively spin silk in all instars.
Feeding may proceed in one direction (trumpet
mine) or in all directions (irregular blotch mine).
Some species construct a firm, silk-lined nidus
within the mine into which the larva often re-
treats when alarmed or not feeding (Fig. 6.12 D).
As many as 4 broods may occur. Hosts include
the Asteraceae, Combretaceae, Ericaceae, Faga-
ceae, Malvaceae, Rhamnaceae, Rosaceae, Ster-
culiaceae, and Tiliaceae (Braun, 1972). Pupa:
Head with vertex usually smooth, rarely with
frontal process. Wings to anterior margin of A4.
A 3 - 7 movable in , A 3 - 6 in $ (Mosher 1916).
Concentrations of irregular, tergal spines on A2
or 37; D1 and 2 approximate on the movable
segments. Cremaster consisting of 2 dorsal hooks
on AIO. Pupation occurs within mine, usually in
a flimsy cocoon.
The Tischeriidae are a small, primarily holarc-
tic family of ca. 80 species, all included in one
genus, Tischeria. The greatest concentration of
species occurs in the United States (47 sp.) with
only a few described from the Neotropical, Ethi-
opian, and Indo-Malayan regions, and none
from the Australian region or Oceania.
Acknowledgments
I am indebted to Vichai Malikul and George
Venable of the Department of Entomology,
Smithsonian Institution, Washington, D. C.,
USA and Biruta Akerbergs Hansen of Liverpool,
88
Pennsylvania for the drawings used in this chap-
ter. Similarly, I wish to thank Victor Kranz and
Carl Hansen of the Smithsonian Photographic
Laboratory and Susann Braden and Walter
Brown of the Smithsonian Scanning Electon
Microscope Laboratory for their photographic
assistance. Ebbe S. Nielsen of CSIRO, Canberra,
Australia graciously provided some illustrations
and notes.
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Puplesis, R . K . (1984a): sisteme molej-maljutok
(Lepidoptera, Nepticulidae) palearkticeskoj fauny.
- Ent. Obozr., 63: 582-597.
- (1984 b): Obzor vidov roda Microcalyptris (Lepi-
doptera, Nepticulidae) iz pystyn Mongolii i SSSR
(A review of the nepticulid moths of the genus
Microcalyptris (Lepidoptera, Nepticulidae) from
the deserts of Mongolia and the USSSR). - Nasek-
omye Mongolii, 9: 484-507.
- (1985): Contributions to the classification of the
palaearctic Nepticulidae (Lepidoptera). - Ent.
Rev., Washington, 63: 149-164. [English transla-
tion of Puplesis 1984 a].
- (1994): The Nepticulidae of eastern Europe and Asia:
western, central and eastern parts. 299 p. Leiden.
Razowski, J. & Wojtusiak, J. (1978 a): Family-group
taxa of the Adeloidea (Lepidoptera). - Polskie
Pismo Ent., 48: 3 - 1 8 .
- (1978 b): Polish genera of the Adelidae (Lepidop-
tera). - Polskie Pismo Ent., 48: 19-33.
Riley, C. V. (1892): The yucca moth and Yucca pollen-
ation. Missouri Bot. Gard., Third Ann. Rept.,
p. 9 9 - 1 5 8 .
Donald R. Davis
Ross, D. A. (1958): The maple leaf cutter, Paraclemen-
sia acerifoliella (Fitch) (Lepidoptera: Incurvarii-
dae), descriptions of stages. Canadian Ent., 90:
541-555.
Rozefelds, A . C . (1988): Lepidoptera mines in
Pachypteris leaves (Corystopermaceae: Pterido-
spermophyta) from the Upper Jurassic/Lower Cre-
taceous Battle Camp formation, North Queens-
land. - Proc. R. Soc. Qd. 99: 7 7 - 8 1 .
Scoble, M . J . (1980): a New incurvariine leaf-miner
from South Africa, with comments on structure,
life-history, phylogeny, and the binominal system
of nomenclature (Lepidoptera: Incurvariidae). - J.
Ent. sth. Afr., 43 (1): 7 7 - 8 8 .
- (1983): A revised cladistic classification of the Nep-
ticulidae (Lepidoptera) with descriptions of new
taxa mainly from South Africa. - Trans. Mus.
Monog. no. 2, Pretoria, xi + 105 p.
Wagner, D. L. & Powell, J. A. (1988): A new Prodoxus
from Yucca boccata: first report of a leaf-mining
Prodoxine (Lepidoptera: Prodoxidae). - Ann. Ent.
Soc. Amer., 81: 547-553.
Wiegmann, . M. (1994): The earliest radiation of the
Lepidoptera: Evidence from 18S rDNA. - Ph. D.
thesis, Univ. Maryland, College Park, ix + 230 p.
Wilkinson, C. & Scoble, M . J . (1979): The Nepticuli-
dae (Lepidoptera) of Canada. - Ent. Soc. Can., Ot-
tawa, Mem. 107, p. 1 - 1 2 9 p.
Wille, J. (1926): Cecidoses eremita Curt, und ihre Galle
an Schinus dependens Ortega. Z. Morph. Okol.
Tiere, 7: 1 - 1 0 1 .
Zimmerman, E. C. (1978): Microlepidoptera. - Insects
of Hawaii, 9: i-xviii, 1 1903 (2 vols).
 7. The Tineoidea and Gracillarioidea
Donald R. Davis & Gaden S. Robinson
(Don R. Davis: non-tineid families; Gaden S. Robinson: Tineidae)
TINEOIDEA.
The Tineoidea are generally recognized to repre-
sent the most plesiomorphic group of ditrysian
Lepidoptera; until 1988 the superfamily included
the groups now assigned to the Gracillarioidea.
The principal autapomorphy of the Tineoidea is
the presence of a slender pair of ventral pseuda-
pophyses within AIO of most females (Robinson
1988; Davis 1988); the telescopic 'ovipositor'
itself is unusually elongated. Other characteris-
tics useful in distinguishing Tineoidea from
Gracillarioidea and most ditrysian Lepidoptera
are the presence of erect scales on the frons, la-
bial palpi with lateral bristles, and haustellum
with short, disassociated galeae (Dugdale 1988).
Head: Generally rough, with mostly erect pili-
form scales (Fig. 7.3 B). Ocelli usually absent,
present in a few Eriocottidae and Psychidae. An-
tennae usually less than length of forewing; scape
with pecten present or absent; flagellum usually
filiform, occasionally pectinate in $ Eriocottidae,
Acrolophidae, Psychidae, and $ Arrhenophani-
dae; usually 1 row of slender scales per segment,
rarely 2, either encircling segment or restricted
to dorsal half. Chaetosemata absent. Pilifers and
mandibles either present or absent. Haustellum
naked, usually reduced, loosely joined and un-
coiled. Maxillary palpi 5-segmented in many Tin-
eidae and in Crepidochares (Eriocottidae), 4-seg-
mented to absent in all other genera. Labial palpi
usually 3-segmented, with stout, erect, piliform
scales (bristles) from apical and lateral surfaces
of second segment (Fig. 7.3 B); bristles absent in
Arrhenophanidae and most Acrolophidae and
Psychidae. Thorax: Metafurca with apophyses
usually free, joined to secondary arms in Arrhen-
ophanidae and Psychidae. Forewings moderately
broad, index (W/L) 0.25-0.47, with Rs4 usually
terminating on costa or apex; Rs4 to termen in
Psychidae, and most Arrhenophanidae and Erio-
cottidae. Chorda with accompanying accessory
cell present or absent. Base of M usually present
and either simple or forked and forming interca-
lary cell. 1A + 2 A usually with basal fork. Spe-
cialized subanal interlocking microtrichia pre-
sent. Hindwing index 0.20.55; base of M usu-
ally preserved, either simple or forked; discal cell
closed. Abdomen: Tergal spines absent. A2 with-
out tergosternal connection. Tympanal organs
absent except on S2 of Harmocloninae (Tine-
idae). Anterior margin of S2 (Figs. 7.4 AE)
with a pair of slender apodemes which usually
continue caudally into S2 as sternal rods (Kyrki
1983); a minute pair of warts (Fig. 7.4 B) usually
near caudal end of rods; two pairs of small tu-
berculate plates present in pleura; ventral pair
typically near or on margin of S2; dorsal pair
near T2; plates often repeated but progressively
smaller on posterior segments. Male genitalia:
Valvae with elongate, internal apodemes; 7 pairs
of genital muscles present (Kuznetsov and Stek-
ol'nikov 1976, 1977). Female genitalia'. Oviposi-
tor often greatly extensible (Fig. 7.4 M), usually
with 2 elongate pairs of apophyses, sometimes
reduced to 1 short posterior pair (Acrolophidae,
Fig. 7.4 L, some Psychidae); a ventral pair of
slender pseudapophyses usually present within
AIO (Bradley 1951); $ of Eriocottidae with an
additional pair (rarely 2 pairs) of dorsal pseuda-
pophyses within A8 (Fig. 7.4 M, N); ostium
opening on S8.
Egg: Oval, flat type, with micropylar axis par-
allel or subparallel to oviposition substrate;
chorion relatively smooth to finely reticulated,
micropylar end finely reticulated. Eggs usually
deposited externally and singly, sometimes in
mass. Larva: Often living within tube or portable
case (Fig. 7.6 A - N ) ; all instars with fully devel-
oped legs and 5 pairs of prolegs; ventral crochets
usually in a uniserial circle, ellipse, or penellipse.
Pupa: Adecticous, incomplete, with appendages
variably appressed to body and with 3 - 6 mov-
able abdominal segments. Maxillary palpi usu-
ally present. Abdomen with tergal spines usually
in rows, sometimes in patches. Pupa extruded
partially from cocoon prior to adult eclosion.
Six families are here recognized within Ti-
neoidea. Family interrelationships were dis-
cussed by Robinson (1988) and Robinson &
Nielsen (1993), who tentatively presented them
as Tineidae + (Eriocottidae + (Acrolophidae +
(Psychidae + Arrhenophanidae))). The Lypusi-
dae have subsequently been removed from syn-
onymy with the Psychidae. Possible synapomor-
phies of the non-tineid families are the strong
tendency of the antennae to bipectination, the
reduction of the maxillary palp (which has at
most 3 segments except for some Eriocottidae),
forewing with Rs4 terminating on the termen
rather than on the costa, male retinaculum aris-
ing from wing membrane between C and Sc
rather than on Sc, female frenulum with more
than 4 bristles, apex of sacculus in male genitalia
with one or more thorn-like sensilla and spinose
membranous lobe ('pulvillus') on inner valve sur-
face. Possible autapomorphies of the acrolophid/
92
psychid/arrhenophanid assemblage (to which the
Lypusidae apparently do not belong) are the ab-
sence of an antennal pecten and the fusion of the
metafurcal apophyses with the secondary furcal
arms, forming a bridge-like structure; on the
other h a n d it is now k n o w n that distinct, albeit
short, galeae are indeed retained in the psychid
ground plan (Fig. 7.3 H). Separation of the Arr-
henophanidae f r o m the Psychidae has been ques-
tioned (Robinson 1988), but they are pheneti-
cally distinct (see account below) and the separa-
tion is here upheld.
Key to the tineoid families (Lypusidae not
included)
1 Forewings with Rs4 terminating on costa.
Antennae filiform. Maxillary palpi 5-seg-
mented in most species Tineidae
Forewings with Rs4 usually terminating at
termen or apex. Antennae often bipecti-
nate. Maxillary palpi usually 4-segmented
or less 2 M o u t h p a r t s (except galeae) fully developed in
2 Antennal pecten present. Metathoracic
furca free; furcal apophyses arising f r o m
lateral ridges of metafurcal stem (Fig.
7.4 H). 9 with 1 (fused) or 2 short, pseuda-
pophyses within dorsal wall of A8 (Figs.
7.4 M - N ) Eriocottidae
Antennal pecten usually absent. Dorsal
pseudapophyses absent within A8 of
Metathoracic furca usually joined as a
bridge to secondary furcal arms (Figs.
7.4 J, K), or with only tendon of vestigial
bridge arising f r o m secondary arms (Fig.
7.41) 3 Galeae short, disassociated, not strongly sclero-
3 abdomen without corethrogyne. Oviposi-
tor short, less than length of enlarged S8
(Fig. 7.4 L); anterior apophyses absent.
Metathoracic bridge vestigial, broken (Fig.
7.41). Larva with lateral pinacula partially
separated f r o m p r o n o t u m (Fig. 7.5 M) . .
Acrolophidae
9 abdomen with corethrogyne on A7. Ovi-
positor usually elongate, exceeding length
of more narrow S8; anterior apophyses
usually present (sometimes absent and ovi-
positor short in apterous ). Metathoracic
furca forming a bridge. Larva with lateral
pinacula either separated or fused with
pronotum 4
4 $ genitalia with non-retractile vesica and
slender, tubular anellus. $ often with bipec-
tinate antennae. Larva with lateral pina-
cula widely separated f r o m p r o n o t u m (Fig.
7.5 N ) Arrhenophanidae
$ genitalia with retractable vesica; anellus
usually m e m b r a n o u s or with separated lat-
eral lobes. 9 antennae filiform or vestigial.
Larva with lateral pinacula containing
spiracle and completely fused to p r o n o t u m
(Figs. 7.5 J, O) Psychidae
Donald R. Davis & Gaden S. Robinson
Tineidae. Very small to medium-sized moths,
forewings 2.5 to more than 25 m m long (Figs.
71. AL). Features for rapid recognition of
many species are: head vestiture of erect piliform
scales; labial palpi with stiff lateral bristles; pro-
boscis very short or even absent, never strongly
sclerotized, the galeae only loosely associated
(Fig. 7.2 A); hind tibiae with erect and elongate
scales on upper surface; adults resting with wings
raised tent-like over the body and with body par-
allel to substrate, moving with characteristic
scuttling run.
Head vestiture of erect piliform scales on occi-
put, vertex and frons; scales grouped into tufts,
forming more or less conspicuous whorls; head
with semi-appressed forward-directed scales in
(e. g.) Setomorpha, with appressed plate-like
scales in Hieroxestinae (Fig. 7.1 L). Head with-
out externally visible ocelli. C o m p o u n d eye un-
modified, with facets of regular size; interocular
index 0.50 (some Machaeropteris) to 2.20 ( M e t -
apherna).
many species (Fig. 7.2 A), but reduction fre-
quent. Pilifers large, or reduced; mandibles usu-
ally large, together often as wide as frons, but
absent in some groups. Maxillary palpi more
strongly developed than in any other ditrysian
group, but reduction occurring frequently, corre-
lated with reduction of the galeae. Fully devel-
oped maxillary palpi 5-segmented, folded; f o u r t h
and fifth segments with densely packed sensilla
trichodea interspersed with microtrichia, fifth
segment usually with conspicuous apical and
subapical processes bearing sensilla basiconica.
tized and, if rolled, rolled haphazardly; elongate
and apparently forming a functional proboscis in
only a very few genera. Labial palpi invariably
3-segmented, in some Hapsiferinae and Hier-
oxestinae distinctly recurved, drooping in some
Meessiinae; second segment in most genera with
u p to 15 coarse lateral bristles arranged in irreg-
ular line and with terminal whorl of u p to 15
bristles, usually with ventral brush of scales but
smooth-scaled in (e. g.) Opogona; terminal seg-
ment slightly to strongly flattened in m a n y taxa
and often swollen (e. g., Tiquadra), most genera
with conspicuous vom Rath's organ. Antenna 0.3
to 1.3 length of forewing; scape smooth-scaled in
most genera, usually stout but distinctly slender
and elongated in Erechthiinae and Hieroxesti-
nae; pecten present in m a n y genera, consisting of
two or three to about forty stout bristles; flagel-
lomeres cylindrical or approximately barrel-
shaped, conspicuously ciliated to finely pubes-
cent, females with fewer and shorter cilia than
males, flagellomeres in most genera completely
covered by one or, rarely, two annuii of overlap-
ping scales.
Thorax: Legs usually smooth-scaled except for
hind tibia which carries elongate suberect scales
The Tineoidea and Gracillarioidea 93

Fig. 7.1. Tineoidea. A - L , Tineidae: A, Morophaga bucephala, Japan (9.5 mm); B, Moerarchis anstralasiella, Aus-
tralia (17 mm); C, Nemapogon granella, Germany (6.8 mm); D, Tiquadra aeneonivella, Colombia (16 mm); E,
Gerontha captiosella, Sri Lanka (10.7 mm); F, Coryptilum klugii, Indonesia (17 mm); G, Monopis monachella,
Germany (7 mm); H, Trichophaga tapetzella, USA (9 mm); I, Ceratophaga vastella, South Africa (12.5 mm); J,
Tinea pellionella, Germany (6 mm); K, Tineola bisselliella, Germany (7.2 mm); L, Opogona nipponica, Japan
(6 mm); M - N , Eriocottidae: M, Eriocottis fuscanella, Germany (8 mm); N, Compsoctena thwaitesii, Sri Lanka
(15.5 mm); O - P , Acrolophidae; O, Amydria effrentella, USA (9 mm); P, Acrolophus popeanellus, USA (13 mm);
QR, Arrhenophanidae: Q, Arrhenophanes perspicilla, Venezuela (24 mm); R, Dysoptus lantalola, Venezuela
(6 mm); S - T Psychidae: S, Kearfottia albifasciella, USA (5 mm); T, Narycia monilifera, Switzerland (5 mm). (Fore-
wing lengths in parentheses.)
94
on upper surface; foretibial epiphysis present in
most genera; tibial spur pattern invariably 0-2-4.
Wings typically moderately broad (forewing
index 0.25-0.30, hindwing index 0.20-0.25),
rarely very narrow. Both wings generally sub-
ovate. Forewing apex produced and upturned in
some Erechthiinae; termen usually convex; with
a wide variety of forewing patterns, from uni-
form dirty brown of some Tinea (Figs. 7.1 J) to
striking metallic transverse strigulae of some
Moerarchis (Fig. 7.1 B); complicated patterns in
some Erechthias make use of particoloured
scales; pattern elements transverse, rarely longi-
tudinal; bright coloration may provide mimetic
resemblance, disruptive patterning, or possibly
aposematism. Marked sexual dimorphism rare,
e.g. Timaea (pattern), Gerontha (Fig. 7.1 E)
(shape), Setomorpha (venation). Hindwing un-
patterned. Scales of forewing often in rough or
raised patches, some taxa with specialised sex-
scales. Typically with full complement of free
forewing veins; Sc invariably present, but
strongly displaced anteriorly in Dryadaula and
Eudarcia; all branches of Rs terminating anterior
to or at apex, with rare exceptions; pterostigma
developed sporadically; R absent in Crypsithyris,
absent or vestigial in Opogona and Amphixystis;
absence of any other branch of R/Rs rare, but
Rs3 occasionally lost; stalking of Rs3 + 4 or
Rs2 + 3 frequent, Rsl always free; more compli-
cated stalking sequences occur, involving Rs2,
Rs3, Rs4 and, in some cases, M l ; all three M
branches usually present, but one may be absent;
stalking of Ml + 2, M2 + 3, and M3 + CuAl oc-
curs, also loss (or fusion) of either CuAl or
CuA2; CuP present in all taxa, but tubular only
proximally and distally; Al and A2 typically
forming a broad basal loop but fused for distal
two-thirds; A2 weak in many genera, rarely un-
developed; venational modification occurs in
some genera (e. g., Monopis) to accommodate a
hyaline spot. Hindwing venation typically com-
plete, all veins free; Se + R and Rs separate and
subparallel; Sc + R displaced anteriorly in Eud-
arcia; stalking of Ml + 2 common; M3 lost in
some genera; full complement of posteriormost
veins CuP, 1A + 2A and 3 A expressed only
in large species with broad hindwings; 3 A absent
in most groups with narrow hindwings. Female
retinaculum consisting of a few semi-erect scales
on Cu. Male retinaculum typically an elongately
triangular tongue with a rolled apex arising from
Se, but exhibiting considerable variation
broad and shallow lobe from Se or from mem-
brane between costa and Se, or triangular tongue
from spur from Se, or absent. Male frenulum a
single bristle, thickened and spatulate in a few
taxa; female frenulum typically consisting of two
or three bristles, but varying from one (e. g.,
Coryptilum) to about 15 (Scardia). Supplemen-
tary wing-coupling devices involve microtrichia
Donald R. Davis & Gaden S. Robinson
or specialised scaling on opposing wing surfaces
in Gerontha and Harmaclona.
Abdomen with S I vestigial; I a quadrangular
frame; without prespiracular bar connecting S II
and I; S II with pair of anterior apodemes but
usually lacking additional sclerotization (venu-
lae) along margins; S II with tympanal organs in
Harmaclona and Micrerethista, with hair-pencils
in male Comodica and Dryadaula. Male abdo-
men in scattered genera with various types of
scent-organs in various positions; many genera
with shallow coremata in pleural membrane of
eighth segment; terminal segmentation highly
modified in Setomorpha and Dryadaula. Func-
tional and evolutionary aspects of male genital
musculature are discussed by Kuznetsov & Ste-
kolnikov (1976, 1977), Robinson (1981) and Ste-
kolnikov & Kuznetsov (1995). Female abdomen
with terminal corethrogyne in some genera (e. g.,
Tinissa); corethrogyne invaginated into three in-
tersegmental pouches in Perissomasticinae. Male
genitalia of diverse form; vinculum and tegumen
not differentiated, saccus usually present, rarely
bilobed; uncus usually simple and bilobed, but
articulated and hook-like in Tineinae; gnathos
usually present, absent in Scardiinae; valval apo-
demes elongate, robust; aedoeagus usually sim-
ple and cylindrical, ductus and bulbus ejacula-
torius entering and connecting at base, thus with-
out coecum penis. Female with ovipositor typi-
cally elongate and telescopic with soft and setose
papillae anales, with ventral pair of sclerotized
rods set in membrane; armoured piercing ovipos-
itor developed in a few genera (e. g., Tiquadra);
eight tergum and sternum typically separate, ter-
gal margins extending anteriorly to articulate
with apophyses anteriores which invaginate from
anteriorly extended corners of sternum; ostium
bursae invariably within margins of eighth ster-
num; bursa copulatrix usually differentiated into
ductus and corpus bursae; ductus seminalis typi-
cally connected to ductus bursae close to ostium;
signa present in a few genera.
Egg of 'flat' type, with micropylar axis parallel
or subparallel to oviposition substrate; chorion
minutely wrinkled and in some taxa with coarser
ridging, either longitudinal ribbing or polygonal.
Larva with few specializations (Figs. 7.5 AB,
H, L), thorax not as strongly sclerotized as in
Psychidae, with only primary setae present; most
taxa with ecdysial lines of head extending be-
yond apex of frontoclypeus and terminating at
epicranial notch; frontoclypeus large, extending
almost to epicranial notch; with six or fewer
pairs of stemmata and three pairs of epipha-
ryngeal setae; hypostoma either narrowly sepa-
rated or contiguous. Thorax with prespiracular
L sclerite typically separated from pronotum
which is usually distinct; prothoracic L-group
trisetose (bisetose in Scardiinae); three pairs of
thoracic legs present, with precoxal plates rarely
fused; four pairs of ventral prolegs and one pair
The Tineoidea and Gracillarioidea
of anal prolegs; crochets invariably present on
prolegs, arranged in uniserial circle or ellipse
(mesal penellipse in Oinophila); D1 and D2 pre-
sent on all segments with D2 ventral; XD1, XD2
and SD forming an almost straight vertical line;
SV group bisetose but on second and third seg-
ments it is unisetose in all groups except Tinei-
nae. Abdomen with D1 and D2 developed on
all segments, D2 caudal to DI; SD setae on all
segments but SD2 absent from A9; L group tri-
setose; SV group trisetose to A6, bisetose on re-
maining segments (A9 unisetose in Tineinae);
anal plate with four pairs of setae; dermal cuticle
with numerous fine microtrichia. Larvae of the
termite commensal Acridotarsa are exceptional
(see Harris, 1968).
Pupa adecticous, obtect; at least two pairs of
coxae visible; first and second abdominal seg-
ments fused in both sexes; 7th, 8th and 9th seg-
ments fused in females, 8th and 9th in males.
Typically with anterior band of posteriorly-di-
rected spines on segments 39, additional pos-
terior band of spines on segments 36 in females
and 3 7 in males; this pattern may be consider-
ably modified. Ventral abdominal surface with
broad fields of posteriorly-directed microtrichia.
Biology. Instead of occupying the typical di-
trysian leaf-feeding role, practically all tineid lar-
vae are fungivorous in the sense that their food
comprises a substantial proportion of fungal tis-
sue. Tineid larvae have traditionally been catego-
rised as detritophagous, lichenivorous or fungiv-
orous. Detritophagy involves tineids in a wide
range of niches in a wide range of biological sys-
tems with habits ranging from the mundane to
the bizarre, from feeding on fallen leaves to feed-
ing on guano in bat-caves, on mammal corpses,
on insect remains in or below spiders' webs, and
on feathers in birds' nests. Some detritophagous
Tineidae are commensals, sharing food resources
with social Hymenoptera, Isoptera, and with
mammals including man. Synanthropic species
include pests of stored food products and of fur-
nishings and fabrics made from wool, fur or
feathers. There is strong correlation between the
higher taxonomic category to which a tineid be-
longs and its larval food.
While many tineid larvae build portable cases
from which they feed (Figs. 7.6 AC), in a man-
ner analogous to that of Psychidae, case-building
is by no means consistent within genera or even
within species, and the definition of 'case' is
blurred by the range of practices observed
among Tineidae. The simplest structure is no
more than a tubular cocoon constructed at the
beginning rather than at the end of larval life.
The tube may be attached to the substrate or
form a tunnel through it. Trichophaga swinhoei
feeds on weathered carnivore faeces in the Horn
of Africa and builds a 4 cm vertical tube within
which it lives; the tube may provide not only
protection but a means of escaping extreme tem-
95
peratures close to the soil surface. Long tubes
that ramify through the lichen host are built by
larvae of Infurcitinea. Short, fixed tubes are built
as shelters for ecdysis by certain Monopis and
Tinea species. Fully portable tubular cases of
varying degrees of sophistication are built by
many Tineinae and Meessiinae.
Pre-pupation behaviour involves firstly move-
ment to the pupation site. Portable cases are sus-
pended well away from the food substrate prior
to pupation. Case- or tube-dwellers pupate in the
case or tube; free-living species spin a simple co-
coon; case- or tube-dwelling species then clean
out their refuge and seal it with a few strands of
silk. Internal feeders partially cut out an opercu-
lum in the surface of the fungus or wood in
which they have fed and then spin a cocoon close
to this exit.
Eclosion involves considerable movement by
the pharate adult; the pupa is wriggled and its
abdomen contracted and expanded to bring the
spines of the abdominal segments to bear against
the walls of the case or cocoon. Once the cocoon
is ruptured, the pupa is protruded clear for about
half its length and the adult emerges.
Some Tineidae have been recorded as forming
mating swarms, notably Nemapogon and some
Scardiinae; at least one Monopis needs several
cubic metres of free space for successful court-
ship. Gyratory dance has been recorded in sev-
eral Tineidae, but this may not have a courtship
function. Female pheromones of Tineidae have
been little studied but responses have been re-
corded to octadecenyl acetate derivatives, nota-
bly E3Z13 18Ac; two pheromones have been
isolated from Tineola, E2-18A1 and E2E13-
18A1.
Oviposition may involve insertion of eggs into
crevices (by species with soft ovipositors) or pen-
etration of host tissue (by species with piercing
ovipositors); eggs may be protected with fine
hair or scales from the corethrogyne or abdomi-
nal apex glued to the pool of mucilage surround-
ing each egg.
Larviparity is known in numerous Andean
Tinea and Indo-Australian Monopis species; fe-
cundity is traded off against egg mortality. Mo-
nopis with conventional reproductive systems
carry 150200 mature and immature ova
whereas larviparous species produce 4060
fully-developed first-instar larvae which are ma-
tured in the grossly enlarged common oviduct.
Classification and information sources. The
family Tineidae encompasses some 3000 de-
scribed species in 320 genera. The group may not
be monophyletic and represent a grade rather
than a clade (Robinson 1988; Robinson & Niel-
sen, 1993). Fifteen subfamilies are currently rec-
ognised. Of these, 13 are probably monophyletic.
Relationships between subfamilies are still ob-
scure. Some 150 genera have yet to be attributed
to subfamilies, but many of these may prove to
96 Donald R. Davis & Gaden S. Robinson

Fig. 7.2. A - G , Psychidae: A, Pseudarbela semperi, Philippines (12 mm, .,)] , P. semperi, Phillipines (20 mm, 9);
C, Dahlica triquetrella, Switzerland (12.3 mm); D, Clania peyeri, China (18 mm); E, Oiketicus kirbyi, Venezuela
(16 mm, ), F, O. kirbyi, Ecuador (28 mm long, 9); G, Thyridopteryx ephemer aefor mis, USA (10.7 mm); HI,
Roeslerstammiidae: H, Roeslerstammia erxlebella, Germany (6.3 mm); I, Agriothera issikii, Taiwan (12 mm); J,
Douglasiidae, Tinagma perdicillum, Russia (5 mm); KM, Bucculatricidae: K, Leacoedemia ingens, South Africa
(6 mm); L, Bucculatrix magnella, USA (6.1 mm); M, B. tridenticola, USA (3.7 mm); NS, Gracillariidae: N,
Caloptilia azaleella, USA (4.2 mm); O, C. syringella, Germany (5.7 mm); P, Acrocercops astaurota, Japan (5 mm);
Q, A. brongniardella, Germany (4.6 mm); R, Macarostola pontificalis, Polynesia (4.8 mm); S, Phyllonorycter blanc-
ardella, Germany (3.8 mm); T, Phyllocnistispopuliella, USA (2.8 mm). (Forewing lengths in parentheses except for
body length in Fig. F.)
The Tineoidea and Gracillarioidea
be synonyms of genera whose systematic posi-
tion is already known. For a broad and detailed
review of the classification and biology of Tine-
idae, see Robinson & Nielsen (1993).
Present knowledge of Tineidae on a worldwide
scale is patchy, and most recent work has been
restricted to particular geographical regions.
There are checklists for the Nearctic (Davis
1983), Neotropical (Davis 1984), Oriental (Rob-
inson & Tuck, 1996) and Australian (Robinson &
Nielsen 1993) regions; the checklist of New
Zealand Tineidae (Dugdale 1988) is cross-refer-
enced to the colour illustrations of Hudson
(1928; 1939). The last monographic treatment of
the Nearctic Tineidae was the unillustrated work
of Forbes (1923); this, in combination with the
earlier monograph by Dietz (1905), is neverthe-
less useful. More than 100 papers on the Tine-
idae of the Palaearctic region have been pub-
lished since 1955 by A. K. Zagulajev, G. Petersen
and R. Gaedike (see Zoological Record, passim).
The contents of very many of these papers were
summarized by Zagulajev (1960; 1964; 1973;
1975; 1979). Afrotropical Tineidae have been
treated by Gozmny & Vri (1973); this work
should be used in conjunction with that of Janse
(1968). For the Oriental region, figures of adults
of some Indian species with descriptions of life
histories were given by Fletcher (1921; 1933); the
Oriental Nemapogoninae, predominantly Indian
species, were reviewed by Robinson (1980 a).
Robinson & Nielsen (1993) provide a detailed
treatment of the Australian Tineidae. Clarke
(1971; 1986) and Zimmerman (1978) dealt with
the Tineidae of the islands of the eastern Pacific;
the latter work contains detailed reviews of bio-
logical information. Robinson (1979) revised the
case-bearing clothes-moths, and the cave-dwell-
ing species of Tineidae (Robinson, 1980 b). The
same author reviewed the fungivorous group
Scardiinae in a series of papers (Robinson 1976;
1981; 1986). Robinson & Tuck (1997) have re-
viewed the Hieroxestinae. Hinton (1956) gave
detailed descriptions of larvae of a number of
species. G. Chauvin and collaborators (1960 to
date) have published numerous papers on tineid
physiology (see Zoological Record, passim),
other recent works are reviewed by Robinson &
Nielsen (1993).
Myrmecozelinae (= Haplotineinae, Phthoro-
poeinae, Rhodobatinae). Autapomorphies: none.
The group was considered by Robinson & Niel-
sen (1993) to be polyphyletic but might contain
a monophyletic core defined by: male genitalia
with aedoeagus strongly curved and basally
boot-shaped; anterior margin of vunculum re-
curved dorsad.
Constituent species are generally large and ro-
bust and cannot be included in any other sub-
family.
36 genera (Myrmecozela, Cephimallota, Rho-
dobates etc.) with 180 species. Widely distributed
97
in the Old World; some New World groups may
also be attributable to the monophyletic 'core' of
this subfamily. Larvae feeding externally, or
tube-dwelling, on plant detritus.
Harmacloninae. This group which was in-
cluded in the myrmecozeline assemblage by Rob-
inson & Nielsen (1993) has been given subfamily
status by Davis (in press a). Autapomorphies:
Unique wing coupling system consisting of
curled, rigid scales along ventral hindmargin of
forewing interlocking into straight, rigid scales
along dorsal subcostal area of hindwing; pretar-
sus lacking arolium and pseudempodial seta;
paired tympanic organs present on abdominal
sternum II; enlargement of sternal apophyses on
abdominal segment II; and aedoeagus with a
basal, midventral keel.
2 genera (Harmaclona, Micrerethista) and 22
species. Pantropical distribution, most diverse in
the Old World, especially southeast Asia and
Australia. Biology poorly known; a single rear-
ing record reports larvae boring in logs of Bu-
chanania latifolia (Anacardiaceae; Fletcher
1933). The Harmacloninae are the only group of
microlepidopterans (in a restricted sense, exclud-
ing cossuids and pyraloids) known to have devel-
oped tympanic organs (discussion and illustra-
tion in 211).
Meessiinae. Autapomorphies: none. The group
was considered by Nielsen & Robinson (1993)
unlikely to be monophyletic.
Constituent species are small, often narrow-
winged; antennal pecten frequently lacking; fore-
wing pattern simple, often consisting of large
spots or blotches, or transverse bands. Most spe-
cies with reduced venation; forewing with Rs
weak and with loss or stalking of Rs and M
branches, the cell narrow and open; hindwing
with Rs basally weak and one M branch lost, cell
narrow and open, anal venation reduced and 3 A
rarely present. Male genitalia of many species
with uncus and gnathos reduced or absent.
Larva lichenivorous.
23 genera (Homosetia, Baeophylla, etc.) with
200 species; a further 2 0 - 2 5 genera, most mono-
basic, have been doubtfully attributed to this
subfamily. Worldwide distribution but diversity
dominated by the Palaearctic species of Eudarcia
(= Meessia) and Infurcitinea. Larvae feeding ex-
ternally or within tube galleries on lichen.
Dryadaulinae (= Archimeessiinae). Autapo-
morphies: transfrontal suture of head reflexed
dorsally, inverted V-shaped; labial palpus
broadly spatulate, lacking terminal bristles; fore-
wing venation with Sc displaced anteriorly, R
arising close to base, A2 weak or absent; hind-
wing with M3 or CuA2 absent, 3 A absent; male
retinaculum a broad and very shallow lobe aris-
ing from Sc; female frenulum a single bristle;
male frenular bristle sharply angled; abdominal
terga I and II entirely fused; male with terminal
abdominal segmentation strongly reduced and
modified, frequently asymmetrical, eighth seg-
98 Donald R. Davis & Gaden S. Robinson

Fig. 7.3. Tineoidea, Head morphology and wing venation: A, Tineidae: Tinea pellionella; C D, Eriocottidae:
C, Crepidochares neblinae, Oc = ocellus; D, Compsoctena thwaitesii, maxilla; E, Acrolophidae, Acrolophus po-
peanellus, FG, Arrhenophanidae: Arrhenophanus perspicilla; G, Maxilla; HI, Psychidae: H, Kearfottia albi-
fasciella\ I, Thyridopteryx ephemeraeformis, caudal view, Mx = maxilla, L = labium; J, Tineidae, Scardia anato-
mella; K, Eriocottidae, Crepidochares neblinae; L, Acrolophidae, Acrolophus popeanellus; M, Arrhenophanidae,
Cnissostages oleagina; NO, Psychidae: N, Thyridopteryx ephemeraeformis', O, Kearfottia albifasciella. (All
scales = 0.5.)
The Tineoidea and Gracillarioidea
ment incorporated into genitalia which are
strongly modified and asymmetrical, gnathos ab-
sent; female with ovipositor strongly attenuated;
apophyses anteriores rudimentary or absent.
One genus {Dryadaula) with 35 species.
Worldwide distribution but speciose only in Neo-
tropical region and Australia. Larva feeding ex-
ternally on lichen or fungus.
Scardiinae (= Tinissinae). Autapomorphies:
male genitalia invariably lacking gnathos; larva
with only two prothoracic L-group setae; D1 se-
tae more widely separated than D2 setae on Al
to A8; L3 absent from A9; mandible with small
ventral tooth and six or more distal teeth.
24 genera (Scardia, Morophaga, Tinissa etc.)
with 112 species (Fig. 7.1 A). Worldwide distribu-
tion, most speciose in Indo-Australian region but
with greatest generic diversity in Neotropics.
Larva boring in bracket-fungi or in bark or
wood permeated by fungal mycelia.
Nemapogoninae. Autapomorphies: male geni-
talia with gnathos transverse and appressed to
diaphragma with membranous connection for
most of its length; valva with costa rolled inward
and invariably cleft longitudinally, dorsal lobe
often a flexible, hirsute digitate process; valvae
fused or articulated at ventral margins; juxta
fused with inner surfaces of valvae and, with
anellus, forming complex support for aedoeagus;
female genitalia with ostium bursae in transverse
fold of S VIII; ventral lip of ostium with elongate
setae; ductus bursae narrow, ornamented with
microtrichia and with thorny internal ring close
to junction with corpus bursae.
7 genera (Nemapogon, Nemaxera, Triaxomera,
etc.) with 82 species (Fig. 7.1 C). Distribution
primarily Holarctic but with one monobasic
Australian genus (Vanna). Larva boring in
bracket-fungi or in bark or wood permeated by
fungal mycelia.
Tineinae. Autapomorphies: male genitalia with
uncus lobes fused and forming an articulated
hook; pupa with only anterior bands of abdomi-
nal spines; larva with bisetose SV-group on
meso- and metathorax, stemmata reduced to a
pair or none.
36 genera (Tinea, Tineola, Trichophaga, Mo-
nopis, Proterospastis etc.) with 320 species (Figs.
7.1 G - K ) . Worldwide distribution but poorly
represented in the Neotropical region. Larvae
feeding externally, or burrowing in loose sub-
strates, on keratin and chitin (wool, feathers, ar-
thropod remains, guano, etc.), often in the pres-
ence of fungal hyphae.
Setomorphinae. Autapomorphies: head with
lamellate semi-appressed scales; head sutures
fused; male eighth sternite a narrow, reflexed, al-
most complete ring; male genitalia with dia-
phragma microtrichiate; female with eighth seg-
ment membranous and lacking rami from the
apophyses anteriores; larva with unidentate
99
mandibles, prolegs with band of minute recurved
spines above crochets on A3 to A6.
Maxillary palpi and galeae are fully developed
in Prosetomorpha (Davis, 1996) or vestigial in
Lindera and Setomorpha (Robinson & Nielsen
1993); the larva has six pairs of stemmata in
Prosetomorpha (Davis, 1996) but none in Lindera
and Setomorpha.
3 genera (Prosetomorpha, Setomorpha and
Lindera) with 9 species. Two cosmopolitan pest
species with worldwide distributions, remaining
taxa Neotropical. Larvae detritophagous, feed-
ing externally or burrowing in loose substrates
or (Prosetomorpha) inhabiting borings in plant
stems made by Coleoptera, on plant or arthro-
pod remains.
Perissomasticinae. Autapomorphies: forewings
usually glossy and unicolorous; labial palpi
short; male genitalia with gnathos absent,
aedoeagus articulating with vinculum, bulbus
ejaculatorius hypertrophied; female with cor-
ethrogyne invariably present on intersegmental
membrane between A7 and A8, invaginated into
one dorsal and a pair of lateroventral pouches.
16 genera (Perissomastix, Edosa, Neoepiscar-
dia etc.) with 200 species. Distribution restricted
to the Old World tropics and subtropics, with
greatest diversity in the Afrotropical region. Bio-
logy unknown.
Hapsiferinae. Autapomorphies: head vestiture
consisting of lamellate scales appressed more or
less loosely to the head; epicranial suture absent,
scale-bases of vertex forming an unbroken field;
second segment of labial palpus with sparse lat-
eral bristles and no terminal whorl; male genita-
lia with valvae fused medially at base, juxta
forming a bridge between faces of valvae,
transtilla present (most taxa), coremata invagi-
nated into ventral intersegmental membrane
(most taxa); in female eighth segment and ovi-
positor elongated, corethrogyne invaginated into
pair of ventral pouches.
11 genera (Hapsifera, Tiquadra, Scalidomia
etc.) with 103 species (Fig. 7.1 D). Worldwide
distribution but majority of genera and species
are Afrotropical. Larvae feeding externally or
burrowing in soft substrates, on vegetable detri-
tus including sawdust used for cultivating mush-
rooms; also recorded from mammal burrows.
Hieroxestinae (= Oinophilidae). Autapomor-
phies: head wedge-shaped in lateral view; head
vestiture consisting of lamellate scales closely ap-
pressed to head, posterior scales of vertex and
scales of occiput directed caudally; scale-bases
on frons in regular transverse lines; antennal
scape elongated; antennal pecten absent; labial
palpi strongly divergent; scale bases on abdomi-
nal sternites in transverse lines; female with sin-
gle bobsled-shaped signum.
6 genera (Opogona, Oinophila, Amphixystis,
Phaeoses etc.) with 272 species (Fig. 7.1 L) (Rob-
inson & Tuck 1997). Pan tropical distribution, de-
100
pauperate in the New World. Larvae feeding ex-
ternally or burrowing in soft or loose substrates,
on dead or decaying plant material, or grazing
fungi in termite mounds or birds' nests; Asym-
plecta are reported to be leaf-miners; Wegneria
are reported to feed on guano.
Erechthiinae. Autapomorphies: male genitalia
with juxta forming a deep pouch that displaces
aedoeagus dorsad; male retinaculum distinctly
broadened at base and comparatively shallow;
ovipositor lacking the ventral rods present in al-
most all other Tineidae; larva with PI and P2 on
head much closer than in other Tineidae, labrum
with only three pairs of marginal setae, abdomi-
nal D2 displaced ventrad and roughly level with
SD-group (larval synapomorphies require con-
firmation). Many species have the forewing apex
upturned and/or one branch of R and one of M
missing in the forewing.
Two genera (Erechthias, Comodica) with 140
species. Indo-Pacific distribution, most speciose
in Australasia and the Pacific; isolated species on
St. Paul's Rocks and Ascension I. Larvae feeding
externally on plant detritus or, rarely, on living
leaf tissue of tough-leaved plants such as palms
and Pandanus; some larvae tunnel in soft sub-
strates including galls on Acacia.
Siloscinae. Autapomorphies: forewing with
erect spicular scales; male genitalia with articu-
lated saccus.
3 genera (Silosca, Organodesma, Autoch-
thonus) with 12 species. Afrotropical distribu-
tion. Biology unknown.
Euplocaminae. Autapomorphies: male antenna
bipectinate, pectinations elongate and strongly
ciliate; labial palpus lacking lateral bristles.
One genus (Euplocamus) with 11 species. Pa-
laearctic distribution including the Himalayas.
Larvae boring in rotten wood and bracket fungi.
Stathmopolitinae. Autapomorphies: female
eighth abdominal segment asymmetrical, with
process on left side.
One genus (Stathmopolitis) with 1 species. Ca-
nary Is. Biology unknown.
Teichobiinae. Autapomorphies: head with semi-
appressed forward-directed lamellate scales, frons
smooth-scaled; mouthparts reduced; antenna at-
tenuated; male genitalia lacking gnathos, juxta
fused with inner faces of valvae; ovipositor of
female with papillae anales sclerotized and
pointed to form piercing apex.
One genus (Psychoides) with three species. Pa-
laearctic distribution. Larva leaf-mining in early
instars then feeding externally on fern sporangia.
Eriocottidae (Figs. 7.1 MN). Small to medium
size moths, with forewings ranging from 4.5 to
23 mm in length. Head (Figs. 7.3 CD): Vesti-
ture rough consisting of mostly long piliform
scales with simple, acute apices. Ocelli present or
absent. Eye small to moderately large; interocu-
lar index 0.61.1; cornea with sparse scattering
Donald R. Davis & Gaden S. Robinson
of interfacetal microsetae. Antenna 0.4-0.7 the
length of forewing; pecten present; flagellum
usually filiform, bipectinate in S Compsoctena,
with one row of slender bidentate scales per seg-
ment, either restricted to dorsal surface or com-
pletely encircling segment, more concentrated in
Compsoctena. Pilifers variable, well developed to
absent. Mandibles vestigial to absent. Haustel-
lum vestigial (Fig. 7.3 D) to elongate. Maxillary
palpi 2-3-segmented and extremely short or elon-
gate and 4-5-segmented. Labial palpi moderately
long, 3-segmented, either upturned or porrect.
Thorax: Metafurca (Fig. 7.4 H) with apophyses
free. Wings moderately broad (Fig. 7.3 k) fore-
wing index 0.23-0.4; microtrichia sparsely scat-
tered over all wing surfaces; of Deuterotinea
brachypterous (Zagulayev 1988); all veins arising
separate from cell; Rs4 terminating either on
costa or termen; forewing with both accessory
and intercalary cells preserved; $ retinaculum
consisting of a broad fold arising ventrally from
costal margin and partially extending over base
of Sc: 1A + 2 A with basal fork. Hindwings
about as broad as forewings; hindwing index
0.32-4; $ frenulum a single stout seta; with
24 smaller frenular setae. Legs with tibial spur
pattern of 0-2-4; epiphysis relatively smooth
(without pecten) in Compsoctena. Abdomen:
Paired sternal apodemes of A2 slender, continu-
ing half way into sternum as sternal rods (Fig.
7.4 B). Coremata and corethrogyne absent. Male
genitalia. Uncus moderately bilobed, fused to
hoodlike tegumen. Vinculum usually short and
V-shaped, rarely Y-shaped (with an elongate sac-
cus). Gnathos slender, U- to V-shaped or re-
duced and widely separated. Subscaphium mod-
erately sclerotized. Transtilla present. Juxta usu-
ally indistinct, slender and weakly sclerotized in
Eriocottis. Valvae typically with a prominent spi-
nose lobe (clasper) from medioventral margin, or
with a more basal, variably developed process
from sacculus (in Crepidochares). Aedoeagus a
simple, slender cylinder without cornuti. Female
genitalia: Segments 8 and 9 + 1 0 forming a slen-
der, elongate, telescoping ovipositor (Fig. 7.4 M)
supported by typically 2 pairs of elongate apo-
physes and 2 (rarely 3) pairs of pseudapophyses
(Davis 1990 a); anterior dorsal pseudapophyses
located entirely within A8 and usually fused for
most of their length (Fig. 7.4 N). Ostium situated
at caudal margin of membranous S8. Ductus
bursae elongate, membranous. Corpus bursae
membranous; signum a single, irregular scleroti-
zation, or absent. Morphology of immature
stages unknown. Some larvae known to con-
struct underground silken tubes (Zagulayev
1988) or to bore into decayed wood, leaves, or
stems (Dierl 1970). Two subfamilies are recog-
nized (Nielsen 1978, Davis 1990 a).
Eriocottinae. Ocelli typically present, rarely re-
duced or absent. Antennae filiform in both sexes.
Haustellum moderate to elongate. Maxillary
The Tineoidea and Gracillarioidea 101

Fig. 7.4. Tineoidea, thoracic and abdominal morphology. AE, Abdominal sterna li: A, Nemapogon granella (L.);
B, Eriocottidae, Eriocottis fuscanella, T P = tuberculate plates, W P = wart-like protuberance; C, Acrolophidae,
Acrolophus popeanellus; D, Arrhenophanidae, Arrhenophanes perspicilla, E, Psychidae, Kearfottia albifasciella,
FK, Metathoracic furcasterna: F, Tineidae, Eudarcia simulatricella; G, Scardia anatomella; H, Eriocottidae,
Eriocottis fuscanella, I, Acrolophidae, Acrolophus popeanellus, P D T = posterior dorsal tendons; J, Arrhenophani-
dae, Arrhenophanes perspicilla, K, Psychidae, Kearfottia albifasciella, LN, 9 genitalia: L, Acrolophidae, Acrolo-
phus pholeter\ , Eriocottidae, Crepidochares neblinae, AAp = anterior apophyses, DPs = dorsal pseudapo-
physes, PAp = posterior apophyses, VPs = ventral pseudapophyses; N, Detail of dorsal pseudapophyses of A8.
(All scales = 0.5 mm.)
102
palpi 3-5-segmented. Forewing with Rs4 termi-
nating usually on costa, or termen (in Deutero-
tinea). Male genitalia with V- to U-shaped
gnathos. Mostly Old World from southern Pale-
arctic or Spain and North Africa (Eriocottis,
10 sp., Deuterotinea, 5 sp.) to Australia (Eucryp-
togona, 1 sp.), with a single genus in South
America (Crepidochares, 5 sp.).
Compsocteninae. Ocelli absent. Antennae pec-
tinate in $, filiform to serrate in $. Haustellum
minute (Fig. 7.3 D). Maxillary palpi 2-3-seg-
mented. Forewings with Rs4 terminating on ter-
men. Male genitalia with gnathos reduced,
widely separated. A single genus ranging from
southern Africa to Taiwan (Compsoctena, ca.
50 sp.).
Lypusidae. Monobasic. The Palaearctic Lypusa
maurella is a unicolorous dark moth, forewing
length some 67 mm; sexes overall similar. Head
dorsally with appressed lamellar scales; antennae
simple, scapus and pedicellus with scale tufts;
proboscis and maxillary palpi rudimentary; la-
bial palpi very short. Forewing Rs4 to termen. In
male genitalia vinculum U-shaped; uncus lobes
slender; prominent, setose 'peniculus' arising
from tegumen/valve articulation area. Female in
A8 with dorsal strengthening ridges confluent
with base of anterior apophyses. Dierl (1996)
gives additional details on adult structure.
The affinities of Lypusa are unresolved. It was
previously associated with the Psychidae, either
as a distinct subfamily (Torre and Strand 1929)
or family (Tutt 1900), but this relationship was
disputed by Sauter and Httenschwiler (1991).
Indeed it is clearly not a psychid as evidenced by
the free arms of the adult metafurca, or even a
tineid (Sauter 1982) because of its unusual larval
case construction. The latter is formed by folding
a leaf. Lypusa was assigned to a family of its own
by Heinemann (1870), and very recently again
by Dierl (1996). A further assessment of the rela-
tionships of the taxon is needed following a thor-
ough study of the immature stages.
Acrolophidae (Figs. 7.1 OP): Small to medium
size moths, with forewings 4 to 26 mm in length.
Head (Fig. 7.3 E): Vestiture usually rough with
dense tufts of piliform scales directed upward
from frons and forward from occiput, to rela-
tively smooth with slender scales possessing
rounded apices. Ocelli absent. Eyes small to
moderately large; interocular index 0.65-1.1;
cornea with sparse covering of interfacetal
microsetae to densely setose. Antennae 0.250.7
the length of forewing; scape smooth, pecten ab-
sent; flagellum filiform to bipectinate in , fili-
form in all $; a single row of slender bidentate
scales per segment. Pilifers and mandibles ab-
sent. Haustellum a minute, setose lobe (Fig.
7.3 E) usually shorter than first maxillary palpal
segment, or absent. Maxillary palpi greatly re-
Donald R. Davis & Gaden S. Robinson
duced, 3-segmented, or absent. Labial palpi mod-
erate to extremely long, curving over head in
some S Acrolophus, 3-segmented; lateral bristles
present or absent. Thorax: Metafurca (Fig. 7.41)
with furcal apophyses short, free; furcal bridge
vestigial with paired tendons arising caudally
from secondary arms. Forewings moderately
broad (Fig. 7.3 L); index 0.29-0.39, with veins
arising separate from cell; Rs4 terminating usu-
ally at apex; forewing with both accessory and
intercalary cells usually preserved; 1 A + 2 A
with moderately short basal fork; retinaculum
consisting of a broad, triangular lobe arising
from underside of costal membrane immediately
anterior to and curling over base of Sc. Hind-
wing index 0.3-0.4; $ frenulum a single stout
seta; with 2 - 7 frenular setae. Legs with tibial
spur pattern of 0-2-4; epiphysis rarely reduced.
Abdomen: Paired sternal apodemes of A2 (Fig.
7.4 C) relatively short with broad bases; sternal
rods absent. Coremata and corethrogyne absent.
Male genitalia: Uncus either acute or bilobed.
Vinculum U- to V-shaped, a slender to broad
ventral ring. Gnathos either fused (V-shaped) or
divided. Transtilla absent. Juxta usually absent,
sometimes present in Acrolophus. Valvae vari-
able, typically elongate and moderately slender,
with elongate costal apophysis at base; basal part
of sacculus fused medially to opposite valva. Ae-
doeagus variable, usually with one or more cor-
nuti. Female genitalia: Segments 9 + 1 0 ex-
tremely short with posterior apophyses ca. equal
in length to A 9 + 10 (Fig. 7.4L); sternum 8
more elongate, much broader and almost com-
pletely sclerotized with ostium opening within
sternum or on caudal margin; ostium usually
flush, sometimes projected into a sclerotized
cone. Anterior apophyses absent (completely
fused within eighth sternum). Ductus bursae usu-
ally short and membranous, rarely elongate and
sclerotized. Corpus bursae usually a single mem-
branous sac, sometimes partially divided and
with numerous spicules.
Egg: Poorly known; white and smooth in
Amydria effrentella (Johnson and Martin 1969),
averaging 0.42 mm long and 0.32 mm wide.
Larva (Figs. 7.5 C - G ) : Body small to large, to
more than 50 mm in length, whitish with dark
suffusion sometimes dorsally and brownish to
black head, notai plates, and pinacula. Fron-
toclypeus broad, extending ca. half the distance
to epicranial notch (Fig. 7.5 E); ecdysial line ter-
minating at epicranial notch. Usually 6 pairs of
stemmata, rarely reduced to 3 pairs. Labrum
with three pairs of epipharygeal setae (Fig.
7.5 G). Hypostoma variable (Figs. 7.5 CD),
broadly joined at midventer in Acrolophus. Pre-
spiracular plate partially separated from preno-
tai plate (Fig. 7.5 M). Coxae well separated.
Crochets arranged in a uniserial ellipse with sides
of prolegs densely covered with small scattered
The Tineoidea and Gracillarioidea 103

Fig. 7.5. Tineoidea, larval morphology. AB, Tineidae, Nemapogon granella: A, head (0.2 mm); B, hypostoma,
ventral (0.5 mm); C - G , Acrolophidae: C, Amydria effrentella, hypostoma, ventral (0.5 mm); D G, Acrolophus
arcanellus\ D, head, ventral, h = hypostoma (1 mm); E, head, dorsal; F, larva, lateral (0.5 mm); G, labrum, ventral
(0.2 mm); H, Tineidae, Nemapogon granella, labrum, ventral; I , Psychidae, Thyridopteryx ephemeraeformis: I,
labrum, ventral, with 4 pairs of epipharyngeal setae, ES = epipharyngeal sclerite; J, larva, lateral (0.5 mm); K,
Head (1 mm); L - O , prothorax: L, Tineidae, Nemapogon granella', M, Acrolophidae, Acrolophus pholeter; N,
Arrhenophanidae, Arrhenophanes perspicilla; O, Psychidae, Kearfottia sp. (Scale lengths in parentheses.)

spines; anal crochets in a semi-ellipse, opened
caudally. Larvae typically constructing long
silken tubes (Fig. 7.6 D) underground or on/in
plant detritus and feeding primarily on plant de-
bris as well as living plants, rarely coprophagous
or mycetophagous (Hasbrouck 1964; Davis
1984; Davis et al. 1986; Davis and Milstrey 1988;
Davis and Hogue 1992). Pupa: Head smooth,
without a frontal ridge. Wings relatively short,
seldom surpassing A4. Dorsum of A38 with a
single, anterior row of dense, ridgelike spines.
Segments A3 7 movable in $ and . Cremaster
reduced, variable, sometimes with small, paired
dorsal and ventral spines or with a dorsal furcate
ridge. Pupation usually in coarse cocoon or in
larval tube.
The family is exclusively New World and pri-
marily Neotropical with ca. 80 species occuring
in North America north of Mexico. The largest
genus is Acrolophus (245 sp.) followed by Amy-
dria (17 sp.), Ptilopsaltis (4 sp.), Exoncotis (3 sp.)
and Drastea (1 sp.). The most diagnostic features
of the family include the incomplete metafurcal
bridge and loss of anterior apophyses in the
adult $, and the partially separated prespiracular
plate in the larva (Robinson 1988, Davis 1990).
Adult Acrolophus have been examined from mid-
Tertiary Dominican amber.
104
Arrhenophanidae (Figs. 7.1 QR): Small to mod-
erately large moths, with forewings 4 to 33 mm
in length. Head (Fig. 7.3 FG): Vestiture gen-
erally rough, with slender scales over vertex
longer and more erect than much shorter scales
over frons. Ocelli absent. Eyes large, interocular
index ca. 1.5. Cornea naked. Antenna 0.30.6
the length of forewing; scape smooth to rough;
pecten absent; flagellum piliform to serrate in $,
serrate to bipectinate in ; a single row of slender
scales per segment over dorsal half. Pilifers and
mandibles absent. Haustellum (Fig. 7.3 G) ex-
tremely reduced to a minute setose lobe less than
length of first maxillary palpal segment. Maxil-
lary palpi reduced to two short, rounded seg-
ments. Labial palpi moderately long, upcurved,
3-segmented; lateral bristles absent. Thorax:
Metafurca (Fig. 7.4 J) with furcal apophyses
joined as a bridge to secondary arms. Forewings
moderately broad (Fig. 7.3 M); index 0.40.47,
with Rsl usually completely fused to Rs2
(stalked in Cnissostages) and Rsl + 2 usually
stalked to Rs3 (separate in Cnissostages and in
unnamed Taiwanese genus); Rs4 terminating
either on apex or termen; accessory and interca-
lary cells usually absent; 1 A + 2 A with basal
fork; S retinaculum consisting of a broad, trian-
gular lobe arising from underside of costal mem-
brane immediately anterior to and curling over
base of Sc. Hindwing index 0.43-0.55; $ frenu-
lum a single stout seta; with 412 frenular se-
tae. Legs with tibial spur pattern of 0-2-4; epi-
physis present, inner surface densely spinose. Ab-
domen: Paired sternal apodemes of A2 (Fig.
7.4 D) short, stout to slender; sternal rods pre-
sent; S8 of S with a bilateral pair of short, broad
apodemes directed anteriorly. Coremata some-
times present on S8 of consisting of small,
paired tufts of short, broad scales. Female with
dense corethrogyne of long piliform scales encir-
cling A7. Male genitalia: Uncus fused to tegu-
men, usually tapering to a slender, rounded apex,
rarely bilobed. Vinculum usually Y-shaped, with
a slender, elongate saccus, rarely V-shaped.
Gnathos absent. Distinct juxta and transtilla ab-
sent. Anellus a sclerotized tube tightly enclosing
aedoeagus, and broadening at base to fuse with
inner surfaces of valvae. Valvae elongate, usually
with a prominent, strongly sclerotized lobe or
process from apex of sacculus. Diaphragma usu-
ally invaginated to form a large membranous sac
(apotheca, Bradley 1951) dorsad to anellus. Ae-
doeagus extremely slender and elongate, usually
with a tenuous, non-retractile, partially sclero-
tized vesica often exceeding length of aedoeagus;
vesica normally coiled inside apotheca prior to
mating, sometimes broken off inside ductus sem-
inalis of $ during copulation. Female genitalia:
Segments 8 and 9 + 1 0 forming a slender, elon-
gate telescoping ovipositor supported by 2 pairs
of rodlike apophyses and 1 pair of pseudapo-
physes. Ostium located at caudal margin of S8.
Donald R. Davis & Gaden S. Robinson
Ductus bursae elongate, usually broader and
more thickened over caudal third. Ductus semi-
nalis joining ductus bursae slightly anterior to its
middle, of variable length, extremely long in
those species with $ possessing greatly length-
ened vesica. Corpus bursae relatively small,
membranous and without signa.
Egg: Undescribed. Larva: Body small to large,
up to 30 mm in length, whitish with yellowish
brown head, light brown to black notai plates
and pinacula. Frontoclypeus extending ca. half
the distance to epicranial notch; ecdysial line ter-
minating well before cranial notch. Six pairs of
stemmata. Labrum with 3 pairs of epipharyngeal
setae. Hypostoma narrowly joined at midventer.
Prespiracular plate completely separate from
prenotai plate (Fig. 7.5 N). Coxae partially fused
at midventer. Crochets arranged in a lateral pen-
ellipse; prolegs without peripheral spines; anal
crochets in a semi-ellipse, opened caudally. Lar-
vae feed on woody bracket fungus (Polyporus)
into which are constructed long, irregular silk-
lined tunnels; a tough, slightly flattened case
(Fig. 7.6 E) is attached to the outer opening of
the tunnel (Busck 1912). Pupa: Head smooth,
without frontal ridge. Wings relatively short, not
surpassing A4. Dorsum of A3 - 8 in $ and A 3 - 7
in 9 with a single anterior row of short spines;
A 3 - 7 movable in $ and A 3 - 6 in Cremaster
reduced to a pair of small tubercules on venter
of AIO. Pupation occurs inside larval case with
the adult emerging from distal slit-like opening.
The Arrhenophanidae are mostly restricted to
the New World where four genera are recognized
(Arrhenophanes, 2 sp.; Cnissostages, 2 sp.; Ecpa-
thophanes, 2 sp., Dysoptus, 2 sp.). An unnamed
genus and two species have recently been re-
ported from Taiwan (Davis 1991). Closely allied
to the Psychidae, they differ from the latter by
the antenna being serrate to bipectinate in the
$; genitalia with sclerotized anellar tube and
aedoeagus often with non-retractible vesica that
is stored in an invaginated apotheca; oviposi-
tor often with prominent lamella postvaginalis;
larva with three pairs of epipharyngeal setae and
the prespiracular plate separate from the prono-
tum; and pupa with only a single anterior row of
tergal spines.
Psychidae (Figs. 7.1 S - T , 7 . 2 A - G ) . Small to
medium size moths, with forewings 4 to 28 mm
in length; fully winged, $ either fully winged,
brachypterous, apterous, or vermiform (Fig.
7.2 F) with all body appendages vestigial or lost.
Head (Figs. 7.3 HI): Vestiture generally rough,
consisting usually of moderately long, slender to
piliform scales directed mostly forward. Ocelli
present in some primitive genera, usually absent.
Eyes small to large; interocular index 0.42.3;
cornea naked. Antennae 0.30.8 the length of
forewing; usually vestigial in vermiform 9; scape
rough, pecten present but often indistinct; flagel-
The Tineoidea and Gracillarioidea
lum filiform to strongly bipectinate in <?, usually
filiform in (when developed) to rarely serrate;
vestiture variable, usually with scales scattered
over dorsal half, sometimes with one distinct row
per segment. Pilifers and mandibles absent.
Haustellum usually absent, present in a few of
the most primitive genera (Davis in press b).
Maxillary palpi usually absent, rarely rudimen-
tary to 3-segmented (in Kearfottia, Fig. 7.3 H,
Lamyristis). Labial palpi often lost, 1-3 seg-
mented in more primitive genera; lateral bristles
absent. Thorax: Metafurca (Fig. 7.4 E) with fur-
cal apophyses joined as a bridge to secondary
arms. Forewings (Figs. 7.3 N - O ) slender to
broad; index 0.3-0.66, forewings normally with
7 - 9 veins arising from cell; Rs4 terminating on
termen; M present in cell; accessory and interca-
lary cells usually absent; 1 A + 2 A either with or
without basal fork; S retinaculum varying from
a broad fold to a more slender coiled flap from
underside of costal membrane immediately ante-
rior to and extending under base of Sc. Hindw-
ing index 0.33-0.76; $ frenulum a single stout
seta; alate ? with 3 8 frenular setae. Legs with
tibial spur pattern highly variable, ranging from
0-2-4 to 0-0-0; epiphysis present or absent. Abdo-
men: Paired sternal apodemes of S2 (Fig. 7.4 E)
elongate, straight to curved, continuing more
than half way into 2 S as sternal rods. A pair of
short coremata sometimes present on S8 of $ in
more primitive genera; S8 usually sclerotized
with a pair of prominent apodemes directed an-
teriorly. Female with dense corethrogyne of long
piliform scales encircling A7, often concentrated
in clusters. Male genitalia. Uncus completely
fused, rounded caudally or represented by short,
setose lobes. Tegumen well developed, hoodlike.
Vinculum U- to Y-shaped; saccus often elongate.
Gnathos absent. Juxta sometimes present and
fused with partially sclerotized anellus in lower
forms. Subscaphium sometimes sclerotized, usu-
ally absent. Valvae relatively slender, elongate,
typically with a broad, setose pad at base (often
referred to as pulvillus, vallum penis, but proba-
bly derived from anellus) as well as an acute or
spinose process from ventral margin or apex of
sacculus. Transtilla usually present. Seven pairs
of genital muscles present (Kuznetsov and Stek-
ol'nikov 1977). Aedoeagus elongate, slender,
usually without cornuti. Female genitalia. Highly
variable depending upon degree of specializa-
tion; segments 8 and 9 + 10 usually long and tel-
escoping, with 2 pairs of apophyses and 1 pair of
pseudapophyses (within AIO) in more primitive
genera, to greatly shortened and with only 1 - 2
pairs of apophyses in most specialized, vermi-
form $ (Davis 1964). Ostium located on S8 or at
caudal margin. Ductus bursae and corpus bursae
usually short and membranous; signum absent;
ductus seminalis sometimes greatly elongated in
most advanced genera.
105
Egg: Poorly known. Oval in shape. Chorion
smooth to finely reticulated; micropylar area
with faint reticulations (Davis 1990). Usually
laid in masses of 1813,000 eggs (Kozhanchikov
1956; Davis 1975, 1987) inside larval case and
mixed with piliform scales from body of ; de-
posited on host substrate in some primitive
forms. Larva (Figs. 7 . 5 1 - K , O): Body usually
between 6 - 5 0 mm long, with head and thorax
well sclerotized and variously pigmented; abdo-
men usually unicolorous, varying from white to
fuscous, or with dark, dorsal plates. Frontoclyp-
eus extending ca. 0.5 to epicranial notch; ecdysial
lines converging at or just dorsad to apex of
frontoclypeus. Six pairs of stemmata. Venter of
labrum with 4 pairs of epipharyngeal setae (Fig.
7.51). Thoracic legs well developed and directed
anteriorly (Fig. 7.5 J); coxae partially to com-
pletely fused at midline; lateral pinacula of Tl,
bearing 3 L setae and spiracle, fused to prono-
tum (Fig. 7.5 O). All prolegs present, moderately
short, with crochets of A36 uniordinal and ar-
ranged in a lateral penellipse; crochets of AIO
in a uniordinal semi-ellipse. All larvae are case
bearers throughout all instars and feed on a
broad range of plant hosts, from lichens, grasses,
to leaves of evergreen and decidous trees and
shrubs; a few have also been reported as insec-
tivorous (Clausen 1940). Structure of larval case
highly divergent (Figs. 7.6 FN), from helical
(Fig. 7.6 H) to greatly elongated, from less than
6 mm long at maturity to 32.7 cm (Oiketicus aris-
tocosma Fig. 7.6 G). Number of instars not well
known, at least 4 or more (Jones and Parks 1928;
Davis 1964; Kozhanchikov 1956) the $ some-
times possessing one more instar than $. Pupa:
Vertex usually with short frontal process. Wings
usually terminating on A3 or 4, variably reduced
in $ to absent. Tergal spines highly variable, typi-
cally present on at least some segments between
Al - 8, ranging from an irregular cluster in many
primitive genera to 2 rows in most advanced
forms with the posterior row usually more slen-
der, elongate, and recurved (Davis 1975). Seg-
ments A 2 - 7 movable in A 2 - 6 in . Cremas-
ter usually present as a single pair of small to
large ventral hooks. Pupation occurs inside lar-
val case. Prior to this the last instar larva atta-
ches the case to some support and then typically
inverts itself with the head pointing to the rear
of the case; pupal extrusion and adult emergence
thus occurring usually through the lower or cau-
dal end.
Nearly 1000 species are known from all major
regions except Antarctica (Heppner 1991), with
more than 85% of these restricted to the Old
World. Although generally rare in Lepidoptera,
parthenogenesis has developed in at least five
genera of Psychidae (Dahlica (= Solenobia),
Scoriodyta, Luffia, Apterona, and Acanthopsyche;
Narbel 1946, Suomalainen 1962, Robinson 1971,
Henderickx 1988, Httenschwiler 1989). The ge-
106 Donald R. Davis & Gaden S. Robinson

Fig. 7.6. Larval cases, tubes, and cocoons. AC, Tineidae: A, Ceratophaga vastella, on horn of Kob antelope
(Kobus kob), Sudan ( 3 0 - 5 0 mm); B, Eccritothrix trimaculella, USA (16 mm); C, Tinea pellionella, Germany
(8 mm); D, Acrolophidae, Acrolophus arboreus, Peru (35 cm); E, Arrhenophanidae, Arrhenophanes perspicilla, Pan-
ama (38 mm); FN, Psychidae: F, Lithopleura monochopis. South Africa (39 mm); G, Oiketicus aristocosma
Lower, Indonesia (20 cm); H, Apterona helix Siebold, Switzerland (5.5 mm); I, Amida quadrangularis, Oman
(37 mm); J, Mahasena polytricha, India (37 mm); K, Clania sp., India (50 mm); L, Oiketicus geyeri, Mexico
(75 mm); MN, Thyridopteryx ephemeraeformis, USA, M, mating with 9 inside pupal case, note extension of
S abdomen (pupa 23 mm); N, 3 , clustered at end of 9 case (50 mm); , Bucculatricidae, Bucculatrix sp.,
Belize, O, last instar ribbed cocoon and third instar, circular moulting cocoon; P, ribbed cocoon, lateral view
(2.8 mm). (Length of cases, cocoons in parentheses.)
The Tineoidea and Gracillarioidea
netics of a few palearctic species of Dahlica and
Talaeporia have been studied extensively by
Seiler and his students (Seiler 1920-1963). Seiler
discovered, for example, that Dahlica triquetrella
has bisexual as well as diploid and tetraploid
parthenogenetic races whose distributions were
closely associated with recent geological history
and biotic changes.
Because of numerous and variously correlated
morphological specializations which have ap-
peared at different levels throughout this family,
attempts to classify the Psychidae have varied
considerably. Previous attempts to partition the
family have resulted in as many as 10 families
(Tutt 1990) to as few as 2 subfamilies (Koz-
hanchikov 1956). Gomez Bustillo (1979) superfi-
cially divided the Psychidae into two paraphy-
letic families: Micropsychidae (later emended by
Gomez Bustillo (1984) to Micropsychiniidae) to
include the "tineid-like" forms and the true
Psychidae, or "bombyx-like" forms. The most
recent classification (Httenschwiler 1989; Sauter
and Httenschwiler 1991) lists 8 subfamilies,
mostly without accompanying keys or diagnoses.
Until these are presented, hopefully within the
format of a cladistic analysis, it will not be pos-
sible to apply this classification on a global basis.
The subfamilies proposed by Sauter and Htten-
schwiler, with representative genera shown in pa-
rentheses, are: Naryciinae (Dahlica (= Soleno-
bia), Diplodoma, Narycia, ?Kearfottia); Talepori-
nae (Bankesia, Pseudobankesia, Taleporia); Pla-
codominae (Placodoma); Typhoniidae (Dissoc-
tena, Dissoctenioides, Melasma, Penestoglossa,
and possibly the Neotropical genera Amiantastis,
Plumana, Pterogyne); Scoriodytinae (Scorio-
dyta)', Psychinae (Bruandia, Luffia, Psyche
(= Fumea)-, Epichnopteryginae (Bijugis, Epich-
nopterix, Rebelia, Whittleia), and Oiketicinae
(Acanthopsyche, Amicta, Apterona, Brachycyt-
tarus, Cryptothelea, Eumeta (= Clania), Oiketi-
coides, Oiketicus, Thyridopteryx).
The Pseudarbela-group, previously given fam-
ily status (Figs. 7.2 A - B ; Clench 1959) is cur-
rently included within Psychidae (Robinson 1988)
but some species are referable to the Limacodi-
dae. Only two features - the elongate basal fork
of 1 A + 2 A in the forewings and the spinose
ridge of the S valvae define this taxon. These
are of minor significance when compared to the
major synapomorphies shared with Psychidae.
The group includes 3 genera (Casana with 1 spe-
cies, Linggana with 1 species, Pseudarbela with
4 species) restricted to the Indo-Australian area.
Females are fully winged, with forewings up to
20 mm in length. Immature stages are unknown;
the larva is believed to be a casebearer, although
this needs confirmation.
GRACILLARIOIDEA
The Gracillarioidea are currently restricted to
four families that share the following features:
107
pupa (1) partially extruded from cocoon prior to
eclosion; (2) with abdominal tergal spines; adult
(3) without pleural lobes on A8 of (4) with
smoothly scaled frons; (5) labial palpi without
lateral bristles; (6) galeae usually elongate and
closely associated; (7) hindwings without interca-
lary cell; larva (8) with early instars mining or
boring into plant host; and (9) crochets of later
instars usually biserial. The first three character-
istics distinguish the Gracillarioidea from the
Yponomeutoidea (Kyrki 1984), and the last six
separate them from the Tineoidea. The presence
of a naked haustellum and a second abdominal
sternum with tineoid apophyses (Kyrki 1983 b),
along with the foregoing pupal characters,
closely associate the Gracillarioidea with the Ti-
neoidea and serve to distinguish both groups
from the higher Ditrysia.
Head: Frons smooth; vertex often smooth,
rough in Roeslerstammiidae, Bucculatricidae,
and some Gracillariidae. Ocelli usually absent,
present only in Douglasiidae. Antennae often as
long as or longer than forewing, ranging from
0.61.75 the length of wing; scape with pecten
present or absent; flagellum filiform; 1 - 2 rows
or annuii of scales per segment. Chaetosemata
absent. Pilifers present. Mandibles usually ab-
sent. Haustellum naked, usually elongate, ca.
12X length of labial palpi. Maxillary palpi re-
duced, 1 4 short segments. Labial palpi usually
3-segmented, reduced to 1 in most Bucculatrici-
dae (Fig. 7.7 F), without erect bristles. Thorax:
Metafurca with apophyses free. Forewings mod-
erately broad to slender, index (W/L) 0.09-0.24;
Rs4 terminating on costa or apex; chorda with
accompanying accessory cell usually absent,
sometimes present in Roeslerstammiidae and
Gracillariidae; intercalary cell absent; 1 A + 2 A
with or without basal fork; subanal interlocking
microtrichia present. Hindwings often lanceo-
late, index 0.1-0.4; intercalary cell absent; discal
cell usually open (m-cu crossvein absent), closed
in Roeslerstammiidae and most primitive Gracil-
lariidae. Abdomen: Tergal spines absent. Anterior
margin of S2 with tineoid apodemes and usually
with sternal rods (Figs. 7.7 N - O , Q); rods absent
in Bucculatricidae (Fig. 7.7 P); a minute pair of
warts and 2 pairs of tuberculate plates usually
present as in Tineoidea. Male genitalia: Uncus
usually absent, slightly bilobed in Roeslerstam-
miidae; genital muscles 3 - 7 pairs (Kuznetsov
and Stekol'nikov 1977, 1987). Female genitalia.
Ovipositor moderately to only slightly extensi-
ble, with 2 pairs of moderate to short apophyses.
Ostium opening usually on S8 or at caudal mar-
gin; opening between S7 and 8 in Roeslerstammi-
idae.
Egg: Oval, fiat type, with micropylar axis par-
allel to oviposition substrate. Chorion relatively
smooth to finely reticulated; micropylar end with
reticulated ridges. Eggs laid singly and cemented
to plant host, with larva boring directly into tis-
sue upon eclosion. Larva: Often hypermetamor-
108 Donald R. Davis & Gaden S. Robinson

Fig. 7.7. Gracillarioidea, Head and abdominal morphology, wing venation: A - , Roeslerstammiidae, Roesler-
stammia erxlebella; B, Base of maxilla and maxillary palpus; CD, Douglasiidae, Tinagma obscurofasciella, O
ocellus; D, Base of maxilla and maxillary palpus; EF, Bucculatricidae: Bucculatrix solidaginiella; F, Ventral
view of labial palpi (LP); G - H , Gracillariidae: G, Caloptilia stigmatella; H, Phyllocnistis insignis(0.25 mm); I,
Roeslerstammiidae, Roeslerstammia erxlebella, J, Douglasiidae, Tinagma obscurofasciella; K, Bucculatricidae,
The Tineoidea and Gracillarioidea
phic; early instars typically internal borers or
miners, often apodal or with legs and prolegs re-
duced; ventral crochets usually in a biserial circle
with inner series incomplete (Fig. 7.81). Pupa:
Adecticous, incomplete, with 4 - 5 movable abdo-
minal segments. Maxillary palpi usually present.
Abdomen with tergal spines usually in a single
anterior row, or in dense patches (Gracillarii-
dae). Pupa extended partially from cocoon prior
to adult eclosion.
Key to the Families
1 Ocelli present Douglasiidae
Ocelli absent 2 slightly bilobed, setose. Tegumen moderately
2 Antennal scape broad, partially covering eye.
Haustellum short, less than 1.5X eye diameter
(Fig. 7.7 E). Cocoon longitudinally ribbed
(Fig. 7.6 P) Bucculatricidae
Antennal scape slender, not covering eye.
Haustellum more than 2.5 X eye diameter. Co-
coon smooth, not ribbed 3 elongate tuft of piliform scales arising from base.
3 Wings slender to lanceolate (Figs. 7.7 L - M ) ;
hindwing index (W/L) 0.10.2; cilia longer
than hindwing width. Flagellomeres with sin-
gle annulus of scales Gracillariidae
Wings broader (Fig. 7.71), hindwing index
0.28-0.4; cilia less than hindwing width. Fla-
gellomeres with 2 rows of dorsal scales . . . .
Roeslerstammiidae
Roeslerstammidae (Figs. 7.2 H - I ) . Small moths,
with forewings 5 to 10 m m in length. Head (Fig.
7.7 AB): Vestiture of frons smooth with broad
scales; vertex rough with dense covering of erect
piliform scales. Ocelli absent. Eyes moderately
large to greatly enlarged, sometimes indented
posteriorly or completely divided by a slender
scaled band; interocular index 0.87-2.0; cornea
naked. Antennae 0.81.5 the length of forewing;
scape with dense pecten; flagellum piliform; two
rows of slender scales per segment dorsally, with
distal row encircling segment. Pilifers well devel-
oped. Mandibles absent. Haustellum moderate
to elongate, usually equalling or exceeding length
of labial palpi. Maxillary palpi reduced to a sin-
gle ovoid segment (Fig. 7.7 B). Labial palpi elon-
gate, 3-segmented, slender, and curved upwards;
lateral bristles absent. Thorax: Metafurca (Figs.
7.8 - B ) with posterior apophyses slender, elon-
gate, free; anteromedial process elongate and
slender, equalling or surpassing length of furcal
stem, and with a prominent ventral keel from
base. Forewings (Fig. 7.71) narrow, index 0 . 2 2 -
0.31; Rs3 and 4 either stalked or fused and termi-
Bucculatrix solidaginiella', LM, Gracillariidae: L, Caloptilia stigmatella; M, Phyllocnistis insignis; NQ, Second
abdominal sterna: N , Roeslerstamiidae, Roeslerstammia
Bucculatricidae, Bucculatrix solidaginiella; Q, Gracillariidae, Caloptilia stigmatella. (All scales = 0.5 mm, except
Fig. H.)
109
nating on costa; accessory cell absent or weakly
present; intercalary cell absent; 1 A + 2 A with
basal fork; $ retinaculum a broad, triangular
flap from costal cell coiling underneath base of
Sc. Hindwing index 0.28-0.4; $ frenulum a sin-
gle stout seta; $ with 2 frenular setae. Legs with
tibial spur pattern of 0-2-4; epiphysis relatively
short, arising from distal third of tibia. Abdomen :
Paired sternal apodemes of A2 (Fig. 7.7 N) elon-
gate, slender, slightly curved inwards, continuing
caudad as well defined sternal rods 2/3 the length
of S2; a transverse, sclerotized bar extends par-
tially across anterior margin of S2. Coremata
and corethrogyne absent. Male genitalia: Uncus
broad dorsally. Vinculum usually Y-shaped with
an elongate saccus, rarely U-shaped. Gnathos
slender, either fused medially or weakly united
by membrane. Subscaphium slender, weakly
sclerotized. Valvae relatively simple, sometimes
with an acute basal or subapical process; an
7 pairs of genital muscles present (Kuznetsov
and Stekol'nikov 1977). Aedoeagus a slender,
often curved cylinder without cornuti. Female
genitalia: Ovipositor short, with 2 pairs of short
apophyses. Ostium opening in intersegmental
membrane between S7 and 8. Antrum variously
sclerotized. Corpus bursae membranous with a
single, large, linear to cruciform signum.
Egg: Undescribed; deposited at the tip of host
leaf (Kyrki 1983a). Larva (Figs. 7.8 G - I ) : Body
pale green, up to 13 mm long, with thoracic legs
and 5 pairs of prolegs. Head brownish; pronotal
and anal plates paler brown to indistinct. Fron-
toclypeus slender, extending 0.8 the distance to
epicranial notch (Fig. 7.8 G). Adfrontal sutures
terminate at epicranial notch. Six pairs of stem-
mata present. Prothorax (Fig. 7.8 H) with 3 pre-
spiracular (L) setae. Legs well developed; coxae
separated. Ventral prolegs with 2 4 - 3 4 crochets
arranged in a posteriorly uniserial or biserial lat-
eral penellipse (Fig. 7.81). Anal prolegs with ca.
15 uniordinal crochets in a semi-circle. First two
larval instars are leafminers (on Aceraceae, Betu-
laceae, Elaeocarpaceae, and Tiliaceae) and the
later instars feed externally, either exposed, or in
webbing on underside of leaf (Kyrki 1983 a), or
within a shelter formed by webbing several
leaves together (Moriuti 1978). Pupa: Frons
smooth. Antennae extend to or exceed caudal tip
of abdomen, in which case they tend to coil
around abdomen (Common 1990). Wings ex-
tending to caudal margin of A6. An anterior row
of tergal spines present on A 3 - 7 . Segments
erxlebella; O, Douglasiidae, Tinagma obscurofasciella; P,
110
A 2 - 6 movable in $. AIO without sclerotized
cremaster but with stout to hooked setae. Pupa-
tion occurs inside a dense silken cocoon among
leaf litter or within a folded leaf edge.
The family is widespread through the Old
World in the Palearctic, Oriental, and Australian
regions where 10 genera and ca. 40 species are
known. A key to six of the Oriental-Australian
genera (Agriothera, Amphithera, Chalcoteuches,
Enchoptila, Sphenograptis, Telethera) and one
Neotropical genus (Dasycarea) was compiled by
Moriuti (1978). Common (1970) also included
two additional Australian genera (Macarangela
and Thereutis) within the family. Kyrki (1983)
listed seven species ranging from western Europe
to Japan within the Palearctic genus Roeslers-
tammia.
Douglasiidae (Fig. 7.2 J). Small moths, with fore-
wings 3.0 to 6.4 mm in length. Head (Fig.
7.7 CD): Vestiture of broad, smooth scales.
Ocelli well developed. Eyes small, interocular in-
dex 0.65-0.7; cornea naked. Antennae 0.6-0.7
the length of forewing; scape usually smooth, oc-
casionally with rudimentary pecten of 2 - 3 pili-
form scales; flagellum filiform, with a single row
of slender scales encircling each segment. Pilifers
rudimentary. Mandibles reduced to a small lat-
eral lobe. Haustellum elongate, more than 2X
length of labial palpi. Maxillary palpi reduced to
a single lobate segment (Fig. 7.7 D). Labial palpi
relatively short, 3-segmented, drooping to
slightly upcurved. Thorax: Metafurca (Fig. 7.8 C)
with posterior apophyses short, free, curved
ventrad; anteromedial process with prominent
ventral keel. Forewings (Fig. 7.7 J) narrow; index
ca. 0.28; Rs with 3 or 4 branches; Rs4 terminat-
ing on costa, stalked with M l ; M either 2 or 3
branched; accessory and intercalary cells absent;
A 1 + 2 A with or without basal fork; $ retinacu-
lum a slender coiled flap under base of Sc.
Hindwing index ca. 0.2; discal cell open; $ frenu-
lum a single stout seta; ? with 1 2 frenular setae.
Legs with tibial spur pattern of 0-2-4; epiphysis
present. Abdomen: Paired sternal apodemes of
A2 (Fig. 7.7 O) elongate, slender; sternal rods
weak. Coremata and corethrogyne absent. Male
genitalia: Uncus absent. Tegumen a triangular
hood. Vinculum slender, often with a short, stout
saccus which is lacking in some species. Gnathos
and transtilla absent. Anellus a partially sclero-
tized ring. Valvae asymmetrical in Klimeschia or
symmetrical in Tinagma, relatively short and
broad and often with a peculiar apical process.
Fig. 7.8. Gracillarioidea. A - F , Metathoracic furcasterna: - B , Roeslerstammiidae: A, Roeslerstammia erxlebella,
B, Agriothera issikii; C, Douglasiidae, Tinagma obscur ofasciella\ D, Bucculatricidae, Bucculatrix soldaginiella;
E - F , Gracillariidae: E, Caloptilia stigmatella; F, Phyllocnistis insignii; GS, Larval morphology: G I, Roeslers-
tammiidae: G, Agriothera elaeocarpophaga, head (0.25 mm); H I. Roeslerstammia erxlebella: H, Prothorax; I,
Abdominal segment 36; J, Douglasiidae, Tinagma balteolellum; K - L , Bucculatricidae, Bucculatrix thur-
Donald R. Davis & Gaden S. Robinson
Aedoeagus a slender cylinder usually without
cornuti. Female genitalia. Ovipositor telescoping,
moderately long; two pairs of apophyses of equal
length. Ostium a sclerotized ring of various out-
line at caudal margin of S8. Ductus bursae slen-
der, mostly membranous, sometimes with a few
minute spicules or a distinct collar. Corpus bur-
sae with a single stellate signum.
Immature stages poorly known. Egg: Unde-
scribed. Larva (Fig. 7.8 J): Small; length ca.
5 - 8 mm. Frontoclypeus higher than wide, ex-
tending 0.6-0.75 to epicranial notch. Six pairs of
stemmata. Body white to yellow with pigmented
pronotal plate, pinacula, and anal plate. Prespi-
racular (L) group of T1 trisetose. Primary setae
extremely long. Thoracic legs present; coxae sep-
arated. Prolegs reduced; crochets apparently ab-
sent. Larvae have been reported as leaf miners
Klimeschia), flower petiole miners and stem bor-
ers (Tinagma) on Boraginaceae, Rosaceae, and
Laminaceae (Heppner 1984). Pupa: Unde-
scribed, reportedly with abdominal tergal spines
(Kyrki 1984) and partially extruded from cocoon
prior to adult eclosion. The family is small with
only 17 species represented in the Palaearctic re-
gion (Gaedike 1974), 7 for the Nearctic (Gaedike
1990), and one from Australia (Common 1990).
Two genera, Klimeschia and Tinagma (= Doug-
lasia) are recognized.
Bucculatricidae (Figs. 7.2 - M ) . Extremely to
moderately small moths, with forewings 2.5 to
7 mm in length. Head (Figs. 7.7 E - F ) : Usually
elongate, with frons projecting well below eye;
shortened, less specialized in Leucoedemia. Vesti-
ture with frons smooth; vertex usually with large,
erect tuft of piliform scales projecting in all di-
rections, rarely smooth. Ocelli absent. Eyes mod-
erately large, interocular index ca. 1.0; cornea
naked. Antennae 0.6-0.9 the length of forewing;
scape enlarged, flattened, with a dense row of
slender scales (pecten) partially covering eye; fla-
gellum filiform, with 2 rows of short, slender
scales encircling each segment; first flagellomere
often notched or strongly curved in Pilifers
usually reduced, well developed in Leucoedemia.
Mandibles absent. Haustellum short, 1.5X eye
diameter or less. Maxillary palpi vestigial, usu-
ally a minute, unsegmented lobe. Labial palpi
drooping, without lateral bristles, usually ex-
tremely short and one segmented (Fig. 7.7 F) or
with three short segments in Leucoedemia. Tho-
rax: Metafurca (Fig. 7.8 D) with apophyses re-
duced to a pair of low dorsal ridges. Forewings
beriella: , Fifth instar (0.5 mm); L, Head (0.2 mm); M - S , Gracillariidae: M - O , Phyllonorycter blancardella: M,
Fifth instar (0.5 mm); N, Third instar, lateral; O, Third instar, ventral (0.5 mm); PQ, Caloptilia azaleella, P,
Head, second instar (0.1 mm); Q, Head, Fifth instar (0.2 mm); R - S , Phyllocnistis insignis: R, Head, third instar
(0.2 mm); S, Head, Fourth instar (0.2 mm). (Scale lengths in parentheses; Figs. G from Moriuti 1978, H - I from
Kyriki 1983, J from Heppner 1987, K - O , R - S from Davis 1987.)
112
(Fig. 7.7 ) lanceolate, index 0.14-0.25, with
apex of forewing usually acuminate; forewing
with retinaculum a broad fold from underside of
fused C and Sc veins; Rs usually 4-branched, rar-
ely 3; Rs4 to apex; Accessory and intercalary
cells absent; Cu usually single branched, rarely
absent; 1A + 2 A without basal fork. Hindwing
index 0.180.21; $ frenulum a single stout seta;
$ with 2 frenular setae; discal cell open in hind-
wing. Epiphysis either present or absent; legs
with tibial spur pattern of 0-2-4. Abdomen:
Paired sternal apodemes of A2 (Fig. 7.7 P) elon-
gate, slender; sternal rods absent. An eversible
scale sac present in most $ Bucculatrix as a mid-
dorsal invagination of membrane between A2
and 3; corythrogyne absent. Male genitalia. Un-
cus usually absent, rarely well developed and
acute. Tegumen and vinculum narrow to broad;
saccus rarely developed. Gnathos usually absent;
well developed, fused medially, and setose in
Leucoedemia. Socii typically well developed and
setose in Bucculatrix. Transtilla usually absent,
sometimes weakly developed. Anellus a membra-
nous cone often with lateral sclerotizations. Dis-
tinct juxta usually absent. Valvae usually elon-
gate and simple, rarely lobed. Three pairs of gen-
ital muscles present (Kuznetsov and Stekol'nikov
1987). Aedoeagus usually an elongate, sinuate
cylinder; cornuti rarely present. Female genitalia:
Ovipositor short, usually with only posterior
apophyses present; anterior apophyses some-
times evident. Ostium opening either along cau-
dal margin of S8 or near center of S8. Ductus
bursae usually sclerotized over caudal portion,
usually elongate and slender. Corpus bursae
abruptly enlarged; signum usually a single large
plate with numerous rows of spines.
Egg: Usually flattened and oval, with a reticu-
lated chorion (Braun 1963). Larva (Figs.
7.8 K L): Hypermetamorphic, typically with 5
instars (Braun 1963); apodal in at least first two
leafmining instars, with thoracic legs and 5 pairs
of prolegs in later externally feeding instars; pro-
thoracic pretarsal claw highly modified in Buccu-
latrix, greatly lengthened; crochets in an incom-
plete circle. Frontoclypeus short, extending 0.5
0.6 to epicranial notch. Tentorial bridge very
slender, deeply arched (Davis 1987). Stemmata
5 - 6 pairs. Prespiracular (L) group of T1 triset-
ose. Most larvae undergo two instars as leafmin-
ers, with the third instar emerging to feed exter-
nally. A flattened moulting cocoon (Fig. 7.8 L) is
sometimes constructed underneath the leaf by
the third instar larva as well as a second moult-
ing cocoon by the fourth instar; the fifth instar
prior to pupation constructs a silken, ribbed co-
coon (Figs. 7.6 OP) typical for the family.
Some species are stem borers or feed inside galls
(Braun 1963, Scoble and Scholtz 1984). The
larva of Ogmograptis scribula forms an elongate
mine in the bark of Eucalyptus (Nielsen and
Common 1991). Nearly 30 families of gym-
Donald R. Davis & Gaden S. Robinson
nosperms and angiosperms have been reported
as hosts, with the Asteraceae, Betulaceae, and
Fagaceae being particularly favored (Braun
1963, Seksyayeva 1994 b). Pupa: Frontal ridge
longitudinal, well developed. Wings and anten-
nae usually extending to A7; A2 or 3 - 7 with
a single row of dorsal spines. Segments A37
movable in S, A 3 - 6 in ? (Mosher 1916). AIO
with cremaster reduced, consisting of minute,
paired, dorsal spines and often a pair of larger,
lateral spines.
Approximately 250 species are known around
the world, with the greatest concentration
(100 sp.) in the Nearctic region (Braun 1963,
Heppner 1991, Seksyayeva 1994 a). With the no-
table exception of the most plesiomorphic taxon,
the monotypic South African Leucoedemia,
nearly all the species are included in the single
genus Bucculatrix . The Australian Ogmograptis
has recently been placed in this family (Nielsen &
Common 1991).
Gracillariidae (Figs. 7 . 2 N - S ) . Small moths with
forewings 210 mm in length. Head (Figs.
7.7 G - H ) : Vestiture variable, usually smooth (in
Gracillariinae) with moderately broad scales
from vertex directed forward and down over
frons; similar in Lithocolletinae except with tufts
of erect filiform and sometimes broader scales
arising from occiput caudad to vertex. Ocelli ab-
sent. Eyes moderately large; interocular index
0.9-1.0; cornea naked. Antennae 0.8-1.75 the
length of forewing; scape usually smooth, with
or without pecten; flagellum filiform, with a sin-
gle row of slender scales completely encircling
each segment. Pilifers well developed. Mandibles
absent. Haustellum usually elongate, 1 - 2 X
length of labial palpi. Maxillary palpi rarely ab-
sent, usually short to minute, 1-4-segmented. La-
bial palpi 3-segmented, usually upturned, to
straight and drooping; segment 2 sometimes with
ventral scale tuft. Thorax. Metafurca (Figs.
7.8 E - F ) with anteromedial process elongate,
relatively slender; furcal apophyses free, short to
moderately long and downcurved, arising well
caudad near secondary arms. Forewings (Figs.
7.7 L - M ) slender; index 0.9-0.25; venation
variable with R/Rs 3- to 5-branched; Rs4 usually
to costa, rarely to apex; M l - to 3-branched; Cu2
sometimes absent; discal cell elongate, extending
0.7-0.8 the length of forewing; accessory cell
vestigial to absent; base of M vestigial to absent;
intercalary cell absent; 1A + 2 A usually present,
sometimes vestigial, occasionally with minute
basal fork; $ retinaculum usually formed of
curved scales from underneath slightly swollen
base of Sc; Hindwing lanceolate; index 0.1-0.2;
$ frenulum usually with 2 frenular setae, rarely 1 ;
stout, composite pseudofrenular setae sometimes
arising near apex of Sc in both sexes (Davis
1991); venation reduced, from 7 to 4 veins; discal
cell often open. Legs with tibial spur pattern usu-
The Tineoidea and Gracillarioidea
Fig. 7.9. Gracillariidae, leaf mines. A, Parectopa robiniella on Robinia pseudoacacia, upperside; , Cremastobom-
bycia ignota on Verbesina occidentalism lower side, flat blotch mine of third instar on left, tentiform blotch of
fourth-fifth instar on right; C, Cameraria guttifinitella on Toxicodendron radicans, upperside; D, Phyllocnistis
vitegenella on Vitus cordifolia, upperside, ca. 8 individual mines shown.
ally 0-2-4; epiphysis rarely absent. Abdomen. S2
(Fig. 7.7 Q) long and narrow, with long slender
sternal apodemes continuing caudad as sternal
rods ca. 0.B the length of sternite. A8 and some-
times 7 of $ frequently with paired coremata; S8
of $ often enlarged, particularly in Lithocolleti-
nae; T8 sometimes with median sclerotization;
corethrogyne absent. Male genitalia. Uncus ab-
sent. Tegumen a relatively elongate, mostly
membranous dorsal hood. Vinculum U- to Y-
shaped, with often a short, rarely elongate sac-
cus. Subscaphium frequently distinct, variably
sclerotized. Gnathos absent. Transtilla present or
absent. Yalvae usually elongate and simple, rar-
ely asymmetrical, sometimes lobed, spined, or
with 1 - 2 pairs of pectinifers. Anellus usually
membranous. Juxta absent. Aedoeagus usually
elongate and slender; cornuti present or absent.
$ genitalia with 4 pairs of muscles, Ml absent
(Kuznetsov and Stekol'nikov 1987). Female geni-
talia. Ovipositor short, with usually 2 pairs of
moderately short, rarely elongate apophyses; an-
terior apophyses sometimes reduced or absent.
Ostium on S8 or at caudal margin; sterigma
sometimes well sclerotized and variously modi-
fied. Ductus bursae usually elongate, slender,
either membranous or partially sclerotized, ex-
panding anteriorly into membranous corpus bur-
sae; signa usually of 1 - 2 pairs, or sometimes ab-
sent.
Egg: Usually oval to elliptical, partially flat-
tened underneath and slightly convex above with
nearly smooth to finely sculptured chorion,
sometimes with upper and lower halves different
in texture; less than 0.5 mm in length. Micropyle
area finely reticulated, often reduced in size.
Larva (Figs. 7.8 MS): Body usually not exceed-
ing 10 mm, variable in color depending upon
species and instar, from white to green, with
some becoming bright red prior to pupation,
with or without darkly pigmented head and scle-
113
rotized plates. Hypermetamorphic development
(Kumata 1978; Davis 1987) with at least 2 dis-
tinct forms and habits: (1) an early, usually sap-
feeding (Trgardh 1913) or flattened (Chambers
1877) form, often comprising the first 25 in-
stars, which possess a depressed, apodal body
with specialized, prognathous mouthparts and
rudimentary spinneret (Figs. 7.8 N - O , P, R),
and (2) a later hypognathous, tissue-feeding or
cylindrical form possessing a more generalized,
eruciform body with unspecialized, chewing
mouthparts and legs (Figs. 7.8 M, Q); or a vari-
ously modified nonfeeding spinning form, with
either a depressed or cylindrical body, mandibles
and legs reduced or absent, but with a functional
spinneret (Figs. 7.8 S). An additional nonfeeding,
quiescent stage is also interposed between the fi-
nal sap-feeding and single spinning instars of
Chrysaster, Dendrorycter, and Marmara (De-
Gryse 1916; Kumata 1978). Moreover, in the lat-
ter two genera the quiescent instar exists in a
pharate condition within the cuticle of the last
sap-feeding instar. Later instar larvae of Gracil-
lariidae are further characterized by 2 prespirac-
ular (L) setae on T1 and crochets, if present, on
A35 and 10 and almost always absent on A6
(present on A 3 - 6 in Artifodina, Prophyllocnistis,
and on A26 in Metriochroa psychotriella, Davis
1994). Number of instars varies from 4 - 1 1 , de-
pending upon genus and sometimes species (Fitz-
gerald and Simeone 1971). Most species are leaf-
miners (Figs. 7.9 - D ) ; others mine the subepi-
dermal layers of new growth stems and fruits or
bore inside stems and galls (Davis 1991). Early
sap-feeding instars with slashing mandibles typi-
cally feed in a horizontal plane, initiating slen-
der, subepidermal serpentine mines while ingest-
ing relatively little solid tissue. In some genera
(e. g. Phyllonorycter) the 'sap-feeding' instars in-
stead initiate a blotch mine by devouring mostly
spongy parenchymal cells (Watson 1956). Tissue-
114
feeding instars possess typical chewing man-
dibles that enable these larvae to feed deeper into
the host, usually ingesting the remainder of the
spongy parenchyma and most of the palisade
cells within the blotch. Later tissue-feeding
instars of the most primitive genera (Gracilla-
riinae) tend to exit the mine and feed externally,
often in a rolled-over leaf. Pupa: Head usually
with a serrated or pointed process (cocoon cut-
ter) from vertex, or smoothly rounded. Antennae
extending to A6 to well beyond AIO. Wings to
A56. Tergal spines variably developed, usually
small to minute in multiple, irregular rows or
dense concentrations on A2 or 3-7 or 8; Phylloc-
nistis often with a few pairs of large tergal hooks
on A 2 - 7 . A 4 - 7 moveable in A 4 - 6 in
(Mosher 1916). Cremaster highly variable, usu-
ally present as small paired spines, often with
recurved apices. Pupation usually in a small,
whitish silken cocoon either outside the host
plant (most Gracillarinnae) or within the mine
(most Lithocolletinae and all Phyllocnistinae).
The Gracillariidae constitute the principal
family of plant mining Lepidoptera. Approxi-
mately 2000 species grouped into ca. 75 genera
are known from all major regions of the world
except Antarctica (Meyrick 1912 a, b; Vari 1961;
Kumata 1963, 1982; Kumata et al. 1988; Davis
1983, 1984). Fossil leafmines of Gracillariidae
(Phyllocnistinae) are known from the early Cen-
omanian of the Dakota Formation in Kansas
and Nebraska (ca. 97 m. y. a.; Davis 1994; La-
bandeira et al. 1994) near the beginning of the
Angiosperm radiation. This currently represents
the earliest fossil record of any ditrysian moth
assignable to family. Three subfamilies are recog-
nized.
Gracillariinae: Adults of most genera rest with
anterior part of body raised at steep angle. Head
usually smooth. Hindwings with base of Rs
arched toward costa and approaching apex of Sc
(Ely 1918; Kumata 1978). Abdomen of $ usually
with S8 membranous; coremata usually present.
Last instar larva with fully developed mandibles
and legs, 5 coxal setae, and A9 with D group
bisetose. Pupation usually outside of mine. Rep-
resentative genera: Acrocercops, Aristaea, Calop-
tilia, Epicephala, Macrostola, Parectopa, Paror-
nix, Philodoria, Stomphastis.
Lithocolletinae: Adults rest with body parallel
to surface or with head end lowered. Head usu-
ally with occipital tufts of piliform scales.
Hindwings with Rs nearly parallel to costal mar-
gin. Abdomen of $ with S8 sclerotized, elongate;
coremata absent. Last instar larva (Fig. 7.8 M)
usually with fully developed mandibles and legs
(both reduced in Cameraria), 4 coxal setae, and
A9 with D unisetose. Pupation normally within
blotch mine (Figs. 7.8 B - C ) . Representative gen-
era: Cameraria, Chrysaster, Cremastobombycia,
Leucanthiza, Phyllonorycter (= Lithocolletis).
Donald R. Davis & Gaden S. Robinson
Phyllocnistinae: Adults rest with body parallel
to surface. Head smooth. Hindwings with Rs
closely parallel to costal margin. Abdomen of $
with S8 membranous; coremata usually present.
All feeding instars sap-feeding, creating elongate
serpentine, subepidermal mines (Fig. 7.9 D); last
instar larva non-feeding with mandibles absent
(Fig. 7.8 S), legs reduced to unsegmented stubs,
A9 with D bisetose. Pupation within slightly en-
larged cavity at termination of mine. Representa-
tive genera: Cryphiomystis, Metriochroa, Phylloc-
nistis, Prophyllocnistis).
A fourth subfamily, Ornichinae, has been pro-
posed (Kuznetsov and Stekol'nikov 1987) for
Callisto and Parornix based on the presence of a
sclerotized anellus and different genital muscula-
ture. Major characters in the larva, pupa, and
adult wing venation of these genera do not sup-
port subfamily distinction but instead place them
within Gracillariinae.
Acknowledgments
The authors wish to thank Vichai Malikul and
Young Sohn of the Department of Entomology,
Smithsonian Institution, Washington, D. C.,
USA and Biruta Akerbergs Hansen of Liverpool,
Pennsylvania and Andr del Campo Pizzini (de-
ceased) for the line illustrations used in this
chapter. We also are grateful to Victor Kranz,
formerly of the Smithsonian Photographic Labo-
ratory and Susann Braden and Walter Brown of
the Smithsonian Scanning Electron Microscope
Laboratory for photographic assistance. Peter
Httenschwiler of Uster, Switzerland was helpful
in reviewing the section on Psychidae.
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Addendum
Previous work on the skeleto-muscular structure
of tineid male genitalia by V. I. Kuznetsov and
A. A. Stekol'nikov has been supplemented by
additional contributions from the same authors
in 1995 and 1997; both morphological and phy-
logenetic issues were addressed.
Norman et al. (1995) listed the SE Asian
Psychidae and showed the taxonomic usefulness
of wing scale characters in the family.
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Common bagworm pests (Lepidoptera: Psychidae)
of oil palm in Malaysia with notes on related
South-east Asian species. Malay. Nat. J., 48:
93-123.
Kuznetsov, V. I. & Stekol'nikov, A. A. (1997): Phylo-
genese relations of some subfamilies of true moths
(Lepidoptera, Tineidae) based on a new study of
functional morphology of male genitalia. Ent.
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362-372.
Stekol'nikov, A. A. & Kuznetsov, V. I. (1995): New ar-
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tera) and correction of the hypothesis concerning
the origin of some male genital appendages. - Ent.
Obozr. 74: 387-394. [In Russian],
 8. The Yponomeutoidea
John S. Dugdale, Niels P. Kristensen, Gaden S. Robinson & Malcolm J. Scoble
The Yponomeutoidea are a heterogeneous as-
semblage of relatively primitive micro-moths.
Kyrki (1984) first recognized apomorphies indi-
cating the monophyly of this group but, as yet,
precise delimination of the superfamily remains
debatable.
The principal yponomeutoid autapomorphy is
the presence, in males, of posterior expansions of
pleuron VIII (Figs. 8.1 F - G ) . The pleural lobes
enclose the genital apparatus and their size is
broadly correlated with that of the latter, the
valves in particular; in the Plutellidae/Acrolepii-
dae and in the Heliodinidae the lobes are quite
small and narrow. In the Bedelliidae and Glyph-
ipterigidae (Fig. 8.1 H) the lobes are not present
as discrete formations, and in these extra ad hoc
hypotheses are needed to justify their assignment
to the superfamily. Another possible yponomeu-
toid groundplan autapomorphy is the presence
of a transverse costa behind the anterior margin
of sternum II (Fig. 8.3 C); this costa is absent in
Ypsolophidae-Ochsenheimeriinae and some Lyo-
netiidae.
Ocelli and chaetosemata present or absent.
Antenna simple, scape with or without pecten or
scale awning. Proboscis unsealed, usually well
developed; maxillary palp small, 1 4 segmented,
sometimes appressed to proboscis base. Labial
palp drooping, porrect or ascending, not mark-
edly elongate and recurved above head; apical
segment with vom Rath's organ basally or near
midlength. Epiphysis present, spurs usually 0-2-
4, occasionally reduced, hind tibiae usually
smooth-scaled. Forewing with pterostigma and
chorda present or absent, Rs4 nearly always
postapical, M-stem in cell usually vestigial or ab-
sent. Female frenulum usually of only two bris-
tles. Sternum II of tineoid type. Abdominal ter-
gal spinosity variably developed, often absent.
Male postabdomen usually with paired ventro-
lateral coremata between pleural lobes and IX;
X almost never forming well developed median
uncus (one known exception is the New Zealand
Cadmogenes, which may not belong in the Ypo-
nomeutoidea); socii often well developed, flank-
ing anal tube, or absent (Glyphipterigidae, also
the Australian/New Zealand 'plutellid' Or-
thenches); gnathos present or absent, sometimes
fused medially with socii; subscaphium strongly
developed and strap-like, or weakly developed.
Valve base with strong apophysis for extrinsic
muscle. The male genital musculature in ypono-
meutoids and its (phenetic-)systematic implica-
tions are discussed by Kuznetzov & Stekolnikov
(1976-77, 1993) and Kuznetzov, Kozlov & Seks-
jaeva (1988). Female anterior apophyses usually
continued into dorsal and ventral costae at base
(Fig. 8.1 D); lamella antevaginalis unsclerotized.
Larva with prothoracic L usually trisetose (in
post-1st. instars), occasionally (almost all Glyph-
ipterigidae, a few Yponomeutidae and Ypsoloph-
idae) bisetose. Pupal abdomen lacking spine
bands (single spines sometimes associated with
setae D1 and SD), segments I - I V immobile,
pupa not protruded prior to eclosion.
The controversial phylogenetic significance of
the tergal spinosity and pupal structure is dis-
cussed in 1 - 2 . The yponomeutoids as now de-
limited comprise the bulk of those non-apoditry-
sian Ditrysia which have non-protruded pupae
(conditions in Glyphipterigidae-Orthoteliinae
may represent an autapomorphic character re-
versal) and whose adults do not have the charac-
teristic gelechioid scaled proboscis.
Heppner (1987) corrected his earlier statements
(1977) about the presence of pupal protrusion in ypo-
nomeutoid families.
The formal classification of the superfamily
has so far varied considerably. The family Ypo-
nomeutidae has at times comprised the taxa now
assigned to the Plutellidae and Ypsolophidae
Ypsolophinae, but thus delimited it is almost cer-
tainly paraphyletic. A phylogenetic analysis of
the superfamily by Kyrki (published posthu-
mously 1990) led to a reclassification, which is
followed by Scoble (1992) and in the present
work (except that the Acrolepia-group is treated
as a distinct family). The tentative nature of this
arrangement must be emphasized, and the mo-
nophyly of the redefined Yponomeutidae is still
in obvious need of further scrutiny. The family/
subfamily position of a number of extra-Hol-
arctic genera (listed by Kyrki 1984) which have
been assigned to the Yponomeutidae s. lat. re-
mains in doubt.
Key to yponomeutoid families
(based on Holarctic taxa)
1 Ocelli present, rarely rudimentary (if rudi-
mentary, maxillary palp normally 34 seg-
mented, porrect, and forewing pterostigma
extending to Rs2) 2
Ocelli absent 6
2(1) Wings pronouncedly narrow, forewing
with only 5 postapical veins arising from
cell (Fig. 8.3 D) Heliodinidae
Wings less narrow, forewing with 6 posta-
pical veins arising from cell 3
120 John S. Dugdale, Niels P. Kristensen, Gaden S. Robinson & Malcolm J. Scoble
3 (2) Head smooth-scaled (Fig. 8.3 A); labial
palp often with transverse light and dark
bars; forewing often with metallic mark-
ings; male genitalia partially enclosed in a
posteriorly expanded tergopleural 'cone',
i. e. distinct lateral lobes not developed
(Fig. 8.1 H)
. . . . Glyphipterigidae (Glypiphteriginae)
Head rough-scaled (Fig. 8.1 B); labial palp
without transverse bar pattern; forewings
usually without metallic markings (such
markings do occur in some S. Hemisphere
taxa); male pleural area VIII/IX with well
developed lateral and/or ventral lobe . . 4
4 (3) Hindwing Rs and M l stalked or coincident
(Fig. 8.1 M) Ypsolophidae
Hindwing Rs and M separate 5 diverse in facies, structure and bionomics. Six
5 (4) Hindwing with M3 and CuAl separate . .
Plutellidae
Hindwing with M3 and CuAl stalked . . .
Acrolepiidae
6(1) Wings greatly narrowed; hindwing vena-
tion markedly simplified, with at most 4
pre-CuP veins 7 and its applicability to tropical/south temperate
Wings less narrowed; hindwing venation
more nearly complete, with at least 6 pre-
CuP veins 8 (1976-77), Moriuti (1977), Hannemann (1977),
7 (6) Forewings simply lanceolate, R s 2 - 4 and
an M branch on common stalk from
pointed apex of discal cell; scape with
dense awning, but not expanded into genu-
ine eye cap Bedelliidae
Forewing apex narrowly produced, Rs2
arising before apex (which is not a single
point) of discal cell, or forewing without
closed discal cell; scape expanded into eye
cap (Fig. 8.4 J) Lyonetiidae
8 (6) Rs3 apical/postapical; monobasic W. Pa-
laearctic taxon, large micro-moth with
broad, angular wings (Fig. 8.2 N), associ-
ated with freshwater plants
Glyphipterigidae (Orthoteliinae)
Rs3 apical/preapical . . . . Yponomeutidae
Yponomeutidae. Autapomorphies few and obvi-
ously homoplasious. Ocelli absent. Pretarsus
('claw') of larval thoracic legs unusually long and
straight; absence of this trait in the endophagous
Argyresthiinae interpreted by Kyrki (1990) as
secondary. Larval proleg crochets usually (the
endophagous argyresthiines and a few ypono-
meutines excepted) bi-/multiserial; polarizing this
character is difficult, however, since while
crochets are uniserial in tineoids a bi-/multiserial
arrangement is predominant in those non-ditry-
sians and gracillarioids which do have crochets.
Other possible groundplan autapomorphies of
the family enumerated by Kyrki are even more
questionable: there is no obvious support for the
interpretation of the relative shortness of the
commonplace yponomeutoid pterostigma (which
does not extend distad beyond R) as derived;
moreover the Scythropiinae have a long pteros-
tigma, while the short kind also occurs within
the Yposolophidae. A difficulty with the inter-
pretation of the particularly strong melanization
of the male sternum VIII as a family autapomor-
phy is that the unspecialized state of this plate
in the subfamily Yponomeutinae must, then, be
secondary. The abrupt narrowing of the apical
hindwing area in front of Se + R (Fig. 8.1 L), a
specialization characteristic of many yponomeu-
tids (Minet 1986, Scoble 1992), also occurs else-
where in the superfamily (Fig. 8.1 MN).
Second segment of labial palp usually without
scale tuft. Larva with prolegs shorter than broad;
on segment IX SV bisetose.
The taxa brought together in this family are
groups are recognized and given subfamily status
in the present classification; their phylogenetic
interrelationships remain unclarified. This classi-
fication, as that of the Yponomeutoidea as a
whole, is primarily based on a study of the Hol-
arctic genera (Moriuti 1977, Kyrki 1984, 1990),
faunas remains to be tested.
Friese (1960), Kuznetzov & Stekolnikov
Gershenzon (1990).
Key to yponomeutid subfamilies
1 Chaetosema present; male hind tibia with-
out spurs, reduced (like the tarsus)
Attevinae
Chaetosema absent; male hind tibia with
spurs 2
2 (1) Antennal scape without pecten Praydinae
Antennal scape usually with pecten . . . 3
3 (2) Proboscis reduced, short and soft; pteros-
tigma extending distad beyond R
Scythropiinae
Proboscis well developed; pterostigma,
when present, not extending distad beyond
R 4
4 (3) Abdominal terga with normal scales only
Argyresthiinae
Abdominal terga with spines (modified
scales (Fig. 8.1 F) 5
5 (4) Male sternum VIII more strongly mela-
nized than VII; segment X processes with
specialized scales Saridoscelinae
Male sternum VIII not more strongly mel-
anized than VII; segment X processes un-
sealed Yponomeutinae
Yponomeutinae. Perhaps a paraphyletic as-
semblage. Uniform melanization of male abdo-
minal sterna VII and VIII diagnostic, but proba-
bly plesiomorphic; this applies also to spinose
abdominal terga (Fig. 8.1 F) shared only with the
following subfamily among yponomeutids.
The Yponomeutoidea 121

Fig. 8.1. Yponomeutidae: A, C - G , I - L , P; Ypsolophidae: , M; Plutellidae: O; Acrolepiidae: N; Glyphipterigi-

dae: H, Q. A, Thecobathra head; B, Ypsolopha head; C, Thecobathra maxillary base with palp; D - , Yponomeuta
genitalia (D female, E male, phallus lower right); FH 'unrolled' male abdominal segments VIIVIII of Metano-
meuta (F), Prays (G) and Glyphipterix (H); I - J , Argyreslhia male genitalia, note modified sensilla (highly magni-
fied in J); K, Argyreslhia moth in resting position; L, Yponomeuta venation; M, Ypsolopha hindwing venation; N,
Acrolepiopsis hindwing venation; O, Plutella larval segments I V - V I I I : P, Yponomeuta larva; Q, Orthotaelia pupa.
- B after Robinson et al. (1994); I - J , M after Moriuti (1977); C , after Friese (1960); F - H after Kyrki
(1984); L after Nielsen & Common (1991); after Zagulyaev (1990); O - P after Heppner (1987); Q after Kyrki &
Itmies (1986).
122 John S. Dugdale, Niels P. Kristensen, Gaden S. Robinson & Malcolm J. Scoble
Chaetosema absent. Antennal scape with pec-
ten. Maxillary and labial palps variable. Spurs
0-2-4. In male genitalia (Fig. 8.1 E) saccus and
segment X processes ('socii' in principal taxo-
nomic treatments) usually prominent, the latter
unsealed. As in the Saridoscelinae the phallus
usually has a basal 'scape'. Larva with cranial
seta VI long, PI below A f 2 - P 2 line.
A small genus group is characterized by plesi-
omorphic traits including 4-segmented maxillary
palp (Fig. 8.1 C; basal segment synscleritous with
stipes, hence palp described as 3-segmented by
Moriuti 1977), long 'basal fork' of forewing anal
veins, and hindwing with all three M-branches
present. This group has been given subtribe rank
(by Moriuti 1988 whose tribe Yponomeutini
equals the present subfamily Yponomeutinae),
but there is no evidence for its monophyly. At
least the genera Niphonympha (3 species, Eura-
sian, Mexico) and Thecobathra (13 species, S. E.
Asian) pertain to this grade; the known larvae
feed on leaves of Fagaceae and Hamamelida-
ceae.
In the bulk of the Yponomeutinae the maxil-
lary palp is only 2-segmented, the forewing anal
fork is very short, and the hindwing has only two
M veins (Fig. 8.1 L). The near-cosmopolitan (not
New Zealand) Yponomeuta ('ermine moths', 80 +
species; Fig. 8.1 P, 8.2 B) mostly have larvae liv-
ing in communal webs on (botanically diverse)
trees and scrubs. Mass occurrence, with forma-
tion of highly voluminous and conspicuous webs
occurs in a number of species, and some have
economical importance. Recent concerted inves-
tigations of the ecology, behaviour, physiology
and evolutionary biology of European species
have been reviewed by Menken, Herrebout &
Wiebes (1992). The more narrow-winged Zelleria
(50 + species) have a characteristic resting posi-
tion with the head end close to the substrate and
the abdomen lifted (as in argyresthiines, Fig.
8.1 K); ecology diverse, comprising webmakers
on broad-leaved trees/scrubs as well as pine nee-
dle miners. The closely related Xyrosaris (Aus-
tralasian/Neotropical) is aberrant in having a
prominent ventral scale tuft on labial palp seg-
ment 2 (recalling that present in some other ypo-
nomeutoid families). Other principal genera in-
clude Kessleria (Huemer & Tarmann 1991), Cal-
lithrinca and Swammerdammia. Ocnerostoma
(Holarctic) is exceptional in that the larvae (nee-
dle miners) have bisetose prothoracic L (proba-
bly related to the endophagy) and uniserial pro-
leg crochets; it belongs to a small group with re-
duced proboscis. Altogether the subfamily com-
prises 360 + species worldwide.
In the following subfamilies the male sternum
VIII is more strongly sclerotized than VII (Fig.
8.1 G), and often of a characteristic V- or Y-like
shape.
Saridoscelinae. Monogeneric, differing from
preceding mainly in male sternum VIII configu-
ration, and in having the socii set with fringed
sensilla recalling those of the Argyresthiinae.
Hindwing with all three M branches.
Saridoscelis (4 described species and at least
twice as many undescribed, Asia; Fig. 8.2 C).
Larvae in communal webs on Ericaceae.
Scythropiinae. Monogeneric. Probable autapo-
morphies are the reduced proboscis and dorsally
hairy hind tibia.
Scape with pecten. Maxillary palp 4-seg-
mented. Pterostigma extending distad from R.
Abdominal terga without spines. Phallus without
basal 'scape'. Socii unsealed.
Larva with cranial seta VI long, PI below
A f 2 - P 2 line.
Scythropia (2 species, European; Fig. 8.2 D).
Larva on woody Rosaceae, initially leafmining,
eventually feeding externaly in a communal web.
Attevinae. Probable autapomorphies are the
presence of chaetosema, the reduction of hindleg
tibia and tarsus (particularly in males where tib-
ial spurs are absent), the presence of two sub-
ventral setae in larval meso- and metathorax,
and the concealment of the labial palp in the
pupa.
Large, colourful (markings often red/orange/
yellow) and probably aposematic micro-moths.
Scape without pecten. Maxillary palp with 3 seg-
ments. No pterostigma. Abdominal terga with-
out spines. Phallus without basal 'scape'. Tegu-
men bearing an unsealed prong (probably an-
other subfamily autapomorphy); socii complex;
gnathos W-shaped.
Larva with cranial seta VI short, PI not below
A 2 - P 2 line.
Atteva (50+ species, pantropical; Fig. 8.2 A)
larvae live in communal webs recalling those of
Yponomeuta, but looser; host families include Fa-
baceae, Simaroubaceae, Lauraceae and Aralia-
ceae.
Praydinae. Probable autapomorphy an en-
larged (broader than long) male sternum VIII
(Fig. 8.1 G). Female anterior apophyses greatly
reduced, without branched costa at base (Mori-
uti 1977). Loss of forewing 'chorda' (mentioned
by Kyrki 1984) parallelled in several Yponomeu-
tinae.
Scape without pecten. Maxillary palp 1-seg-
mented. Pterostigma not extending distad from
R. Abdominal terga without spines. Phallus
without basal 'scape'. Socii unsealed. Larva with
cranial seta VI short, PI not below Af2P2 line.
Prays (25 + species, Old World; Fig. 8.2 E).
Larvae on diverse woody plants, endophytic
(leaf miners or borers in flower buds or shoots)
in all instars, or older larvae feed externally in
light webs; P. citri and P. oleae are pests on citrus
and olives respectively.
Argyresthiinae. Principal autapomorphies are
in male genitalia (Fig. 8.11J): vinculum more
or less produced laterally and lacking a median
saccus; socii set with peculiar fringed sensilla
The Yponomeutoidea 123

Fig. 8.2. Hyponomeutidae: A - F ; Ypsolophidae: G - H ; Plutellidae: I - K ; Acrolepiidae: L; Glyphipterigidae:

M - N. A, Atteva punctella; B, Yponomeuta padella; C, Saridoscelis sp.; D, Scythropia crataegella; E, Prays fraxi-
nella; F, Argyresthia brockella; G, Ypsolopha dentella, H, Ochsenheimeria bubalella\ I, Proditrix tetragma; J, Eido-
phasia messingiella; K, Plutella xylostella; L, Acrolepiopsis assectella; M, Glyphipterix haworthana; N, Orthotaelia
sparganella. Uniform magnification; scale (milimetres) in A.

(somewhat similar formations occurring in Sari- Argyresthia (150 + species, near-cosmopolitan;

doscelinae). Hindwing M l and M2 long-stalked
(shorter stalks occur in a few Yponomeutinae).
Larvae usually endophagous, rarely in loose
webs; with simple pretarsi of thoracic legs (auta-
phomorphic character reversal?) and uniserial
crochets on prolegs. Resting position of adult
characteristic (Fig. 8.1 K), with head end lowered
and abdomen lifted (as in Yponomeutinae-Ze/-
leria).
Scape with pecten. Maxillary palp 1-seg-
mented. Pterostigma not extending distad from
R. Abdominal terga without spines. Phallus
without basal 'scape'.
Larva with cranial seta VI short, PI below
A f 2 - P 2 line.
Fig. 8.2 F) are narrow-winged moths with di-
verse, often metallic forewing patterns, larvae
bud-, fruit-, leaf- and twig-miners on a variety of
conifers and woody dicots, some attaining pest
status.
Friese (1969), Gershenzon (1990).
Putative clade Ypsolophidae +
(Plutellidae + Acrolepiidae + Glyphipterigidae +
(Heliodinidae + Bedelliidae + Lyonetiidae))
The monophyly of this entity is weakly sup-
ported. Kyrki (1990) enumerated three possible
autapomorphies: Ventral scale tuft on second la-
bial palp segment (Fig. 8.1 B; absent from many,
124 John S. Dugdale, Niels P. Kristensen, Gaden S. Robinson & Malcolm J. Scoble
if not most, Glyphipterigidae and from all mem-
bers of the heliodinid/bedelliid/lyonetiid assem-
blage). Larval prolegs (Fig. 8.10) longer than
wide (not applicable to endophagous glyphipteri-
gids and lyonetiids with reduced prolegs, nor to
certain genera currently placed in Plutellidae).
Larva with only one SV seta on IX, a trait recur-
rent in several endophytic micro-moth larvae.
(Note: two setae are present in some ypsolophids
and heliodinids; it remains unclear whether this
is a genuine plesiomorphy or a character re-
versal.)
Ypsolophidae. Probable autapomorphies (Kyrki
1990) include the stalked (or coincident) hind-
wing Rs and Ml (Fig. 8.1 M), the anteriorly
deeply bilobed male tegumen, the membranous
and strongly spinose tubular anellus, the 2 cor-
nuti/cornutus groups in phallus, the markedly
elongate female apophyses, the exit site of the
ductus seminalis on ductus bursae close to os-
tium, the signum configuration (elongate plate
with two transverse ridges), and loss of pupal
cremaster setae.
Ocelli usually well developed. Pterostigma,
when present, not extending distad beyond vein
R.
Larval proleg crochets, when present, uni-
serial.
The apparently close relationship between the
two constituent subfamilies has only recently
been recognized. The Ypsolophinae have mostly
been associated with the phenetically similar Plu-
tellidae; the family distinction is based on the as-
sumption (which needs corroboration, see be-
low), that a taxon centred on Plutella is cladisti-
cally closer to the Glyphipterigidae and the Heli-
odinid/Bedelliid/Lyonetiid assemblage. The Och-
senheimeriinae were consistently associated with
the tineoid families until Kyrki's 1984 study, and
their status deserves continued scrutiny; it can-
not be ruled out that they are actually cladisti-
cally subordinate in the ypsolophines.
Ypsolophinae. Monophyly not well supported.
Kyrki (1984) suggested the phallic coecum to be
an autapomorphy.
Ypsolopha (Holarctic) is the principal genus
with 120 + species, some of which have the fore-
wing termen sinuate or even markedly falcate
(Fig. 8.2 G); larvae in open webs on leaves of
diverse (usually woody) plants.
Kuznetzov & Stekolnikov (1976-77), Moriuti
(1977), Zagulyaev (1990).
Ochsenheimeriinae. A compact group of moths
with characteristic habitus (Fig. 8.2 H). Possible
autapomorphies include the 'shaggy' vestiture of
the head and labial palp, the small size of the
compound eyes (probably reflecting the moths'
diurnal activity, as do the well developed ocelli),
absence of pterostigma, the strongly sclerotized
anal cone of the male, the asymmetrical ostium
bursae of the female, and the reduced crochet
complement on the larval prolegs (a single row,
or complete absence).
Larvae initially leaf miners in Poaceae
(mainly), Cyperaceae and Juncaceae; later in-
stars are stem borers.
10+ valid species. Ochsenheimeria and the
closely related Aridomeria are Palaearctic;
O. vacculella, occasionally a serious cereal crop
pest in the Old World, has been casually intro-
duced into N. America.
Davis (1975, 1987), Karsholt & Nielsen (1984),
Zagulyaev (1988, 1990), Kuznetznov & Stekolni-
kov (1993).
Putative clade Plutellidae + Acrolepiidae +
Glyphipterigidae + (Heliodinidae + Bedelliidae +
Lyonetiidae)
The monophyly of this assemblage was suggested
by Kyrki (1990) on the following: loss of the
gnathos and reduction of the tegumen (which
dorsally at most forms a narrow band) in the
male genitalia; protruded pupal thoracic spira-
cle; larval feeding on herbaceous plants. How-
ever, there are serious difficulties with this
scheme: contrary to the statements of Kyrki &
Itmies (1986) a gnathos is actually present in
members of the Plutellidae, Heliodinidae and Ly-
onetiidae (e. g. Common 1990, Kuroko 1964).
And as noted by Kyrki himself, the lyonetiids do
not have protruded pupal spiracles, and their
host spectrum includes many woody plants; ac-
ceptance of the suggested phylogeny therefore
requires, that these traits are interpreted as char-
acter reversals.
Plutellidae. As delimited by Kyrki (1990) the
family included the Acrolepia group of genera
which has often been ranked as a separate fam-
ily. The alleged autapomorphy of the family thus
circumscribed is the structure of the female post-
vaginal lamella, which consists of two setose
lobes; however, a somewhat similar lamella (with
the lobes more or less distinctly united by mela-
nized cuticle, see Moriuti 1977) is widespread in
Yponomeutidae and Ypsolophidae, and it may
be plesiomorphic at this level. Kyrki further
mentioned the large-meshed cocoon structure,
but the latter also occurs in some yponomeutids
(Heppner 1987). Both traits are absent in some
S. Hemisphere taxa currently included in the Plu-
tellidae. In the absence of good synapomorphies
of the Plutella and Acrolepia groups they are here
treated as separate families.
According to Kyrki (1984) the Plutella group
in the restricted sense is characterized by male
genitalia with 'teguminal processes' curved, sur-
rounding anal tube; the processes in question are
actually gnathal.
Ocelli usually distinct, rudimentary in some
S. hemisphere genera currently placed in the
The Yponomeutoidea
family. Pterostigma extending beyond vein R.
Female retinaculum a subcostal scale-patch.
Larval prolegs usually slender (Fig. 8), with
uniserial crochets.
Plutella (40 + species, cosmopolitan) larvae
mostly feed on Brassicaceae, living in loose webs
and skeltonizing leaves. The genus includes the
diamondback moth P. xylostella (= maculi-
pennis) (Fig. 8.2 K), an ubiquitous pest species
which is one of the few micro-moths renowned
for long-range migrations; its biology and con-
trol are reviewed by Talekar & Shelton (1993).
Other principal genera include Leuroperna, Rhi-
gognostis (Kyrki 1988), Eidophasia (Fig. 8.2 K).
The monobasic Embryonopsis from Indian
Ocean subantarctic islands is brachypterous in
both sexes. Proditrix (Fig. 8.21; New Zealand,
also known from Tasmania) species are note-
worthy for their large size (some exceeding
50 mm wingspan) and their larval adaptations
for feeding in intermittently waterfilled sites in
crowns of large monocots belonging to several
families (Dugdale 1987), or, in the case of the
Tasmanian taxon, in a dicot (Richea, Epacri-
dacreae) with a similar growth form. Their pro-
legs are blunt, with partially biserial crochets and
the cocoon is non-meshed, so the systematic
placement is tentative. In this and other New
Zealand 'Plutellidae' non-meshed cocoons have
anterior and posterior openings, and the larval
skin is protruded from the latter; this is most
probably an autapomorphic condition.
Kuznetzov & Stekolnikov (1976-77), Moriuti
(1977), Zagulyaev (1990).
Acrolepiidae. As noted above this taxon has re-
cently been included in the Plutellidae; like the
bulk of the latter they have retained ocelli. Prob-
able autapomorphies (Kyrki 1984) include details
in the male genitalia (reduction of the tegumen,
teguminal processes and gnathos, basal widening
of the phallus) and stalking of hindwing
Ml + M2 and M3 + CuAl (Fig. 8.1 N). Females
lack the ventral costa at the base of the anterior
apophysis, but this apomorphy is recurrent in
some S. Hemisphere taxa currently placed in Plu-
tellidae.
Adults are crepuscular/nocturnal. Larvae are
leaf miners or borers in stems, flower buds, and
seeds. While the main pest species (including the
leek moth Acrolepiopsis assectella, Fig. 8.2 L,
studied in detail by Frediani, 1954) are Liliaceae
feeders, the host spectrum also includes a variety
of dicot families such as Asteraceae, Lamiaceae
and Solanaceae.
95 + species; worldwide, but more than half
are Palaearctic. Acrolepia, Acrolepiopsis and Dig-
itivalva.
Kuznetzov & Stekolnikov (1976-77), Gaedike
(1970, 1983, 1986), Zagulyaev (1990).
Glyphipterigidae. This family long comprised an
assemblage of unrelated diurnal micro-moths,
125
including choreutoids and immoids. Heppner
(1977, 1985, 1987, 1991) assigned the Glyphip-
terigidae s. str. to the Copromorphoidea with
which they share prominent protuberances bear-
ing the larval spiracles (particularly on thorax
and VIII) and the dorsocaudal displacement of
the larval spiracle VIII. However, the different
abdominal sternum II configurations (Kyrki
1983, Minet 1983, contra Heppner 1977) and the
non-apical vom Rath's organ contradict a close
relationship between Glyphipterigidae s. str. and
copromorphoids. The placement of the former
within the Yponomeutoidea received additional
support with Kyrki & Itmies' arguments (1986)
for their close affinities to Orthotelia, whose male
has retained typical yponomeutoid pleural lobes.
Probable autapomorphies of the family as now
delimited (Kyrki 1990) include the absence of an
antennal pecten/scale flap, absence of segment X
('teguminal') processes; larvae endophagous
(hence with shortened prolegs) and primarily
monocot feeders, their cranial setae F l , Afl, Af2,
A2, P2, Ol, G2 (= 03 auct.) and S02 short, and
the abovementioned larval spiracular traits.
Kyrki & Itmies (1986) mentioned two addi-
tional possible family autapomorphies in larval
structure, viz. the absence of an adfrontal 'punc-
ture' and the presence of sclerotized shields on
segments VIII and IX. The larval prothoracic L
group is almost always bisetose, but a New
Zealand glyphipterigine has retained a vestigial
third seta (Dugdale, in Heppner 1987). The pu-
pal thoracic spiracle is borne on a prominent
process.
Orthoteliinae. Monobasic. Orthotelia sparga-
nola (Fig. 8.2 N) is a rather large and broad-
winged, crepuscular micro-moth without ocelli.
Vertex with tufts of raised narrow scales. Labial
palpi prominent, upcurved. Larva stem- and
leafboring in freshwater plants belonging to sev-
eral monocot families; prolegs with crochets in
two transverse rows (on anal pair a single). Pupa
(Fig. 8.1Q) with prominent, bifid frontal process
and segments V - V I I I with spiny transverse
ridges; pupal skin protruded from foodplant as
adult emerges, an unusual condition within the
Yponomeutoidea. W. Palaearctic.
Kyrki & Itmies (1986).
Glyphipteriginae. A probable autapomorphy is
the more or less conical male segment VIII (Fig.
8.1 H) with an apparently unusually enlarged ter-
gum, now interpreted (Kyrki & Itmies 1986) as
being fused with the pleural lobes, which are not
distinguishable as such (Kyrki 1984). Kyrki & It-
mies also considered the vestigial forewing M-
stem and CuP, as well as the approximation of
the hindwing M3 and CuA (Fig. 8.3 B), to be
probable subfamily autapomorphies; these spe-
cializations are commonplace elsewhere among
yponomeutoids, but Orthotelia exhibits the plesi-
omorphic states. The diurnal activity of adult
glyphipterigines, and their striking habit of
126 John S. Dugdale, Niels P. Kristensen, Gaden S. Robinson & Malcolm J. Scoble

Fig. 8.3. Glyphipterigidae: - C ; Heliodinidae: D; Bedelliidae: E; Lyonetiidae: F - J . A, Glyphipterix head; B,

Glyphipterix venation; C, Glyphipterix sternum II, arrow indicates 'venula'; D, Heliodines venation; E, Bedellia
hindwing venation; F, Leucoptera larval head; G, Lyonetia 'ovipositor' apex; H, Lyonetia venation; I, Leucoptera
mines on Laburnum, J, Lyonetia head. A after Robinson et al. 1994; , D, H after Nielsen & Common 1991; C
after Minet (1983); E, G after Kuroko (1964); F after Davis (1987); I after Hering, E. M., Biology of Leaf Miners
(1951, Junk); J original (B. Rubaek del.).

rhythmically raising and lowering the back-
folded wings in repose, may be further derived
groundplan traits.
Moths mostly small, wingspan range 4,5
30 mm (Heppner 1985), usually ca. 10 mm.
Ocelli prominent (Sericostola excepted, Heppner
1990), chaetosemata absent; head smooth-scaled
(Fig. 8.3 A). Labial palpi usually upturned,
sometimes with prominent ventral scale tuft on
segment 2. Forewing outline variable, in many
Glyphipterix termen concave; pattern (Fig.
8.2 M) usually with light transverse lines or bars,
often with prominent metallic markings. Hind-
wings narrower than forewings. Males often with
coremata on VIII, sometimes on valvae (not in
both locations at the same time, Heppner 1985);
anal cone long, prominent.
Larval proleg crochets sparse, in lateral penel-
lipse or transverse band, or completely absent.
Larvae mostly borers/miners in leaves or flow-
ering stems of Poaceae, Cyperaceae, Juncaceae,
Zingiberaceae and Araceae, but also of some di-
cot families including Urticaceae and Crassula-
ceae. The mesh cocoon is spun in the cavity made
by the larva, or away from the food plant.
383 species (Heppner 1991). Many of the 20 +
genera are monobasic, and about two-thirds of
the species are placed in the cosmopolitan genus
Glyphipterix.
Arita (1987), Arita & Heppner (1992), Diako-
noff (1986), Kuznetzov & Stekolnikov (1976-
77), Heppner (1982, 1985), Kuznetsov (1990).
Clade Heliodinidae + Bedelliidae + Lyonetiidae
The monophyly of this assemblage was suggested
by Kyrki (1990) on the basis of the following
probable synapomorphies of its constituent fami-
lies: maxillary palp reduced, at most with 2 seg-
ments (parallelled in subordinate taxa elsewhere
in the superfamily). At most 8 branches present
in forewing Rs/M/CuA group (the identity of in-
dividual branches cannot be identified, but it is
assumed that one and the same branch has been
The Yponomeutoidea
lost in all families; Kyrki's inference that there
are "at most 4 M and CuA veins" may well be
correct, but it cannot be excluded that it is an
Rs branch, which has been lost). Pupa without
distinct maxillary palp, with strong lateral ridges,
and with immovable abdominal segments. The
consistent absence of a ventral scale tuft on labial
palp segment 2 may also be seen as a groundplan
autapomorphy (parallelled in Plutellidae-Acro-
lepiinae and many Glyphipterigidae-Glyphipteri-
ginae). Kyrki also considered an open discal cell
in the hindwing to be synapomorphic of these
families, but some Lyonetiidae-Cemiostominae
do have the cell closed (Kuroko 1964). All mem-
bers of this assemblage have very narrow wings,
and the simplifications of the venation (which
may be very pronounced in subordinate taxa) are
evidently related to this; similar modifications
are recurrent in narrow-winged members of
other micro-moth superfamilies. Kyrki noted
that particularly the assignment of the Lyonetii-
dae to this assemblage is problematical, since
they have (retained?) a non-protruded pupal tho-
racic spiracle and at least the lack of ocelli could
be taken as indication that they are most closely
related to, or even subordinate in, the Ypono-
meutidae.
Relationships within this assemblage remain
unresolved. As noted by Kyrki a sistergroup rela-
tionship between bedelliids and heliodinids is
suggested by characters in the larval spinneret
and thoracic chaetotaxy as well as pupation
mode; on the other hand the head scaling, ab-
sence of ocelli and the hind tibial vestiture sug-
gest a sistergroup relationship between bedelliids
and lyonetiids.
Kyrki also mentioned a very prominent
"fringe" on the underside of the larval labrum as
a possible bedelliid/lyonetiid synapomorphy, but
the epipharyngeal armature is actually somewhat
differently developed in these taxa (Jayewick-
reme 1940).
Heliodinidae. Probable autapomorphies (Kyrki
1984): entire head scaling smooth, and forewing
CuP vestigial (Fig. 8.3 D) (states also encoun-
tered in Glyphipterigidae-Glyphipteriginae);
abdomen non-spinose; pupa with stiff lateral and
dorsal bristles. Many taxa previously included in
the family (Meyrick 1914) have no close relation-
ships to Heliodines and are now placed in various
families in the Gelechioidea, or even (Schrecken-
steinia) in the Apoditrysia.
Small moths with ocelli but no chaetosemata;
male antennae typically thickened, scape without
pecten/scale flap. Hind tibia smooth scaled on
dorsal side. Forewings with metallic markings
(Fig. 8.4 A). Female frenulum a single bristle, ret-
inaculum subcostal, of short recurved bristles.
Larval spinneret unusually long and VI unde-
veloped on thorax (similarities with Bedeliidae,
Kyrki 1990); prolegs in uniordinal circle.
127
Adults are mostly diurnal. Larvae are leaf
miners, borers in stems, buds/fruits (even in gall
midge galls) or feeding, sometimes in a commu-
nal web, externally on leaves. Hosts include a
variety of dicot families (Chenopodiaceae, Ny-
taginaceae, Araliaceae, Onagraceae and Ai-
zoaceae). Pupation in larval shelter or externally,
in some taxa in a loose silk webbing.
50 + species, the great majority of which are
in the New World; stated by Heppner (1991) to
be absent from the Afrotropical region, but rep-
resented here by Hierophanes. About a dozen
genera, Heliodines being the largest and most
widespread.
Meyrick (1914), Heppner (1987), Fal'kovich
(1990), Common (1990).
Bedelliidae. A monogeneric taxon previously in-
cluded in the Lyonetiidae, but as noted above the
evidence concerning its affinities is ambiguous.
Probable autapomorphies (Kyrki 1984): Fore-
wing Rs2, Rs3, Rs4 and ?M1 + 2 on common
stem from pointed apex of discal cell. Male seg-
ment VIII without pleural lobes. Larva with only
2 lateral setae on abdominal segments IVII.
Pupa with conical process on frons. The non-spi-
nose adult abdomen (Kuroko 1962) and the
shortening of larval prolegs III and VI relative
to those of I V - V (Common 1990) may be con-
sidered additional family autapomorphies, the
former parallelled elsewhere among yponomeu-
toids.
Small greyish moths. Head with tuft of pili-
form scales on vertex, face smooth-scaled; ocelli
and chaetosemata absent; scales on scape form-
ing dense pecten, but no true eye cap. Hind tibia
hairy on dorsal side.
Larva with long spinneret and undeveloped
thoracic VI setae (similarities with Heliodinidae),
and prominent epipharyngeal fringe (similarity
with Lyonetiidae) (Kyrki 1990).
Larvae leafminers, feeding on several plant
families, e. g. Poaceae, Liliaceae, Urticaceae and
Convolvulaceae. Initial mine a narrow gallery.
Later instars blotch miners and eroders, which
may make several mines and even change leaf;
frass is ejected from the mines. An open silk net-
work is spun outside the mined leaf. Pupation in
a loose web below or above the mine, without
formation of a genuine cocoon.
Bedellia (18 described species; Fig. 8.4 B) is
distributed in all regions except the Neotropical;
there is a small radiation of the genus in Hawaii,
where at least 14 species occur. The very wide-
spread B. somnulentella is occasionally a pest on
Ipomoea.
Zimmerman (1978), Davis (1987), Kuznetzov,
Kozlov & Seksjaeva (1988).
Lyonetiidae. Probable autapomorphies (Kyrki
1990): scape slightly flattened, scales forming
eye-cap (Fig. 8.3 J). Forewing apices markedly
128 John S. Dugdale, Niels P. Kristensen, Gaden S. Robinson & Malcolm J. Scoble
Fig. 8.4. Heliodinidae: A; Bedelliidae: B; Lyonetiidae: CD. A, Heliodines nodosella\ B, Bedellia
C, Lyonetia clerckella; D, Leucoptera laburnella. Uniform magnification; scale (milimetres) in A.
produced. Larva with head (Fig. 8.3 F) flattened,
cranial setae F l , Afl, A , LI and G2 (= 03)
short, labrum at most with four pairs of dorsal
setae, and abdominal setae D1 and D2 approxi-
mated, D1 more laterad than D2. Two first lar-
val instars without prolegs, and frequently also
without thoracic legs (Davis 1987, Common
1990). Pupal thoracic spiracle non-protruded
(plesiomorphy or autapomorphic character re-
versal?).
Scaling on vertex variable, on face consistently
smooth; ocelli and chaetosemata absent. Hind
tibia hairy on dorsal side.
Larval epipharynx with prominent fringe
(similarity with Bedelliidae, Kyrki 1990).
Larvae leaf- (occasionally twig-) miners. Pupa-
tion outside the mine.
Kuroko (1964), Bradley & Carter (1982), Em-
met (1985), Davis (1987), Kuznetsov, Kozlov &
Seksjaeva (1988), Seksyaeva (1990), Mey (1994).
Lyonetiinae. Probable autapomorphies (Kyrki
1984): extreme narrowing of the wings and abdo-
minal terga. Female 'ovipositor' (Fig. 8.3 G)
piercing, strongly sclerotized, pointed and
equipped with small lateral/ventral teeth. The
non-spinose abdomen may also be considered
autapomorphic (but very homplasious, cp. Be-
delliidae above).
Vertex always with tuft of piliform scales (Fig.
8.3 J) Antennae long, exceeding forewings. Fore-
wing often with dark markings in basal half.
Larval mines may be narrow galleries
throughout, or gallery-blotch mines. Pupa usu-
ally in a flimsy cocoon suspended by silk threads,
but occasionally naked (Davis 1987). Hosts are
woody dicots belonging to a dozen (partly unre-
lated) families.
Cosmopolitan; Heppner (1991) counted 89 de-
scribed species (only 1 Afrotropical). Lyonetia is
the principal genus; L. clerckella (Fig. 8.4 C) is
occasionally a pest on rosaceous fruit-trees in the
Palaearctic. Stegommata in the Australian region
are Proteaceae miners.
somnulentella\
Cemiostominae. Probable autapomorphies
(Kyrki 1984): forewing with tornai patch of
raised metallic scales. Larval head with adfrontal
ridges subparallel, reaching epicranial notch
(Fig. 8.3 F). The adult's spinose abdomen is not
here considered apomorphic.
Vertex with tuft of piliform scales, or smooth-
scaled like the face. Antenna not longer than
forewing. Forewing mostly with uniform white/
greyish ground colour and a characteristic
pattern (Fig. 8.4 D) of brownish and black
bands/bars in the apical half (strikingly similar
to that of GracillariidaePhyllocnistinae). In
resting moths the apical black bars in the fringes
are believed to mimic the antennae of an insect
head (Emmet 1985; this phenomenon is recurrent
in several other groups of micro-moths).
Larval leaf mines are blotches or gallery-
blotches, twig mines narrow galleries through-
out. Some blotch miners have a characteristic
habit of smearing the viscous faeces in concentric
rings on the upper epidermal wall of the mine
(Fig. 8.31). Pupation in a usually fusiform co-
coon, spun on the hostplant or among leaf litter.
Principal foodplants belong to Fabaceae, fol-
lowed by Rubiaceae, but about a dozen dicot
host families have been recorded, and even the
monocot family Liliales.
120+ species, in all regions. Main genera in-
clude Paraleucoptera, Perileucoptera (with coffee
pests: P. coffeella in the Neotropics and Masca-
rene Isles, P. meyricki in Africa) and Leucoptera
(L. malifoliella is a Eurasian orchard pest,
L. psophocarpella a pest of Psophocarpus in
New Guinea).
Acknowledgments
Comments from Mr. Frank Hsu (Heliodinidae)
and Mr. Ole Karsholt on our manuscript drafts
are gratefully acknowledged.
The Yponomeutoidea
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Agassiz (1996) monographed the British Ypono-
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 9. The Gelechioidea
Ronald W. Hodges
Gelechioidea are a large group with 1,425 genera
and 16,250 described species. They are generally
distributed throughout the world, becoming less
numerous at higher latitudes. Representation of
families and subfamilies within geographic re-
gions varies greatly: Cosmopteriginae are ex-
tremely speciose in one of four genera in Hawaii
(Zimmerman 1978); Oecophorinae are the pre-
dominant Lepidoptera in Australia (Common
1990), the second most species numerous taxon
in New Zealand (Dugdale 1988), but absent from
Hawaii before the advent of man (Zimmerman
1978); Gelechiidae far outnumber Oecophori-
dae-Elachistidae in America north of Mexico
(Hodges 1983); and Stenomatinae are extremely
abundant, 1,095 species, in the Neotropical re-
gion (Becker 1984), but have only nine species in
the Antilles (Duckworth 1969). Gelechioidea
have been very poorly collected except for very
few areas. They may be the least well described
large superfamily of Lepidoptera. In the Nearctic
region approximately 30% of Gelechiidae are de-
scribed, 10% of Scythridinae (Landry 1991),
20% of Coleophorinae, 25% of Momphinae,
6 0 - 7 0 % of Cosmopterigidae, 30% of Elachisti-
dae (unpublished); for the Neotropical region
Stenomatinae may be 50% described, Gelechi-
idae < 20% described (V. O. Becker, unpub-
lished); for all of Subsaharan Africa only Gelech-
iidae and Lecithoceridae of South Africa have
been studied in any detail (Janse 1949-63), sug-
gesting that the 1,027 species of Gelechioidea
(Vri & Kroon 1986) are but a small part of the
total. The oecophorine fauna of Australia is
about 50% described (Common, unpublished).
Adults have forewing lengths of 2.0 mm
(Siskiwitia falcata) to 35 mm (Cryptophasa setio-
tricha) with those of most species 5 15 mm.
Most are active at dusk or night; many genera
are diurnal, and they often are very brightly col-
ored. Alpine and arctic species may be diurnal
but are very somberly colored. Larvae feed on
dead/decaying plant or animal tissue, virtually
any living plant tissue, and sometimes on living
invertebrates. Most species are concealed exter-
nal feeders; they usually construct a silk shelter
or tie leaves. Some are root or stem gall makers,
leaf miners, or case bearers (Bradley 1987; Pow-
ell 1980). Pupation usually occurs in the larval
chamber, but the pupa may be fully exposed. It
remains in situ on eclosion of the adult. Eggs
normally are laid singly (Common 1990).
Except for the Palearctic region very little is
published on the immature stages. Most reports
in the literature are from rearings of field-col-
lected larvae or pupae. Few species have been
reared through a complete cycle, and very few
voucher specimens document this information.
One apomorphy defines Gelechioidea: the
haustellum has overlapping scales on the dorsal
surface from the base extending variously to 1/2
its length (Common 1970, 1990; Hodges 1986;
Minet 1990); however, this state also occurs in
Pyraloidea and in Choreutoidea among ditry-
sians. Pyraloidea have paired tympanic organs at
the base of the abdomen that never are present
in Gelechioidea; and Choreutoidea have naked,
minute 1/2-segmented maxillary palpi, a state
that does not occur in Gelechioidea. The pupa
remains in its chamber or in place on adult eclo-
sion, a behavioral character shared with Ypono-
meutoidea (Common 1990).
Characters common to most Gelechioidea are
1) maxillary palpus 4-segmented, scaled and
folded over/parallel with base of the haustellum
(probable apomorphy); 2) labial palpus upturned
and third segment long and acute; 3) head
smooth scaled; 4) chaetosemata absent; 5) dorsal
surface of hind tarsus with long, slender scales;
6) pupal antennae meeting mesially before their
apexes (probable apomorphy); and 7) larval ab-
dominal segments 1 - 8 with setae L1/L2 closely
approximated or on the same pinaculum (Com-
mon 1970, 1990; Hodges 1978, 1986; Minet 1990,
1991; Nielsen 1989).
Gelechioidea have the following characters:
Head smooth scaled; frons evenly convex, occa-
sionally with processes; haustellum usually de-
veloped, rarely absent, scaled on anterior sur-
face, at least at base; labial palpus three seg-
mented, usually upturned, rarely porrect with
third segment slightly upturned, third segment
normally acute; maxillary palpus four segmented
(rarely one-three segmented), scaled, folded over
base of haustellum or downturned and parallel
with base of haustellum; ocellus present or ab-
sent; antenna simple or ciliate, rarely pectinate
or bipectinate, pecten present or absent; chaeto-
semata absent. Forewing broad to narrowly lan-
ceolate, pterostigma present or absent, cell open
or closed, 10 to 12 veins present, CuP present or
absent, 1 A and 2 A forked basally; retinaculum
subcostal in male, subcostal to subcubital in fe-
male, sometimes with secondary wing coupling
structures. Hindwing broad to linear, fringe
short to over five times width of wing; cell closed
or open; six to nine veins present; frenulum a
single acanthus in male, usually three acanthi in
female; cubital pecten sometimes present; base of
wing sometimes with hair pencils. Wings rarely
132
reduced or non-functional in , then lanceolate/
linear or short lobes. Legs: epiphysis present; tib-
ial spurs 0, 2, 4; hindtibia usually with numerous
long, slender scales on dorsal surface. Abdomen
frequently with spiniform dorsal setae; some-
times with secondary hair pencils from lateral or
ventral surface in male; segments seven and eight
sometimes modified in male; S2 with tineoid ven-
ulae and apodemes, sometimes venulae or apo-
demes, infrequently neither. Male genitalia sim-
ple and symmetrical to highly specialized by re-
duction, modification, and asymmetry; aedeagus
sometimes ankylosed, rarely unsclerotized; val-
vae sometimes reduced and replaced functionally
by modifications of preceding segments. Female
ovipositor usually adapted for external place-
ment of eggs but sometimes for piercing and in-
ternal placement of eggs; ostium bursae ventro-
mesial, rarely lateral or dorsal. Egg normally
flat, rarely upright in Coleophoridae and
Scythridinae (Common 1990); chorion ridges
frequently present; texture rough (Peterson 1967)
or smooth. Larva sometimes with submental pit,
almost always with six stemmata, rarely three or
none; prothorax with three L setae, rarely two;
setae LI and L2 usually approximate on abdo-
men, L2 anterior to dorsal of LI; A8 usually
with SD anterodorsad to dorsad of spiracle; A9
with D2 setae on separate pinacula, SD setae
often hairlike; anal comb sometimes present in
Gelechiidae; crochets uniserial, in a circle, el-
lipse, or two transverse parallel rows, rarely very
few, rarely a mesoseries. Pupa obtect, frontocly-
peal suture present or absent, almost always
without dorsal spines, maxillary palpus usually
visible, labial palpus and prothoracic femur
sometimes exposed, antennae usually touching
mesially before apexes, abdominal segments five-
seven movable in male, five and six in female,
lateral condyles sometimes present on segments
5/6 and 6/7; segment nine sometimes with paired,
ventrolateral, anteriorly directed lobes; segment
10 with recurved setae or cremaster.
Classification. Fracker (1915) proposed Gelechi-
oidea, but he did not indicate characters to de-
fine them. In his key to families the included taxa
arise in different places; Elachistidae and Col-
eophoridae were associated with the tineoid
series but without superfamily assignment. He
stated (1915: 45) "The grouping and the division
into families is principally a compilation from
Walsingham and from published and unpub-
lished work of August Busck. At the same time
we believe that nothing in the larval structure
precludes this arrangement and that it is a more
natural classification than any published more
than a decade ago."
Mosher (1916) recognized the pupae of Gel-
echioidea as having "... a distinct epicranial su-
ture, with the caudal portions of the antennae
lying adjacent on the meson and usually separat-
Ronald W. Hodges
ing at their distal ends to expose the metathora-
cic legs." She included Coleophoridae in Ypono-
meutoidea.
Subsequently, Forbes (1923), McDunnough
(1939), Common (1970, 1990), Bradley (1972),
Hodges (1978), Kuznetsov and Stekol'nikov
(1978, 1984), Minet (1986, 1990), and Nielsen
and Common (1991) have proposed classifica-
tions; however, either no characters were given
or each taxon was descriptively characterized
without an indication of relationship across the
superfamily.
A phylogenetic analysis using Hennig 86
(Farris 1988) based on 38 characters provides the
basis for my discussion concerning relationships
among gelechioid higher taxa. No previous clas-
sification of Gelechioidea has been based on an
analysis of characters and character states using
parsimony. Other cladograms resulting from dif-
ferent interpretations of characters (different
character state matrixes) and methods of analy-
sis (Mickevich & Mitter 1981, Mickevich and
Weiler 1990), are possible; however, most clades
are stable among the many trees examined dur-
ing the analysis.
Within Gelechioidea several characters are ho-
moplasious; a relatively large number of charac-
ters are polymorphic in the sense that a character
has more than one state within a taxon (Micke-
vich & Mitter 1981) (appendix 3); and homology
has not been determined for several genital struc-
tures. These facts likely are the basis for continu-
ing problems in classification of parts of the su-
perfamily and make the resulting classification
tentative. Discovery of additional characters and
reinterpretation of existing ones should result in
a more nearly stable classification. Concurrently,
Gelechioidea exhibit an extremely wide array of
morphological diversity. Likely, much additional
diversity will be discovered as new species and
the immature stages of nearly all taxa are col-
lected and studied.
Analytical procedure. Characters and states detailed
in appendix 1 and coded in appendix 2 were used in the
analysis. Terminology follows Braun (1924), Common
(1990), Heinrich (1920), Klots (1970), Kyrki (1983),
Minet (1982), Mosher (1916), Stehr (1987). Plesiomor-
phic states are indicated by " O . "
Characters were polarized on the basis of knowl-
edge of the Gelechioidea and other Lepidoptera; a hy-
pothetical outgroup consisting of the plesiomorphous
character states was used. Three characters were
weighted: 1) valva divided or not (character 1), 2) pupa
with/without lateral condyles (character 31), and 3)
pupa with/without paired ventromesial lobes on A9
(character 32). An "mh*" run yielded one tree with a
length of 201, consistency index of 37, and retention
index of 65. The "bb*" command yielded 2,456 trees
with the same length and consistency and retention in-
dexes as for the "mh*" run.
Taxa shifted association among the trees because
polymorphic characters were coded as "?," several
characters are homoplasious, and states for several
The Gelechioidea
characters of immature stages are unknown. The
following discussion of alternative associations of
some taxa caused by homoplasious or polymorphic
characters led to selection of a single tree. 1. Depressa-
riinae/Elachistinae: a) sister-groups to each other, b)
Depressariinae the sister-group to clade Elachistinae/
Agonoxeniae/Hypertrophinae/Deuterogoniiae/Aeolan-
thinae, or c) forming a trichotomy among Depressarii-
nae, Elachstinae, and Agonoxeninae/Hypertrophinae/
Deuterogoniinae/Aeolanthinae. " a " was selected be-
cause of the synapomorphies in the gnathos and uncus
shared by Depressariinae and Elachistinae. 2. Xyloryc-
tinae/Scythridinae: Because they share three apomor-
phies (larva with secondary SV setae on Al8, larva
with a pore posterad of SVI on A l - 8 , and larva with
a pinaculum ring around SVI on A l - 8 ) , they may
associate in four diverse ways, a) forming a trichotomy
with Stenomatinae/Aeolanthinae and Chimabachidae/
/Pexicopiinae, b) associated with Ethmiinae in clade
Stenomatinae//Aeolanthinae, c) Xyloryctinae/Scy-
thridinae sister group to clade Chimabachidae/- /Pexi-
copiinae. d) derived taxa within clade Chimabachidae/
Scythridinae/Xyloryctinae. " a " was discarded because
Xyloryctinae/Scythridinae share many character states
with the clade ending with Pexicopiinae; " b " was dis-
carded because Xyloryctinae/Scythridinae lack lateral
condyles in the pupa, an apomorphy of clade Steno-
matinae/-/Aeolanthinae; and " d " was discarded be-
cause the apomorphy defining the clade is weak, "c"
was selected to emphasize the significance of the three
apomorphies. 3. Physoptilinae: a) sister-group to Gel-
echiinae/-/Pexicopiinae, b) sister-group to any one of
Gelechiinae, Dichomeridinae, or Pexicopiinae. " a " was
selected because the retinaculum of Physoptilinae fe-
males is state 7, whereas that of Gelechiinae/-/Pexi-
copiinae is state 4, which I construe as the most signifi-
cant apomorphy of the clade. 4. Gelechiinae/Dichom-
eridinae/Pexicopiinae: a) Gelechiinae sister-group to
Dichomeridinae + Pexicopiinae, b) Dichomeridinae
sister-group to Gelechiinae/Pexicopiinae, c) Pexicopii-
nae sister-group to Gelechiinae/Dichomeridinae. " a "
was selected because Gelechiinae have character states
that could give rise to Dichomeridinae - Pexicopiinae
through loss or modification. Dichomeridinae have
two apomorphies, 1) 9 with secondary corpus bursae
and 2) pupal terga 1/2 and 2/3 with setose, raised, me-
sial/submesial prominences on opposing margins; and
Pexicopiinae usually have only one pair of apodemes
on sternum 2 (apomorphy within clade), often have an
antennal pecten, and never have an anal fork.
The proposed classification (fig. 9.1) is a major
rearrangment of family and subfamily-group
concepts. My use of family names reflects some
conservatism based on the tentativeness of this
analysis but recognizes what probably are reli-
able associations. For this reason all taxa in clade
2 are included in Elachistidae and the taxa in
clade 24 in Coleophoridae. The list of valid and
synonymic family/subfamily-group names to-
gether with authors and dates of publication will
help associate information in previous literature
with recognized taxa (appendix 4).
Key to Families and Subfamilies
1 Gnathos present, fused with tegumen later-
ally, expanded mesially (fig. 9.4 C), dor-
133
somesial surface usually finely scobinate . .
(Oecophoridae) 2
- Gnathos present and fused or articulated
with tegumen or absent; when present and
fused with tegumen then a sclerotized band
(fig. 9.4.B), mesial region downturned and
laterally compressed (fig. 9.4 D), or with a
mesial spinose bulb (fig. 9.4 I) 3
2 Abdominal terga with spiniform setae on
posterior margin of each segment (fig.
9.5 H) Stathmopodinae
- Abdominal terga usually without spiniform
setae, spiniform setae in broad band when
present (fig. 9.5 G) Oecophorinae
3 Pupa with lateral condyles between abdomi-
nal segments 5/6 and 6/7 (fig. 9.7 F)
(most Elachistidae) 4
- Pupa without lateral condyles between ab-
dominal segments 12
4 Valva almost always with some setae that
are thickened and have bifid or multifid
apexes Stenomatinae
- Valva without setae having such modified
apexes 5
5 Gnathos fused broadly with tegumen, apex
with stout, toothlike projections
Ethmiinae, part
- Gnathos articulated with tegumen, with
finely spinose mesial bulb (fig. 9.4 I), some-
times arms separate and each arm ending in
finely spinose bulb (fig. 9.4 K) or toothlike
projections (fig. 9.4 L), or gnathos absent 6
6 Gnathos present 7
- Gnathos absent 9
7 Larva with secondary setae in SV group on
A46 (fig. 9.7 I); pupa with paired leglike
structures arising from ventral surface of
segment 9 or 9/10 (fig. 9.7 G) Agonoxeninae
- Larva without secondary setae in SV group
on A 4 - 6 (SVI, SV2, SV3 only); pupa with-
out paired leglike structures arising from
ventral surface of segment 9 or 9/10 . . . .8
8 Hindwing narrowly lanceolate, cell open, Rs
and Mi long stalked (fig. 9.6 N)
Elachistinae
- Hindwing subquadrate, cell usually closed,
Rs and Mi arising separately from cell (fig.
9.6 K) Depressariinae
9 Hindwing with Rs and Mi stalked
Aeolanthinae
- Hindwing with Rs and Mi separate . . . 10
10 Abdominal sternum 2 with paired venulae
and apodemes (fig. 9.5 E) Deuterogoniinae
- Abdominal sternum 2 with paired venulae
only (fig. 9.5 D) 11
11 Valva with free sclerite at apex of sacculus,
socii present Hypertrophinae
- Valva without free sclerite at apex of saccu-
lus, socii absent Ethmiinae, part
12 Juxta present 13
- Juxta absent 36
134 Ronald W. Hodges

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115-0.1
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13 Abdominal terga 26 with spiniform setae Gnathos absent 17

(fig. 9.5 G) 14 17 Juxta a mesial plate . . . Blastobasinae, part
Abdominal terga without spiniform setae . . Juxta with paired distal lobes (fig. 9.4 A) . .
33 Momphinae
14 Gnathos with mesial region downturned 18 Gnathos present 19
and laterally compressed (fig. 9.4 F) Gnathos absent Deoclonidae, Syringopainae
Lecithoceridae 19 Forewing with CuP present (fig. 9.6 H) . 22
Gnathos otherwise 15 Forewing with CuP absent (fig. 9.6 F) . . 20
15 Valva with costa developed as a free lobe 20 Gnathos bearing a mesial, unarticulated
that appears to arise from saccular part of hook (fig. 9.4 J)
valva (fig. 9.4 A) . (Coleophoridae, part) 16 Autostichidae, Symmocinae, part
Valva otherwise 18 Gnathos otherwise, without hook . . . . 21
16 Gnathos fused to tegumen, bearing mesial, 21 Gnathos bearing a mesial, finely spiculose
finely spinose bulb (fig. 9.4 G) structure Deoclonidae, Deocloninae
Coleophorinae Gnathos with mesial region slightly ex-
The Gelechioidea 135

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4-9 a 4-3 : 4-9 114-0: : 4-5 ::13-6 :I=13-3 * 10-0.1.: 4-6
: I " . .13-15 1: 2-0
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autapomorphy
synapomorphy
parallelism
reversal
polymorphy

Fig. 9.1. Hypothetical Relationships among Families and Subfamilies of Gelechioidea.

panded (fig. 9.4 H) 9.6 H); abdominal sternum 2 with a pair of

. . . . Batrachedridae, Batrachedrinae, part apodemes Epimarptinae
22 Gnathos fused with tegumen laterally (fig. Forewing with R5 to anterior margin (fig.
9.4 C) 23 9.6 F); abdominal sternum 2 with a pair of
- Gnathos articulated with tegumen laterally venulae + a pair of apodemes
(fig. 9.5 A) 26 Batrachedrinae, part
23 Frenum present (fig. 9.6 G, J) 28 Forewing with CuP present 29
(Schistonoeidae) 24 - Forewing with CuP absent
- Frenum absent 25 Autostichidae, Symmocinae
24 Abdominal sternum 2 with a pair of apo- 29 Haustellum present (Autostichidae, part) 30
demes Schistonoeinae - Haustellum absent
- Abdominal sternum 2 with a pair of venulae Coleophoridae, Pterolonchinae
+ a pair of apodemes Oeciinae 30 Forewing with CuA2 directed posterobasally
25 Antenna with pecten on scape and joining CuP, CuP strong at margin; ab-
Coleophoridae, Blastobasinae, part dominal tergal setae spiniform
- Antenna without pecten Holcopogoninae
Xyloryctidae, Xyloryctinae Forewing with CuAi and CuA2 stalked and
26 Gnathos with mesial region slightly ex- directed posteriorly from cell or separate,
panded (fig. 9.4 A) or not (fig. 9.4 H) . . . . CuP very weak on margin; abdominal tergal
(Batrachedridae) 27 setae slender, sometimes deciduous
- Gnathos with a mesial, unarticulated hook Autostichinae
(fig. 9.4 J) 28 31 Gnathos a single or paired spinose bulb(s)
27 Forewing with Rs to outer margin (fig. 32
136
Gnathos present, without spinose bulb(s), or
absent 33
32 Hindwing with Rs and stalked beyond
end of cell Peleopodidae
Hindwing with Rs and Mi arising separately
from cell Amphisbatidae
33 Abdominal sternum 2 with paired venulae
and apodemes (fig. 9.5 E) 34
Abdominal sternum 2 with paired apodemes
only (fig. 9.5 F) 35
34 Forewing with CuP present
Gelechiidae, Physoptilinae
Forewing with CuP absent . Glyphidoceridae
35 Transtilla present Chimabachidae
Transtilla absent Xyloryctidae, Scythridinae
36 Female retinaculum consisting of anteriorly
directed scales on R (fig. 9.6 F), gnathos
usually present (absent in Isophrictis group)
(Gelechiidae, part) 37
Female retinaculum consisting of anteriorly
directed scales on CuA (fig. 9.6 I), anteriorly
directed scales between CuA and R in a row,
or posteriorly directed scales/setae arising on
Sc + anteriorly directed scales between Sc
and R (fig. 9.6 C); gnathos absent
(Cosmopterigidae) 39
37 Abdominal sternum 2 with a pair of venu-
lae; forewing usually with CuA] und CuA2
stalked and directed posteriorly from end of
cell; female with secondary bursa arising
from corpus bursae; pupa with raised, se-
tose, mesial or submesial prominences on
opposing margins of terga 1/2, 2/3, 3/4 . . .
Dichomeridinae
Abdominal sternum 2 with a pair of venulae
+ a pair of apodemes or a pair of apodemes
(if a pair of venulae only, then forewing with
CuAi and CuA2 separate); female lacking
secondary bursa from corpus bursae; pupal
terga without prominences 38
38 Abdominal sternum 2 with a pair of venulae
+ a pair of apodemes (apodemes very short
in Anacampsis group); antenna without pec-
ten (some Aristotelia, Bryotropha, Metzneria,
Monochroa with pecten usually a single,
slender scale, Apatetris group with full pec-
ten of broader scales) Gelechiinae
Abdominal sternum 2 with a pair of apo-
demes; antenna often with a pecten of sev-
eral slender scales Pexicopiinae
39 Female retinaculum with series of posteri-
orly directed slender scales or setae arising
on base of Sc 40
Female retinaculum without series of poste-
riorly directed slender scales or setae arising
on base of Sc Antequerinae
40 Male genitalia asymmetrical, uncus two
separate lobes, right lobe almost always
larger than left lobe, aedeagus ankylosed
with diaphragma; labial palpus usually very
slender and sickle shaped; forewing often
brightly colored with well-defined pattern,
Ronald W. Hodges
without patches of raised scales
Cosmopteriginae
Male genitalia symmetrical or asymmetrical,
uncus present and symmetrical or absent,
aedeagus often ankylosed with saccus; labial
palpus never extremely slender, often stout;
forewing often monochromatic gray, black,
or brown, often with patches of raised scales
Chrysopeleiinae
Family-group statements
Apomorphic characters, possible apomorphic
characters, and untreated characters (often of
larvae and pupae) are provided for each taxon.
Appendices 2 and 3 give the states of characters
used in the parsimony analysis. When the family-
group concept is new, no literature may exist for
the entity.
Elachistidae. Defined by the apomorphy of pupa
with lateral condyles on abdominal segments 5/6
and 6/7, which character occurs as a polymorphy
in Elachistinae; ventral surface of A9 often with
two submesial, slightly swollen zones of recurved
setae. Highly diverse, worldwide taxon; 8 sub-
families, 165+ genera, 3270+ species. (Minet
1990).
Stenomatinae (fig. 9.2 A). All species, except
some Agriophara, have some setae on valva with
prominent, multilobed apexes (only some Anar-
sia (Gelechiidae) have such setae elsewhere in
Gelechioidea); uncus uni-, bi-, trilobed, or ab-
sent; abdominal support diffuse/diffuse with very
short apodeme/sclerotized ridge at 45 degree an-
gle with anterolateral margin of second sternum
(), well-developed apdeme, rarely sclerotized
area with lateral ridge ($); hindwing with Rs bent
toward Sc near end of cell; antenna long cili-
ated. Larva: frons < , = , > 1/2 distance to epicra-
nial notch; thoracic legs approximate or distant;
Al with 2(3) SV setae; A 3 - 6 : SD setae without
pinaculum rings; crochets complete, biordinal el-
lipse, numerous. Pupa: beginning development
of "legs" Rectiostoma xanthobasis (Duckworth
1971: 31), R. fernaldella (Keifer 1936: 356), Cer-
conota monella, C. palliata (USNM), fully devel-
oped in Timocratica melanocosta (Becker 1982:
262); antennal scape usually with prominent
ridges. Larvae feed on turgid, living (16 families,
35% Myrtaceae) and wilted plant tissue. Mainly
New World, esp. Neotropics; Madagascar, India,
Ceylon, New Guinea, Australia, New Zealand;
30 + genera, 1,200 + species (Anadasmus, An-
taeotricha, Chlamydastis, Phylomictis, Stenoma,
Timocratica).
Becker (1982, 1984 a, 1984 b), Busck (1934),
Common (1990), Duckworth (1964, 1967, 1970,
1971), Hodges (1978), Minet (1990), Powell
(1980), Stehr (1987).
Ethmiinae (fig. 9.2 B). Variable in most charac-
ters; hindwing Rs and M2 separate at base; phal-
The Gelechioidea 137

Fig. 9.2. Gelechioidea. A - , Elachistidae: A, Cerconota monella [11 mm*]; B, Ethmia semilugens [10 mm]; C,
Agonopterix umbellana [10.5 mm]; D, Perittia eremonoma [4.5 mm]; E, Tetanocentria sp., [8.5 mm]; F, Hypertropha
tortriciformis [6.5 mm]; G, Deuterogonia chionoxantha [5.5 mm]; H, Aeolanihes callidora [8.5 mm]; I - J , Xylorycti-
dae: I, Xylorycta thrasycosma [8 mm]; J, Scythris mixaula [9 mm]; K, Chimabachidae: Cheimophila salicella
[8.5 mm]; L, Glyphidoceridae: Glyphidocera democratica [7 mm]; M - N , Schistonoeidae: M, Oecia oecophila
[4.5 mm]; N, Schistonoea fulvidella [5.5 mm]; OP, Oecophoridae: O, Alabonia geoffrella [9.5 mm]; P, Stathmopoda
lychnacma [5.5 mm]; Q, Lecithoceridae: Tisis chalybaeella [8.5 mm]; R - S , Batrachedridae: R, Epimarptisphilocoma
[5 mm]; S, Homaledra heptathalama [8 mm]; T-U, Deoclonidae: T, Deoclona yuccasella [8 mm]; U, Syringopais
temperatella [7 mm]; VX, Coleophoridae: V, Coleophora eleagrtisella [8 mm]; W, Mompha deflnitella [6.5 mm]; X,
Blaslobasis glandulella [7.5 mm]. [*= length of forewing]
138 Ronald W. Hodges

Fig. 9.3. Gelechioidea. A, Coleophoridae: Pterolonche sp. [10 mm*]; B - D , Autostichidae: B, Holcopogon bubul-
cella [6 mm]; C, Stoeberhinus testaceus [6.5 mm]; D, Aprominta designatella [5.5]; E. Peleopodidae: Durrantia ama-
bilis [8.5 mm]; F, Amphisbatidae: Psilocorsis cirrhoptera [9 mm]; GI, Cosmopterigidae: G, Walshia miscecolorella
[7 mm]; H, Euclemensia bassettella [6.5 mm]; I, Triclonella xuthocelis [7 mm]; J - M , Gelechiidae: J, Physoptila sp.
[5 mm]; K, Chinodes retiniella [8.5 mm]; L, Dichomeris mulsa [9 mm]; M, Pectinophora gossypiella [8 mm].
[* = length of forewing]

lobase recurved; gnathos usually broadly at-
tached to tegumen, when present. Larva: A 3 - 6
or A l , 2, 79 with secondary SV setae in New
World taxa, but assamensis and nigroapicella
groups in Palearctic with secondary setae on
Al9. Pupa with "legs" in section I of Ethmia,
absent in section II, extremely poorly developed
in sphaerosticha and in undescribed Pyramido-
bela from Mexico (USNM). Larval hosts mainly
Boraginaceae and Hydrophyllaceae (70%), but
also Papaveraceae, Ranunculaceae, Rosaceae,
Sabiaceae, Scrophulariaceae. Most numerous in
tropical-subtropical, seasonally arid areas, esp.
northern Neotropics; 35 genera, 250 species
(Pseudethmia).
Common (1990), Keifer (1936), Minet (1990),
Powell (1973, 1980, 1985), Sattler (1967), Stehr
(1987).
Depressariinae (fig. 9.2 C). Maxillary palpus
usually four segmented; hindwing with Rs and
Mi separate; fore- and hindwings broad. Larva:
stemmata with gap between 5/6, T1 normally
with 3 L setae (rarely 2), Al with 2/3 SV setae,
A9 with SD usually hairlike. Larvae leaf tiers,
seed feeders, stem borers on dicots (17 families,
including Apiceae, Asteraceae, Betulaceae, Cory-
laceae, Fabaceae, Fagaceae, Malvaceae, Rosa-
ceae, Rutaceae, Salicaceae, Urticaceae). World-
wide, but absent from many island groups; 80 +
genera, 600 + species (Agonopterix, Depressaria,
Exaeretia, Semioscopis, Tonica).
Clarke (1941, 1963, 1978), Common (1964,
1990), Hannemann (1953), Hodges (1974),
L'vovskii (1981), Minet (1990), Palm (1989), Pa-
tocka (1989), Powell (1980), Stehr (1987), Toll
(1964).
The Gelechioidea
Elachistinae (fig. 9.2 D). Maxillary palpus usu-
ally one segmented, rarely two segmented (an
apomorphy in Gelechioidea); hindwing with Rs
and Mi long stalked, distant from costal margin.
Fore- and hindwings narrow. Larva: head prog-
nathous, flattened, recessed into 1, 5 or 6 stem-
mata, frontoclypeus extending to epicranial
notch; tarsal claw with bladelike pretarsal seta;
A l with 1 SV seta; A9 with SD setiform. Pupa
with abdominal segments often fused, permitting
no movement, sometimes with condyles on A4/
5, 5/6. Larvae leaf miners (rarely stem miners) on
Boraginaceae, Commelinaceae, Cyperaceae, Jun-
caceae, Lamiaceae, Loniceraceae, Poaceae (70%
on monocots). Worldwide, except eastern Poly-
nesia; 15 genera; approximately 250 species,
mainly Elachista (Cosmiotes, Biselachista, Dicra-
noctetes, Mendesia, Perittia, Stephensia).
Braun (1948), Common (1990), Falkovitsch
(1981), Hodges (1978), Kuroko (1982), Minet
(1990), Powell (1980), Traugott-Olsen & Nielsen
(1977), Wagner (1987).
Agonoxeninae (fig. 9.2 E). Highly variable:
Secondary wing coupling, when present, 1) stout
setae in anterior margin of hindwing, 2) series of
recurved setae on anterior margin of hindwing,
3) few stout setae extending from costal margin
of hindwing appearing as an extension of Sc.
Apex of hindwing produced in Glaucacna iridea.
Gnathos an articulated band with a mesial bulb
bearing parallel rows of short spines (Agonox-
ena), a pair of articulated rami, each bearing spi-
nules at its apex (Parametriotini). Larva with
two flattened setae at apex of tarsus (Agonox-
ena)\ crochets in mesoseries (Blastodacna curvili-
neella)', T1 with L setae on shield (Agonoxena,
unknown genus/species from S. Africa, U S N M )
or separate; surface often with abundant second-
ary setae. Pupa with/without "legs." Larvae bor-
ers, miners, or external feeders on Arecaceae, Eu-
phorbiaceae, Proteaceae, Rosaceae, Theaceae,
Tiliaceae. Pupation away from larval habitat, but
not in ground. Worldwide; 23 genera; 95 species
(.Agonoxena, Blastodacna, Chrysoclista, Microco-
lona, Pammeces, Prochola, Spuleria, Tetano-
centria, Zaratha).
Bottimer (1926), Bradley (1966), Clarke
(1965), Common (1990), Hodges (1978), Kasy
(1976), Kuznetsov, N . J . (1915), Riedl (1969),
Stehr (1987).
Hypertrophinae (fig. 9.2 F). Valva with free
sclerite at distal end of saccalus a supposed apo-
morphy but occurring also in Cryptophasa bal-
teata (Xyloryctinae). Larva with mesial crochets
of AIO widely spaced or absent. Larvae external
feeders, within silk shelter, on Myrtaceae, esp.
Eucalyptus. Australia, Tasmania, New Guinea
(two species); 11 genera, 50 + species (Eupselia,
Hypertropha, Thudaca).
Common (1980, 1990), Diakonoff (1954),
Hodges (1974, 1978), Minet (1990).
139
Deuterogoniinae (fig. 9.2 G). Defined by tegu-
men undeveloped and membranous mesially (an
autapomorphy within Gelechioidea), very lightly
sclerotized uncus separated from tegumen by
broad membrane. Hindwing with outer margin
slightly excavated posterad of apex. Immature
stages unknown; one adult emerged from a cyni-
pid gall on Quercus. Palearctic region: eastern
Europe, Japan, Taiwan, and Thailand; 2 genera
(Deuterogonia, Paradeuterogonia) 4 species.
Fujisawa (1991), Hodges (1978), Toll (1964),
Saito (1987, 1989).
Aeolanthinae (fig. 9.2 H). Valva broadly at-
tached to vinculum, usually parallel with vincu-
lum for basal 1/2, then directed posteriorly as a
right angled lobe (an apomorphy within Gelechi-
oidea); aedeagus ankylosed with juxta. Larva
leaf roller on Cedrela (Meliaceae). India, Western
China; 1 genus (Aeolanthes), 20 + species.
Clarke (1955 b), Fletcher (1933).
Xyloryctidae. Sister-group to clades 1136. De-
fined by 3 apomorphies: 1) larval segments
Al8 with pinaculum ring around SD; 2) a
pore posterad/ventroposterad of SD; and 3)
segments 3 7 with secondary SV setae.
Worldwide: 86 genera, 1200 + species.
Xyloryctinae (fig. 9.2 I). Within Xyloryctidae
possess autapomorphy, abdomen with band of
spiniform setae on posterior part of terga 26. c
antenna simple, ciliate, bipectinate. Larva: head
with bordered submental plate (a submental pit
in some Cryptophasa), subgenal sclerite triangu-
lar, short, stemmata with gap between 1/2, 4/5,
5/6; A l with 3 SV setae; A 3 - 6 with 5 - 7 SV se-
tae. Pupa: abdominal terga often with crenulate
raised ridge near anterior margins, becoming
spined in some species; metathoracic legs ex-
posed distally. Wings reduced in Thyrocopa apa-
tela. Larval hosts in 21 plant families (47 % Pro-
teaceae + Myrtaceae), some on lichens. Subsa-
haran Africa, Indoaustralia, Polynesia; 60 + gen-
era, 500 + species (Cryptophasa, Lichenaula,
Metathrinca, Ommatothelxis, Pansepta, Phtoner-
odes, Scieropepla, Telecrates, Uzucha, Xylorycta).
Common (1990), Duckworth (1973), Hodges
(1978), Moriuti (1982), Powell (1980), Zimmer-
man (1978).
Scythridinae (fig. 9.2 J). Sister-group to Xylo-
ryctinae with derived states for characters 8 (ae-
deagus ankylosed), 27 (antennal pecten present),
and ocellus separate from eye (when present),
each a parallelism within Gelechioidea. Possibly
two subunits defined by presence (Scythris + ) or
absence (Arotrura + ) of frenum. S genitalia rela-
tively simple to extremely highly modified by re-
duction and fusion of structures. $ with ductus
seminalis arising broadly from posterior-most
part of corpus bursae, an autapomorphy within
Gelechioidea. Larva: stemmata 3/4/5 close to-
gether, L2 ventrad of LI on same pinaculum.
140 Ronald W. Hodges

gnathos uncus

transtilla

costal lobe of valva

gnathos

gnathos
vinculum

uncus

gnathos uncus
manica
^aedeagus

gnathos

gnathos gnathos

Fig. 9.4. Structural Features of Gelechioidea. A, ventral view of male genitalia, left valva removed, Mompha
franclemonti; B - D , F - L , ventral/ventrolateral view of tegumen, gnathos, and uncus: B, Diurnea fagella; C,
Lactistica longimana; D, Deltoplastis figurata', F, Eridachtha prolocha', G, Coleophora sacramenta', H, Batrachedra
enormis; I, Depressaria juliella; J, Symmoca oenophila; K, Parametriotes bipunctella, L, Chrysoclista linneella; E,
lateral view of aedeagus and sclerotized manica, Anoncia brunneipes.

Larvae feed externally on buds and leaves, or are
leafminers, on 20 plant families including Apia-
ceae, Asteraceae, Cactaceae, Caryophyllaceae,
Chenopodiaceae, Convolvulaceae, Fabaceae,
Hydrophyllaceae, Lamiaceae, Malvaceae, Ona-
graceae, Poaceae, Polygonaceae, Portulacaceae,
Solanaceae. Worldwide; 26 genera, 700 + spe-
cies; many undescribed taxa (Areniscythris, Apos-
tibes, Aratura, Eretmocera, Mapsidius, Neo-
scythris, Rhamphura, Scythris).
The Gelechioidea
Bengtsson (1984), Clarke (1965), Falkovitsch
(1981), Landry (1991), Powell (1976 a, 1980),
Stehr (1987).
Chimabachidae (fig. 9.2 K). Sister-group to
clades 1 2 - 3 6 . Defined by 1) larval metathoracic
tibia and tarsus swollen (an autapomorphy
within Gelechioidea), 2) D 1 setae on A 9 closer
to each other than the D 2 setae (possible autapo-
morphy), 3) transtilla present (parallelism), 4)
plesiomorphic states for characters 8, 27, 35, and
5) ocellus, when present, distant from eye (paral-
lelism with Scythridinae). Larvae general feeders
on Berberidaceae, Betulaceae, Cornaceae, Erica-
ceae, Fagaceae, Myricaceae, Rosaceae, Salica-
ceae, Styracaceae; make shelters within leaves or
flower buds; feed on developing flowers, fruit,
and leaves. Palearctic region from western
Europe to Japan, introduced into northwestern
North America; 2 genera, 6 species (Cheimoph-
ila, Diurnea).
Hodges (1974), M a c K a y (1972), Minet (1990),
Palm (1989), Patocka (1989), Raine (1966), Saito
(1979), Toll (1964).
Glyphidoceridae, new family. Type genus: Glyphi-
docera Walsingham 1892 (fig. 9.2 L). Sister-
group to clades 1 3 - 3 6 ; defined by two paral-
lelisms: 1) forewing with C u A i and C u A 2 stalked
and downcurved from posterodistal angle of cell;
2) forewing with Rs terminating on the outer
margin. Possible apomorphy, AIO of larva with
D 2 setae stout, strong, directed posteroventrally
from anal plate (similar state in undescribed
Agonoxeninae from Africa ( U S N M ) . Valva with
slender, posteriorly directed lobe from base of
costal margin; ostium bursae broad, ductus bur-
sae broadly triangular, often sclerotized; ductus
seminalis arising from side of corpus bursae near
base and coiled; signa patches of inwardly di-
rected denticles. Larva with short frontoclypeus
( < 1 / 2 distance to epicranial notch); stemmata
with gaps between 1/2, 4/5, 5/6; A l with 2 SV
setae; A l 8 with seta L 2 dorsad or slightly post-
ered orsad of L I , S D 2 very small, directly ante-
rad of S D and on some pinaculum; A 9 with D 1
anterad of D2, S D setiform. Larva of G.juni-
perella on Juniperus (Cupressaceae). New World;
1 genus, 49 species. [Pseudodoxia and Psaltica
may belong to Glyphidoceridae.]
Adamski & Brown (1987), Clarke (1969 b),
Stehr (1987).
Schistonoeidae, new family. Type genus: Schisto-
noea Forbes 1931. Sister-group to clades 1536;
defined by the parallelism, frenum present. $
frenulum with two acanthi. Immature stages un-
known. Pan tropical; 2 genera, 2 species.
Oeciinae, new subfamily. Type genus Oecia
Walsingham 1897 (fig. 9.2 M). Defined by two
parallelisms: 1) retinaculum consisting of ante-
riorly directed scales between Sc and R and pos-
141
teriorly directed scales/setae on Sc; 2) haustellum
absent. Gnathos broadly fused with tegumen lat-
erally, ending in unturned lobe; signum 7/8
thumb tackshaped spines circling anterior part
of corpus bursae; forewing lanceolate, cell very
weakly closed, Ri from 1/2 length of cell, R2
from 5/6 length of cell, R3 nearly from end of
cell, R4 and R5 stalked, R5 to outer margin just
beyond apex, M i - C u A i separate and evenly
spaced, M 3 from end of cell, CuA2 from about 4/5
length of cell; hindwing slightly narrower than
forewing, Sc and Ri fused from base, Rs
straight, Rs and M i long stalked, M2 curved an-
teriorly on basal 1/3, M2, M3, C u A i , and Q 1 A 2
separate, cell open between Rs + M | and M2.
Pantropical, possibly transported from West In-
dies; 1 genus, 1 species.
G o z m n y (1963, 1975), Zimmerman (1978).
Schistonoeinae, new subfamily. Type genus:
Schistonoea Forbes 1931 (fig. 9.2 N). Defined by
four parallelisms: 1) transtilla present; 2) abdo-
minal support on sternum two a pair of apo-
demes; 3) hindwing with Rs running into Sc be-
yond base; 4) hindwing with outer margin
slightly excavated posterad of apex. Gnathos a
pair of very slender, slightly overlapping lobes;
signum a deeply invaginated band; forewing lan-
ceolate, Ri from 1/2 length of cell, R2 and R3
fused, R4 and R5 stalked, R5 to anterior margin,
M i and M2 separate, distant, one vein missing of
M3 to C u A 2 (M3 and C u A i fused (likely), C u A i
and C u A 2 fused, or CuA2 absent), C u P weak at
margin; hindwing broadly lanceolate, apex
slightly produced, closure of cell broadly V-
shaped between Rs + M i and M 2 , Sc and R
anastomosed shortly beyond base to 2/5 wing
length, Rs and Mi stalked then diverging to mar-
gins, M2 separate and distant from Rs + M i , M3
and C u A i connate at end of cell, C u A 2 from 3/4
length o f cell. West Indies; 1 genus, 1 species.
Oecophoridae. Sister-group to clades 1736. De-
fined by two polymorphic parallelisms and one
reversal: 1) Second abdominal sternum with a
pair of venuale or a pair of venulae + a pair of
apodemes; 2) abdominal terga without/with spin-
iform setae; 3) hindwing with Rs/Mi separate
(reversal). Broad-based, unarticulated gnathos,
tapering to slender or rounded apex, dorsomesial
surface often with series of small teeth or small,
slender projections distinctive. Worldwide; 326
genera, 3,150 species.
Oecophorinae (fig. 9.2 O). Defined by abdomi-
nal terga lacking spiniform setae or with spini-
form setae in broad band. Most characters vari-
able, labial palpus often very slender, sickle
shaped, long. Larvae mainly on dead plant tis-
sue, leaf litter/bark of dead trees (fungi?), esp.
Myrtaceae in Australia. Worldwide, dominant
Lepidopteran group in Australia; 300 genera,
3,000 species (Barea, Borkhausenia, Chezala,
Chrysonoma, Diocosma, Endrosis, Eulechria,
142 Ronald W. Hodges

A B
gnathos ^ J )

Fig. 9.5. Structural Features of Gelechioidea. A, ventral view of male genitalia, valvae and aedeagus removed,
Dichomeris marginella; B, ventral view of vinculum and base of valvae, Cosmopterix montisella; C, ventral view
of vinculum, Coleotechnites edulicola; D F, abdominal sternum 2: D, Mompha franclemonti; E, Gelechia rhombelli-
formis; F, Pectinophora gossypiella; G I, abdominal terga: G, Deltoplastis ocreata; H, Stathmopoda pedella; I,
Mompha franclemonti, JL, base of hindwing: J, Blastobasis glandulella; K, Coleophora atromarginata; L, Ti-
myra pastas.
The Gelechioidea
Garrha, Hofmannophila, Inga, Orophia, Oxystola,
Philobota, Pleurota, Tanyzancla, Wingia).
Clarke (1941, 1963), Common (1990), Hodges
(1974), Leraut (1989, 1993), L'vovskii (1981),
Meyrick (1922), Patocka (1989), Stehr (1987),
Toll (1964).
Stathmopodinae (fig. 9.2 P). Defined by auta-
pomorphy, abdominal terga with spiniform setae
on posterior margins. Wall of aedeagus with
ventrodistal, sclerotized projection (possible apo-
morphy); signa paired, sclerotized plates with
mesial, inwardly directed flange, often secondary
signa consisting of rows of inwardly directed
conical projections. Most species very slender
winged, with scales closely appressed to surface
of head. Larvae feed in fruits and flowers, on
dead plant tissue, predatory on scale insects and
aphids. Mainly pan tropical/subtropical; 26 gen-
era, 150 species (Hieromantis, Lactistica, Oede-
matopoda, Pseudaegeria, Snellenia, Stathmo-
poda, Tor tilia).
Common (1990), Hodges (1978), Kasy (1973),
Walsingham (1889).
Lecithoceridae (fig. 9.2 Q). Sister-group to clades
18 36; defined by apomorphy, gnathos fused to
tegumen, mesial region downturned and laterally
compressed, and five parallelisms: 1) frenum pre-
sent, 2) forewing with CuP absent (rarely pre-
sent), 3) hindwing with outer margin slightly ex-
cavated posterad of apex, 4) larva with second-
ary setae, 5) larva with 2 L setae on prothorax.
Head usually smooth scaled, sometimes tufted or
loosely scaled. Larvae feed on detritus. Southern
Palearctic, Africa; mainly Indoaustralia; 90 gen-
era, 500 + species (Atrichozancla, Ceuthoma-
darus, Crocanthes, Dragmatucha, Eridachtha,
Frisilia, Homaloxestis, Idiopteryx, Lecithocera,
Odites, Tisis, Torodo ra). Torodorinae [defined by
apomorphy, uncus thorn shaped, tapering to
slender, acute apex (fig. 9.4 D)]; Lecithocerinae
[defined by uncus with narrow base and broad,
bilobed apex (fig. 9.4 F) and slender sclerite ex-
tending from base of tegumen to mesial surface
of valva]; and Ceuthomadarinae (defined by ab-
sence of haustellum, apophyses anteriores usu-
ally as long as bursa copulatrix, and slender scle-
rite extending from base of tegumen to mesial
surface of valva) proposed for Palearctic fauna,
but additional diversity extant in Australasian
region.
Clarke (1955 a, 1965), Common (1990), Goz-
mny (1978), Janse (1949-54, 1958-63), Kana-
zawa (1986), Moriuti (1982), Piskunov (1981),
Sattler (1973).
Batrachedridae. Sister-group to clades 20-36;
defined by one polymorphy, abdominal support
either pair of venulae or pair of apodemes, and
two parallelisms: 1) abdominal terga with spini-
form setae in two submesial patches, 2) hindwing
143
with Mi and M2 stalked. Worldwide; 6 genera,
100 + species.
Epimarptinae (fig. 9.2 R). Defined by three
parallelisms: 1) abdominal support a pair of apo-
demes, 2) hindwing with Mi and M2 fused, 3)
$ antenna notched. Immature stages unknown.
Western India; 1 genus, 1 species (Epimarptis).
Meyrick (1914).
Batrachedrinae (fig. 9.2 S). Defined by abdo-
minal second sternum with venula or venula +
apdeme (polymorphy); hindwing with Rs/Mi
separate (reversal); and hindwing with M3 and
Cu fused (polymorphy). Mi and M2 often
fused and arising from Rs; tegumen often with
strong, mesial sclerotized ridge; larva with sub-
mental pit present or absent, T1 with 2 or 3 L
setae, Al with 1 - 3 SV setae; A9 usually with
SD hairlike. Larvae feed on plant tissue (Amar-
yllidaceae, Arecaceae, Bromeliaceae, Cypera-
ceae, Juncaceae, Pinaceae, Salicaceae), fern spo-
rangia, scale insects, or scavenge. Worldwide; 5
genera, 100 + species (Batrachedra, Chedra, Ho-
maledra).
Clarke (1963), Hodges (1966), Zimmerman
(1978).
Deoclonidae, new family. Type genus: Deoclona
Busck 1903. Sister-group to clades 2236; de-
fined by gnathos an articulated band with a me-
sial bulb bearing parallel rows of short spines
(parallelism), and female retinaculum with ante-
riorly directed scales on CuA or with anteriorly
directed scales between CuA and R (poly-
morphy). New World and Palearctic; 3 genera,
4 species.
Deocloninae, new subfamily. Type genus:
Deoclona Busck 1903 (fig. 9.2 T). Defined by
gnathos articulated with mesial, spinose bulb(s)
(parallelism), abdominal terga with spiniform se-
tae in band or two submesial patches (poly-
morphy), retinaculum consisting of anteriorly
directed scales on CuA or on R (polymorphy),
forewing with CuP absent (parallelism), hind-
wing with outer margin slightly excavated post-
erad of apex (parallelism). Larva: T1 with 3 L
setae, Al with 2 SV setae, A9 with SD setiform.
Pupa with paired, submesial, ridged protuber-
ances on T l . Larva in dry seed pods of Fabaceae,
Liliaceae, or fruit mummies of Rosaceae. New
World, California to northern Argentina; 2 gen-
era, 3 species (Proclesis).
Busck (1939), Clarke (1969 b), Keifer (1936).
Syringopainae, new subfamily. Type genus
Syringopais Hering 1919 (fig. 9.2 U). Defined by
4 parallelisms 1) gnathos absent, 2) vinculum
weakly sclerotized mesially, 3) hindwing with
M1/M2 stalked, 4) larva with a submental pit and
a reversal, hindwing with cell open. Larva: fron-
toclypeus reaching epicranial notch, Al with 2
SV setae, A 3 - 6 SD with adjacent, very small
socket, A9 with SD setiform. Larva leaf miner
144
on Triticum (Poaceae). Palearctic region, Medi-
terranean to India; 1 genus, 1 species.
Hering (1919).
Coleophoridae. Sister group to clades 26-36. De-
fined by hindwing with Rs/Mi separate (rever-
sal), hindwing with M3/CUA1 separate or con-
nate or stalked (polymorphy), antennal pecten
present (parallelism). Males of Blastobasinae +
Momphinae + Coleophorinae have the valva di-
vided with the costa developed as a free lobe that
often appears to arise from the sacculus (synapo-
morphy). Worldwide; 43 genera, 1418 species.
Coleophorinae (fig. 9.2 V). Defined by gnathos
fused with tegumen laterally and bearing a me-
sial, spinose bulb (apomorphy), and five paral-
lelisms: 1) uncus absent; 2) Aedeagus ankylosed;
3) $ frenulum with two acanthi fused distally, ap-
pearing as a single acanthus; 4) retinaculum
consisting of anteriorly directed scales between
Sc and R + posteriorly directed scales/setae on
Sc; 5) forewing with CUAI/CUA2 stalked and di-
rected toward posterior margin from end of cell.
Aedeagus membranous; valva sometimes ap-
pearing like Oecophorinae with well-developed
sacculus and prominent costal area not a discrete
lobe to the base. Larva: frontoclypeus nearly to
epicranial suture; A l , A2 with 2 SV setae; A9
with SD setiform; prolegs highly reduced,
crochets usually two transverse, uniordinal rows.
Pupa: labial palpus present, antennae often ap-
proximate mesially (sometimes meet), diverge
distally; segments 9/10 blunt distally, with short
lateral projection. Larvae mostly leaf miners or
seed feeders on 34 plant families in Coniferae
(rare), Monocotyledonae, and Dicotyledonae;
those of most species case makers. Worlwide, but
mainly Holarctic; 13 genera, 1,050 species;
Nearctic fauna extremely rich but poorly de-
scribed (Augasma, Coleophora, Goniodoma, Met-
riotes).
Capuse (1971, 1972, 1973, 1975), Common
(1990), Falkovitsch (1972), Powell (1980), Ra-
zowski (1990), Sattler & Tremewan (1974, 1978),
Stehr (1987), Toll (1952, 1962), Vives Moreno
(1988).
Momphinae (fig. 9.2 W). Defined by paral-
lelism gnathos absent. Juxta ending in two small
lobes closely associated with aedeagus; signa
paired with lightly sclerotized, circular base and
heavily sclerotized rod. Larva: T1 with 2 L setae,
tarsus sometimes with scalelike seta; Al with
1 - 3 SV setae; A9 with SD setiform; plantae
separate, distant or mesial, almost fused,
crochets uniordinal transverse circle, open el-
lipse, two parallel rows. Pupa with labial palpus
present, antennae touching mesially. Larvae feed
on living tissue, mainly Onagraceae, also Astera-
ceae, Fabaceae, Fagaceae, Halogoridaceae, Rub-
iaceae. Mainly Holarctic, very few in Neotropics,
New Zealand; 6 genera, 60 species (Anchimom-
Ronald W. Hodges
ha, Mompha, Moriloma, Synallagma, Urodela,
Zapyrasta.
Clarke (1963), Riedl (1969), Stehr (1987), Za-
gulyaev & Sinev (1981).
Blastobasinae (fig. 9.2 X). Defined by two par-
allelisms: 1) forewing with pterostigma, 2) larva
with submental pit. Aedeagus nearly membra-
nous with sclerotized, ventral band; anellus
finely or coarsely setose before aedeagus. Larva:
frontoclypeus 1/2-3/5 distance to epicranial
notch; A l - 6 with 3 SV setae; A l - 8 , SD setae
often with pinaculum rings, L2, LI vertical; A9,
SD seta hairlike or setiform. Pupa: T1 femur
visible; AIO without cremaster or recurved setae.
Larvae mainly scavengers, sometimes on living
plant tissue, sometimes opportunistic predators.
Worldwide, best developed in New World; 22
genera, 300 species (Auximobasis, Asaphocrita,
Blastobasis, Calosima, Holcocera, Hypatopa, Pi-
gritia, Tecmerium).
Adamski and Brown (1989), Clarke (1963),
Common (1990), Hodges (1978), Piskunov
(1981), Powell (1976 b), Stehr (1987).
Pterolonchinae (fig. 9.3 A). Defined by three
parallelisms: 1) abdominal support a pair of apo-
demes, 2) abdominal terga with spiniform setae
in band across segments, 3) haustellum absent
and two reversals: 1) $ retinaculum consisting of
diffuse, anteriorly directed, slightly upturned
scales, 2) hindwing with M3/CUA1 separate.
Larva with no/few crochets; pupa with antennae
approximate but separated mesially. Pterolonche
a root borer in Centaurea (Asteraceae). Palearc-
tic (circum Mediterranean) and South Africa, in-
troduced into Nearctic; 2 genera, 8 species (Ana-
thyrsa).
Hodges (1978), Minet (1988), Vives Moreno
(1986).
Autostichidae. Sister-group to clades 2936. De-
fined by gnathos an articulated band with an un-
articulated mesial hook (parallelism) and abdo-
minal terga with spiniform setae in band across
segment (parallelism). Larvae feed on dead/de-
caying plant/animal material. Palearctic, Aus-
tralasia, Polynesia, Nearctic; 67 genera, 308 spe-
cies.
Holcopogoninae (fig. 9.3 B). Defined by $ reti-
naculum a series of anteriorly directed scales be-
tween Sc and R and posteriorly directed scales/
setae on Sc (parallelism). Hindwing with Rs and
Mi long stalked (possible autapomorphy), lan-
ceolate; spiniform setae on abdominal terga
prominent. Larva with stemmata reduced or un-
developed, Al 8 without pinaculum ring
around SD or pore posterad of SD, A9 with 2
extra D setae. Larva of Holcopogon bubulcellus
feeds on cattle dung. Palearctic region, mainly
Mediterranean and eastward in xeric areas; 7
genera, 18 species (Arragonia, Charadraula, He-
ringita, Holcopogon, Ilionarsis.)
The Gelechioidea 145

M3+C11A CuA2

Fig. 9.6. Wings of Gelechioidea. A - , G, I, base of forewing: A, Timyra pastas, 9; B, Depressaria clnereocostella,
V; CD, Cosmopterix montisella, 9; E, Blastobasis glandulella, 9; G, Scythris basilaris, 9; Stathmopoda pedella,
9; F, H, M, forewing: F, Polyhymno luteostrigella, 9; H, Deuterogonia chionoxantha, 9; M, Dichomeris marginella,
<J; J, base of hindwing, Scythris basilaris, 9; K L, NP, hindwing: K, Deuterogonia chionoxantha, 9; L, Syringo-
pais temperatella, 9; N, Epimarptis philocoma, 9; O, Batrachedra enormis, 9; P, Dichomeris marginella, S-
146
Amsel (1942), Benander (1937), Gozmny
(1964, 1967), Hodges (1978), Vives Moreno
(1987).
Autostichinae (fig. 9.3 C). Defined by 1) $ reti-
naculum consisting of anteriorly directed setae/
scales starting in cell between base of CuA and
R and extending to area between Sc and R (par-
allelism), 2) larva with pore posterad of SD on
Al 8 (parallelism), and 3) larva with pinaculum
ring around SD on Al 8 (parallelism). Hind-
wing with Rs/Mi stalked/connate/separate;
broadly lanceolate to subquadrate; abdominal
terga with fine, slender, sometimes deciduous se-
tae. Larva: frontoclypeus 2/5 distance to epicra-
nial notch, Al with 2 SV setae, A l - 8 with LI/
L2 vertical, A9 with SD setiform. Pupa: labial
palpi minutely visible, antennae approximate/
meet mesially, cremaster present. Larvae on li-
chens, decaying vegetable matter. Eastern Pale-
arctic, Australasia, Polynesia; 18 genera, 120 +
species (Apethistis, Autosticha, Pachnistis, Pro-
cometis, Ptochoryctis, Stoeberhinus).
Common (1990), Hodges (1978), Le Mar-
chand (1947), Zimmerman (1978).
Symmocinae (fig. 9.3 D). Defined by forewing
with CuP absent (parallelism), $ with retinacu-
lum a series of anteriorly directed scales between
CuA and R (parallelism), second abdominal ster-
num with venula, venula + apdeme, or ap-
deme (polymorphy), forewing with R5 terminat-
ing on outer margin or on costa (polymorphy).
Gnathos a slender lateral band becoming a me-
sial hook, distinctive; spiniform setae on abdomi-
nal terga sometimes deciduous, slender. Larva:
Frons less than 1/2 distance to notch; Al - 8 with
L1/L2 vertical, SD followed by a pit, pinaculum
ring absent; A9 with SD setiform. Pupa: labial
palpus and T1 femur exposed; antennae meet
mesially, separate distally. Larvae scavengers on
dead plant tissue. Palearctic, circum Mediterra-
nean east to China in xeric areas, Nearctic; 42
genera, 170 species (Amselina, Aprominta, Con-
quassata, Dysspastus, Kurtomesis, Mylothia, Oe-
goconia, Symmoca.)
Gozmny (1957, 1959, 1963, 1964), Hodges
(1978), Stehr (1987).
Peleopodidae (fig. 9.3 E). Sister-group to clades
30-36; defined by gnathos with mesial, spinose
bulb (sometimes paired) (parallelism), and abdo-
minal terga with scales only or with spiniform
setae in band (polymorphy). genitalia often
asymmetrical; male second sternum with ap-
deme only, with venula and apdeme; forewing
with CuP present; larva: T1 with 3 L setae, L pi-
naculum fused with shield, Al with 2 SV setae,
A9 with SD setiform. Immature stages poorly
known, feed on Arecaceae, Malpighiaceae, Plat-
anaceae, Polygonaceae, Verbenaceae. New
World and Oriental region; 6 genera, about 25
species (Acria, Antoloea, Durrantia, Letogenes,
Peleopoda, Pseuderotis).
Ronald W. Hodges
Bourquin (1957), Duckworth (1970), Hodges
(1974), Kuznetsov & Stekol'nikov (1984, as Acri-
inae), Minet (1986, 1990).
Amphisbatidae (fig. 9.3 F). Sister-group to clades
31-36; defined by 1) gnathos an articulated
band with mesial, spined bulb (parallelism), 2)
second abdominal sternum with venula only,
venula and apdeme, or apdeme only (poly-
morphy), 3) 9 frenulum three acanthi (reversal),
4) $ retinaculum diffuse, anteriorly directed
scales on CuA, or anteriorly directed scales be-
tween CuA and R (polymorphy). Uncus broad
based, tapering to broadly rounded or acute
apex; signum absent or a linear band with al-
ternating, lateral, toothed extensions; larva with
SD hairlike on A9; antennae meet mesially on
pupa. Larva (Amphisbatis) case bearer on Aster-
aceae and Ericaceae; few others leaf tiers/rollers
on Aceraceae, Betulaceae, Cornaceae, Fabaceae,
Fagaceae, Juglandaceae, Myricaceae, Oleaceae,
Rosaceae, Salicaceae, Tiliaceae, Ulmaceae; most
unknown. Mainly New World and Palearctic; 11
genera, 65 species (Amphisbatis, Doina, Eupragia,
Machimia, Psilocorsis).
Hodges (1974), Minet (1990), Stehr (1987),
Toll (1964).
Cosmopterigidae. Sister-group to clade 34. De-
fined by 1) gnathos absent (parallelism), 2) ae-
deagus ankylosed (parallelism), 3) $ retinaculum
series of anteriorly directed scales between CuA
and R or series of anteriorly directed scales on
R (polymorphy), 4) hindwing with M3 and CuAi
almost always separate (reversal). Worldwide;
106 genera, 1628 species.
Common (1990), Fletcher (1933), Hodges
(1978), Kuroko (1982), Riedl (1969), Stehr
(1987), Zagulyaev & Sinev (1981), Zimmerman
(1978).
Chrysopeleiinae (fig. 9.3 G). Defined by hind-
wing with cell open (reversal). Uncus present or
absent (not two large, asymmetric lobes separate
to base); larva: Al with 1/2 SV setae, Al - 8 with
L1/L2 nearly horizontal, A9 with SVI setiform,
crochets uniordinal circle/ellipse (sometimes
open laterally); pupa: extremely compact, anten-
nae meet mesially, no cremaster, usually no
cremastral setae. Larvae on living plant tissue,
leaf and bark miners, stem/root gall makers,
flower/seed feeders. Worldwide except Oceania;
22 genera, 270 species (Ascalenia, Cholotis,
Ithome, Leptozestis, Perimede, Periploca, Sorha-
genia, Stilbosis, Walshia).
Clarke (1965), Hodges (1964, 1969, 1978),
Kasy (1968 a, 1968 b, 1969), Mosher (1916),
Riedl (1969), Zagulyaev & Sinev (1981).
Cosmopteriginae (fig. 9.3 I). Defined by 3 par-
allelisms 1) vinculum weakly sclerotized or open
mesially, 2) $ genitalia asymmetric, 3) antennal
pecten present and one polymorphy, hindwing
with Rs and Mi stalked or separate. Larva: Al
The Gelechioidea
with 2/3 SV setae, A 3 - 6 usually with 3 SV setae
(rarely 4-10), A9 with SD setiform or hairlike,
A l - 8 with L1/L2 nearly horizontal to nearly
vertical, crochets uni/biordinal circle (sometimes
open laterally); pupa: extremely compact to very
slender, T3 legs usually not exposed distally. Lar-
vae leaf miners/tiers, stem and root borers/gall
makers, seed feeders, aquatic, scavengers on at
least 26 plant families (including Asteraceae,
Cyperaceae, Fabaceae, Pandanaceae, Poaceae,
Rosaceae, Typhaceae, Zamiaceae, see Zimmer-
man, 1978: 1129). Worldwide; 80 genera, 1350
species (Anoncia, Asymphorodes, Cosmopterix,
Hyposmocoma, Labdia, Leptozestis, Limnaecia,
Macrobathra, Persicoptila, Pyroderces, Stagma-
tophora, Trachydora, Triclonella, Trissodoris).
Clarke (1965), Common (1990), Gaedike
(1967), Hodges (1962, 1978), Riedl (1969), Zagu-
lyaev & Sinev (1981), Zimmerman (1978).
Antequerinae (fig. 9.3 H). Defined by 9 frenu-
lum with three acanthi (reversal), retinaculum
usually a row of anteriorly directed scales be-
tween CuA and R (parallelism), and hindwing
with closure of cell directed at 45 0 angle toward
base of wing from M2 (parallelism). Aedeagus
free, valva a single lobe, costal/saccular margins
not developed nor differentiated; larva: Al with
one SV seta, A9 with SD setiform, crochets
sparse, a uniordinal circle; pupa unknown.
Larva of Euclemensia parasitoids of armored
scale insects. North America, England; 4 genera,
8 species (Antequera, Euclemensia, Panealia,
Ramphis).
Hodges (1978).
Gelechiidae. Defined by gnathos a pair of lateral,
articulated, symmetric sclerites with an articu-
lated, mesial hook (synapomorphy); forewing
with CuP absent (parallelism); and three poly-
morphies: 1) hindwing with R1 and Sc united
from base of wing or R1 running into Sc beyond
base of wing, 2) hindwing with closure of cell
perpendicular with long axis of wing or directed
at 45 angle toward base of wing from M2, 3)
appendix appendicular absent or present.
Hindwing usually with outer margin excavated
posterad of apex. Larvae usually leaf rollers/
tiers, also within seeds, stem/root galls, mining
leaves/needles, recorded from 82 plant families
(Gymnospermae and Angiospermae). World-
wide; 507 genera, 4,530 + species.
Benander (1937), Clarke (1969 a, 1969 b),
Common (1990), Hodges (1986), Janse (1949-
1954, 1958-1963), Meyrick (1925), Moriuti
(1982), Mosher (1916), Patocka (1987 a, 1987 b,
1987c, 1987d, 1987e, 1988), Piskunov (1981),
Powell (1980), Sattler (1973, 1979), Stehr (1987),
Weber (1948).
Physoptilinae (fig. 9.3 J). Sister-group to clade
35. Defined by 3 parallelisms 1) aedeagus anky-
losed, 2) 9 retinaculum of anteriorly directed se-
tae starting in cell between base of CuA and R
147
and extending to area between Sc and R, 3) pupa
with two rows of spines on terga 47 (USNM).
$$ and 9 have area of hindwing between Rs
and CuA lacking broad scales, except on veins,
scales very slender, wing appears transparent
(possible autapomorphy). Characters of larva
unknown; larva stem borer in Lecythidaceae. In-
dia, Java, Borneo, Australia; 1 genus, 7 species
(Physoptila).
Clarke (1963), Common (1974), Meyrick
(1914).
Gelechiinae (fig. 9.3 K). Defined by abdominal
support a pair of venulae and a pair of apodemes
(Anacampsis and allies often with very short,
broad venulae) (parallelism). Several different S
genital types typified by Anarsia, Aristotelia, Chi-
onodes, Gelechia, Gnorimoschema, Hypatima,
Isophrictis, Recurvara. Larva: frontoclypeus 1/2
distance-nearly to epicranial notch, Al usually
with 2 SV setae, A9 with SD setiform on hair-
like, anal fork usually absent, crochets absent/
few/many in circle or 2 rows; pupa: antennae
usually touch mesially, diverge distally, A7 often
with fringe of hairs on posterior margin, cremas-
ter usually absent. Larval habits and hosts nearly
as for family. Worldwide; 480 genera, 3,400 spe-
cies (Anarsia, Aproaerema, Apatetris, Aroga, Bat-
taristis, Bryotropha, Caryocolum, Coleotechnites,
Compsolechia, Deltophora, Exoteleia, Filatima,
Keiferia, Kiwaia, Lita, Megacraspedus, Metzne-
ria, Mirificarma, Ornativalva, Palumbina,
Phthorimaea, Polyhymno, Protolechia, Pycnos-
tola, Rhynchopacha, Schizovalva, Scrobipalpa,
Scrobipalpula, Stegasta, Stomopteryx, Telphusa).
Benander (1937), Common (1990), Hodges
(1986), Janse (1949-1954, 1958-1963), Meyrick
(1925), Minet (1979), Piskunov (1981), Po-
volny & Sustek (1987), Sattler (1973).
Dichomeridinae (fig. 9.3 L). Defined by 1)
pupa with raised, setose mesial/submesial promi-
nences on terga 1/2, 2/3, 3/4 (apomorphy), 2) cor-
pus bursae with prominent secondary bursa
(apomorphy), 3) abdominal support a pair of
venulae (reversal), 4) frenum absent (paral-
lelism), 5) forewing with CuAi/CuA2 stalked and
downcurved from end of cell, 6) hindwing with
R1 running into Sc beyond base of wing (paral-
lelism), 7) appendix appendicular present (paral-
lelism). Larva: frontoclypeus 1/2-3/4 distance to
epicranial notch, Al with 2 SV setae, A9 with
SD hairlike, anal fork usually present, prolegs
noticeably long, slender, crochets usually biordi-
nal in two sparse rows; pupa: antennae usually
touch mesially, cremaster usually absent. Larvae
leaf tiers on many plant families. Worldwide, but
not endemic to New Zealand or Hawaii; 4 gen-
era, 1,000 + species (Brachmia, Dichomeris, Hel-
cystogramma, Scodes).
Hodges (1986), Povolny (1980, 1983).
Pexicopiinae (fig. 9.3 M). Defined by abdomi-
nal support a pair of apodemes (rarely venulae
+ apodemes) (polymorphy). Ocellus usually pre-
148 Ronald W. Hodges

ocellus

antennal
pecten

condyles

submental pit

posterior
pinaculum pore

SV
spiracle 0

Fig. 9.7. Structural Features of Gelechioidea. A, hindwing, Perittia eremonoma, <?; - C , lateral view of head: ,
Dichomeris simpliciella; C, Sitotroga cerealella; D, base of antenna, Dichomeris punctipennella, <$; EG, posterior
end of pupa: E, Physoptila pinguivora\ F, Cerconota attornila; G, Parametriotes bipunctella; HJ, larva: H, ventral
view of head, Blastobasis glandulella; I, lateral view of segment A6, Parametriotes bipunctella; J, lateral view of
part of segment A6, Autosticha pelodes.

sent, antennal pecten often present; larva: fron-
toclypeus 3/5 distance to epicranial notch, Al
with 2 (1) SV setae, crochets 1, 514 in uniordi-
nal lateral penellipse, A9 with SD usually seti-
form, AIO without anal fork; pupa: antennae
meet mesially, usually diverge apically, cremaster
absent, cremastral spines usually present. Larvae
internal feeders in seeds/developing seed pods
(Bombacaceae, Malvaceae, 1 on Poaceae), most
life histories unknown. Mainly pantropical, but
The Gelechioidea
also Palearctic, New Zealand; 21 genera, 110 spe-
cies (Pectinophora, Pexicopia, Sitotroga, Thio-
trie ha).
Common (1958), Hodges (1986).
Acknowledgments
I thank Drs. V. O. Becker, I. F. B. Common, J.-
F. Landry, J. Minet, S. Passoa, J. A. Powell, K.
Sattler, . A. Sols, and . Vandenberg for re-
view of the manuscript and my wife, E. R. S.
Hodges, for the line drawings.
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Addendum
A number of major contributions to gelechioid
systematics have appeared after the completion
of this chapter. Fetz (1994) provided a massive
amount of novel factual information on larval
structure of Oecophoridae (as previously delim-
ited), but the significance of the phylogenetic
conclusions presented is limited by the restricted
taxonomic coverage of the material studied. A
large proportion of the Australian oecophorines
were reviewed by Common (1994, 1997). The
classification within the Elachistidae-Elachisti-
nae were discussed by Traugott-Olsen (1995
1996), while Albrecht & Kaila (1997) examined
the systematic significance of wing venation in
this taxon. There are regional 'fauna' treatments
of Oecophoridae (as previously delimited) (Han-
154
nemann 1995, 1997), Elachistidae-Elachistinae
(Bland 1996; see also Kaila 1997 for an entry
into recent literature on Nearctic taxa), Xyloryc-
tidae-Scythridinae (Bengtsson 1997), Coleophor-
idae-Coleophorinae (Baldizzone 1994, Emmet et
al. 1996). N e w regional checklists cover the
gelechioid faunas o f Australia (Nielsen et al.
1996) and Europe (Karsholt & Razowski 1996).
Albrecht, A. & Kaila, L. (1997): variation of wing ve-
nation in Elachistidae (Lepidoptera: Gelechioidea):
methodology and implications to systematics. -
Syst. Ent. 22: 185-198.
Baldizzone, G. (1994): Coleophoridae dell'Area Irano-
Anatolica e regioni limitrofe (Lepidoptera). -
Assoc. nat. piemont. Mem. 3: 1 - 4 2 3 .
Bengtsson, B. A. (1997): Scythrididae. - Microlepidop-
tera of Europe 2, 1 - 3 0 1 .
Bland, K.P. (1996): Elachistidae. In Emmet, A . M .
(ed.) The Moths and Butterflies of Great Britain and
Ireland 3, 339-410.
Common, I. F. B. (1994): Oecophorine genera of Aus-
tralia. I. The Wingia group. Monogr. Aust. Lepid.
3: [i-xvi], 1 - 3 9 0 .
(1997): Oecophorine genera of Australia. II. The
Chezala, Philbota and Eulechria groups. - Monogr.
Aust. Lepid. 5: [i-xvi], 1 - 4 0 7 .
Emmet, A. M., Langmaid, J. R., Bland, K. P., Corley,
M. F. V. & Razowski, J. (1996): Coleophoridae. In
Emmet, A. M. (ed.) The Moths and Butterflies of
Great Britain and Ireland 3, 126338.
Fetz, R. (1994): Larvalmorphologische Beitrge zum
phylogenetischen System der ehemaligen Oecopho-
ridae (Lepidoptera, Gelechioidea). - Neue ent.
Nachr. 33: 1 - 2 7 3 .
Hannemann, H. J. (1995): Kleinschmetterlinge oder
Microlepidoptera IV. Flachleibmotten (Depressari-
idae) Tierwelt Deutschlands 69, 1 192.
(1997): Kleinschmetterlinge oder Microlepidoptera
V. Oecophoridae, Chimabachidae, Carcinidae, Eth-
miidae, Stathmopodidae. Tierwelt Deutschlands
10, 1 - 1 6 3 .
Kaila, L. (1997): A revision of the Nearctic species of
Elachista s. 1. II. The argentella group (Lepidoptera,
Elachistidae). Acta zool. fennica 206: 1 - 9 3 .
Traugott-Olsen, E. (1995-1996): Phylogeny of the
subfamily Elachistinae s. str. I - I V . - Shilap 23:
153-180, 257-290, 417-449; 24: 129-149.
Appendix 1. Characters and character states used to
classify Gelechioidea.
1. Valva undivided (0); divided, costa developed as a
free lobe that appears to arise from the saccular
part of the valva (fig. 9.4 A) (1).
2. Juxta present (fig. 9.4 A) (0); absent (fig. 9.5 A)
(1). [The juxta is a sclerotized plate in the
ventromesial wall of the anellus (Klots, 1970); it is
not attached to the vinculum.]
3. Transtilla absent (0); present (fig. 9.4 A) (1).
4. Gnathos a sclerotized band that is generally uni-
formly wide, fused to tegumen, sometimes incom-
plete mesially (fig. 9.4 B) (0); fused to tegumen, a
sclerotized band that is expanded mesially (fig.
9.4 C) (1); fused to tegumen, a sclerotized band
with mesial region downturned and laterally com-
pressed (fig. 9.4 D, F) (2); fused to tegumen, a scle-
rotized band with a mesial bulb bearing parallel
Ronald W. Hodges
rows of short spines (fig. 9.4 G) (3); an articulated
band with mesial region slightly expanded (fig.
9.4 H) (4); an articulated band with an unarticu-
lated mesial hook (fig. 9.4 J) (5); an articulated
band with a mesial bulb bearing parallel rows of
short spines (fig. 9.4 I), sometimes with arms of
gnathos separate and each ending in spinose bulb
bearing short spines in parallel rows (fig. 9.4 K)
(6); a pair of articulated rami, each bearing spi-
nules at its apex (fig. 9.4 L) (7); a pair of lateral,
articulated, symmetric sclerites with an articulated
mesial hook (fig. 9.5 A) (8); absent (9). [The ante-
romesial part of the gnathos in primitive Gelechi-
oidea is finely scobinate, having a series of dia-
mond-shaped projections. This surface structure is
present in many Oecophoridae. The highly modi-
fied gnathos with a mesial knob and parallel rows
of very small spines likely is a development from
the condition in Oecophoridae and probably has
occurred more than once.]
5. Uncus present (fig. 9.4 A) (0); absent (fig. 9.4 G)
(1).
6. Vinculum a slender band, generally and uniformly
sclerotized, saccal region expanded or not (fig.
9.4 A) (0); weakly sclerotized or open mesially (fig.
9.5 B) (1).
7. Sicae (Heinrich, 1920) absent (fig. 9.4 A) (0); sicae
present (fig. 9.5 C) (1).
8. Aedeagus free (0); ankylosed (fused with sur-
rounding diaphragma) (1).
9. Male genitalia symmetric (0); asymmetric (1).
10. Abdominal second sternum; venula only (fig.
9.5 D) (0); venula + apdeme (fig. 9.5 E) (1); ap-
deme only (fig. 9.5 F) (2); neither venula or ap-
deme present (3).
11. Abdominal terga without spiniform setae (0); with
spiniform setae in band across segment (fig. 9.5 G)
(1); with spiniform setae in two submesial patches
(fig. 9.5 I) (2); with row of spiniform setae on pos-
terior margin of most segments (fig. 9.5 H) (3).
12. Female frenulum with three acanthi (fig. 9.5 J) (0);
two acanthi (fig. 9.5 L) (1); one acanthus (fig.
9.5 K) (2).
13. Female retinaculum consisting of diffusely distrib-
uted, anteriorly directed, slightly upturned scales
(fig. 9.6 A) (0); anteriorly directed scales on CuA
(fig. 9.61) (1); anteriorly directed scales between
CuA and R (usually in a row) (fig. 9.6 B) (2); ante-
riorly directed scales starting in cell and extending
to area between SC and R (3); anteriorly directed
scales between CuA and R (usually in a row), then
on R farther out from base (fig. 9.6 E) (4); anteri-
orly directed scales on R (fig. 9.6 F) (5); anteriorly
directed scales between Sc and R (fig. 9.6 C, D)
(6). [Posteriorly directed, slender scales/setae on Sc
near base of forewing almost always present.]
14. Fren um absent (0); present (fig. 9.6 G, J) (1). [Fre-
num = secondary wing coupling structure con-
sisting of anterodistally directed, strong, slender
scales on ventral surface of forewing on/near pos-
terior margin and posteriodistally directed, strong,
slender scales on dorsal surface of hindwing on/
near anterior margin.]
15. Forewing with Rs terminating on outer margin
(fig. 9.6 A) (0); R5 terminating on costa (fig.
9.6 F) (1).
16. Forewing with CuAi/CuA2 separate and directed
toward outer margin (fig. 9.6 F) (0); CuAi/CuA2
The Gelechioidea 155

stalked/connate/approximate and directed toward
posterior margin at origin from end of cell (fig.
9.6 M ) (1).
17. Forewing with C u P present (fig. 9.6 H) (0); C u P
absent (fig. 9.6 F, M) (1).
18. Forewing with pterostigma absent (fig. 9.6 M) (0);
pterostigma present (fig. 9.6 F) (1).
19. Hindwing with Sc/Ri united f r o m base of wing
(fig. 9.6 K) (0); Ri running into Sc beyond base of
wing (fig. 9.6 P) (1).
20. Hindwing with R s / M , separate (fig. 9.6 K.) (0); Rs/
Mi stalked (including connate or approximate
then diverging) (fig. 9.6 N, P) (1).
21. Hindwing with M , / M 2 separate (fig. 9.6 P) (0);
M1/M2 stalked (fig. 9.6 L) (1); M , / M 2 fused (fig.
9.6.0) (2).
22. Hindwing with M 3 / C u A i separate (fig. 9.6 N) (0);
M j / C u A i connate or stalked (fig. 9.6 P) (1); M 3 /
C u A i fused (fig. 9.6 O) (2).
23. Hindwing angle of closure of cell by vein M 2 - M 3
( ? M i - M 2 ) perpendicular with long axis of wing,
or within 10 toward base (fig. 9.6 K) (0); directed
at 4 5 angle toward base of wing f r o m M2 (fig.
9.7 A) (1); directed toward apex (fig. 9.6 P) (2);
open (fig. 9.6 N) (3).
24. Hindwing outer margin rounded/straight (fig.
9.6 L) (0); outer margin slightly excavated just
posterad of apex (fig. 9.6 P) (1).
25. Hindwing without scales or stout setae on costal
margin (fig. 9.6 N) (0); with scales or stout setae
on costal margin (fig. 9.6 O) (1).
26. Ocellus present (fig. 9.7 B) (0); absent (fig. 9.7 C)
(1).
27. Antennal pecten absent (fig. 9.7 B) (0); present (fig.
9.7 C) (1).
28. Antennal notch ( J ) absent (0); present (fig. 9.7 D)
(1)
29. Male genitalia with appendix appendicular absent
(fig. 9.4 A) (0); present (fig. 9.5 A) (1).
30. Haustellum present (0); absent (1).
31. P u p a without lateral condyles on segments 5/6, 61
7 (0); with lateral condyles on segments 5/6, 6/7
(fig. 9.7 F) (1).
32. P u p a without ventrolateral projections on segment
8/9 (0); with anteriorly directed, ventrolateral pro-
jections on segment 9 (sometimes 8/9) ("pupal
legs") (fig. 9.7 G) (1).
33. P u p a without row of spines on terga 4, 5, 6, 7 (fig.
9.7 F) (0); with row of spines on terga 4, 5, 6, 7
(fig. 9.7 E) (1).
34. Late instar larva without submental pit (0); with
submental pit (fig. 9.7 H) (1).
35. Larva without secondary SV setae on A 3 - 9 (SVI,
SV2, SV3 setae only) (0); with secondary SV setae
on A 3 - 9 (fig. 9.7 I) (1).
36. Male genitalia with manica m e m b r a n o u s (0); man-
ica sclerotized (fig. 9.4 E) (1).
37. Larval abdominal segments A l - 8 without a pore
posterad of S D (0); with a pore posterad or post-
eroventrad of SD (fig. 9.7 J) (1).
38. Larval abdominal segments Al8 without a pina-
culum ring a r o u n d SD (0); with a pinaculum ring
a r o u n d S D (fig. 9.7 J) (1).

Appendix 2. Character states for family/subfamily groups of Gelechioidea.

Missing data and polymorphic characters are represented by a "?."

12345 67890 12345 67890 12345 67890 12345 678

hypothetical 00000 00000 00000 00000 00000 00000 00000 000

Aeolanthinae 0009? 10102 00200 10001 01700 10000 110?? 0??

Agonoxeninae 00??? 00000 0??01 0?00? 70377 17000 11001 000
Amphisbatidae 00060 0000? 00?01 00000 01000 10000 00000 000
Antequerinae 01090 00101 00201 01000 00100 70000 00000 100
Autostichinae 00? 50 00001 1130? ?000? 010?0 10000 00000 Oil

Batrachedrinae 00040 0000? 2???1 0?000 1230? 17000 00070 000

Blastobasinae 10000 00?01 1??01 70100 01000 11700 00010 0??
Chimabachidae 00100 00002 0?200 00000 07100 ?0000 00000 000
Chrysopeleiinae 0109? ?01?1 0??01 0100? 0030? 77070 00000 000
Coleophorinae 10131 00100 22601 1000? 70000 11000 00000 000

Cosmopteriginae 01090 10111 0??01 01001 ?0?0? 71000 0000? 100

Deocloninae 00?60 00001 ?1?01 01001 01310 10000 00000 000
Depressariinae 00?60 0000? 0020? 70000 0??00 ??000 10000 000
Deuterogoniinae 00090 00001 01200 00000 OHIO 11000 ????? 0??
Dichomeridinae 0?080 ?1??0 07501 1101? 07710 70710 00000 000

Elachistinae 00160 0000? 07101 00001 00100 11000 10000 000

Epimarptinae 00040 00002 21100 00001 20300 10100 00??? 0??
Ethmiinae 00?0? ?0000 07201 00?00 0??00 1?000 17001 000
Gelechiinae 0108? ????1 00501 01??? ???1? ?00?0 00000 000
Glyphidoceridae 00000 00001 01701 11001 01170 10700 00000 000

Holcopogoninae 00?50 00001 1160? 00001 01000 10000 00000 000

Hypertrophinae 0009? 00000 07707 00700 0?000 10000 11001 000
Lecithoceridae 00020 0000? 1101? 71001 ?1?10 10000 00001 Oil
Momphinae 10190 00000 21101 00000 70000 11000 00000 000
Oeciinae 00000 00001 1161? 00001 00300 10001 00??? 0??

Oecophorinae 00? 10 00?0? ???0? 00000 0??00 17000 00070 00?

Peleopodinae 00060 ?00?? 0?201 00001 0??00 10000 00000 000
156 Ronald W. Hodges

Pexicopiinae 01080 0007? 07501 01077 07017 77070 00000 000

Physoptilinae 00080 00101 00311 00000 OHIO 10000 00177 000
Pterolonchinae 00050 00002 17001 00000 00000 11001 00000 000
Schistonoeinae 00100 00002 11211 00011 01710 10000 00777 077
Scythridinae 0000? 70177 07770 00000 00700 71000 00071 111
Stathmopodinae 00010 00000 37101 00000 07300 10000 00007 000
Stenomatinae ?0?0? 70707 07700 70707 07700 10000 17000 000
Symmocinae 00750 00007 11277 71007 07000 17000 00000 010
Syringopaidae 00090 10007 11001 00000 11300 10000 00710 000
Xyloryctinae 00700 00007 17200 00777 07100 10000 00771 Oil

Appendix 3. Polymorphie characters. States are indicated in square brackets and separated by a "." States indi-
cated in parentheses occur infrequently to rarely within the taxon.
Aeolanthinae 5[0.(1)] 23[0.2]
Agonoxeninae 3[0.t] 4[(6).7] 5[0.1] 12[0.1] 13[1.2.5.6 ] 17[0.(1)] 20[0.1] 21[(0).l] 24[0.(1)] 25[0.1] 27[0.1]
Amphisbatidae 10[0.1.2.] 13[0.1.2]
Antequerinae 2[0.1] 26[0.1]
Autostichinae 3[0.1 ] 15[0.1 ] 16[(0). 1 ] 20[0.1 ] 24[0.1 ]
Batrachedrinae 6[0.(1)] 10[<?0,?1] 12[1.(2)] 13[1.6] 14[0.1] 17[0.1] 250.(1)] 27[0.1] 34[(0).l]
Blastobasinae 8[0.1] 12[0.1] 13[2.4] 16[0.1] 28[0.1] 37[(0).l] 38[(0).l]
Chimabachinae 12[0.1 ] 22[0.1 ] 26[(0). 1 ]
Chrysopeleiinae 5[0.1] 6[0.(1)] 9[0.1] 12[0.(1)] 13[1.(2)] 20[1.(0)] 25[(0).l] 26[0.1] 27[0.1] 29[0.(1)]
Coleophorinae 20[0.1 ] 21 [0.2]
Cosmopteriginae 12[0.1] 13[1.2.6] 21[0.1.2] 23[0.(1)] 25[0.(1)] 26[0.1] 35[0.(1)]
Deocloninae 3[0.1] 11[1.2] 13[1.5]
Depressariinae 3[0.1] 10[(1).2] 15[0.1] 16[0.1] 22[0.1.2] 23[0.1] 26[0.1] 27[0.1]
Deuterogoniinae none
Dichomeridinae 2[(0).l] 6[0.(1)] 8[0.(1)] 9[0.1] 12[0.(1)] 20[0.1] 22[0.1] 23[1.2] 26[0.1] 28[0.(1)]
Elachistinae 10[0.2.3] 12[(0).1.(2)]
Epimarptinae none
Ethmiinae 3[0.1] 5[0.1] 6[0.(1)] 12[0.(1)] 18[0.(1)] 22[0.1] 23[0.1] 27[0.(1)] 32[0.1]
Gelechiinae 5[0.(1)] 6[0.1] 7[0.1] 8[0.1] 9[0.1] 18[0.(1)] 19[0.1] 20[0.1] 21[0.1.2] 22[0.1] 23[(0).l] 25[0.(1)]
26[0.1] 29[0.1]
Glyphidoceridae 13[1.5] 24[0.1] 28[0.1]
Holcopogoninae 3[0.1] 15[0.1]
Hypertrophinae 5[0.1] 12[0.(1)] 13[1.(2)] 15[0.1] 18[0.(1)] 22[0.1]
Lecithoceridae 10[2.3] 15[0.1] 16[0.1] 21[0.(1.2)] 23[(0).l]
Momphinae 21 [0.1]
Oeciinae 15[(0).l]
Oecophorinae 3(0.1] 8[0.1] 10[0.1.2] 11[0.1] 12[0.1] 13[1.2] 15[0.1] 22[0.1.2] 23[0.1] 27[0.1] 34[0.(1)] 38[0.(1)]
Peleopodinae 6[0.(1)] 9[0.(1)] 1 0 ^ 2 , ? 1.(2)] 12[0.(1)] 22[1.2] 23[0.1]
Pexicopiinae 9[0.(1>] 10[(1).2] 12[0.(1)] 19[0.1] 20[0.1] 22[0.1] 25[0.1] 26[0.(1)] 27[0.1] 29[0.1]
Physoptilinae none
Pterolonchinae 12[0.1]
Schistonoeinae 23[0.1]
Scythridinae 2[0.1] 5[0.1] 6[0.1] 9[0.1] 12[0.1.2] 13[1.2] 14[0.1] 18[0.(1)] 23[(0).l] 26[0.1] 34[0.1]
Stathmopodinae 12[0.(2)] 22[0.1] 35[0.(1)]
Stenomatinae 1[0.(1)] 3[0.(1)] 5[0.(1)] 6[0.1] 8[0.1] 10[<J(2).3,?2] 12[(0).l] 13[1.(2)] 16[0.(1)] 18[0.1] 20[(0).l]
22[1.(2)] 23[1.2] 32[0.(1)]
Symmocinae 3[0.1] 10[0.1.2] 14[0.(1)] 15[0.1] 16[0.1] 20[(0).l] 22[0.1] 27[0.(1)]
Syringopainae 10[c?3,92]
Xyloryctinae 3[0.(1)] 10[1.(2)] 12[(0).l] 18[0.1] 19[0.1] 20[0.1] 22[0.1] 33[0.(1)]

Appendix 4. Valid and synonymic family/subfamily-group names of Gelechioidea 1 2 .

Elachistidae Bruand, 1850: 50, extended concept
Stenomatinae Meyrick, 1906: 50
Ethmiinae Busck, 1909: 91
Azinidae Walsingham, 1906: 177
Depressariinae Meyrick, 1883: 123
Epigraphiinae Guene, 1845: 306, revised status
Cryptolechiidae Meyrick, 1883: 124
Haemilidae Bruand, 1850: 38
Enicostominae Heslop, 1938: 82 [nomenclaturally invalid]
Thalamarchellidae Turner, 1939: 335
Semioscopidinae Kloet & Hinks, 1945: 131 [nomenclaturally invalid]
Elachistinae Bruand, 1850: 50
The Gelechioidea

Cycnodiidae Busck, 1909: 92

Aphelosetiidae Hampson, 1918: 387
Agonoxeninae Meyrick, 1926: 245
Blastodacninae Clarke, 1962: 125 [nomenclaturally invalid]
Paramelriotinae Capuse, 1971: 55
Hypertrophinae Fletcher, 1929: ii
Deuterogoniinae Spuler, 1910: 349, new synonymy, revised status
Aeolanthinae Kuzntzov & Stekol'nikov, 1984: 6, new synonymy, revised status
Xyloryctidae Meyrick, 1890: 23, revised status, new concept
Xyloryctinae Meyrick, 1890: 23
Cryptophasidae Kirby, 1897: 303
Uzuchidae Hampson, 1918: 386
Scythridinae Rebel, 1901: 179, new synonymy, revised status
Butalidae Heinemann & Wocke, [1876]: 436 [nomenclaturally invalid]
Chimabachidae Heinemann, 1870: 130, revised status
Diurneinae Toll, 1964: 161
Glyphidoceridae, new family
Schistonoeidae, new family
Oeciinae, new subfamily
Schistonoeinae, new subfamily
Oecophoridae Bruand, (1850): 45, revised concept
Oecophorinae Bruand, (1850): 45
Dasyceridae Meyrick, 1883: 126
Metachandidae Meyrick, 1911: 275
Herrichiinae Toll, 1956: 172
Pleurotinae Toll, 1956: 174
Coelopoetinae Hodges, 1978: 9, new synonymy
Stathmopodinae Meyrick, 1913: 310
Tinaegeriidae Hampson, [1893]: 206
Ashinagidae Matsumura, 1929: 82, new synonymy
Lecithoceridae Le Marchand, 1947: 153
Lecithocerinae Le Marchand, 1947: 153
Timyridae Clarke, 1955: 21
Ceuthomadarinae Gozmny, 1978: 51
Torodorinae Gozmny, 1978: 189
Batrachedridae Heinemann & Wocke, [1876]: 524, new concept
Epimarptinae Meyrick, 1914: 776, new synonymy, new status
Batrachedrinae Heinemann & Wocke, [1876]: 524
Deoclonidae, new family
Deocloninae, new subfamily
Syringopainae, new subfamily
Coleophoridae Bruand, 1850: 54, new concept
Coleophorinae Bruand, 1850: 54
Augasmidae Heinemann & Wocke, [1876]: 526
Haploptilidae Barnes & McDunnough, 1917: 184
Eupistidae Fletcher, 1929: iii
Metriotinae Cpuse, 1971: 56
Aplotinae Cpuse, 1971: 56
Tolleophorinae Cpuse, 1971: 59
Falkovitshiinae Cpuse, 1972: 269
Momphinae Herrich-Schffer, 1857: 58, new synonymy, revised status
Lavernidae Wocke, 1871: 318
Psacaphorinae Spuler, 1898: 269
Blastobasinae Meyrick, 1894: 22, new synonymy
Pigritiinae Dietz, 1910: 3

Holcocerinae Adamski (in Adamski & Brown), 1989: 16

Pterolonchinae Meyrick, 1918: 299, new synonymy, new status
Autostichidae Le Marchand, 1947: 153, revised status, new concept
Autostichinae Le Marchand, 1947: 153
Symmocinae Gozmny, 1957: 326, new synonymy, revised status
Holcopogoninae Gozmny, 1967: 271, new synonymy, revised status
Peleopodidae Hodges, 1974: 93
Acriinae Kuzntzov & Stekol'nikov, 1984: 22
Amphisbatidae Spuler, 1910: 441, new status
Cosmopterigidae Heinemann & Wocke, [1876]: 520
Chrysopeleiinae Mosher, 1916: 104
Walshiidae Hodges, 1962: 7
Antequerinae Hodges, 1978: 16
158 Ronald W. Hodges

Cosmopteriginae Heinemann & Wocke, [1876]: 520

Diplosaridae Meyrick, 1915: 339
Hyposmocomidae Hampson, 1918: 386
Scaeosophidae Meyrick, 1922: 48
Gelechiidae Stainton, 1854: 75
Physoptilinae Meyrick, 1914: 777, revised status
Gelechiinae Stainton, 1854: 75
Anacampsidae, Bruand, 1850: 40
Litdae Bruand, 1859: 648
Palumbininae Chapman, 1902: 1963
Anomologidae Meyrick, 1926: 308
Stomopteryginae Heslop, 1938: 80 [nomenclaturally invalid]
Hypatiminae Kloet & Hincks, 1945: 129 [nomenclaturally invalid]
Apatetrinae Le Marchand, 1947: 151
Aristoteliinae Le Marchand, 1947: 152
Chelariinae Le Marchand, 1947: 153
Chrysoesthiidae Paclt, 1947: 99 [nomenclaturally invalid]
Metzneriinae Piskunov, 1975: 857
Anarsiidae Amsel, 1977: 234
Dichomeridinae Hampson, 1918: 386
Brachmiinae Heslop, 1938: 80 [nomenclaturally invalid]
Pexicopiinae Hodges, 1986: 7
1
Only names that have been treated as family or subfamily groups are included.
2
Based on strict priority the valid name for the superfamily is Epigraphioidea, for the Elachistidae is Epigraphii-
dae, for the Depressariinae is Epigraphiinae, for the Gelechiidae is Anacampsidae, and for the Gelechiinae is
Anacampsinae. Inasmuch as there has been no usage for the senior synonyms as the valid group names in more
than 50 years, a petition to conserve Gelechioidea, Elachistidae, Depressariinae, Gelechiidae, and Gelechiinae
has been prepared and sent to the International Commission Zoological Nomenclature.
 10. The Zygaenoidea
Marc E. Epstein, Henk Geertsema, Clas M. Naumann & Gerhard M. Tarmann
Like many other superfamilies within the di-
trysian Lepidoptera, the Zygaenoidea are not
characterised by clearly defined autapomorphies.
At first glance it appears that the superfamily has
been formed as a waste-paper-basket taxon to
include a number of families each of which is
well defined by one or several autapomorphies
and which otherwise share a great number of ple-
siomorphic characters. Minet (1986, following
Common, 1970) assumes that the retractile head
of the larva (at least in the late instars) and the
position of the 2nd abdominal spiracle of the
pupa, which is covered by the wings, form auta-
pomorphies of the superfamily. The latter char-
acter has so far only been verified for the Epipyr-
opidae, the Megalopygidae, the Limacodidae,
the Heterogynidae and the Zygaenidae. Both
characters could easily have developed several
times independently, but can be used as a diag-
nostic character to define the superfamily until
further detailed studies have been done.
The Limacodidae, Megalopygidae, Soma-
brachyidae, Aididae and Dalceridae make up the
'limacodid-group families'. These families, or
groups presently within them, were originally
placed within Zygaenoidea until Brock (1971)
moved them to Cossoidea. Brock's arrangement
has been followed by Fletcher & Nye (1982), the
Zoological Record and Freina & Witt (1990).
Heppner (1984, 1995) divides the limacodid
group between Zygaenoidea (megalopygids and
somabrachyids) and Cossoidea (limacodids and
dalcerids). Placement of limacodid-group fami-
lies in Cossoidea has been criticized, however,
both on the weakness of the adult characters and
on the strength of evidence of immature-stage
characters for Zygaenoidea in the earlier sense
(Common, 1975; Kuznetsov & Stekol'nikov,
1981; Minet, 1986; Epstein, 1996). Monophyly of
the limacodid-group families is supported by a
number of synapomorphic characters in the
adults and immature stages including the pres-
ence of prolegs or crochets on abdominal seg-
ments 2 - 7 , a pupa with a sculptured eye flange,
and in the adults, the presence of dense sensilla
trichodea on all legs of females and absence of
ocelli (Epstein, 1996). Brock (1990) no longer
ascribes to the arrangement in his earlier work,
including the "Eucleoidea" and Zygaenoidea in
the 'Zygaeniformia', apart from the 'Cossifor-
mia'. We include the limacodid-group families in
the Zygaenoidea.
Imagines small to medium-sized, only rarely
(subfamily Chalcosiinae of the Zygaenidae)
large; coloration ranging from cryptic grey, black
or brown to highly conspicuous or aposematic
moths, with significant metallic sheen in some of
the families with diurnal habits. Head with
maxillary palpi small, rudimentary or lacking;
proboscis well developed, rudimentary or absent,
ocelli and chaetosemata present or absent, anten-
nae bipectinate, serrate or filiform. MS and CuP
usually present in both wings, Ml slightly ante-
rior, M2 slightly posterior of the insertion of MS
in the distal part of discal cell, Se + R in hind-
wing not approximated to Rs beyond discal cell,
frenulum normally present (absent in Himan-
topteridae, Anomoeotidae, Chrysopolominae
and some Dalceridae), pterostigma absent; legs
with or without tibial epiphysis and tibial spurs.
Abdomen with or without dorsal spines; tympa-
nal organs absent. In some families the imagines
(and/or larvae) are chemically protected or pro-
duce poisonous secretions that have defensive
properties; in some families resistance to cyanide
has been observed (see under Zygaenidae).
Larva stout, with Verrucae and setae, or ap-
pearing naked with smooth or granular cuticle;
ventral surface often slug-like, with short prolegs
or suckers and crochets in mesoseries; feeding ex-
posed on foliage, sometimes leaf-mining or bor-
ing; living as ectoparasites on Homoptera in two
families (Epipyropidae, Cyclotornidae), or re-
ported to be found in the nests of ants and/or
termites (Cyclotornidae, Himantopteridae); in
some families (Heterogynidae, Zygaenidae) cu-
ticular cavities of the larval epidermis are used
for the storage of defensive secretions. In these
cavities secretions with concentrations of the cy-
anoglucosides linamarin and lotaustralin are
stored. In some groups the larvae possess pain-
fully stinging setae (Limacodidae). The larvae
frequently resemble those of lycaenid larvae.
Pupa weakly sclerotized, appendages mostly
free, abdomen with fine dorsal spines; spiracles
on 1st abdominal segment visible (except some
Epipyropidae); cocoon firm or soft, pupa usually
protruding when the moth emerges.
Zygaenoid families occur world-wide, but
some appear to be endemic to certain zoogeo-
graphical regions (Dalceridae to the Neotropics,
Cyclotornidae to Australia). Some families have
been intensively studied because of their interest-
ing life habits, reproductive behaviour or chemi-
cal ecology (Epipyropidae, Zygaenidae).
Key to families of the Zygaenoidea:
1 Hindwing extremely narrow, tailed, resem-
bling that of Nemopteridae (Neuroptera)
Himantopteridae
160 Marc E. Epstein, Henk Geertsema, Clas M. Naumann & Gerhard M. Tarmann

Hindwing otherwise 2 epiphysis and tibial spurs; forewing without reti-

2 Ocelli present 3 naculum but with an elongate area of micro-
Ocelli absent 4 trichia along inner margin; hindwing with a short
3 Wings translucent, proboscis reduced, tib-
ial spurs absent . . . Anomoeotidae (part)
Wings often bright or metallic, proboscis
well developed, tibial spurs present or ab-
sent Zygaenidae (part)
4 Chaetosemata present 5 genitalia with tegumen broad, without uncus,
- Chaetosemata absent 10
5 Forewing narrow, elongated and small . . 6
Forewing otherwise 7 valva short, stout, with pointed apex; vinculum
6 Antennae bipectinate (male) or biserrate
(female) Phaudinae (Zygaenidae)
- Antennae filiform Lacturidae
7 Rsl + (Rs2 + Rs3) or Rsl + Rs2 (African
or Asian) 8 slightly sclerotized; antrum translucent, ductus
- Rsl, Rs2, Rs3 pectinate off from Rs4 or
R + R1 and Rs4 + (Rs2 + Rs3) (New
World) 9 ondary accessory gland found in Fulgoraecia ex-
8 Tibial spurs present (wings absent in fe-
male Somabrachys) . . . Somabrachyidae
Tibial spurs absent. Anomoeotidae (part)
9 Rsl, Rs2, Rs3 pectinate off from Rs4 . . .
Megalopygidae
R + R1 and Rs4 + (Rs2 + Rs3) Aididae
10 Rsl + Rs2 and Rs3 + Rs4 stalked or
fused Dalceridae
Rsl and Rs2 independent (not stalked)
(wings of females can be absent) . . . . 11
11 Wings smoky black, all forewing veins free
of discal cell, chorda absent, wings of fe-
males absent, male antennae quadripecti-
nate Heterogynidae
Various wing colours, forewing veins
branched or free from discal cell, chorda
present or absent, male antennae never
quadripectinate 12
12 R veins unbranched or branched from end
of forewing discal cell 13
R veins branched beyond discal cell . . 14
13 1 A + 2 A without basal fork, tibial spurs
absent Epipyropidae
- 1 A + 2 A with strong basal fork, tibial
spurs present Cyclotornidae
14 Rs2 and Rs3 stalked off from Rsl or Rs4
(rarely fused) Limacodinae
Rs2 and Rs3 stalked or independent off
from Rsl Chrysopolominae
Epipyropidae (Figs. 10.1 A, B; 10.3 A, J). Imag-
ines small, blackish brown to dark grey, with tri-
angular forewing. The family is especially inter-
esting because of the ectoparasitic life habits of
the larvae which feed on a number of homopter-
ous hosts and appear to be host-specific.
Head smoothly scaled or hairy, antenna less
than half length of forewing, bipectinate in both
sexes, pectinations less pronounced or reduced in
females; maxillary palpi absent, proboscis ab-
sent, labial palpi very small or vestigial; ocelli
and chaetosemata absent. Foretibia without
frenulum consisting of a single straight bristle in
both sexes; MS in discal cell of forewing present,
with an additional areole or accessory cell, radial
and medial veins arising directly from the cell.
Abdominal segments 8 - 1 0 not extensile. Male
but with lateral seta-bearing lobes; arms of
gnathos short, stout and not fused medially;
narrow, sometimes with saccus; aedeagus short,
stout, without cornuti. Female genitalia with
small to very small papillae anales, apophyses
posteriores and apophyses anteriores, sterigma
bursae short and slender, corpus bursae small
and membraneous, without signum; large sec-
igua.
The egg is either flat-discoid or ovoid, with a
thin chorion through which the development of
the embryo can be observed; may be coated by
black secretion from secondary accessory gland
similar to Cyclotornidae, as reported by Com-
mon (1990) and (Epstein, 1996). The first instar
larva is most unusual and very different from
later instars: triunguloid, slender, tapering poste-
riorly, head and thorax comprising about half of
the entire length, labium large, spinneret long,
tapering; thoracic legs long, prolegs present or
absent; abdominal segments 2 - 7 with ventral
protuberances, each bearing long, ambulatory
setae, segment 10 with a pair of long, fine, down-
ward-curved setae. Mature larva onisciform,
strongly convex dorsally and flat ventrally; head
very small, retractile into the prothoracic seg-
ment; spinneret prominent, the six stemmata
grouped together; thoracic legs small, with only
three segments, and with a sharp backward-
curved apical claw; ventral prolegs present on
abdominal segments 36 with 5060 crochets in
a uniordinal circle. Pupa short and stout, weakly
sclerotized, cremaster absent; spiracle on 1st ab-
dominal segment either hidden (Common, 1990)
or visible (Epstein, 1996).
The adults have a rapid, erratic flight and are
attracted to artifical light, usually soon after
dusk. Parthenogenesis has been reported in at
least two species (Agamopsyche threnodes from
northern Australia and Epipomponia nawai from
Japan), but most appear to reproduce bisexually
(Yamamoto et al., 1987: 268; Common, 1990).
The females lay numerous eggs up to 3000 in
the Afro tropical species of Fulgoraecia (= Epi-
pyrops) (Common, 1990 referring to Kirkpatrick,
1947). The eggs are laid either in large batches
or in small groups on the food plant of the ho-
mopterous host. The larvae live as ectoparasites
on Homoptera and cling to silken threads spun
on the surface of the host. The epipyropid-host
The Zygaenoidea 161

Fig. 10.1: Zygaenoidea families (length of bar: 10 mm). A. Epipyropidae: Epipomponia nawai. B. Epipyropidae:
last instar larva of Epipomponia nawai on its host, Tanna japnica (Fulgoridae). C. Cyclotornidae: Cycloiorna
diplocentra. D. Himantopteridae: Semioptila hilaris. E. Anomoetidae: Thermochrous stenocraspis. F. Somabrachyi-
dae: Somabrachys aegrota. G. Megalopygidae: Megalopyge lanata. H. Lacturidae: Gymnogramma rufiventris. J.
Limacodidae, Limacodinae: Chrysamma purpuripulcra. K. Limacodidae, Chrysopolominae: Chrysopoloma venata.
L. Dalceridae: Dalcera abrasa. M. Heterogynidae: Heterogynis cf. penella.
162 Marc E. Epstein, Henk Geertsema, Clas M. Naumann & Gerhard M. Tarmann
relationship appears to be very close. The larvae
are usually covered with wax secreted by the epi-
dermis, which remains on the cuticle until pupa-
tion. At first glance it is difficult to identify the
larva with this camouflage. Prior to pupation the
larva leaves the host by using a silken thread; it
usually spins a white, ovoid, silken cocoon, heav-
ily impregnated with wax from the larval body.
Pupation takes place near the ground, on a leaf,
a twig or a stem of grass. Near the anterior end
of the cocoon a 'frog-mouth' slit is found which
is used by the protruding pupa shortly before
emergence of the moth.
Distribution mainly tropical to subtropical
world-wide, apparently most diverse in the Ori-
ental region; approximately 3 5 - 4 0 species are
known. Principal genera include Epipyrops, Epi-
pomponia and Heteropsyche.
References: Common (1970, 1990), Epstein
(1996), Holloway, Bradley & Carter (1987), Kato
(1940), Krampl & Dlabola (1983), Davis (1987),
Yamamoto et al. (1987), Lindner (1990).
Cyclotornidae (fig. 10.1 C). Imagines small,
mainly grey-coloured, with tineoid appearance.
Head smoothly scaled, antenna filiform in both
sexes but slightly thickened in male, scapus with
a tuft of scales which probably represent a pec-
ten; maxillary palpi and proboscis absent; labial
palpi short and drooping, with three segments;
ocelli and chaetosemata absent. Foretibia with-
out epiphysis, but with short tibial spurs on mid
and hindleg (0-2-4); forewing with a small mem-
braneous retinaculum on the proximal part of Sc
in male and a group of raised scales near CuA
in female, and with a group of microtrichia on
the underside of the forewing base forming an
additional wing-locking system; frenulum con-
sisting of a stout single bristle in male and several
thinner bristles in female; wing venation only
slightly reduced; MS present, branched distally
in forewing; R and Rsl4 of forewing stalked
or arising freely from discal cell; CuP present;
1 A + 2 A with strong basal fork, with a short
cross vein, thus forming an anal cell near base of
wing, and with the lower branch giving rise to a
short vein, representing the distal part of 2 A,
never reaching the wing margin. MS of hindwing
present, but not branched distally; upper half of
discal cell open, Se + R strong, Rs weak but
separate throughout its length; sometimes one
branch of M or CuA absent; 1A + 2 A anasto-
mosed basally, 2 A vestigial distally; 3 A vestigial.
Abdomen stout in both sexes, with tortricoid-like
apodemes on the second sterni te; segments 8 10
not extensile in female. Male genitalia strongly
sclerotized, with broad uncus; lateral lobes of
tegumen may represent socii, with a pair of
pointed gnathos arms, valva broad, rounded, the
ventroproximal margin firmly fused to form the
juxta, aedeagus short and slender, without cor-
nuti. Female genitalia with small papillae anales,
very short apophyses posteriores and poorly de-
fined apophyses anteriores; sterigma heavily scle-
rotized; ductus bursae short and membraneous,
corpus bursae membraneous, without signum
(Common, 1990).
Egg dark brown, ovoid, slightly flattened,
with a series of longitudinal ridges and less
prominent transverse ribs. The coloration is dark
due to a brown secretion from the very large ac-
cessory glands of the female (Common, 1990).
As in the Epipyropidae and Limacodidae, the
first instar larva differs from later instars. It is
oval in shape, very flat, with thoracic legs and a
large strongly sclerotized head capsule; the fron-
toclypeus extends to the epicranial notch; six
closely grouped stemmata present on each side;
antenna long; mandibles long and curved,
sharply pointed (used to pierce the host cuticle);
spinneret well developed; cuticle of body granu-
lose, spiracles positioned dorsally; ventral pro-
legs present on abdominal segments 3 - 6 . The
mature larva is broadly oval in shape, flattened
dorsoventrally, with a small retractile head, body
extended laterally, with pointed segments; small,
closely set anal prolegs, crochets arranged in uni-
ordinal mesoseries, body and lateral protuber-
ances clothed with fine, secondary setae. Pupa
flattened dorsoventrally, well sclerotized, with-
out cremaster, spiracle on 1st abdominal segment
visible (Common, 1990).
Imagines of Cyclotornidae have not often
been observed. The females lay large numbers of
eggs (according to Common (1990) 1400 by a
species closely related to Cyclotorna monocentra)
on twigs and bark of trees infested with Cicadel-
linae (Homoptera). The first instar larvae are ex-
tremely active and follow the routes of Iridomyr-
mex ants in order to find their cicadellid first
hosts on which they live as ectoparasites. Later
the first instar larva leaves the host and spins a
flat shelter on or under loose bark where it
moults into a brightly coloured, flat larva. This
is collected by Iridomyrmex ants and taken to
their nest, where it feeds upon the ant larvae,
yielding an anal secretion that is eagerly de-
voured by the ants. When mature the larva spins
its cocoon in the crevices of or under loose bark
of a nearby tree. The cocoon has a 'frog-mouth'
slit through which the pupa protrudes before the
emergence of the adult (Common, 1970, 1990).
Endemic to Australia. Only one genus (Cyclo-
torna) with approximately 12 known species.
References: Common (1970, 1990), Dodd
(1912).
Himantopteridae (fig. 10.1 D). Small to medium-
sized moths with rounded forewings and narrow,
tailed hindwings, the moths thus resembling
imagines of the Nemopteridae (Neuroptera). In
some Oriental species the hindwings may be
broader at the base. The family was erected by
Rogenhofer (1884). Kirby (1897) treats it under
The Zygaenoidea
the name Thymaridae. The group was transfer-
red by Alberti (1954) as a subfamily into the Zy-
gaenidae. Fletcher & Nye (1982) have once more
raised it to family rank. The systematic status of
the group is still unclear. There are arguments
placing this family within the Zygaenoidea (e. g.
the presence of dorsal abdominal spines which
are present in some other Zygaenoidea families).
According to Jordan (1907), Gaede (1926) and
Pinhey (1975, citing Vri) the larva is supposed
to have some symbiotic interactions with ter-
mites. Recent observations of a larval swarm of
Himantopterus fuscinervis in Malaysia did not re-
veal any association with termites (Barlow &
Carter, 1996).
Head covered with dense, long hairy scales,
frons flat; proboscis, maxillary and labial palpi
reduced; ocelli absent but a prominent pair of
chaetosemata present on the postero-dorsal part
of the head; antenna bipectinate in male, biser-
rate with terminal hair-tufts in female. Thorax
densely covered with hairy scales, wing-locking
microtrichia on dorsal part of metathorax and
on the forewing underside absent; epiphysis and
tibial spurs absent; venation of forewing partly
reduced, anterior part of discal cell extended,
hindwing with strongly reduced venation; frenu-
lum and retinaculum absent. Abdomen very
hairy, with dorsal spines, and sometimes caudal
hair-tufts. Male genitalia with short uncus and a
double subuncus, tegumen large and broad, val-
vae small, without processes, vinculum strong,
with prominent saccus; aedeagus very slender,
straight or curved, broader proximally and
pointed distally. Female genitalia without ovi-
positor (i. e. segments 8 - 1 0 not extensile), some-
times with strongly sclerotized 8th segment, duc-
tus bursae slender, translucent, corpus bursae
globular, translucent, signa absent, ductus semi-
nalis inserting into proximal part of corpus
bursae.
The description of the larva is summarized
from the account of H. fuscinervis by Barlow &
Carter (1996): head retractile into thorax, body
with secondary setae primarily on Verrucae as
spines or scattered slender or plumose setae; ab-
dominal segments 1 and 8 with subdorsal glan-
dular protuberances packed with minute decidu-
ous setae and final instar with segments 27
having pocket-like structures postero-dorsad of
spiracles, also filled with deciduous setae (similar
to those described in Somabrachys, below); pro-
legs on abdominal segments 3 - 6 and 10, with
those on segments 36 with triordinal crochets
in a double mesoseries and segment 10 with sin-
gle biordinal mesoseries; two subventral protu-
berances on abdominal segment 2.
Very little is known on the biology of the Hi-
man topteridae (see above). Jordan (1907) states
that the imagines have an erratic flight and are
mainly found resting on twigs and leaves. The
larva of Himantopterus dohertyi has been re-
163
ported to feed on Quercus griffithii and Q. robur
(Fletcher, 1931). H. fuscinervis was observed to
swarm on the trunk of Shorea platyclados (Dipt-
erocarpaceae) and later to defoliate the crown;
the larva forms a cocoon in the ground (Bar-
low & Carter, 1996).
Approximately 40 species are known from the
Afrotropical and Oriental regions. Principal gen-
era include Himantopterus, Semioptila and Dora-
topteryx.
References: Alberti (1954), Barlow & Carter
(1996), Fletcher (1931), Gaede (1926), Hering
(1937b), Jordan (1907), Pinhey (1975).
Anomoeotidae (Figs. 10.1 E, 10.3 B). Imagines me-
dium-sized, uniformly coloured with translucent,
apically rounded wings and geometroid habitus.
This group was treated within the zygaenid sub-
family Phaudinae by Gaede (1926), Bryk (1936)
and Pinhey (1975), within the Chalcosiidae by
Kirby (1892) and as a subfamily of the Zygaeni-
dae by Hering (1937 c) and Alberti (1954).
Fletcher & Nye (1982) raised it to family rank.
Like that of the Himantopteridae the systematic
position of this family is unclear. The biology
and larval and imaginai characters are very uni-
form in the various subgroups. It is possible that
this group will have to be transferred to another
superfamily, once the biology is better known.
Head covered with hair-like scales, without
projections and extensions; proboscis, maxillary
and labial palpi absent; ocelli present in the ge-
nus Dianeura but absent in all other genera;
chaetosemata present; antenna strongly bipecti-
nate in male, weakly bipectinate or biserrate in
female. Thorax hairy, legs without epiphysis and
tibial spurs; medial stem of forewing well devel-
oped or slightly reduced, a number of cross veins
linking the subcosta to the costa are found in
several species; Sc sometimes fused with R dis-
tally; radial veins stalked, thus forming either
(Rsl + Rs2), [R + (Rsl + Rs2)], or [Rsl + (Rs2
+ Rs3)] configurations, Rs4 usually reduced or
anastomosed with M l ; medial veins free from
discal cell or sometimes stalked, CuP well devel-
oped distally but sometimes reduced proximally;
subcosta of hindwing anastomosed with cranial
part of discal cell, the latter extended distally;
medial stem present or rarely reduced, all veins
free from the cell, M l sometimes absent; CuP
mainly well developed distally, sometimes re-
duced proximally; frenulum and retinaculum ab-
sent. Abdomen hairy, with dorsal spines. Male
genitalia (with the exception of the genus Dia-
neura which may possibly not belong to the same
monophylum as the Afrotropical species) with-
out prominent uncus but with well-developed
subunci, tegumen very strongly sclerotized, hel-
met-like, broad and connected to the vinculum
by a strong ring-like configuration; a prominent
saccus present only in Staphylinochrous\ valva
short and stout, sometimes nearly completely re-
164 Marc E. Epstein, Henk Geertsema, Clas M. Naumann & Gerhard M. Tarmann
duced; aedeagus either tube-like without or with
cornuti, or strongly curved with a broad basal
part and a pointed apical part similar to that in
the Castniidae. Female genitalia without oviposi-
tor, 8th segment strongly sclerotized; antrum
strongly sclerotized, ductus bursae short and
translucent, or long and strongly sclerotized; cor-
pus bursae translucent, with or without signum;
ductus seminalis inserting into ductus bursae
either near the ostium or, more frequently, near
the opening of the ductus into the corpus bursae.
Biology not properly known. According to
Piatt (1921) the larva of the South African spe-
cies Anomoeotes levis feeds on Bauhinia (Caesal-
piniaceae) and Gardenia (Rubiaceae).
The family includes about 40 species and oc-
curs in the Oriental (only the genus Akesina) and
Afrotropical regions (with the principal genera
Dianeura, Staphylinochrous, Anomoeotes and
Thermochrous).
References: Alberti (1954), Bryk (1936), Gaede
(1926), Hering (1937 c), Naumann (1988 a),
Piatt (1921).
Megalopygidae (Figs. 10.1 G, 10.3 K). Presently
considered to be limited to the New World and
found primarily in the tropics, Megalopygidae
consists of subfamilies Megalopyginae and Trosi-
inae. Aididae, mostly regarded a megalopygid
subfamily until recently, is considered to be a
family. Somabrachyidae, which at times has been
a megalopygid subfamily, is placed here as a sep-
arate family.
Imagines small to medium-sized, forewing
markings ranging from elaborate patterns or
spots to unmarked, at times brightly coloured,
grey or simply white; fore- and hindwings api-
cally rounded; wing scales piliform or cleft, often
tripartite. Head densely scaled; male antenna bi-
pectinate to the tip; pectinations in female an-
tenna as in males or shorter or serrate; proboscis,
maxillary and labial palpi reduced; ocelli absent
and chaetosemata present. Thorax densely
scaled; mesonotum with longitudinal depressions
(Hopp, 1934) and mesepimeron broadly sclero-
tized; epiphysis absent and tibial spurs 0-2-2; fe-
male fifth tarsomere of each leg with sensilla
trichodea beneath, intermixed with scales (Ep-
stein, 1996). Forewing with R and R s l - 4 form-
ing a pectinate branch (Aididae and Limacodi-
dae have Rs2 and Rs3 stalked); medial stem pre-
sent, Ml arising near upper edge of discal cell at
quite some distance from the origin of M2. In
the hindwing Se + R anastomosed with cranial
part of discal cell or independent from near base
(male Megalopyginae); medial stem present.
Frenulum present, partly reduced or absent. Ab-
domen hairy, short and tapering distally. Male
genitalia with a short, narrow uncus, and small
digitate valvae; pair of lobes ventrad of the val-
vae may be homologous with juxtal lobes (as
found in Aididae and Dalceridae) or ventral por-
tions of subdivided valvae (Epstein, 1996). Fe-
male abdomen with a dense tuft of piliform
scales on A7; papillae anales laterally appressed;
utriculus of the receptaculum seminis not pro-
nounced; anterior and posterior apophyses well
developed.
Eggs are covered with pilose scales originating
from the end of the female abdomen. Larva with
three rows of spined scoli (laterodorsal, lateral
and subspiracular); variably pilose (more so in
the Megalopyginae than in Trosiinae) (Holloway,
Bradley & Carter, 1987); digitate lobes near the
spiracles; six pairs of prolegs as well as the anal
claspers, often only bearing crochets on abdomi-
nal segments 36 and on the claspers, but some-
times with crochets on segments 2 and 7; unique
in the limacodid group is the presence of spongy
membraneous pads on all prolegs located to the
anterior of the crochets or covering the proleg
on segments 2 and 7 when crochets are absent;
anal fork often present (Epstein 1996). Pupa
with an eye-plate similar to that in the Limacodi-
dae and Dalceridae; cocoons either hard, with an
apparent hatch, or soft with an operculum hid-
ing the hatch; species with plumose larvae incor-
porate setae in the cocoon mesh. The Megalo-
pygidae (Megalopyginae and Trosiinae) contain
242 species primarily in Central and South
America, but also in North America (Becker,
1995; Davis, 1983).
Larvae are reported as pests of various palms,
cacao, guava, ferns (Genty et al., 1978; Miller
et al., 1995); spines cause erucism in humans
(Delgado, 1978). Adult Podalia bolivar i will feed
on a solution of water and honey absorbed on
cotton (Miller et al., 1995).
Principal genera include Podalia, Norape,
Megalopyge, Mesoscia and Trosia.
References: Becker (1995), Davis (1983), Ep-
stein (1996), Hopp (1927, 1934), Holloway,
Bradley & Carter (1987), Miller et al. (1995).
Somabrachyidae (Fig. 10.1 F). Most authors have
treated Somabrachyidae as a distinct family
within the Zygaenoidea. They share a number of
characters with megalopygids, but most can be
argued to be symplesiomorphies in relation to
the Zygaenoidea or to the limacodid group (Ep-
stein, 1996).
Imagines small to medium-sized; body hairy,
covered with piliform scales, wings triangular in
males and narrowly rounded in females, some
females (Somabrachys) wingless. Wings generally
fawn-coloured in males and almost translucent
in winged females, forewing densely covered with
hair-like scales and some darker spots; wing ex-
panse 1050 mm. Head small and densely
scaled, vertex densely and roughly scaled with
distinct frontal protuberance (autapomorphy) in
Somabrachys but less developed in Psycharium,
antenna short, bipectinate in males and filiform
in females; proboscis and maxillary palpi absent,
The Zygaenoidea
labial palpus reduced to two segments; ocelli ab-
sent and chaetosema present (not always clearly
discernable) latero-posteriad of antennal inser-
tion, eyes large, longer than broad. Thorax large,
densely covered with setiferous scales, legs
densely scaled, epiphysis of foretibia absent but
proximal claw present in females of Somabrachys
and in male of undescribed taxon from South
Africa, tibial spurs 0-2-2 but small. Wings absent
in some females (see below). Forewing with in-
distinct retinaculum, frenulum formed by spi-
ralled bristles in Somabrachys and three to six
bristles in Psycharium; venation only slightly re-
duced; forewing in males triangular with concave
costal edge, in females narrowly rounded, M pre-
sent, sometimes branched distally, Sc and R
closely appressed, R short and merging into Sc
in Psycharium, Rsl separate anterior of apex,
Rs2 + Rs3 stalked with Rsl, Rs4 free from up-
per angle of cell, CuP present and 1 A + 2 A with
basal fork; hindwing with unbranched M; Sc +
R and Rs arising separately, but closely, from
upper angle of cell; CuP, 1 A + 2 A and 3 A pre-
sent, 1 A + 2 A not forked basally. Abdomen
short in male, larger in female, densely covered
with setiferous scales and transverse row of spiny
setae on posterior margin of segments. Male gen-
italia with tegumen large and bulbous, uncus
stout with downcurved medial projection,
gnathos absent, valva triangular, simple, without
projections; aedeagus simple, often flexed along
proximal part, cornuti present. Female genitalia
(Psycharium) with weakly sclerotized ovipositor
lobes; antrum simple, corpus bursae without
signa, pseudobursa elongate, receptaculum sem-
inis complex and sebaceous glands robust, Peter-
sen's glands absent.
Egg flat, ovoid, with thin chorion, eggs laid in
clusters and covered with irritant setiferous
scales originating from abdomen of female moth.
Larva with rows of Verrucae on thorax and ab-
domen. Head retracted into enlarged prothoracic
lobe, except when feeding. Head with stemma 5
disjunct, two stemmatal setae, spinneret and la-
brum all as in Megalopygidae. Psycharium with
a variety of Verrucae, dorsal Verrucae on meta-
thoracic and abdominal segments 1 8 consisting
of eversible urticating spines, Verrucae on lateral
thoracic and abdominal segments consisting of
tactile setae; abdominal segment 1 with ventral
pair of gibbosities. Somabrachys have Verrucae
with both urticating and tactile setae; abdominal
segments 1 8 each with two Verrucae and a shal-
low, recessed pouch above the spiracle; the lower
verruca, pouch and spiracles with thin, detacha-
ble hair-like scales; abdominal segments 19
with three Verrucae below spiracle. Thoracic legs
large in relation to prolegs, especially in Psycha-
rium. Prolegs present on abdominal segments
27, those on segments 2 and 7 without
crochets. Crochets in later instars arranged in
uniordinal lateroseries and a shorter anterior un-
165
iordinal mesoseries. Early instar prolegs with
two spatulate setae on anterior margin in
Psycharium. Late instar prolegs with up to seven
stout, spinose SV setae, anterolateral to crochets
(Psycharium) or with normal hair-like setae lat-
eral to crochets (Somabrachys). Anal plate nor-
mal size, with numerous tactile setae; anal comb
present. Pupa with distinct eye piece similar to
that in Limacodidae, pupa short and stout with
a delicate, weakly sclerotized cuticle, appendages
distinct and loosely fused to body; pupation in a
loosely woven cocoon on the ground, pupal case
protruding after emergence of moth.
The adults are nocturnal; males with erratic,
rapid flight, winged females also good fliers,
males frequently attracted to artificial light, ap-
terous females of Somabrachys (and undescribed
taxon from South Africa) fast walkers. After ovi-
position, wings of dead female folded over emp-
tied abdomen shielding egg cluster. Eggs hatch
after about 10 days, larval development from
four to six months. After cocoon formation, lar-
vae aestivate for some months, pupating about
three weeks before emergence of the moth. Lar-
val movement sluggish, positively phototropic
when feeding. Larva initially gregarious and not
feeding but after first moult dispersing, usually
on silken thread, to start feeding. Faecal pellets
ejected forcebly by anal comb. Larvae of Soma-
brachys polyphagous, larvae of Psycharium feed-
ing on Restionaceae, Cyperaceae, Bruniaceae
and the exotic Pinus radiata (Coniferae).
Somabrachyidae are found in the Palaearctic
(southern Spain, North Africa and eastern Medi-
terranean Somabrachys) and the Afrotropical
(South Africa Psycharium and possibly Mada-
gascar Boisduvalodes) zoogeographical re-
gions. Although Boisduvalodes is presently
placed in Limacodidae, it may well prove to be
a member of Somabrachyidae. Many new taxa
from South Africa await formal description.
References: de Freina & Witt (1990), Epstein
(1996), Hampson (1920), Hering (1933), Hopp
(1927), Jordan (1916), Oberthr (1911, 1922),
Powell (1916), Seitz (1912), Viette (1980).
Aididae. This small taxon has frequently been
placed as a subfamily of Megalopygidae until a
recent phylogenetic analysis found it to be a sep-
arate family and sister group to the limacodid +
dalcerid clade (Epstein, 1996). Possible autapo-
morphies: male antenna bipectinate from half to
two-thirds distance from base; veins R and Rsl
stalked in distal third of wing beyond discal cell
and near costa; larval spinneret tubular, with
amorphous tip; cocoons with 1 or 2 pairs of
holes in outer mesh (Epstein, 1996).
Imagines medium-sized, often grey on fore-
wings with contrasting red hindwings. Labial
palpus either two- or three-segmented, porrect,
reaching as high as dorsum of eye; proboscis and
maxillary palpi absent. Forewing cell with stem
166 Marc E. Epstein, Henk Geertsema, Clas M. Naumann & Gerhard M. Tarmann
of medial vein from near base, branched into V
near distal end; veins Rs2 and Rs3 stalked from
Rs4; scales on forewing and legs often spatulate;
epiphysis absent on foretibia, tibial spurs 0-2-2;
ventral sensilla trichodea on 5th tarsomere of fe-
male without scales interspersed. Male genitalia
with uncus gradually tapering from tegumen,
apex down-curved, gnathos absent; aedeagus
slightly curved, narrowed at tip; valvae broad,
entire and somewhat triangular with a rounded
apex; juxtal process (valvella) each side of aedea-
gus with upturned, clawed end; vinculum over
half breadth of tegumen, saccus weakly devel-
oped. Female genitalia with papillae anales not
appressed, lobes flattened, somewhat divided
into dorsal and ventral regions, with dorsal por-
tion fused medially; both anterior and posterior
apophyses longer than 8th abdominal segment;
corpus bursae oval and membraneous; sperma-
theca connects with short ductus seminalis near
ductus bursae; two large sebaceous glands fused
at base, where connected to common oviduct.
Aidids have secondary accessory glands with two
lobes occupying a large proportion of the 8th ab-
dominal segment; similar to those found in Dal-
ceridae, except they are medially connected and
without external ducts; connection of secondary
glands to common oviduct unclear (Epstein,
1996).
Larva head with two stemmatal setae and
stemmata similar to Megalopygidae; labrum with
stout spines on anterior margin. Dorsum of
Aidos amanda unicoloured green; Verrucae with
thick, black urticating setae up to 11 per cluster
in dorsal and subdorsal rows on thorax and ab-
domen; urticating setae sunken and appressed to
a common point when in repose and splayed out
into rosettes when protruded; spiracle on Al
midway between position of other spiracles and
subdorsal row of spines; abdomen with two tac-
tile lateral setae each on a pinaculum in a hori-
zontal row below each spiracle; an additional
row of ventral pinacula found between proleg
bases. Thoracic legs moderate in length; prolegs
on abdominal segments 27 and the claspers as
in megalopygids, although without membra-
neous pads or plumose setae; crochets in V-
shaped row on prolegs 36, linear on the clasp-
ers, and absent on segments 2 and 7; anal fork
reduced or absent (Epstein, 1996). Pupa similar
to that of dalcerids, except mandibles vestigial
and fractures along eye suture occur during eclo-
sion; frons often with a projecting hump; spirac-
ular sensilla absent.
Larvae reported on Annona and quinine yield-
ing Cinchona ledgeriana; reports on guyaba ap-
pear to be based on misidentification (Epstein,
1995). Cocoon spun between leaves, with pre-
formed escape hatch, and one or two pairs of
small, round holes in outer silk mesh, giving ap-
pearance of emergence holes of parasitic Hyme-
noptera (Epstein, 1995).
Neotropical. Two genera, Aidos, Brachycod-
illa, with a total of 6 species.
References: Epstein (1995), Epstein (1996),
Hopp (1934).
Limacodidae (Figs. 10.1 J, K, 10.3 C, H). Imag-
ines small to medium-sized, often brightly col-
oured, with stout body and triangular forewings.
Limacodid moths have a characteristic resting
position with the axis of the body held at an
acute angle to the substrate, the body being sup-
ported by the extended legs and the wings
pressed closely against the flanks. The Afrotropi-
cal Chrysopoloma group, usually treated as a
separate family, is now placed in Limacodidae
based on information from the immature and
adult stages (Epstein, 1996). This placement
should be viewed as preliminary, since definitive
status must await a higher classification for the
Limacodidae.
Limacodinae (fig. 10.1 J). Comprising the non-
chrysopolominae limacodids this assemblage
may prove to be paraphyletic. Head with thick,
dense scales; antenna about half as long as fore-
wing, in males often bipectinate only in the basal
half or third of its length or in a variety of con-
figurations including filiform, unipectinate or bi-
pectinate to apex; usually dentate or filiform in
the females, although sometimes bipectinate to
the apex; maxillary palpi and proboscis usually
reduced, sometimes vestigial, but also well devel-
oped in some primitive species; labial palpi short
and porrect, usually with three segments; ocelli
and chaetosemata absent. Thorax densely scaled
with slender hair-like scales; foretibia without
epiphysis, tibial spurs 0-2-4, 0-2-2 or rarely ab-
sent; fifth tarsomere of female usually with a
dense mat of ventral sensilla trichodea on a re-
cessed pad without scales interspersed. Forewing
with retinaculum beneath Sc in male and at base
of CuA in female; frenulum consisting of a single
large bristle in male and three or more smaller
bristles in female, an additional area of wing-
locking microtrichia present at proximal part of
the dorsum of forewing underside; venation only
slightly reduced; forewing without chorda, MS
present and sometimes branched distally, Sc + R
often anastomosed in distal half of wing, Rs2 +
Rs3 usually stalked from Rs4 or Rsl, sometimes
in between Rsl and Rs4 or Rs4 is "split back"
to connect with discal cell; CuP present and 1 A
+ 2 A with basal fork. Hindwing with MS pre-
sent in discal cell but rarely branched distally;
Se + R and Rs anastomosed basally for a short
distance or, less frequently, R forming a short
oblique cross vein between Sc and Rs, sometimes
with a few vestigial veins or pseudoneuria from
Sc + R to costa; Rs and Ml usually stalked be-
yond discal cell, while occasionally there is a
cross vein formed between the two veins at the
end of the discal cell; CuP and 1A + 2 A and
also 3 A present, 1 A + 2 A not forked basally.
The Zygaenoidea
Abdomen covered with long, woolly, piliform or
slender lamellar scales; with tortricoid-like apo-
demes on sternum 2; segments 8 - 1 0 not exten-
sile in female. Male genitalia with stout uncus
opposed by a stout gnathos, without socii; arms
of gnathos fused medially to form a strongly
sclerotized and broadly curved hook, valva often
simple, without sclerotized projections, but may
be deeply divided or have processes or lobes; ae-
deagus usually simple, tube-like, often flexed
along its proximal half, cornuti present or ab-
sent. Female genitalia with two setose ovipositor
lobes forming a disc-like structure, probably an
adaptation related to oviposition; antrum sim-
ple, but ductus bursae often very long and spi-
ralled, ductus seminalis branching off subbasally;
corpus bursae membraneous, with or without
signa; 8th abdominal segment in female with a
pair of rather unusual lateral lugs (Common,
1990; Holloway, 1990).
Egg flat, discoid or ovoid, laid singly or in
overlapping groups or rows; a few species are
known to cover the eggs with specialized pili-
form scales from the posterior end of the female
abdomen. Larva in a variety of forms and colour
patterns, with urticating species having spines on
scoli or on sunken Verrucae, species with elon-
gate and sometimes detachable tubercles, and
smooth ('gelatine') species. Head with stemma 6
most isolated, equidistant to 1 or 5; only one
stemmatal seta present. Labrum often with a
large number of bristles covering anterior mar-
gin, more extensive than in related families; spin-
neret shape variable, usually widest at apex, al-
though sometimes changing ontogenetically, ta-
pering distally (post-first instar) or narrow and
more tubular (first instar). TI with primary setae
only (as in dalcerids); T2, T3, and abdominal
segments 19 with several dorsal and subdorsal
rows of scoli or elongate tubercles, smooth with
tactile setae, or rarely with gelatinous warts; lat-
eral setae positioned below the spiracles and tac-
tile; thoracic legs small to minute; suckers on ab-
dominal segments 18 or 1 - 7 , with fungiform
setae between the lateral margin and the median
(as in dalcerids); crochets normally absent but
are known to occur in the African genus Pantoc-
taenia on segments 2 - 7 and on the anal claspers
in late instars (added after the first instar, pre-
sumably as in dalcerid larvae); frass-flipping se-
tae occurring singly or in small groups, or absent
(Epstein, 1996).
Larvae are known to undergo significant on-
togenetic change between first and later instars.
Lineages with smooth larvae in late instars ('gel-
atine caterpillars') have very different first in-
stars, with setae including dilated or branched,
warts, or spines found in many forms. Lineages
with spiny larvae ('nettle caterpillars') or hairy
larvae ('monkey slugs') in late instars undergo
less ontogenetic change, retaining setose tuber-
cles in first instars in the form of scoli or hairy
167
tubercles (Dyar, 1899; Epstein, 1996). The larvae
appear to lack abdominal prolegs, however they
possess a highly reduced form of prolegs modi-
fied with elastic cuticle and suckers. This is based
on the presence of crochets on abdominal seg-
ments 27 in Pantoctaenia (and in the Dalceri-
dae). Ventral suckers found on abdominal seg-
ments 1 and 8 are not considered to be derived
from prolegs, but rather from warts as found in
megalopygid and aidid larvae. The specialized
ventrum enables the larva to adhere to the fo-
liage, often aided by semifluid silk or fluid
secreted with the silk from the spinneret, which
enhances surface tension. The spinneret also
serves a cleaning function by removing debris
from the ventrum (Epstein, 1996). Pupa short
and very stout, with a delicate, only slightly scle-
rotized cuticle and with the appendages not or
only very slightly fused to one another and to
the body; pupal maxilla often with elaborate
lobes that hold the legs in place below the eyes.
The adults of most species are nocturnal and
have an erratic, rapid flight. They are frequently
attracted to artificial light, only a few species be-
ing diurnal. Males exhibit several unusual pos-
tures, including raising forelegs or resting fore-
legs on midlegs, or curling the abdomen above
the body. Females have been observed to feed on
sugar water (Epstein, 1996). The first instar lar-
vae of several genera (e. g., Parasa) do not feed
at all, but after a period of about two days un-
dergo their first moult before commencing to
feed on a wide range of trees and shrubs; a few
also on Poaceae. The larvae of some limacodid
species are pests of coconut and oil palms, ba-
nana, tea, coffee, cacao, rice, sugar-cane and
other economically important plants. Cocoons
usually hard and often with calcium oxalate that
is secreted through the anus by the prepupa dur-
ing the final stages of construction (Ishii et al.,
1984); lid on cocoon, formed from a line of
weakness rather than a preformed hatch, is only
evident after eclosion (Schremmer, 1990; Ep-
stein, 1996).
Limacodinae are found in all zoogeographical
regions, but reach their highest diversity in the
tropics. There are around 1000 species described.
Principal genera include Susica, Thosea, Narosa,
Scopelodes, Parasa, Eue lea, Acharia (= Sib ine),
Miresa, Perola and Apoda (= Limacodes, Coch-
lidion).
Chrysopolominae (Fig. 10.1 K). Imagines sim-
ilar to those of other Limacodidae, being round-
winged, coarsely scaled, hairy and of moderate
size. Head smoothly scaled, maxillary palpi and
proboscis absent, labial palpi with three seg-
ments, slightly curved upwards as in the Lima-
codinae; antenna bipectinate in male, serrate or
filiform in female; ocelli and chaetosemata ab-
sent. Thorax hairy, foretibia without epiphysis,
tibial spurs 0-2-4 or 0-2-2; female fifth tarsomere
with recessed pad similar to, though more ovoid
168 Marc E. Epstein, Henk Geertsema, Clas M. Naumann & Gerhard M. Tarmann
than, in other Limacodidae (Epstein, 1996); wing
venation with MS present in both wings, forked
distally in forewing; radial stem present in fore-
wing and, together with the medial stem, divid-
ing the discal cell into three parts; forewing Rs2
and Rs3 stalked or independent, and branching
from Rsl; M2 and M3 free from discal cell or
stalked; CuP present, 1 A + 2 A with basal fork;
hindwing with Se + R partly anastomosed with
discal cell, at least for a short distance, or with a
short cross vein; Rs and Ml free from cell or
stalked; discal cell longer distally, M2 and M 3
free from cell; 1 A, 2 A and 3 A separate; frenu-
lum and retinaculum absent. Abdomen rough
and hairy. Male genitalia with a single stout un-
cus with fine setae, tegumen broad, valva simple,
not very strongly sclerotized, or stronger sclero-
tized and stout proximally, but very slender and
finger-like distally; vinculum without saccus;
transtilla well developed, the proximal part
doubled in Ectropa but fused to form a single
plate in Chrysopoloma; anellus prominent, some-
times with a double dorsal and a single ventral
process, but also without such processes; aedea-
gus simple, forming a slender, often downward-
curved tube with stronger sclerotizations
ventrally, sometimes with tooth-like crests and a
group of fine cornuti (Hering, 1937 a); gnathos
absent (autapomorphy). Female genitalia with
only slightly sclerotized papillae anales, a
strongly sclerotized sterigma and a well-devel-
oped bursa copulatrix that may have a distal ap-
pendix (Hering, 1937 a); with lateral lugs on 8th
abdominal segment, as in Limacodinae (Ep-
stein, 1996).
Larva green with long, black hairlike setae
having white tips, one seta in dorsal row per seg-
ment and two setae below spiracles, length of
each seta over half body width. Cuticle with spi-
nulose texture, evenly distributed over dorsum.
Labrum without multiple spines on anterior
margin that are commonly found in Limacodi-
nae. One stemmatal seta on the head, as in other
limacodids and dalcerids; stemmatal arrange-
ment more similar to other limacodids. Crochets
absent; thoracic legs as in Limacodinae (Epstein,
1996). Pinhey (1975) describes a pink smooth,
slug-like larvae as very similar to those of the
Limacodinae. Pupa with maxillary lobe parallel
to and adjoining labial palpus, and lateral exten-
sion of maxilla adjacent to eye (similar to many
in Limacodinae); mandibles very small. Sculp-
tured eye piece intact on exuvia after eclosion, as
in Dalceridae. Cocoon smooth, oval and hard,
with circular lid, as in Limacodinae; constructed
just below ground surface (Epstein, 1996).
The subfamily is restricted to the Afrotropical
region and includes about 30 species. Principal
genera include Chrysopoloma, Ectropa and Scoti-
nocerides.
References: Cock, Godfray & Holloway
(1987), Common (1970, 1990), Epstein (1996),
Hering (1937 a, 1955), Holloway (1990), Hollo-
way, Bradley & Carter (1987), Janse (1964), Pin-
hey (1975), Viette (1980).
Dalceridae (Figs. 10.1 L, 10.3 F - G ) . The Dalcer-
idae were first considered to form a separate
family by Dyar (1898), and subsequent authors
have usually placed them near to Limacodidae
and/or Megalopygidae within the Zygaenoidea.
Miller (1994) regards them to be closer related to
the other zygaenoid families and suggests that
Dalceridae, Limacodidae, Megalopygidae and
possibly Epipyropidae might form a monophy-
letic group because of the highly specialized cov-
ering of parts of the pupal eye. Epstein (1996)
regards them as the sister group to the Limacodi-
dae based on a number of synapomorphies in-
cluding fungiform ventral setae, absence of
crochets in early instars, highly reduced prolegs
and thoracic legs, a brushlike spinneret, and the
presence of a single stemmatal seta in the larva;
and absence of chaetosemata, reduced sclerite on
mesepimeron, mesonotum without a medial sul-
cus, and presence of a gnathos in the adult.
Imagines small to medium-sized, body hairy,
wings broadly rounded, with bright white, yel-
low or orange ground colour. Head small and
densely scaled, antenna short, broadly bipecti-
nate in males, narrowly bipectinate in females;
labial palpi short, porrect, with two segments,
first segment small and short, second much
larger; ocelli and chaetosemata absent, proboscis
vestigial, densely covered with small spines, eyes
large. Thorax large, strongly scaled, legs densely
scaled, epiphysis of foretibia absent, tibial spurs
absent (Miller, 1994); tarsomeres with large spurs
on ventral surface; fifth tarsomere in female cov-
ered with scales beneath, with a narrow distal
band of sensilla trichodea (Epstein, 1996). Fore-
wing ovoid to rounded, more so in females, stem
of media present in discal cell, R either free or
stalked with Rsl and Rs2, Rs3 and Rs4 often
stalked; the pairing of Rsl + 2 and Rs3 + 4
is considered to form an autapomorphy of the
Dalceridae (Miller, 1994). Subcostal retinaculum
present in males which have a frenulum, forming
a narrow, elongate rolled flap arising from Sc
and forming a groove through which the frenu-
lum fits. Hindwing colouration generally similar
to that of forewing; 3 A present, Sc + R and Rs
free, anastomosed or connected by a small
bridge in discal cell. Male frenulum consisting of
a long bristle formed by the fusion of 10-30
acanthae. In some genera the frenulum is absent.
Frenulum in females formed by 15 or more long,
narrow, unfused bristles. Abdomen densely
scaled, colouration similar to that of hindwings.
Male genitalia characterized by reductions and
fusions, mainly of the valvae; tegumen and soci-
uncus generally well developed, frequently fused,
uncus bilobed, sometimes with medial or lateral
points or hooks, socii sometimes hairy pads or
The Zygaenoidea
only small lateral hairy areas on sociuncus,
gnathos generally well developed, variously
shaped; vinculum usually elongate and slender,
extended anteriorly to form a long saccus, fre-
quently bearing processes of various forms; val-
vae extremely reduced, immobile and frequently
fused with juxta, anellus and/or vinculum; juxta
usually present, rod-like, forming an external
support structure of the aedeagus, anellus usu-
ally fused with valvae, juxta and/or vinculum,
saccus generally elongate; aedeagus usually long
and slender, vesica in some Acraga spp. with
small patches of cornuti. Female genitalia with a
weakly sclerotized sterigma, ostium bursae nar-
row to broad, ductus bursae relatively long, with
one or two strong bends, corpus bursae medium
to small, ovoid to spheroid, without signa; duc-
tus seminalis arising from first half of ductus
bursae; receptaculum seminis (spermatheca) with
a long duct; apophyses anteriores absent, apo-
physes posteriores well developed, papillae an-
ales large and setose, divided into upper and
lower lobes. As in some other Zygaenoidea, the
Dalceridae have a second pair of accessory
glands, but they have been independently
evolved, and open on the outside of the papillae
anales (Miller, 1993). These unique characteris-
tics appear to be autapomorphic for the family.
The eggs are up to 1 mm in size, oblong,
smooth and often yellow. They are laid singly
and dispersed amongst the vegetation. One fe-
male may produce several hundred eggs which
are coated with a fast-drying fluid which appar-
ently forms a protective layer. Larva with dor-
sum covered with gelatinous warts secreted by
setae beneath. Head with one stemmatal seta
present; labrum with dense, anterior bristles on
mesal portion only; spinneret wider at apex than
at base; in post-first instars a brushlike comb
along apical margin, with deeply divided bristles
and dorsal surface reduced medially, revealing a
narrow longitudinal silk pore. Prothoracic shield
translucent with only uncoated hair-like setae
present; two (first instar) or three (later instars)
lateral setae below spiracle. Dorsal and subdor-
sal setal rows covered with gelatinous warts on
T2 and T3 and abdominal segments 1 - 9 ; often
brightly coloured pigmented patterns on cuticle
beneath gelatinous warts. Three tactile setae pre-
sent below gelatinous warts on T2 and T3, lat-
eral abdominal region, between gelatinous warts
and ventrum, spinulate; spiracles closer to
ventrum than in Limacodidae; three tactile lat-
eral setae posterior to each spiracle. Thoracic
legs small; ventral surface of thorax and abdomi-
nal segments 1 - 9 with cuticle translucent and
flexible throughout, except for a narrow band of
spinulose cuticle on anterior portion of Tl; ab-
dominal segments 19 with ventral fungiform
setae in a horizontal row of three, from lateral
margin to meson; ventral crochets absent in early
instars and appear during the antepenultimate
169
instars on abdominal segments 3 - 6 , with
crochets added to segments 2 and 7 in the final
two instars in some species (Stehr & McFarland,
1987); shallow suckers on segments 18, al-
though only segments 2 - 7 are modified from
prolegs; crochets absent on anal segment, which
has hair-like setae, shagreened texture, and me-
dial frass flipping setae near the anus present or
absent (Stehr & McFarland, 1987; Epstein,
1996). Pupa with a sculptured flange lateral to
the eye (similar to Limacodidae and Megalopygi-
dae) that remains connected to eye piece at eclo-
sion; two or three setae present on frons; cremas-
ter hooks present on end of abdomen. Cocoons
are spun between leaves with an outer portion of
diffuse silk and an oval inner chamber. Gelati-
nous warts of the larva are sloughed off in the
inner chamber during construction (e. g., Dalcer-
ides ingenita, Epstein pers. observ.). The pupa ex-
tends out of one side of the chamber, just prior
to eclosion (Dyar, 1925 and pers. observ.). The
flight of the moths is described as weak, flutter-
ing, erratic and undulating; they are mainly noc-
turnal and are occasionally taken at light, but
usually only in small numbers.
The translucent gelatinous warts, which char-
acterize the larvae, are discharged from setae on
segments T 2 - A 9 . This secretion has defensive
properties against ants (Epstein et al., 1994).
Such tubercles have also been reported from one
limacodid genus (Holloway et al., 1987).
Dalceridae are almost endemic to the Neo-
tropics (about 50 species), but the range of one
species (Dalcerides ingenita) extends northwards
to Arizona. Principal genera include Dalcera,
Acraga and Dalcerides.
References: Epstein (1996), Epstein et al.
(1994), Miller (1993, 1994).
Lacturidae (Figs. 10.1 H, 10.3 E). According to
Common (1990) the distribution of this group
of genera is primarily tropical or subtropical. It
includes over 100 species occurring in the Aus-
tralian, Oriental and Afrotropical regions and in
Central and South America (Heppner, 1995).
The adults are brightly coloured and appear to
be mainly nocturnal. Like some Zygaenidae the
adults are extremely resistant to cyanide. Al-
though previously included within the Ypono-
meutoidea, Kyrki (pers. comm.) discovered a
number of characters which indicate a close rela-
tionship to Zygaenidae and he and Common
(1990) and also Nielsen (pers. comm.) include
this group within the Zygaenidae. Athough these
insects share many characters (especially of the
larval morphology) with the Zygaenidae, it ap-
pears to be doubtful that they are closely related
to this group and that they should be included
in this family. The general appearance, filiform
antenna, the presence of a forewing 'chorda', the
ultrastructure of the wing scales, the slug-like
larva and a 'frog-mouth' opening of the cocoon
170 Marc E. Epstein, Henk Geertsema, Clas M. Naumann & Gerhard M. Tarmann
clearly separate them from the Zygaenidae.
There are similarities in the female genitalia with
those in the subfamily Phaudinae of the Zygaeni-
dae (spirally arranged ductus bursae), but this
character is also shared with the Limacodidae
and some other groups. The presence of prolegs
on abdominal segments 3 6 and 10 is similar
to Zygaenidae and other non-limacodid group-
families in the Zygaenoidea. Having compared
them with the other groups dealt with here, we
accept Heppner & Inoue's (1992) treatment of
the Lactura-group as a distinct family Lacturi-
dae; see also Heppner (1995).
The larvae feed fully exposed during the day
and might be distasteful or poisonous. They are
brightly coloured and slug-like and produce a
defensive secretion. They pupate in a flimsy,
silken cocoon, which has a broad 'frog-mouth'
opening, similar to those in the Limacodidae and
related groups (Common, 1990).
Principal genera are Lactura (East Asia, New
World), Gymnogramma (southern Africa) and
Anticrates (Old World).
References: Common (1990), Heppner (1995).
Heterogynidae (Fig. 10.1M). A small family con-
sisting of only one genus. The imagines have
some phenotypic similarities with the subfamily
Procridinae (Zygaenidae), and with some Psychi-
dae and Lymantriidae. Male imagines have api-
cally rounded, dark, though translucent wings
covered with hair-like scales. The females are
wingless and resemble the larvae in habitus and
colouration. The systematic status of the group
is not completely clear at present. Minet (1986)
considers the Heterogynidae to form the adel-
photaxon of the Zygaenidae, a supposition that
is corroborated by the fact that the larval cuticle
of the former contains cavities similar though
not identical to those of the two zygaenid sub-
families Chalcosiinae and Zygaeninae (Nau-
mann, unpublished).
Head of imago without proboscis and with re-
duced maxillary and labial palpi; ocelli and chae-
tosemata absent; male antennae strongly quadri-
pectinate, pointed distally; female antennae re-
duced (Zilli, 1988). Male thorax hairy, legs with
small epiphysis on foretibia and short tibial
spurs on mid- and hindtibia; forewing with fully
developed medial stem, all veins arising freely
from the discal cell; radial veins sometimes partly
reduced; CuP well developed distally but some-
times reduced proximally; hindwing with Se + R
free and not anastomosed with discal cell,
though sometimes connected with the cell by a
short cross vein, discal cell extended distally,
with complete medial stem; frenulum and reti-
naculum well developed; with an additional area
of small wing-locking microtrichia on the under-
side of proximal part of forewing dorsum and on
metathorax, similar to that in most Zygaenidae;
both wings covered with hair-like scales. Male
abdomen very hairy. Male genitalia with strongly
sclerotized, short-hooked uncus and subuncus,
tegumen broad, strongly sclerotized, forming a
ring with the equally broad and strongly sclero-
tized vinculum, saccus absent; valvae rounded, of
medium size, without processes; aedeagus tube-
like, stronger sclerotized dorsally, very translu-
cent ventrally, vesica without cornuti but with a
large number of minute spines. Female genitalia
very simple, bursa copulatrix hardly defined,
merging with the 'vestibulum' into which the re-
ceptaculum seminis, the glandulae sebaceae and
the laterally attached paired oviducti open, with
a characteristic appendix ('papilla') which ap-
pears to take over the function of the corpus bur-
sae in other taxa (Daniel & Dierl, 1966; Nau-
mann, 1988 a).
Egg yellow to whitish green, ovoid, surface
slightly ribbed, laid in rows inside or on top of
the cocoon where the wingless female remains
after copulation. The brightly coloured larva,
with retractile head and prolegs on abdominal
segments 3 - 6 and 10, feeds on different genera
of Fabaceae, including Sarothamnus, Onobrychis
and Cytisus, and spins a loose cocoon on twigs
and stems of grasses. Imagines emerge during the
early morning when the males begin immediately
to search for females. The females remain inside
the cocoon and are hardly distinguishable from
the similarly coloured pupa. Copulation takes
place on top of or inside the cocoon, the male
then inserting its abdomen into the cocoon.
In North African populations oviviparous
hatching of first instar larvae has been observed.
Some populations also seem to be at least partly
parthenogenetic (Naumann, 1988 a; Tarmann,
pers. obs.).
The family is endemic to the western part of
the Mediterranean, where it is represented by
one genus (Heterogynis) comprising about 10
species.
References: Daniel & Dierl (1966), de
Freina & Witt (1990), Naumann (1988 a), Zilli
(1988), Zilli & Racheli (1989, 1992).
Zygaenidae (Figs. 10.2 A - J , 10.3 D, L - P ) . The
Zygaenidae form a group of subfamilies with
well-defined autapomorphies but with rather
large differences between them. Three subfamil-
ies, i. e. the Procridinae, the Chalcosiinae and the
Zygaeninae appear to form a monophylum,
while the relationships of the remaining Phaudi-
nae and Callizygaeninae still remain doubtful.
Zygaenidae are distributed world-wide, being
most diverse in subtropical and tropical Asia and
in the Palaearctic regions. There are more than
1000 species known, and many more still to be
described. The Zygaeninae are restricted to the
Palaearctic, Himalayan and Afrotropical re-
gions. The Chalcosiinae (with the exception of
the western Palaearctic genus Aglaope) are re-
stricted to the eastern Palaearctic and Indo-Aus-
The Zygaenoidea 171

Fig. 10.2: Zygaenidae subfamilies (length of bar: 10 mm). A. Zygaenidae, Phaudinae: Phauda cantonensis. B.
Zygaenidae, Callizygaeninae: Callizygaena durata. C. Zygaenidae, Procridinae: Balataea octomaculata. D. Zygae-
nidae, Procridinae: Adscita subtristis. E. Zygaenidae, Chalcosiinae: Eterusia aedea. F. Zygaenidae, Chalcosiinae:
Aglaope infausta. G. Zygaenidae, Zygaeninae: Pryeria sinica. H. Zygaenidae, Zygaeninae: Orna pectinicornis. J.
Zygaenidae, Zygaeninae: Zygaena trifolii.

tralian regions. The Procridinae have a world-
wide distribution including Australia, Tasmania
and the temperate regions of southern America
and southern Africa. The status of the two South
East Asian subfamilies Phaudinae (approxi-
mately 50 species) and Callizygaeninae (approxi-
mately 12 species) is uncertain.
Head smooth-scaled or hairy, sometimes pro-
jected dorsally (most Chalcosiinae, some Pro-
cridinae), with flat vertex or with a prominent
dorsal crest (most Chalcosiinae); antenna with-
out pecten, maxillary palpi short, with one or
two segments; proboscis usually well developed
and unsealed, sometimes reduced, labial palpi
short, smooth-scaled and mainly porrect or
slightly ascending; ocelli present in Zygaeninae,
Chalcosiinae and Procridinae, absent in Phaudi-
nae; chaetosemata present, of group-specific size
and shape. Imagines of the Procridinae, Chalcos-
iinae and Zygaeninae produce a vitreous, whit-
ish, yellowish white or yellow liquid or foam be-
tween the inner margin of the eye and the base
of the proboscis when disturbed. Epiphysis of
foretibia present or absent, tibial spurs 0-2-4, 0-
2-2, or absent; forewing with a well-developed
retinaculum on basal part of Sc forming a strong
hook in male and with a group of raised scales
at base of CuP in female; a small, elongated area
of wing-locking microtrichia present on metascu-
tum and on forewing underside (Tarmann, un-
published). MS in forewing present or absent,
the radial and medial veins often free from the
discal cell, sometimes stalked, partly reduced or
anastomosed; CuP completely present in Zygae-
172 Marc E. Epstein, Henk Geertsema, Clas M. Naumann & Gerhard M. Tarmann
ninae, Chalcosiinae, Procridinae, only present
distally in Phaudinae; 1 A + 2 A normally with
basal fork. Hindwing with Sc + R free, anasto-
mosed with anterior margin of discal cell, or with
a short cross vein from cell to Se + R at approxi-
mately one-third of wing length; with three or
less medial veins which may either be free from
the cell or stalked; 1 A, 2 A and 3 A usually pre-
sent, forming three well-developed veins, 2 A
sometimes, rarely also 3 A, reduced. Hindwing of
male (and sometimes also female) in Chalcosii-
nae with an S-shaped tuft of scales arising from
inner angle of wing, pointing backwards toward
the flanks of the abdomen (Haase, 1887; Tar-
mann, 1992). Abdomen with tortricoid-like apo-
demes on sternite 2. In male (and sometimes fe-
male) Chalcosiinae the pleura of abdominal seg-
ment 2 (sometimes 1 and 2) have a characteristic
fold where the scale tuft arising from the hind-
wing is stored; this fold bears a presumably
scent-producing gland and a field of very regu-
larly arranged short setae. In the Artona-group
of the Procridinae (most of the South East Asian
and nearly all Australian and Papuan procridine
species) abdominal segments 2 and 7 have addi-
tional lateral protuberances which appear to be
involved in predator deterrence and which pro-
duce a strong smell repellent to the human nose
(Tarmann, 1994 and unpublished). In some
Chalcosiinae the 8th abdominal sternite of the
male is strongly sclerotized and bears hooks
and crests.
Male genitalia with a large number of special-
izations, i. e. symmetrical or asymmetrical (e. g.,
in the American Procridinae genus Harrisind),
uncus bilobed with hairs and setae in Zygaeni-
nae, double or single in Chalcosiinae, single and
slender, without hairs and setae in Procridinae,
and single and hairy in Phaudinae; gnathos usu-
ally absent, but present in some Chalcosiinae
(e. g., Agalope); valva small to large, with or
without strongly sclerotized structures, with or
without pulvinus, with or without a defined sac-
culus, tegumen broad or narrow, sometimes
nearly divided into two parts (some Procridinae);
vinculum narrow or broad, sometimes with sac-
cus; juxta broad and triangular or rectangular,
aedeagus strongly sclerotized with a well-defined
and very spiny lamina dorsalis and lamina
ventralis in Zygaeninae, tube- or hook-shaped in
Chalcosiinae, usually forming a simple tube in
Procridinae and Callizygaeninae, and an S-
shaped tube (similar to some Limacodidae) in
Phaudinae; vesica with or without cornuti. Fe-
male genitalia with abdominal segments 8 - 1 0
extensile in Chalcosiinae (thus forming a func-
tional ovipositor), not so in Zygaeninae, most
Procridinae and Phaudinae. Papillae anales
small to large, covered with long or small setae,
apophyses posteriores short or long, apophyses
anteriores prominent to vestigial; the sclerotiza-
tion of abdominal segment 8 weak or strong to
very strong with a weakly or strongly sclerotized
antrum which is usually situated medially (sym-
metrical), but rarely asymmetrical laterally
(American genus Harrisina of Procridinae); an-
trum with or without sclerotizations, ductus bur-
sae translucent or strongly sclerotized, inflated in
many Procridinae to form a huge praebursa
which may contain spines, crests, ribs and ap-
pendices; in the Phaudinae the ductus bursae is
twisted, sometimes spiral (similar to Limacodi-
dae); deviation of ductus seminalis proximal (Zy-
gaeninae, Phaudinae, Callizygaeninae, some
Chalcosiinae) or distal near the opening of the
corpus bursae (most Chalcosiinae, all Procridi-
nae); corpus bursae usually weakly sclerotized,
sometimes with an appendix, signa present or ab-
sent, a pseudobursa is developed in Zygaeninae,
some Chalcosiinae and in the Callizygaeninae; in
the Procridinae the utriculus and lagena of the
spermatheca (receptaculum seminis) are (at least
externally) fused (Naumann, 1988 a; Tarmann,
1994), again a character state not found in the
Callizygaeninae. In the Zygaeninae and Procridi-
nae an additional gland opening into the oviduc-
tus communis near the ooporus is found, termed
Petersen's glands in the Zygaeninae (fig. 10.3 P;
Bode & Naumann, 1987). In the Zygaeninae it
produces a sticky secretion which provides a thin
coating to the eggs during oviposition; this gland
is reduced in some Australian Procridinae (e. g.,
Pollanisus and Hestiochora).
The eggs of the Zygaenidae are ovoid, slightly
flattened, with a transparent, finely reticulate or
dentate chorion, which may bear fine longitudi-
nal ribs; the colour is white, yellow, greenish or
yellowish white.
Larvae of the Zygaeninae, Chalcosiinae and
Procridinae are stout and broad, with the head
capsule usually retractile under the slightly ex-
tended prothoracic segment; antenna long, only
primary setae present on head capsule, six stem-
mata on each side; body cuticle roughened or
spinulose and covered with dense secondary se-
tae arranged into dorsal, subdorsal, lateral and
subventral Verrucae. The prothoracic legs bear
an apical pair of spatulate setae, equivalent in
size to the claw; the ventral prolegs are short,
with crochets arranged in a uniordinal mesose-
ries on abdominal segments 3 - 6 and 10. The lar-
vae of the Phaudinae and Callizygaeninae are
similar in shape to those of the Limacodidae,
brightly coloured and sluggish (Malicky, 1995,
Yen unpublished); a proper anatomical and mor-
phological analysis has yet to be done.
The pupa is stout and sometimes dorso-
ventrally flattened, with moderately sclerotized
cuticle and with appendages only weakly fused
to one another and to the body; abdominal seg-
ments 28 or 3 8 have a single transverse row
of dorsal spines, cremaster vestigial; abdominal
segments 1 2 are fused in both sexes, also 810
in male and 710 in female (Tremewan, 1985).
The Zygaenoidea
Representatives of the Procridinae, Chalcosii-
nae and Zygaeninae are cyanogenic, biosynthe-
sizing the cyanoglucosides linamarin and lot-
australin in their body tissues (Fig. 10.30). Both
secondary compounds are well known from a
number of plant families, but are not plant-de-
rived in the insects. Davis & Nahrstedt (1979,
1982, 1987) have demonstrated that the moths
biosynthesize the two cyanoglucosides them-
selves starting from the amino acids valine and
isoleucine (Fig. 10.30). Members of the three
subfamilies are also able to release hydrocyanic
acid (HCN) by enzymatic break-down of these
two cyanoglucosides. They can also detoxify hy-
drocyanide by using the enzyme -cyanoalanine-
synthetase (Witthohn & Naumann, 1984).
In the Chalcosiinae and Zygaeninae most seg-
ments of the larval cuticle bear two types of cu-
ticular cavities (Fig. 10.3 N; Franzi, 1980; Po-
volny & Weyda, 1981; Franzi & Naumann, 1984,
1985). One is regularly arranged to form antero-
dorsal, postero-dorsal, antero-subdorsal and
postero-subdorsal pouches (Fig. 10.3 N), which
are connected to the exterior through morpho-
logically highly specialized cuticular opening
structures (Franzi & Naumann, 1984, 1985). The
second type of cavity is much smaller and also
connected to the exterior through a structurally
different type of opening mechanism. In the cu-
ticular cavities a highly viscous defensive secre-
tion is stored, consisting mainly of linamarin and
lotaustralin and a large proteinaceous fraction
(Witthohn & Naumann, 1984).
The adults are mainly diurnal; only a few spe-
cies are known to be attracted to light. The flight
habits vary considerably between the groups;
whereas most Palaearctic Zygaeninae and
Procridinae are sluggish and fly slowly, most spe-
cies of the tropical regions and especially nearly
all Chalcosiinae have a rapid and restless flight
in full sunshine. The high degree of chemical pro-
tection has enabled some zygaenid moths to
form part of complex and mainly unstudied
mimicry relationships (Chalcosiinae, Zygaeni-
nae). Furthermore, in at least some species of
Zygaena a complex dual mate-locating strategy
involving both chemical and optical cues has
been developed (Naumann, 1988 b, 1990;
Prinz & Naumann, 1988; see below).
Zygaenid moths and larvae are usually highly
resistant to hydrocyanide. In some species this
resistance is strong, the central American
Procridinae Pyromorpha cuchumatana survives
up to 30 minutes in a fresh cyanide jar without
any reaction (Tarmann, 1984). The eggs are de-
posited singly, in rows, clusters, or in overlap-
ping patches, and are sometimes covered with
scales. In the Australian Procridinae genera Pol-
lanisus, Onceropyga and Hestiochora the eggs are
covered with long urticating scales originating
from a special abdominal hair-tuft of the female
(Tarmann, 1994). The larvae feed exposed on
173
leaves of numerous herbs, shrubs and trees,
sometimes skeletonizing the leaves in the early
instars, but in many Procridinae of the genera
Adscita and Jordanita, leaf-mining and boring
habits of the larva are also known. The same ap-
plies as a rare exception to the early instars of the
larva of Zygaena carniolica from north-eastern
Turkey (prov. Kars) where strong winds and
rains prevail during much of the summer season.
This habit has not been observed in other pop-
ulations of this species and appears to be a local
adaptation to severe climatic conditions (Nau-
mann, unpublished). Similar behaviour has been
observed in Iranian and Central Asiatic species
of the subgenus Mesembrynus (Tremewan and
Naumann, unpublished).
Pupation takes place either on the ground in
flimsy, silken cocoons or in parchment-like firm
cocoons, some exposed on leaves or on stems,
sometimes on or under rocks, or in stems and
roots of the host plant. The wall of the cocoon is
impregnated with whewellite crystallites (calcium
oxalate monohydrate) which are formed in the
Malpighian tubules of the larva (Hepp, 1920,
1922; Hofmann, 1994; Naumann, 1977 b and un-
published data).
Key to the subfamilies of the Zygaenidae
(not considered: Callizygaeninae, which have
recently been excluded from the Procridinae and
treated as a separate subfamily (Tarmann, 1994))
1 Males (and sometimes also females) with a
characteristic scent brush arising from pos-
terior base of hindwing, resting in a partic-
ular fold of the pleura of the first and/or
second abdominal segments; chaetosemata
fused; females with functional ovipositor
formed by abdominal segments 810 . . .
Chalcosiinae
- Without such scent brushes; females with-
out evertible and movable ovipositor
formed by the abdominal segments 8 - 1 0
(except in the Central American genus
Gonioprocris of the Procridinae) 2
2 Male genitalia with a single uncus . . . . 3
- Uncus of male genitalia always bifid (not
fused); aedeagus with characteristic,
strongly spined dorsal and ventral laminae;
most genera with characteristic Zygaena-
like wing pattern; internal female genitalia
with a complex, two-chambered secondary
accessory gland (Petersen's glands)
Zygaeninae
3 Utriculus and lagena of spermatheca (re-
ceptaculum seminis) fused e x t e r n a l l y . . . .
Procridinae
- Ductus bursae usually strongly twisted,
moths reddish brown to brownish black
with characteristic lateral appendages of
the abdomen, at least in the terminal seg-
ments Phaudinae
174 Marc E. Epstein, Henk Geertsema, Clas M. Naumann & Gerhard M. Tarmann
Procridinae (Figs. 10.2 C, D). Small to me-
dium-sized (wingspan 10-30 mm), forewings
narrow and apically rounded, or triangular and
apically pointed, hindwings rounded, square or
triangular; there is a single species with apterous
females (Pollanisus calliceros from Australia).
Most are unicolorous, but brightly coloured spe-
cies are also known. A very brilliant, sometimes
metallic sheen on the forewing upperside, head,
thorax and abdomen is typical for some genera
(e. g., Adscita, Jordanita, Pollanisus); antenna
usually bipectinate in male and bipectinate, bi-
serrate or filiform in female, usually with pointed
apex, but also clubbed in some groups (e. g., sub-
genus Zygaenoprocris of Adscita). About 500
species on all continents. Principal genera are
Jordanita, Adscita, Illiberis, Artona, Harrisina
and Pyromorpha.
Callizygaeninae (Fig. 10.2 B). The Procridinae
have been divided into two tribes by Alberti
(1954), viz. the Procridini and the Callizygaenini.
More recent investigations, based on the mor-
phology of the internal female genitalia, and on
the male genitalia, have shown that the Callizy-
gaenini do not form part of the same monophy-
lum as the Procridinae and are now considered
to be a separate subfamily, the Callizygaeninae,
which are possibly closer to the Zygaeninae than
to the Procridinae (Tarmann, 1994). Only one
genus (Callizygaena) with 12 species in South-
East Asia.
Phaudinae (Fig. 10.2 A). Forewings narrow,
elongated and small, hindwings rounded. Mainly
yellowish brown or light brown species with
hairy body, sometimes with a pair of long hairs
at the caudal end of the abdomen. The antenna
is pointed distally, bipectinate in male and biser-
rate in female. The Phaudinae are restricted to
South-East Asia. Principal genera are Phauda
and Alophogaster.
Chalcosiinae (Fig. 10.2E, F). Very variable in
size, small to very large (10-120 mm wingspan),
wings usually broad and rounded, but examples
with exceptionally narrow wings are known. The
colouration is sometimes very striking, with met-
allic and brilliant parts, but also uniform in some
species; many Chalcosiinae are involved in com-
plex mimicry rings with other lepidopterous fam-
ilies such as the Arctiidae, Ctenuchidae, Agaristi-
dae, Hesperiidae, Nymphalidae, Lycaenidae, Pie-
ridae and Papilionidae. Antenna usually strongly
bipectinate in both sexes, pectination of the last
apical segments sometimes reduced, the tip of the
antenna thus resembling a clubbed antenna. The
Chalcosiinae are nearly exclusively restricted to
South-East Asia (both palaearctic and oriental)
with a single genus (Aglaope) in the western Pa-
laearctic. They comprise about 300 species. Prin-
cipal genera are Cyclosia, Chalcosia, Elcysma,
Agalope, Aglaope, Heteropan and Arbudas.
Zygaeninae (Figs. 10.2 G - J ) . Uniform in size
(2035 mm wingspan), with narrow, apically
rounded or elongate wings, forewing signifi-
cantly larger than hindwing, forewing ground
colour usually dark with striking red, yellow or
white patterns, patches or streaks; the antennae
are usually filiform and slightly clubbed distally;
only in two genera (Pryeria and Orna) the male
antenna is bipectinate and the female antenna is
biserrate or filiform. The subfamily is found in
the Palaearctic (2 genera), Oriental (1 genus) and
Afrotropical (5 genera) regions and comprises
about 130 species. Important genera are Pryeria,
Orna, Neurosymploca, Praezygaena, Reissita
and Zygaena.
Biology of the Zygaeninae. During the last 15
years the chemical ecology and the defensive and
reproductive biology of the Zygaeninae have
been studied intensively. In the following section
we provide an overview of the biology of this
group of insects which has justifiably been called
the "old-world Heliconius".
Like other zygaenid moths the Zygaeninae
store large amounts of the two cyanogenic gluco-
sides linamarin and lotaustralin (fig. 10.3 N). The
insects biosynthesize these compounds them-
selves (fig. 10.3 O), as can be concluded from the
fact that the more primitive genera of the Zygae-
ninae feed on acyanogenic foodplants, the Cel-
astraceae. In the larvae these compounds are
stored as a viscous or highly sticky proteinaceous
liquid in specialized cavities of the cuticle. The
cyanogenic compounds are transferred into the
body cavity every time the larva prepares to
moult, thus indicating that the cyanogenic com-
pounds represent a costly resource. It appears to
be more efficient for the insect to transfer them
from the cuticle to the body cavity before moult-
ing than to lose them with the exuviae.
The most derived genus of the subfamily, Zy-
gaena, consists of 95 species that are restricted to
the Palaearctic region with major concentrations
of species in the western and south-western Pa-
laearctis. Many species of this genus (subgenera
Agrumenia and Zygaena) are oligophagous feed-
ers on cyanogenic foodplants, mostly Astragalus,
Coronilla, Lathyrus, Lotus, Onobrychis and Vicia,
while the larvae of another group of species (sub-
genus Mesembrynus) feed on acyanogenic Apia-
ceae (e.g. Eryngium, Peucedanum, Pimpinella),
with a few on Asteraceae and Lamiceae. Imag-
ines of Zygaena have a very characteristic apose-
matic wing-pattern (fig. 10.2 J). Due to the com-
bination of chemical protection and aposematic
wing patterns Zygaena moths are carefully
avoided by vertebrate predators (birds, small
mammals and reptiles). In contrast to other zy-
gaenine genera, Zygaena moths (burnets) have
an extremely extended period of diurnal activity
during which much time is spent feeding at flow-
ers, preferably pinkish violet scabious (Scabiosa,
Knautia), but also on Centaurea, Cirsium and
Carduus. Both optical and chemical cues are used
to locate these flowers.
The Zygaenoidea 175

Cyanide metabolism in Zygoeno

-OH
- = N-OH
* y-^
COOH
COOH
2-methyl-propion 2 - m e t h y l - p r o p i o n i tril e
L- voline N-hydroxy-L-
aldoxime
valine
r
CN /CN

hydroxy-
0 glue
propionitrile
I inamarin

O NH, NH2

N
C^^COOH
L- asparagine - cyan-L-alanine

H S - A rC 0 0 H
V N

L-cysleine

Fig. 10.3: Epipyropidae: A, J; Anomoeotidae: B; Megalopygidae: K; Limacodidae: C, H; Dalceridae: F - G ; Lacturi-

dae: E; Zygaenidae: D, LP. A, Heteropsyche sp. venation; B, Anomoeotis felderi venation; C, Pseudanapaea trigona
venation; D, Onceropyga anelia venation; E, Lactura sp. venation; F, G, Dalcerides ingenitus larva (F: note jelly-like
tubercles, G: venter showing crochets on A 2 - A 7); H, Phobetron pithecium larva; J, Heteropsyche sp. larva; K, No-
rape ovina larva; L, Levuana iridescens (Procridinae) larva; M, Zygaena sp. (Zygaeninae); N, Zygaena trifolii larva,
section through dorsal integument showing cuticular cavities (sh: sensory hair, br: basal ring, sc: sensory cell, ep:
epidermis, cavi: cavity type I, cavil: cavity type II); O, Zygaena, cyanide metabolism pathways in Zygaenidae; P,
Zygaena trifolii female, internal genitalia in lateral view, caudal end to the right (bcop: corpus bursae (bursa copu-
latrix), db: ductus bursae, ds: ductus seminalis, glpet: Petersen's glands, glrec: gianduia receptaculi, glseb: glandulae
sebaceae, lag: lagena, ob: ostium bursae, ovcom: oviductus communis, psb: pseudobursa). A, C, D, E, J after Com-
mon (1990): , O original (C. M. Naumann del.); F, G, , (after Stehr 1987); M after Gerasimov (1952); after
Naumann & Povolny (1987); after Naumann (1988).
176 Marc E. Epstein, Henk Geertsema, Clas M. Naumann & Gerhard M. Tarmann
Two different mate-locating systems have been
observed in Zygaena trifolii, one of the better-
studied species of the genus from the behavi-
oural/ecological aspect. Firstly, freshly emerged
virgin females are found by the males during the
morning (9.3011.00 a. m.) by using exclusively
optical cues (wing pattern, spot colouration and
specimen size are evaluated in this process). Dur-
ing this time the females rest on top or above
their cocoons rather high on the stems of grasses
or other vegetation, while males fly at this height
through the habitat irrespective of the direction
of the prevailing wind currents. During this
period males do not respond to natural or artifi-
cal conspecific sex pheromones. As a result of
experiments involving coloured dummies it may
therefore be concluded that during this period
males find their mates by the exclusive use of
optical cues.
A second mate-locating period occurs during
the late afternoon (16.00-18.00 p.m.). By this
time females willing to mate rest in the low vege-
tation and start to emit sex pheromones. The lat-
ter have been chemically identified and artificial
sex pheromones have been used for experimental
studies of the reproductive biology of the moths.
During this period males approach the females
against the direction of the prevailing wind and
use the concentration gradient of the female sex
pheromone as an important long-distance cue to
locate them. At close range the males also exploit
optical cues and again evaluate the size, coloura-
tion and wing pattern of the female. The most
attractive colours have a spectral reflective
pattern extremely similar to that of the flowers
of the preferred imaginai food plants listed
above, which again have identical reflection pat-
terns to those of the female forewing spots.
Irrespective of the mode by which mating has
been induced, copulation continues until the
following morning when the sexes will separate.
The complicated laminae of the male genitalia
described above are used to prevent the untimely
separation of the pair. Sperm transfer occurs
only after the afternoon mating period has
ceased, i. e. during the night. Thus, females en-
tering copulation during the morning hours can-
not mate with other males until the next morn-
ing. Shortly after separation the female will start
ovipositing in the low and dense vegetation
where it is well protected and unlikely to be
found by competing males before the next mat-
ing period. This behaviour has therefore been
considered to be a very effective way of mate
guarding.
The early-morning mate-finding phase where
only optical cues are employed is considered to
be of adaptive value for males, because this be-
haviour secures exclusively virgin females which
will deposit about 80% of their life-time pro-
ductivity of eggs during the few days following
their first copula. Females mating with males
during the morning hours have the advantage
that their mates are most probably also good
nectar exploiters: these males use the same opti-
cal cues for mate locating as they do for flower
finding. Thus a male which can quickly find a
freshly emerged female is likely to be also good
at finding a nectar source.
The late-afternoon mate-locating system is
used mainly during the second half of the flight
period, when virgin females become scarcer, and
when more and more females that have mated
previously tend to enter a second copula, which
may occur because of the need to replenish
sperm or to obtain a second proteinaceous sper-
matophore. It appears that the protein transfer-
red during mating is used not only to sustain the
various life functions of the female, but also to
continue the production of eggs in the ovarioles.
Males can also increase their individual fitness
by mating a second, third or fourth time when
there are no more virgin females available. They
can still increase their individual fitness by up to
20-30% by mating with previously mated fe-
males.
Flower visits by males are induced by a combi-
nation of (violet-blue) colour cues and flower
scents including S(-)-verbenone and are extended
as soon as sugar is found in the flower. This has
been carefully worked out during field tests. The
nectar of Scabiosa and Knautia has been found
to contain trimethyl-pentadecanon. Another
chemically closely related compound which may
be derived from this compound, trimethyl-penta-
decanol, has been shown to form the main com-
pound dispersed by the male scent brushes lo-
cated in the dorsal intersegmental fold of the 8th/
9th abdominal segments. When pairing takes
place these scent brushes are displayed immedi-
ately before the male is accepted by the female
and it is suggested that they serve to mediate be-
tween the two sexes and to convey to the female
information about the efficiency of the male as a
nectar exploiter.
Next to their complicated and fascinating re-
productive biology, of which only an overview
has been provided here, Zygaena moths are also
remarkable for their complicated and unique lar-
val diapause, which occurs only in a morphologi-
cally highly specialized instar involving complex
cuticular adaptations which are thought to re-
duce oxygen and water consumption. In most
species and populations of Zygaena trifolii (and
probably most other Zygaena spp.) there is at
least one obligate larval diapause instar (usually
L4). Diapause may be extended over several sub-
sequent years (up to eight years have been ob-
served in captivity!). However, most larvae com-
mence feeding after the first hibernation, but not
before another moult has taken place which 're-
turns' the larva to physiological and develop-
mental activity. Some larvae may enter a second
diapause in their second year after feeding dur-
The Zygaenoidea
ing one larval instar only, again in a special dia-
pausing instar. Such 'repetitive' diapauses may
occur several times, thus extending the number
o f larval instars considerably. Diapause m a y also
extend over more than one winter season, up to
several years. O n the other hand, bivoltine pop-
ulations o f some Zygaena species are known in
the southern part of the European range (west-
ern and south-western Mediterranean). Diapause
patterns are primarily controlled genetically and
diapause induction is primarily triggered by day-
length, while diapause termination is stimulated
by temperature rise after hibernation. Recent ex-
periments have revealed that hormonal mecha-
nisms are closely related with the diapause be-
haviour o f Zygaena. The complex diapause
pattern o f Zygaena m o t h s is considered to repre-
sent an effective spreading o f survival risk in an
environment with unpredictable winter condi-
tions as is the case in large parts o f the range
o f Zygaena.
References: Alberti (1954); Bode & N a u m a n n
(1988); C o m m o n (1970, 1990); Franzi & N a u -
mann (1984, 1985); G a e d e (1926); H e p p (1920,
1922); Hering (1925); H o f m a n n (1994); H o f -
mann & Tremewan (1996); Jordan (1907/1908,
1909, 1913); N a u m a n n ( 1 9 7 7 a , 1977b, 1985,
1988 a, 1988 b, 1990); N a u m a n n , Feist, Richter &
Weber (1984); N a u m a n n & Tremewan (1984);
Ockenfels, N a u m a n n , Schmitz, Schmidt &
Francke (1990); Povolny & Weyda (1981);
Prinz & N a u m a n n (1988); Tarmann (1984, 1994);
Tremewan (1985); Wipking (1985, 1988, 1986,
1990); Wipking & N a u m a n n (1992); Wipking &
N e u m a n n (1988); Wipking, Viebahn & N e u m a n n
(1995); Witthohn & N a u m a n n (1984).
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 11. The Cossod/Sesioid Assemblage
Edward D. Edwards, Patricia Gentili, Marianne Horak, Niels P. Kristensen & Ebbe S. Nielsen
(E. D. Edwards & E. S. Nielsen: Sesioidea-Castniidae; P. Gentili & M. Horak: Cossoidea;
. P. Kristensen: Introduction, Sesioidea-Brachodidae & Sesiidae)
The Cossoidea and Sesioidea belong to the non-
obtectomeran grade within the Apoditrysia, i. e.,
their pupae have retained the spinosity and con-
siderable mobility of the abdomen, and they are
protruded from their shelter prior to eclosion;
the suggestion (see 12) that the two together
are the sister group of the Zygaenoidea deserves
further scrutiny. The monophyly of the Cos-
soidea + Sesioidea is indicated by two possible
synapomorphies in their larval ground plans
(Minet 1991): prothoracic shield with oblique,
melanized stripe (running between D1 and D2),
and spinules on larval integument irregularly dis-
tributed, tending to concentrate dorsally/antero-
dorsally on most segments. The moths in this as-
semblage are phenetically very diverse; it com-
prises the largest heteroneurous Lepidoptera
outside the Macrolepidoptera (sensu Minet
1991). The Sesioidea-Brachodidae apart, the taxa
treated here are all covered in vols. 2, 6, 10, 14
of Die Groschmetterlinge der ErdelThe Macro-
lepidoptera of the World/Les Macrolpidoptres
du Globe (A. Seitz ed.).
SESIOIDEA
The present concept of the superfamily is due to
Minet (1986, 1991), according to whom the inter-
relationships of the constituent families may be
represented as Brachodidae + (Sesiidae + Cast-
niidae). Probable superfamily autapomorphies:
ocular diaphragm most strongly pigmented ante-
riorly; patagia large, extending ventrad beyond
the anteroventral corners of the pronotum; pos-
terior tendons of metafurcal apophyses elon-
gated caudad or dorsocaudad. The lastmenti-
oned specialization is less pronounced in the
Brachodidae than in the Sesiidae + Castniidae,
whose monophyly is supported by a suite of ad-
ditional apomorphies enumerated by Minet
(1986): distally thickened antennae; antennal
apex with bundle of long hairs; mesothorax with
'parepisterna' (sensu Brock 1971) partly conceal-
ing anterior margin of basisternum; primary
metafurcal arms with anterior and posterior ten-
dons widely separated, the latter approaching
the laminae of the secondary arms; larval pro-
thoracic shield with SD2 close to SD and XD2.
The characteristic crochet configuration on the
larval prolegs (two transverse uniordinal rows),
reported from all sesioid families, is probably an-
other superfamily autapomorphy.
The vast majority of sesioid moths are diurnal
and have prominent ocelli. Proboscis unsealed.
Epiphysis present, spurs 0-2-4. Egg flat. Larvae
usually borers or root feeders, leaf feeding being
recorded only from a brachodid subgroup. Pu-
pae (in family ground plans) with two transverse
rows of dorsal spines on I I - V I (VII in males),
and one row on the following segment (or seg-
ments).
Key to sesioid families
I. Wings narrow, with uniquely specialized
coupling apparatus (Fig. 11.2 E) in addi-
tion to commonplace frenulum/retinacu-
lum; facies (Figs. 11.1 B - D ) usually wasp-
like, wings often partly hyaline . . Sesiidae
Wing coupling by frenulum/retinaculum
only; facies not wasp-like 2
2(1) Antennae with apical/subapical thickening;
wings broad; larger moths with butterfly-
like facies (Figs. 11.2,3) Castniidae
Antennae without such thickening; smaller
moths with 'commonplace micromoth' fa-
cies (Fig. 11.1 A) Brachodidae
Brachodidae. A probable family autapomorphy
is the strong reduction/loss of the ventral arms
of the laterocervicalia (Minet 1991).
Small to medium-sized, robust micro-moths
which mostly (but not exclusively) are diurnal.
Cephalic vestiture variable; chaetosemata absent;
ocelli absent in Hoplophractis, otherwise promi-
nent. Antennae in males uni- or bipectinate or
lamellate, in females usually filiform. Maxillary
palps 13 segmented, usually minute but occa-
sionally enlarged; labial palps upcurved, segment
2 long. Forewing (Fig. 11.2 A) with chorda and
vestigial M-stem sometimes present; pterostigma
absent; spinarea present; no veins stalked; CuP
tubular near margin, or reduced/absent. Hind-
wing with M3 and CuAl separate or stalked;
CuP tubular near margin, or reduced/absent; fe-
male frenulum with 2 bristles.
Few brachodid immatures have been de-
scribed. Larva with prothoracic L bisetose; abdo-
minal segments IVIII with LI and 2 on same
pinaculum.
About 100 species, distributed in all regions
except the Nearctic (Heppner 1991). In a genus
group centred on Brachodes (= Atychia) (Fig.
II.1 A), the described larvae are concealed feed-
182 Edward D. Edwards, Patricia Gentiii, Marianne Horak, Niels P. Kristensen & Ebbe S. Nielsen
Fig. 11.1. Brachodidae: A; Sesiidae: BD. A, Brachodes appendiculata;
formis; D, Synanthedon spheciformis. Scale 1 cm.
ers on monocots; other principal genera are Mis-
cera and Sagalassa. Another genus group,
centred on Phycodes has exophytic leaf-feeding
larvae with prolegs better developed than in the
Brachodes group; recorded food plants belong to
Fabaceae, Melastomaceae and Moraceae. The
moths in the Phycodes group are characterized
by peculiarly enlarged plate-like scales on the
head and thorax; the head is smaller than in
brachodines, extraordinarily small in the mono-
basic Neotropical Hoplophractis, where it ap-
pears partly hidden below the pronotum and is
devoid of ocelli. The two groups were given sub-
family rank by Heppner & Duckworth (1981),
but whereas the Phycodes group seems obviously
monophyletic, there is little support for the mo-
nophyly of the Brachodes group; if the Cossoidea
are indeed the sister group of the Sesioidea it
seems unlikely that the exophytic larval type rep-
resents the ancestral brachodid condition. Pseu-
docossus was transferred to this family from the
Cossidae by Minet (1991).
Principal references: Gartner (1864; life his-
tory of Brachodes), Razowski (1981), Heppner &
Duckworth (1981), Gentili (1989; Pseudocossus);
Common (1990).
Sesiidae. Probable autapomorphies: Retinaculo-
frenate wing coupling supplemented by unique
modification of adjacent wing margins (Fig.
11.2 E): posterior margin of forewing bent
ventrad and engaging with dorsad bent fore mar-
gin of hindwing, both wing margins set with
rows of stiff scales which interlock; female frenu-
lum with single bristle; forewing behind CuA
only with short, unbranched remnant of
1 A + 2 A (often lost); hindwing (Fig. 11.2 B), ac-
cording to Comstock (in Beutenmller 1901),
with medial stem displaced forward, at base in-
separable from Sc + R + Rs; apical vein
Rs + M l . The wasp-like wing proportions, with
proximally markedly narrowed forewings, ap-
pear to be another groundplan autapomorphy.
umilili!
B, Tinthia myrmosaeformis; C, Sesia api-
Putative family autapomorphies in mesopleural
structure (Kozlov 1989) deserve further atten-
tion: pleural ridge anteriorly produced ('looped'),
at base of pleural wing process; mesepimeron
with more or less complete Y-shaped suture.
Rarely small, usually medium-sized to large
micro-moths. Head smooth-scaled; ocelli promi-
nent; chaetosemata present as setose band on
posterior head margin, or absent. Antennae
(Figs. 11.2C, D) variable, sometimes pectinate/
bipectinate in males, also usually (exception Sesi-
inae-Osminiini) ventrally 'ciliate' in males.
Maxillary palp small, 13 segmented; proboscis
(Fig. 11.2 C) in ground plan well developed, re-
peatedly reduced. Labial palp usually prominent,
upcurved or porrect; basal segments with vari-
able, sometimes extensive vestiture, distal seg-
ment consistently smooth-scaled. Forewing with-
out chorda and M-stem (except, occasionally, a
short distal vestige); spinarea present; Rs3 and 4
usually stalked (connate or sessile in some Tin-
thiinae), position of Rs4 relative to apex variable;
hindwing CuP present or absent; 1A and 2 A
fused or (Sesiinae except Paranthrenini) separate,
3 A present or absent. Genitalia very variable.
The resemblance of sesiid moths to stinging
Hymenoptera furnishes a classical example of
Batesian mimicry. The scale/hair vestiture often
has a metallic sheen, and the body and legs are
usually conspicuously (yellow, red or white)
banded. More or less extensive wing transpar-
ency is characteristic of most sesiids, but pig-
mented scales are always retained along the wing
margins and on the veins. It has been suggested
(Kristensen 1974) that in the sesiid ground plan
the entire wing vestiture comprised an upper
layer of pigmented 'normal-type scales' and a
lower layer of near-transparent 'primitive-type
scales' (see 21). Wing transparency could, then,
have evolved independently within the two sub-
families, usually through initial loss of the upper
layer followed by a more or less extensive loss of
the lower-layer scales; however, in the examined
The Cossoid/Sesioid Assemblage
Fig. 11.2. Brachodidae: A; Sesiidae: B - . A, Sagalassa, wing venation; B, Podosesia, hind wing venation; C,
Paranthrene, head; D , tinthiine, antennal apex; E, sesiid wing linking mechanism, diagram; F, Synanthedon male
genitalia (phallus upper right); G, Melittia larva. A - F after Heppner & Duckworth (1981), G after Heppner
(1987).
Sesiinae-Sesiini, pigmented 'normal-type scales'
are present in the newly emerged moth, but on
the greater part of the wing surface they have
an unusually low cuticular collar around their
sockets. Undoubtedly there is much homoplasy
in scaling characters: in most tribes there are
genera which include species with 'normal' vesti-
ture as well as species with extensive wing trans-
parencies. A configuration with forewings nor-
mally scaled (or almost so) and hindwings exten-
sively transparent (Fig. 11.1 B) is also recurrent.
Larvae (Fig. 11.2G) semiprognathous, nearly
always borers. Prothoracic L trisetose; abdomi-
nal segments I - V I I I with LI and 2 on same pi-
naculum. Pupa with stout spines on terminal seg-
ment.
Adult sesiids are rarely observed in nature,
though several are flower-frequenting. The iden-
tification and synthesis, since the 1970s, of the
female pheromones of a number of sesiids not
only enhanced monitoring the occurrence of pest
species, but have also led to a breakthrough in
the inventory of local faunas of these moths, in-
cluding the discovery of several new taxa (review
by Prola & Beer 1995; see also 2 - 1 3 ) . The larval
host spectrum of sesiids includes many dicot
families and both woody and herbaceous forms;
most are trunk-, branch-, stem- or root-borers,
but seed-boring has been reported from a few
tropical Carmenta (Sesiinae-Synanthedonini),
and may be more widespread than hitherto be-
lieved (Harms & Aiello 1995). Almost all species
have narrow host ranges and several are pests.
183
Synanthedon coccidivora from Venezuela is aber-
rant in being carnivorous, feeding on coccids
(Duckworth 1969).
The foundations for a modern phylogenetic
classification of the Sesiidae were laid by Nau-
mann (1971), whose division of the family into
only two subfamilies is adopted here. The group-
ings here given subfamily and tribal ranks were
established from studies on the Holarctic fauna,
and it remains to be seen how well they accom-
modate the family's tropical representatives; they
apply well to the (admittedly depauperate) Aus-
tralian fauna (Duckworth & Eichlin 1974).
Tinthiinae. Probable autapomorphies: chaeto-
semata absent; hindwing 3 A lost; antennae non-
clavate, apex without scale tuft (Fig. 11.2 D);
gnathos absent. While the last-mentioned traits
were considered by Naumann to be plesiomor-
phic, Minet (1986) argued that the polarity
should be reversed, referring to conditions in
Castniidae.
Meso-'preepimeron' simple; hindwing CuP at
least partly developed; male vinculum without
posterior processes and without saccus.
83 species worldwide (Heppner 1991). Three
tribes: Tinthiini (Fig. 11.1 B), Pennisetiini and
(Spatenka et al., 1993) Similipepsini. Pennisetia
species are borers in Rubus (Rosaceae) and the
genus includes pest species in both the Palearctic
and N. America.
Sesiinae. Probable autapomorphies: chaeto-
semata linear, restricted to posterior head mar-
gin; mesopleuron with anterodorsal part of epi-
184 Edward D. Edwards, Patricia Gentiii, Marianne Horak, Niels P. Kristensen & Ebbe S. Nielsen
meron ('preepimeron' sensu Shephard 1930)
forming unique bag-like protuberance which
conceals the pleural suture; hindwing CuP con-
sistently vestigial or absent (its position remain-
ing demarcated by scale line, Fig. 11.2 B); male
vinculum with posterior processes and promi-
nent saccus.
Antennae variously davate, with apical scale
tuft (Fig. 11.2 D); hindwing 3 A and male
gnathos present in ground plan. Prominent mel-
anized sensilla with (group-specific) variably
shaped apex developed on valve in most lineages,
in Synanthedonini (Fig. 11.2 F) also on elongate
'scopula androconalis' flanking anal cone; they
are not a sesiine groundplan trait.
1040 species worldwide (Heppner 1991); 110
genera were listed by Heppner & Duckworth
(1982), but the generic splitting of tropical sesi-
ines seems unjustifiably excessive (Duckworth &
Eichlin 1974). Six tribes are recognized: Par-
anthrenini, Cissuvorini, Sesiini, Osminiini, Melit-
tiini and Synanthedonini. The two first-men-
tioned constitute the subfamily Paranthreninae
of Duckworth and Eichlin, but synapomorphies
of the remaining Sesiinae have not been iden-
tified (absence of basal anastomosis of hindwing
1 and 2 A also present in some Paranthrenini
(Naumann 1971). Sesia includes the familiar Ho-
larctic 'hornet moths' (Fig. 11.1 C) whose larvae
are borers in Salicales. The almost worldwide
(not W. Palaearctic) Melittia with conspicuous
hair tufts on the hind tibiae/tarsi, includes some
of the largest sesiids; their larvae are borers in
Cucurbitaceae, some species attaining pest
status. The Synanthedonini (Fig. 11.1 D) com-
prise the bulk of the family's species; they are
small to medium-sized, slender moths. Several
pests belong here, including the Nearctic 'peach
tree borers' Synanthedon pictipes and S. exitiosa,
and the European 'currant borer' S. tipuliformis,
which has been casually introduced into many
parts of the World.
Principal recent works with references to the
extensive systematic literature on the family in-
clude Duckworth & Eichlin (1974, 1977), Hepp-
ner fe Duckworth (1981; with a checklist of the
World's species), Eichlin (1986), Eichlin & Duck-
worth (1988), Spatenka et al. (1993), Arita (1994)
and Lastuvka & Lastuvka (1995).
Castniidae. The broad wings, diurnal habit,
clubbed (often abruptly clubbed) antennae, the
bifurcate saccus, the form of the stem of M in
both wings, enlarged phallobase, recurved aedea-
gus and very extensible ovipositor have been
used to characterise the Castniidae although
each is found elsewhere in the Lepidoptera.
Rammert (1993) suggested that the trough-
shaped tergum I might be an autapomorphy of
the Castniidae. It seems probable that the family
is monophyletic. In recent years there has been
no disagreement as to the included taxa. The
family has been previously linked with the but-
terflies and the Cossidae, and with some other
day-flying moth families such as the Agaristinae
(Noctuidae). Modern structural studies suggest
no close relationship with these groups.
Medium to large moths (Figs. 11.3, 4, 5 A);
wingspan ranging from 24190 mm. Cephalic
vestiture variable, vertex with lamellar scales di-
rected forwards or with lamellar and piliform
scales directed forwards, frontoclypeus with lam-
ellar or piliform scales; ocelli present, large and
raised; chaetosemata absent; antennae about half
to three quarters length of fore wing with
smoothly scaled flagellum, broadly clubbed api-
cally, apiculus elongate or small to minute some-
times with tuft of long scales; pilifers present;
proboscis naked, well developed, reduced or rar-
ely vestigial; maxillary palpi small, 2 - 4 seg-
mented; labial palpi 3 segmented, distal segments
occasionally fused, erect or ascending, short or
of moderate length. Thoracic terga with closely
appressed lamellar scales mixed with piliform
scales; parapatagia and tegulae well developed;
legs smoothly scaled, epiphysis present, some-
times reduced or spine-like; spurs 0-2-4 usually
asymmetrical and well developed; tarsal spines
present; pulvilli on pretarsus sometimes inflated,
half length of tarsal claw, arolium present, of
variable shape, pseudempodial seta present. Fore
wing (Fig. 11.6 A) broad, pterostigma absent;
chorda generally present, all 5 R/Rs veins pre-
sent, sometimes all free, sometimes with 2 or 3
veins stalked; stem of M well developed lying
close to the cubital stem, anterior branch vestig-
ial, M2M3 cross-vein sometimes weak or ab-
sent, CuP present or absent, 1 A + 2 A with long
fork; retinaculum in male membranous, beneath
Sc; metathoracic wing locking device present. In
hind wing frenulum a single composite bristle in
male, 316 bristles in female, Sc + R separate
from or shortly fused with Rs near base, stem of
M present in cell lying close to cubital stem,
M 1 - M 2 cross-vein absent (cell open), CuP pre-
sent or absent, 1 A + 2 A usually not forked at
base, 3 A present. Specialised scale patches some-
times present on one or both wings. Male genita-
lia (Figs. 1 1 . 6 D - F ) with uncus broad, narrow
or bifurcate, separate or fused with tegumen,
gnathos present, arms not fused, saccus bifurcate
with arms projecting into eighth abdominal seg-
ment, valva rounded or obtuse and excavate near
sacculus, without complex ornamentation, ae-
deagus recurved, often asymmetrical, with en-
larged phallobase. In female genitalia (Fig.
11.6G) segments VIIIX extensible and may
form a long tapering ovipositor; ovipositor lobes
may be lightly or heavily sclerotised and may be
blade-like and broad or long and acute, posterior
apophyses may be very long; ductus bursae may
be short and straight or long, loosely coiled or
long, tightly spiralled, it may be membranous or
sclerotised, with or without spinose projections;
The Cossoid/Sesioid Assemblage
Fig. 11.3. Castniidae: A, Tascina nicevillei; , T. metallica; C, Athis inca; D, Castnia eudesmia. (A, B, The Natural
History Museum, London; C, D, J. Green, CSIRO Entomology, Canberra). Scales: 25 mm.
corpus bursae oval to long and bulbous, signa
may be present or absent.
Egg (Fig. 11.5 B) of 'flat' type, broadly fusi-
form with 4 or 5 prominent longitudinal ridges,
micropyle at one end of long axis, sometimes
with numerous transverse striae. Corybantes py-
lades has round dorsolateral^ flattened eggs, but
other Corybantes species have fusiform eggs.
Larva (Fig. 11.6B) stout, cylindrical; head prog-
nathous, deeply retractable into prothorax, with
6 stemmata on each side, frontoclypeus one third
to two thirds distance to epicranial notch, seta
O l midway between stemmata 1 and 2 or be-
tween 2 and 3; prothorax with large sclerotized
shield, SD2 near to SD and XD2, SD group
sometimes with additional secondary setae, L
group tri- or bisetose on TI T3 (sometimes with
additional secondary setae), SV group uni- or bi-
185
setose; T2, T3 and II - V I sometimes with coarse
spinules dorsally and ventrally; 1st instar with
uniordinal crochets in two transverse rows, last
instar larva with multiserial crochets or crochets
absent. Pupa (Fig. 11.6C) long cylindrical, well
sclerotised; head with clypeo-labral suture pre-
sent, proboscis about half length of wings, maxil-
lary palps small sclerites or absent, abdomen
with 2 rows of dorsal teeth on segments IIVII
in male and IIVI in female and segments VII
(in female), VIII and IX (in both sexes) with an
anterior row of dorsal teeth; without cremaster.
Most Castniidae have cryptically coloured
fore wings and brightly coloured hind wings,
sometimes with iridescent scales. Some species of
Castniini mimic butterfly or moth species with
which they fly. The Castniinae fly mostly in the
heat of the day, from about 11 am to 2 pm, and
186 Edward D. Edwards, Patricia Gentiii, Marianne Horak, Niels P. Kristensen & Ebbe S. Nielsen
l ,
Fig. 11.4. Castniidae: A, Synemon plana; B, S. nupta.
(J. Green, C S I R O Entomology, Canberra). Scales:
10 mm.
species of Synemon fly in bright sunshine. Some
species, eg. Eupalamides cyparissias, may be cre-
puscular and occasionally Synemon can be ob-
served flying at sundown. In one Australian spe-
cies (Synemon plana) the female has somewhat
Fig. 11.5. Castniidae: A, resting adults of Synemon magnifica:
1981).
reduced wings but can still fly. A population of
Synemon selene is known only from females and
is suspected to be parthenogenetic. Some South
American species are regarded as threatened due
to clearing of native vegetation and several Aus-
tralian temperate grassland species are endan-
gered or rapidly declining in numbers (Edwards
1993). Castniid eggs are laid singly and inserted
into crevices, between plant parts or between
plant and soil. The larvae feed internally in plant
tissue or in the soil on subterranean plant parts
retaining frass within the excavated galleries. All
confirmed foodplant records are monocotyle-
dons in the families Arecaceae (Palmae), Bromel-
iaceae, Cyperaceae, Ecdeiocoleaceae, Lomandra-
ceae, Marantaceae, Musaceae, Orchidaceae, and
Poaceae (Gramineae). A single record of Ery-
nium (probably Eryngium) in the Apiaceae
(quoted by Miller (1986)) is unconfirmed. Larvae
are normally solitary and the larval stage may
take from 4.5 months (Telchin licus) to 2 years
(,Synemon magnifica). Some South American
species are minor pests of crops: Telchin licus in-
fests sugarcane, Castniomera atymnius (as hum-
boldti) bananas, and Eupalamides cyparissias oil
palms. Larvae of the Chilean Castnia eudesmia
are gathered for fish bait (Miller 1987). Pupation
occurs in a cell incorporating plant fragments
within plant tissues (Castniini) or in a gallery in
the soil where an escape tunnel is prepared by
the larva (Synemonini). The pupa is mobile and
moves along the tunnel to protrude into the open
, eggs of S. plana (from Common and Edwards
The Cossoid/Sesioid Assemblage 187

Fig. 11.6. Castniidae: A, wing venation (Synemon plana); B, larva, C, pupa (Synemon magnifica), DF, male
genitalia (5. collecta); D, lateral view, E, tegumen and uncus dorsal view, F, saccus; G, female genitalia (S. plana).
(A, G from Common 1990, B - C from Common and Edwards 1981; D - F by S. P. Kim, CSIRO Entomology,
Canberra). Scales: A - C , 5 mm; D - F , 0.5 mm; G, 1 mm.

prior to eclosion. No information is available on nae (sometimes spelled Tasciniinae, = Neocastni-

the biology of the Tascininae.
The family distribution strongly indicates a
Gondwanan origin and is of unusual interest
with the subfamily Tascininae (in South-East
Asia) being the sister group of the Castniinae
(containing the South American and the Austra-
lian species). None are known from the Indone-
sian islands east of Sumatra or from New
Guinea. The Castniidae are found in tropical,
subtropical, and warm temperate environments
but not in cool temperate areas. They are absent
from the southernmost parts of South America
and from Tasmania and New Zealand. Miller
(1986) recognized two subfamilies, the Tascini-
inae) and the Castniinae.
Tascininae are a small subfamily with the sin-
gle genus Tascina Westwood (= Neocastnia) with
three species known from Burma, Malaysia, Sin-
gapore, Indonesia (Sumatra and Borneo) and the
Philippines (Palawan). Very few specimens are
known and little structural information is avail-
able on them. The Tascininae have antennae with
long apiculus and the proboscis is vestigial. The
fore wing has an acute apex, the cell is open and
R to Rs2 are stalked and Rs3 and Rs4 are also
stalked. The accessory cell in the hind wing is
reduced. There are no descriptions of the genita-
lia of either sex.
188 Edward D. Edwards, Patricia Gentiii, Marianne Horak, Niels P. Kristensen & Ebbe S. Nielsen
Castniinae have antennae with a moderate,
short or absent apiculus and usually have a func-
tional proboscis. The fore wing usually has a
rounded apex, the cell is usually closed and the
radial veins are separate at their base or vari-
ously stalked. The accessory cell in the hind wing
is not reduced. The male genitalia have a bifur-
cate saccus and the aedeagus recurved. The fe-
male genitalia usually have a very long extensible
ovipositor. The Castniinae contain two tribes,
the Castniini and the Synemonini. The Synemo-
nini, with about 43 species (23 named) in the sin-
gle genus Synemon, are confined to mainland
Australia. The Castniini (Miller 1986) are found
in Central and South America, from northern
Mexico and the West Indies to central Chile and
northern Argentina. There are 32 genera recog-
nised containing about 134 species (Miller 1995)
or 81 species (Lamas 1995). The Synemonini
possess the characters of the Castniini but the
epiphysis is smaller, the bifurcate saccus reduced
in length, and the aedeagus is less recurved. Nev-
ertheless the close similarity of all the Australian
Castniidae suggest that the Synemonini will
prove to be monophyletic.
Principal taxonomic works are: Dalla Torre
(1913), Houlbert (1918), Miller (1986; cladistic
study of the genera of the Castniini with a listing
of species and available biological information),
Common (1990; account of Synemonini), Ed-
wards (1996, Synemonini checklist), Lamas
(1993; Castniini literature list), Miller (1995;
Castniini checklist), Lamas (1995; same, with ad-
ditions/corrections). Information on the early
stages of the Castniini may be found in Miller
(1986, 1987) and on the Synemonini in Common
and Edwards (1981).
COSSOIDEA
The Cossidae and Dudgeoneidae are pheneti-
cally similar but none of the suggested apomor-
phies for the two families have been confirmed.
As there is no better supported alternative hy-
pothesis for their relationships they are here
treated as members of the Cossoidea, following
Common (1990), Minet (1991) and Nielsen &
Common (1991). The recent discoveries of male
retardines have revealed that these butterfly-like
moths are subordinate within the Cossidae. A
relatively plesiomorphic wing venation with the
M-stem forked in both wings in most cossoids
suggests that they are among the earliest
branches of the Apoditrysia. They are distributed
worldwide with the exception of New Zealand.
Based primarily on the presence of a paler,
membranous area across the posterior part of
the mesepimeron Brock (1971) also included the
exophagous Limacodidae, Dalceridae, Megalo-
pygidae and Chrysopolomidae in the Cossoidea,
together with the Cossidae, Metarbelidae and
Ratardidae. This could not be confirmed by sub-
sequent authors and the exophagous group are
currently considered to belong to the Zygae-
noidea (Minet 1986, 1991; Common 1990; see
1 10). Minet (1991) mentions a pair of asym-
metrical pits in the lower part of the frontoclyp-
eus (Fig. 11.IOC, D), together with strong reduc-
tion or loss of proboscis, as synapomorphies of
Dudgeoneidae and Cossidae, but the first char-
acter could not be found in a number of Austra-
lian Cossinae and New World Hypoptinae and
Cossulinae. Boring and feeding in woody tissue
could also be a possible synapomorphy for the
cossoids.
Adults small to very large, usually robust;
ocelli absent except in a few generalized Cossi-
nae; chaetosemata absent; proboscis unsealed,
reduced or absent; maxillary palps minute, at
most 4 segmented, antennae usually bipectinate
in male; mesophragma long, bilobed, reaching
the thoraco-abdominal articulation (Fig. 11.11);
metafurcal stem short; retinaculum in male a
lobe from Se or from between costa and Sc (usu-
ally absent in Metarbelinae and often lost in Hy-
poptinae and some Cossinae); M present and
usually forked in cell of both wings; fore wing
usually with chorda in cell; 1 A + 2 A in fore
wing deeply forked at base, fork shorter in hind
wing; 3 A present in hindwing; S2 of tortricoid
type. Larvae stem- and woodboring or feeding
on bark from short tunnel.
Key to families and subfamilies of Cossoidea
1 Tympanal organ present at base of abdo-
men (Fig. 11.10 A); moths reddish brown
with silvery white spots on fore wings (Fig.
11.7H) Dudgeoneidae
Tympanal organ absent at base of abdo-
men; moths with variable coloration . . . .
(Cossidae) 2
2 (1) Antennae bipectinate at base, serrate to
tip (Fig. 11.7 B); aedeagus fan-like (Fig.
11.8 D); tibial spurs 0-2-2, short
Zeuzerinae
Antennae filiform, uni- or bipectinate to
tip; aedeagus tube-like; tibial spurs long 3
3 (2) Antennae uni- or bipectinate to tip, asym-
metrical; tibial spurs asymmetrical; re-
stricted to the New World . . . Cossulinae
Antennae filiform or bipectinate, symmet-
rical, sometimes with unpaired midventral
lamellar expansion (Cossus, Chileco-
madia)\ tibial spurs symmetrical 4
4 (3) Hind tibia and first tarsomere inflated;
moths greyish brown, with reticulated
pattern Cossinae
Hind tibia and first tarsomere not inflated
5
5 (4) Female genitalia short, not telescopic (Fig.
11.8 L); restricted to the New World . . . .
Hypoptinae
Female genitalia short, telescopic, oviposi-
The Cossoid/Sesioid Assemblage
Fig. 11.7. Cossidae: A G; Dudgeoneidae: H., Culama australis (Cossinae); B, Endoxyla amphiplecta (Zeuzerinae);
C, Cossula magnifica (Cossulinae); D, Lebedodes naevius (Metarbelinae); E, Ratarda excellens (Ratardinae); F,
Hypopta superba, G, Langsdorfia lunifera (Hypoptinae); H, Dudgeonea actinias (A, B: Nielsen & Common 1991:
figs. 41.47 A, C; E by M. Owada). Scales = 5 mm.
tor lobes 8-shaped (Fig. 11.8 K); Old World
tropics 6 11.IOC, D); antennae less than half length of
6 (5) Wings broad, of almost equal area; butter-
fly-like moths (Fig. 11.7E) . . . Ratardinae
Wings broad, but not of equal area; never
butterfly-like (Fig. 11.7 D) . Metarbelinae
Cossidae. Minet (1986) suggests that the family
is defined by an autapomorphy of the prothorax,
namely medially adjacent, well projecting para-
patagia (see Schultz 1914: fig. 8), noting that the
character is weakly developed in some Cossinae.
Among Australian Cossidae medially approxi-
mated parapatagia are well-developed in Culama
and present, but much smaller than the patagia,
in Mioses. Schoorl (1990) lists high, flap like par-
apatagia as pertaining to a hypothetical cossid
groundplan (as 'supposed plesiomorphies of
Cossidae'), but gives in his cladogram for the
cossid subfamilies the character 'mesepimeron
rather high' as the synapomorphy linking all
subfamilies, though stating subsequently that the
mesepimeron is often lowered within the family.
Schoorl (1990) has excluded a large number of
taxa from the Cossidae mainly on the basis of
thorax and wing base structure. Our study of the
Australian and southern South American taxa in
question did not confirm his conclusions which
seem to be based on too narrow a selection of
characters.
Small to very large, usually robust moths;
wingspan ranging from 9 to 236 mm. Cephalic
vestiture variable, vertex sometimes with a longi-
tudinal crest of erect scales; often with a pair of
189
small pits on lower part of frontoclypeus (Fig.
forewing, usually bipectinate, rarely filiform, ser-
rate or unipectinate in male, usually filiform in
female, with intercalary sclerite except in Zeuzer-
inae; pilifers absent; maxilla (proboscis) an un-
sealed small triangular lobe or further reduced;
maxillary palps very small, sometimes 4-seg-
mented, usually much further reduced; labial
palps short, 3-segmented, upturned and reaching
to middle of frons or extremely reduced; medial
segment the longest or subequal to basal seg-
ment; apical segment oval or pyriform, vom
Rath's organ terminal. Thoracic vestiture vari-
able, hairs only or mixed with scales (most Cossi-
nae, Zeuzerinae), or upright scales forming tufts
and crests or collars on pro- or metathorax
(found in Hypoptinae, Cossulinae and Metarbe-
linae); pronotum variable in shape in dorsal
view; parapatagia usually well developed, promi-
nent, medially approximated. Epiphysis present
(except in Acousmaticus and some Metarbelinae
and Ratardinae), ranging from less than half
length of tibia to as long as tibia (Cossulinae),
inserted halfway along the tibia (Cossulinae,
Metarbelinae) or in basal third (Cossinae, Hy-
poptinae, Zeuzerinae); tibial spurs 0-2-4 or 0-2-
2, usually symmetrical and well developed, some-
times asymmetrical; tarsal spines present or ab-
sent; pretarsus usually complete; ungues well de-
veloped; pulvilli always present but sometimes
reduced to minute oval lobes; arolium and
pseudempodium present or absent. Fore wing
(Fig. 11.8 A) elongate (except Ratardinae),
190 Edward D. Edwards, Patricia Gentiii, Marianne Horak, Niels P. Kristensen & Ebbe S. Nielsen

Fig. 11.8. Cossidae: A, D - G , I - L ; Dudgeoneidae: B, C, H. Wing venation and genitalia: A, , wing venation
of: A, Endoxyla, B, Dudgeonea; CG, male genitalia with aedeagus of: C, Dudgeonea sp., D, Xyleutes strix
(Zeuzerinae), E, Hemipecten niveogrisea (Cossulinae), F, Metarbelodes umtaliana (Metarbelinae), G, Hypopta wat-
soni (Hypoptinae); HL: female genitalia of: H, Dudgeonea sp., I, Chilecomadia moorei (Cossinae), J, Hemi-
pectrona nigripennata (Cossulinae), K, Marshalliana bivittata (Metarbelinae), L, Givira nais (Hypoptinae) (A, B:
Nielsen & Common 1991: figs. 41.48 B, C).

strong, pterostigma rarely present; chorda usu-
ally present (except Metarbelinae and some Hy-
poptinae), accessory cell often protruding be-
yond cell (Zeuzerinae and some Cossinae); con-
figuration of radial veins varying from entirely
free to all radial sectors on common stalk (Para-
cossus), R from before accessory cell (Hypopti-
nae, some Cossinae and Zeuzerinae) or from ac-
cessory cell (Cossulinae, most Cossinae and
Zeuzerinae); M forked in cell, but anterior
branch sometimes reduced or absent, posterior
branch ending between M2 and M3 or M3 and
C u A l ; medians beyond cell usually separate and
equally spaced, sometimes M l displaced towards
radiais to stalked with Rs4, M2 sometimes dis-
placed towards M3; CuP present or absent,
sometimes obsolete except towards wing margin;
1 A + 2 A sometimes connected with CuP by a
transverse vein or fused for a short portion
towards the wing margin. In hind wing (Fig.
11.8 A) frenulum a single composite bristle in
male, 230 bristles in female (usually 57), usu-
ally absent in Metarbelinae and often lost in Hy-
poptinae; Sc + R independent from Rs or con-
The Cossoid/Sesioid Assemblage
Fig. 11.9. Cossidae: Male genitalia with aedeagus of
Ratarda excellens (Ratardinae).
nected by one or two transverse veins at the base
and/or near the end of cell; Rs and M1 separate,
connate or stalked; CuP present, sometimes ob-
solete in basal half. Male genitalia (Figs.
11.8DG) with uncus prominent, broad or
beak-like, separate or fused with tegumen; socii
present or absent; gnathos present or absent,
branches usually fused in a transverse median
plate (= bulla), which is often prominent; vincu-
lum and tegumen separate or fused into a ring;
saccus present, from a slight concavity to a digi-
tiform invagination; valva with intravalvar mus-
cle (m7) present, well sclerotized, very often sim-
ple but different processes can be present;
transtilla present or absent; juxta usually a plate,
often with two dorsal digitiform processes; ae-
deagus medium sized to small, usually without
coecum (present in some Cossinae). In female
genitalia (Figs. 11.81-L) segments V I I I - X par-
tially retractable into previous segment (tele-
scopic ovipositor) except in Hypoptinae; apoph-
yses posteriores very long in Cossinae and Zeuz-
erinae (long telescopic ovipositor); ovipositor
lobes not sclerotized and usually narrow (broad-
ened, 8-shaped in Metarbelinae and Ratardinae;
triangular in Hypoptinae); sterigma often con-
nected with ventral arm of apophyses anteriores;
ductus bursae short; corpus bursae oval; signa
rarely present.
Egg of 'flat' type in Zeuzerinae, Metarbelinae,
Hypoptinae (Minet 1986) and the plesiomorphic
Australian cossine Culama (Common 1990: fig.
2.8), upright in other Cossinae; oval to elliptical,
shape modified by oviposition site; chorion from
smooth to dimpled, striate or with vertical and
horizontal ribs. Surcossus perlaris, the only spe-
cies in a genus of unknown affinities and known
only from females, has intraovarial eggs with a
hardened chorion, suggesting a probable case of
parthenogenesis, yet to be tested, and there is
one reference to parthenogenesis in Chilecomadia
191
valdiviana (Elgueta 1932). Larva stout and usu-
ally cylindrical (slightly flattened in Zeuzerinae);
head relatively small, semiprognathous, with 6
stemmata on each side, with the frontoclypeus to
only one-half or less of the distance to the epicra-
nial notch; prothorax with large, dorsal sclero-
tized shield, often humped and rugose (Zeuzeri-
nae), with SD2 nearly caudad to SD; L group
trisetose on TI, T2 and T3, LI usually on a sepa-
rate pinaculum and closer to L3 than L2 on T2
and T3; ventral prolegs stout with irregularly
uni-, bi- or triordinal crochets arranged in an el-
lipse or more rarely in a penellipse or transverse
bands. Pupa long, cylindrical, well-sclerotized;
head with lateral part of fronto-clypeal suture
distinct and clypeo-labral suture visible, with ex-
tremely short proboscis, short antennae, maxil-
lary palps minute or absent, frequently with ce-
phalic projection; abdomen with segments III
VII in male and IIIVI in female free, I I - V I I
in males and IIVI in females with two
transverse rows of dorsal spines (in Macrocyttara
also I), VII in female and VIII in both sexes with
one row; without cremaster but spines on VIII
also ventrally.
Adult cossids, with the exception of the Ratar-
dinae, are exclusively nocturnal in activity. The
eggs are usually laid in groups inserted under
loose bark and in crevices with an extensible ovi-
positor, or, more rarely, on the surface as an egg
mass (Hinton 1981). There is not sufficient infor-
mation available to correlate oviposition behavi-
our with ovipositor morphology. Egg numbers
can be vast, with over 18000 eggs laid by a fe-
male of Endoxyla encalypti (Nielsen & Common
1991, as Xyleutes encalypti). After an initial dis-
persal phase, sometimes windblown on silk
threads, larvae bore into branches or trunks of a
great diversity of hosts, pushing frass and debris
to the outside and usually boring deeper as the
larvae mature in 14 years. Occasionally they
live gregariously in galleries beneath the bark
(Common 1990) or feed externally on roots
(Nielsen & Common 1991). Phragmataecia is a
borer in large Gramineae (Kalshoven 1981). The
larva of Azygophleps albo vit tata is able to dia-
pause in the soil and reinfest new plants after the
dry season (Carter & Deeming 1980). Pupation
takes place in the gallery or, more rarely, in the
ground, and the pupa is extruded before emer-
gence of the adult.
The Cossidae are here considered to comprise
the Cossinae, Zeuzerinae, Cossulinae, Metarbeli-
nae, Ratardinae and Hypoptinae. The Pseudo-
cossinae, as delimited by Heppner (1984), are
polyphyletic, with Pseudocossus possibly belong-
ing to the Brachodidae (Minet 1991) and Chile-
comadia to the Cossinae. The Pseudocossinae are
mainly based on the alleged shared presence of
'prototympanal' organs in the abdominal base,
which could be confirmed neither for Pseudocos-
sus (Minet 1983) nor for Chilecomadia. The Chile-
192 Edward D. Edwards, Patricia Gentiii, Marianne Horak, Niels P. Kristensen & Ebbe S. Nielsen
Fig. 11.10. Dudgeoneidae: A, B; Cossidae: C, D. Tympanal organs (A, ), and frontoclypeal pits (C, D): A, B,
abdominal tympanal organs of: A, Dudgeonea sp.; B, Acritocera negligens; C, D, paired pits on frontoclypeus of
Endoxyla lituratus, D showing detail of C.
Fig. 11.11. Cossidae: Mesophragma and metathorax
of Chilecomadia valdiviano: A, lateral view; B, poste-
rior view; c coxa, em epimeron, es episternum,
m - metafurca, mp - mesophragma, s - subalare,
sc scutum, si scutellum.
comadiinae (Schoorl 1990) are predominantly
defined by plesiomorphies and are here regarded
as generalized Cossinae for they have an inflated
hind tibia and first tarsomere. Much more infor-
mation, particularly about the neotropical fauna
(approximately 40% of cossid species) and about
immatures, will be needed to elucidate the rela-
tionships among the subfamilies.
Cossinae (Figs. 11.7 A, 11.81). Antenna bipec-
tinate in male up to tip or filiform in both sexes,
male in some species with midventral lamellar
expansion (Cossus cossus)-, labial palps well de-
veloped; tibia and first tarsomere of hind leg di-
lated; tibial spurs usually 0-2-4, rarely 0-2-2; un-
cus broadly uni- or bilobed, socii present in few
plesiomorphic genera; gnathos arms usually
fused medially, often with well-developed bulla;
digitiform processes of the juxta usually present;
aedeagus tube-like, coecum present or absent; fe-
male with long telescopic ovipositor (Fig. 11.81).
The dilated tibia and first tarsomere of the hind
leg is a probable apomorphy for the Cossinae,
already present in such plesiomorphic genera as
Culama and Chilecomadia. In the cossine hind-
wing Rs and Ml are usually connate or stalked,
a character generally used to differentiate Cossi-
nae from Zeuzerinae, but Chilecomadia has
widely separate Rs and M l , probably illustrating
the plesiomorphic condition for the subfamily as
well as for the family. Cossinae (ca. 100 species
in ca. 35 genera; Parahypopta, Dyspessa, Prio-
noxystus, Comadia) are present on all continents
and most diversified in the Old World, with some
generally plesiomorphic genera in Australia,
southern South America and India {Catopta
(with ocelli and socii), Macrocyttara, Culama,
and Chilecomadia (with socii)). The southern
South American Surcossus might belong here,
also the Australian Mioses with well-developed
ocelli, though they both lack dilated tibiae and
first tarsomeres on their hind legs.
Zeuzerinae (Figs. 11.7B, 11.8 A, D, 11.10C,
D). Large, heavy-bodied moths; antenna without
intercalary sclerite and in male bipectinate for
not more than basal three-quarters, remainder
serrate, female antenna very short, bipectinate at
base; labial palps extremely short; tibial spurs 0-
2-2, very short and sometimes concealed in pock-
ets of the first tarsomere; fore wing usually long
and narrow; accessory cell large and distally pro-
truding beyond cell (Fig. 11.8 A); R usually aris-
ing from accessory cell; Rs3 + Rs4 stalked; Rs
and Ml veins separate in hind wing; gnathos, if
present, without bulla; juxta a bilobed plate with
simple, digitiform processes; aedeagus short,
The Cossoid/Sesioid Assemblage
wide, apical and expanded into fan-like mem-
brane; female with long telescopic ovipositor.
The partially bipectinate antenna without an in-
tercalary sclerite and the apically fan-like aedea-
gus are possible apomorphies for the subfamily.
Zeuzerinae (ca. 190 species in 30 genera; Zeuz-
era, Eulophonotus, Phragmataecia, Xyleutes,
Psychonoctua, Azygophleps) are most diversified
in tropical lowlands. They have a worldwide dis-
tribution apart from southern South America
and are well represented in Australia, with over
100 species of Endoxyla, reaching a wingspan of
230 mm in the females of the largest species.
Cossulinae (Figs. 11.7C, 11.8 E, J). Antenna
uni- or bipectinate, asymmetrical; prothorax and
metathorax often with a collar of upright scales;
epiphysis roughly from middle of fore tibia and
as long as fore tibia; tibial spurs 0-2-4 or 0-2-2,
asymmetrical, long; R from accessory cell; uncus
beak-like (Fig. 11.8 E), wide at base; bulla pro-
truding; aedeagus tube-like, apex membranous
usually with large spine; female with short, tele-
scopic ovipositor (Fig. 11.8 J). Schoorl (1990) ex-
cludes the Cossulinae from the Cossidae on the
basis of low or only moderately high patagia, the
absence of a pale band across the posterior part
of the mesepimeron and larval crochets arranged
in two transverse bands instead of a complete
circle. The pale band across the mesepimeron,
found also outside the cossoids, is absent also in
Culama which nevertheless belongs to the Cossi-
nae on the strength of dilated tibia and first tar-
somere of the hind leg. The asymmetrical anten-
nae are a possible apomorphy for the subfamily.
The Cossulinae (ca. 50 species in 5 genera; Cos-
sula) are restricted to the New World, concen-
trated in tropical and subtropical Central and
South America. Austrocossus from southern
South America seems to have affinities with this
subfamily.
Metarbelinae (Figs. 11.7D, 11.8F, K). Anten-
nae in both sexes bipectinate; retinaculum and
frenulum usually absent (present in some Sala-
gena and Teragra (Janse 1932)); epiphysis
roughly from middle of fore tibia; accessory cell
absent; anterior branch of M in cell absent; Rs2
to Rs4 stalked; CuP in forewing obsolete; uncus
beak-like, tip sometimes bifid (Squamura) or
wide and bilobed (Marshalliana, Fig. 11.8 F); so-
d i very small or absent; bulla present or absent;
valvae small; aedeagus tube-like; female with
short, telescopic ovipositor with broad, 8-shaped
ovipositor lobes and expanded membrane be-
tween TVII and TVIII (Fig. 11.8 K). Larva with
rugulose head which is not overlapped by pro-
thorax, with wide gap between ocelli 2 and 3;
bark-feeder, sheltering in short tunnel during day
(Gardner 1945; Roepke 1957; Holloway 1986).
We follow Minet (1986) in including the Metar-
belinae among the Cossidae primarily on the ba-
sis of their high parapatagia, whilst Schoorl
(1990) treats them as a separate family. The 8-
193
shaped ovipositor lobes are apomorphic and
provide an obvious link with the Ratardinae.
The Metarbelinae (ca. 150 species in 20 genera;
Metarbela, Salagena, Indarbela, Squamura)
range from Africa across Arabia to South East
Asia (alleged New World Arbela species are Hy-
poptinae).
Ratardinae (Figs. 11.7 E, 11.9). Delicate but-
terfly-like moths, wingspan 4 3 - 5 4 mm; ocelli
absent; antennae filiform or bipectinate, short,
up to 1/3 length of forewing; labial palps short;
parapatagia rudimentary, patagia probably atro-
phied; epiphysis reduced or absent; tibial spurs
0-2-2; wings rounded and of almost equal area;
fore wing with chorda absent, R separate or
stalked with Rsl or absent (in male of Ratarda
excellens R + Rsl stalked but R vestigial only
(Owada 1993)), stalk Rs2 + [Rs3 + Rs4] present;
M in cell present (but only one branch) in Suma-
tratarda, absent in Ratarda and Callosiope, Ml
separate or shortly stalked with radiais, M2 dis-
placed towards M3, CuP weak or absent; hind-
wing with Se + R connected with Rs by cross
vein at base of cell, M in cell present or absent,
posterior edge of cell protruding, Ml separate,
M2 displaced towards M3, CuP obsolete or ab-
sent; frenulum and retinaculum present or ab-
sent; male (Ratarda excellens) (Fig. 11.9) with
uncus fused with tegumen, beak-like, apex bifid;
tegumen and vinculum fused; gnathos present,
bulla a delicate bridge; sacculus ending in a free
recurved process; juxta simple; aedeagus tube-
like, cornuti present; female with short, tele-
scopic ovipositor with membrane between TVII
and TVIII extended, with broad, 8-shaped ovi-
positor lobes. The recent description of male Su-
matratarda (Kobes & Ronkay 1990) and the re-
cognition of Shisa as a junior synonym of Ra-
tarda (Owada 1993) allows reconsideration of the
taxonomic position of the group based on male
genitalic structure. The slender butterfly-like
build with the rounded wings is highly apomor-
phic, and Ratarda was originally described in Zy-
gaenidae and later assigned family status by
Hampson (1898). Included in Cossoidea, Bom-
bycoidea or Calliduloidea by different authors
(for complete references see Minet (1987)) the
group's status as a subfamily within the Cossidae
(as in Janse (1932)) is based on the similarity
with Metarbelinae in the genitalia of both sexes,
in particular the apomorphic 8-shaped oviposi-
tor lobes. The Ratardinae comprise the above
mentioned 3 genera with 10 species ranging from
India across South East Asia to Taiwan. Reviews
of Ratardinae are provided by Hering (1926),
Holloway (1986), Heppner & Wang (1987),
Kobes & Ronkay (1990) and Owada (1993), but
their biology and immature stages are unknown.
The adults are probably day-flyers but have also
been taken at light.
Hypoptinae (Figs. 11.7 F, G, 11.8 G, L). Anten-
nae shortly bipectinate in both sexes; tibial spurs
194 Edward D. Edwards, Patricia Gentiii, Marianne Horak, Niels P. Kristensen & Ebbe S. Nielsen
0-2-4 or 0-2-2; accessory cell small or absent, dis-
tally never protruding beyond cell; hindwing
often with CuP and 1 A + 2 A distally linked by
crossvein or partially fused; uncus beak-like (Fig.
11.8G), slender at base; bulla rarely present;
juxta ventrally invaginated in an apdeme, with-
out digitiform processes; aedeagus tube-like; fe-
male with short, non-telescopic ovipositor with
triangular ovipositor lobes (Fig. 11.8 L); TIX
usually a distinctive sclerite; ostium in emargina-
tion of SVII. Schoorl (1990) characterizes the
Hypoptinae by a configuration of the 3rd and
4th axillary sclerites in the hindwing unique
within the Cossidae, and this isolated position is
supported by the unusual female genitalia and
the shape of the juxta. The Hypoptinae (ca. 170
species in 20 genera; Hypopta, Givira, Langsdor-
f s ) are restricted to the New World and are most
diversified in the subtropical xerophytic areas of
Central and South America. Acousmaticus may
belong here.
Phylogenetic aspects are discussed by Kuznet-
sov & Stekolnikov (1981), Minet (1986, 1991)
and Schoorl (1990; extensive study based on ex-
ternal adult morphology, mainly thorax and
wing base structures, with a comprehensive bibli-
ography of the group). Principal taxonomic
works for Palearctic region: Daniel (1955-1965),
Wiltshire (1990); Afrotropical region: Janse
(1925) (Metarbelinae), Viette (1951), Clench
(1959); Oriental region: Arora (1976), Roepke
(1955 (New Guinea), 1957); Australian region:
Turner (1902, 1945); Nearctic region: Barnes &
McDunnough (1911); Neotropical region: Ureta
(1957), Clench (1957), Gentili (1989). Morpho-
logical and biological information, especially of
pest species, is found in Postner (1978), Zagul-
yaev (1978) and Skinner (1985) for Europe, Pin-
hey (1975) for Africa, Forbes (1923) for North
America, Inoue et al. (1982) for Japan, Holloway
(1986) for Borneo, Kobes & Ronkay (1990) for
Sumatra, Common (1990) for Australia and Hol-
loway, Bradley & Carter (1987) worldwide. Stehr
(1987) summarises larval morphology for North
American species, Gardner (1945) describes im-
mature stages of Indian cossids, Common & Ed-
wards (1981) compare castniid and cossid imma-
tures, and Mosher (1916) includes cossid pupae
in her comparative study.
Dudgeoneidae. Adults (Fig. 11.7H) medium-
sized, wing-span 2 8 - 7 2 mm. Cephalic vestiture
of short, erect, lamellar scales, frons with tuft of
forwardly directed scales; a pair of small asym-
metrical pits on lower part of frontoclypeus; an-
tennae about half length of fore wing, unipecti-
nate or rarely bipectinate to tip in both sexes,
with intercalary sclerite present; pilifers promi-
nent; proboscis absent, maxillary palps scaled, 3-
segmented; labial palps 3-segmented, upturned,
reaching to base of antennae, medial segment the
longest, apical segment elongated, vom Rath's
organ shallow. Thoracic vestiture of appressed
scales, with a bifid crest on posterior part of
metathorax; pronotum a slender bar; parapa-
tagia not developed. Epiphysis present, long, ap-
prox. 1/2 length of fore tibia, inserted in middle
of fore tibia; hind tibia with dorsal crest of long
lamellar scales; tibial spurs 0-2-4, long, asymmet-
rical; tarsal spines reduced; pretarsus complete;
ungues well developed; pulvilli present, slender;
arolium present; pseudempodium absent. Fore
wing (Fig. 11.8 B) reddish brown with circular,
silvery white spots; chorda short, strong, acces-
sory cell distally slightly protruding beyond cell;
R from well before accessory cell, radial sectors
separate or Rs2 + Rs3 short-stalked or Rs3 +
Rs4 connate; M forked in cell, both branches
well-developed, posterior branch ending between
M2 and M3; Ml displaced towards radial veins;
CuP well developed; 1 A + 2 A separate from
CuP. In hind wing (Fig. 11.8 B) frenulum in male
a single strong bristle, in female several finer
bristles; Se + R separate from Rs; Rs and Ml
connate or shortly stalked; M present and forked
in cell, anterior branch often weak; CuP present;
1 A + 2 A shortly forked at base; 3 A present.
Abdomen with paired simple tympanal organs
on SII (Fig. 11.10 A, B) (Minet 1983). Male geni-
talia (Fig. 11.8 C) with uncus prominent, beak-
like, wide at base, fused with tegumen; socii ab-
sent; gnathos well-developed, branches distally
fused in a prominent pointed bulla; vinculum
and tegumen fused; saccus a digitiform invagi-
nation; valvae short and rounded, ear-like, sim-
ple without processes; juxta a bilobed plate with-
out digitiform processes; aedeagus tube-like,
large, coecum present. In female genitalia (Fig.
11.8H) ovipositor short, telescopic, with slightly
sclerotized, setose, non-piercing ovipositor lobes;
apophyses anteriores as long as apophyses pos-
teriores and their ventral arm connected with tri-
angular sclerotization posterior to ostium bur-
sae; ductus bursae short and almost undifferenti-
ated; corpus bursae elongated, without signa.
The morphology of egg, larva and pupa is un-
known, but Turner (1902) briefly described the
biology of Dudgeonea actinias. The larvae are
stem-borers which excavate tunnels to the out-
side bark prior to pupation and the pupae are
protruded from the larval tunnels through the
remaining bark before eclosion. The only known
host plant is Canthium attenuatum (Common
1990).
The monophyly of the Dudgeoneidae, con-
sisting of the genus Dudgeonea with 6 described
species from Africa, Madagascar, India, South
East Asia, New Guinea and northern Australia,
is undisputed, based on the presence of an abdo-
minal tympanal organ (Fig. 11.10 A) figured in
detail by Minet (1983), and also the sclerotized
ovipositor lobes. Dudgeonea was originally de-
scribed as a cossid and Berger (1957) gave the
group family status. Brock (1971) provisionally
The Cossoid/Sesioid Assemblage
transferred the family to the Sesioidea, based on
unspecified similarities, and Minet (1983) placed
it in the Pyraloidea on the strength of the tympa-
nal organs (Minet 1983). Given the numerous
similarities in the morphology of the adult and
the very similar biology Common (1990) in-
cluded the family again in the Cossoidea. Minet
(1991) subsequently confirmed this assignment
on the basis of two synapomorphies linking Cos-
sidae and Dudgeoneidae, namely absent or
strongly reduced proboscis and a pair of small
pits in the lower part of the frontoclypeus. In
Dudgeonea the pits are asymmetrical and rela-
tively close to each other, as in Xyleutes and the
Australian Endoxyla (Fig. 11.IOC, D). Further
work will be needed to elucidate whether these
frontoclypeal pits are a cossoid apomorphy or
characterise only a subordinate group within the
superfamily. The structure of the female genitalia
indeed suggests a closer relationship with the
Cossulinae and Austrocossus, and apart from the
tympanal organs the group would have all the
characteristics to be considered a cossid subfam-
ily. However, dudgeoneids have retained pilifers
which have so far not been reported from cos-
sids. Archaeoses from Australia (with pilifers)
and Acritocera from Fiji (with tympanal organs,
Fig. 11.10B) might belong to the dudgeoneids.
Acknowledgements
NPK thanks Dr. M. Scoble, The Natural History
Museum, London, for information, Prof. Dr. C.
Naumann, Zoologisches Forschungsinstitut und
Museum Alexander Koenig, Bonn, for con-
structive criticism and Mr. G. Brovad, Zoologi-
cal Museum Copenhagen for photography.
PG und wish to thank the following col-
leagues for information, material, photos, advice
and valuable comments on the manuscript: Dr.
D. R. Davis, Smithsonian Institution, Washing-
ton; Mr. E. D. Edwards and Dr. E. S. Nielsen,
CSIRO, Canberra; Dr. J. D. Holloway, Interna-
tional Institute of Entomology, London: Dr. J.
Minet, Museum National d'Histoire Naturelle,
Paris; Dr. M. Owada, National Science Museum,
Tokyo. Illustrations were provided by Mr. J. P.
Green (photography), Mrs. E. Alyta (printing),
Mrs. H. Geier (SEM) and Mr. C. Davies (pro-
duction of plates), all CSIRO Division of Ento-
mology, Canberra. Melbourne University Press
kindly gave permission for reproduction of fig-
ures from 'The Insects of Australia'.
EDE & ESN thank Drs. M. Horak and M.
Matthews, CSIRO Divison of Entomology, Can-
berra for comments on drafts the castniid manu-
script and Mr. D. Carter and the Photographic
Unit at The Natural History Museum, London
for photographs of Tascina. Mr. J. Green and
Ms. K. Smith, CSIRO Division of Entomology
prepared the other photographs.
195
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nac. Chile 27: 129-153.
Viette, P. (1951): Contribution l'tude des Cossidae
(premire note). Les Cossidae de Madagascar
(Lpidoptres). - Naturaliste Malgache 3: 133
138.
Wiltshire, . P. (1990): An illustrated, annotated cata-
logue of the Macro-Heterocera of Saudi Arabia. -
Fauna of Saudi Arabia 11: 91250.
Zagulajev, A. K. (1978): 19. Family Cossidae - Car-
penter moths, pp. 177-186. In: Medvedev, G. S.
(ed.), Keys to the Insects of the European part of the
USSR. Vol. IV, part 1. 710 pp. Nauka, Leningrad.
[Translated from Russian for U. S. Department of
Agriculture, Amerind Publishing, New Delhi, 1987.]
 12. The Tortricoidea
Marianne Horak
The Tortricoidea are a monophyletic and homo-
geneous group containing the single family Tor-
tricidae. Cochylini, Chlidanotinae and Olethreu-
tinae previously have been treated as distinct
families but are now generally regarded as subor-
dinate taxa within the Tortricidae (R. L. Brown
1987; Razowski 1987 a; Common 1990; Horak &
Brown 1991).
Tortricidae. Small to medium-sized moths, wing-
span usually from 7 to 35 mm, rarely up to
60 mm (Ceracini). Head (Figs. 12.2, 12.3): Vesti-
ture of erect lamellar scales on vertex (except few
day-flying taxa with appressed scales), usually
with large tuft protruding anteriorly between an-
tennae, frons with longish scales oriented down-
wardly in upper half, with short, appressed, up-
wardly oriented scales on lower half; postin-
terocular suture usually absent (present in chli-
danotine Trymalitis climacias); frontoclypeus
without vertical bulge between tentorial pits;
ocelli present (reduced or obsolete in most
Schoenotenini); chaetosemata well-developed;
antenna of middle of forewing or slightly be-
yond, nearly always filiform, often slightly ser-
rate or dentate but very rarely pectinate; small
intercalary sclerite present; each flagellar seg-
ment with two rings of scales (the rings often in-
complete) in plesiomorphic condition (Figs.
12.3 A, C, D) (most Tortricinae, Chlidanotinae),
reduced to one in Olethreutinae (Fig. 12.3 B) and
some Sparganothini; sensory setae ('cilia' in tax-
onomic literature) from short and evenly distrib-
uted to partially very long, especially in male;
base of flagellum in male sometimes notched or
expanded (Epitymbiini, Archipini, Eucosmini);
pilifers present (Fig. 12.2 B); proboscis well-de-
veloped, unsealed at base, maxillary palpi small,
ranging from 4 scaled segments to a naked un-
segmented remnant; labial palpi well-developed,
3-segmented, porrect to ascending, with second
segment sinuate, third segment elongate in por-
rect palpi and usually short and blunt in up-
turned palpi, and a vom Rath's organ with the
opening usually subapical. Thorax. Vestiture of
either appressed scales or with a single or bifid
dorsal posterior crest of raised scales; prothorax
with a pair of flap-like scaled patagia, pronotum
not developed into parapatagia; mesophragma
at most weakly bilobed; metafurcal stem long.
Legs: Epiphysis present in middle of fore tibia;
tibial spurs 0-2-4; mid tibia nearly always with
modified scaling of a spiny appearance (Fig.
12.4 A); expandable hair-pencil in several
olethreutine groups on the hind tibia (Fig.
12.4 B), in many Euliini, Schoenotenini and Atte-
riini on the fore femur; tarsal spines usually pre-
sent ventrally towards apex of segments 14, of
some taxonomic significance (Yasuda 1972); pre-
tarsus complete, with two simple claws, a median
broad arolium, a bristle-like pseudempodium
and two setose pulvilli. Fore wing (Fig. 12.6):
Relatively broad, subtriangular to subrectangu-
lar; costa in some groups strongly convex be-
yond base and in male folded over the wing con-
cealing specialized scent scales (costal fold, Fig.
12.4C, D); pterostigma normally absent; micro-
trichia present only on small area beneath base
of dorsum; retinaculum a small membranous
hook from Sc in male; frequent wing pattern ele-
ments are derivations from five dark oblique fas-
ciae (numerous Tortricinae) and costal strigula-
tion (most Olethreutinae, many Chlidanotini and
Phricanthini); in plesiomorphic condition all
veins present and separate beyond distally closed
discal cell with Rs4 running to termen, and
chorda and unforked M present in cell (Anacru-
sis, many Olethreutinae (Fig. 12.6 B)) (forked in
pupal integument of some New Zealand Archi-
pini (Dugdale 1966) and reportedly forked in
some adult Ceracini (Horak 1984 a)); more de-
rived tortricid forewings without chorda or M
within cell, and Rsi + Rs4 or more rarely Rs2 +
(RS3 + Rs4) and/or M 3 + CuAi stalked; CuP
usually present as tubular vein at least distally
(absent in Cochylini); 1 A + 2 A with large basal
fork; vestiture usually of appressed scales, but
raised scales in most Polyorthini, some raised
tufts in most Phricanthini and some Tortricini,
and in many Schoenotenini veins fringed with
partially raised scales. Hind wing (Fig. 12.6): As
broad as fore wing; frenulum a single large com-
posite bristle in male, usually 3 finer bristles in
female; anal margin in male often modified,
sometimes forming a roll containing specialized,
presumably scent-producing scales (many
Olethreutinae), often associated with expandable
hair-pencils on hind tibia; sex scales sometimes
present as brands on wing surface or within
pockets; cubital pecten often present and con-
sisting of a row or tuft of hairscales on or just
behind basal portion of CuA (most Olethreuti-
nae (Fig. 12.6 B), very few Tortricinae); recurrent
vein rarely present at base of costa; M never pre-
sent in cell and cell closed by only vestigial cross-
veins; in most plesiomorphic condition fused Sc
+ Ri connected near base by cross-vein Ri with
Rs, but base of Rs frequently obsolete or very
rarely (some Sparganothini) stalked with Sc +
Ri; Rs and Mi normally closely approximated
200 Marianne Horak

Fig. 12.1. Tortricinae ( A - J ) , C h l i d a n o t i n a e ( M ) a n d Olethreutinae (T) (all males except H): A, Phricanthes
asperona (Phricanthini), B, Acleris rhombana (Tortricini), C, Epitrichosma neurobapta (Schoenotenini), D, Aethes
smeathmanniana (Cochylini), E, Archips podana (Archipini), F, Epitymbia alaudana (Epitymbiini), G , Sparganothis
pilleriana (Sparganothini), H, Anacrusis stapiana, female (Atteriini), I, Eulia ministrana (Euliini), J, Cerace xantho-
cosma (Ceracini), K, Polylopha epidesma (Polyorthini), L, Trymalitis sp., Australia (Chlidanotini), M, Thaumato-
grapha excellens (Hilarographini), N, Cryptaspasma sordida (Microcorsini), O, Gatesclarkeana sp., Sulawesi
(Gatesclarkeanini), P, Bactra verutana (Bactrini), Q, Sorolopha cyclotoma (Olethreutini), R, Anathamna sp., New
G u i n e a (Enarmoniini), S, Crocidosema plebejana (Eucosmini), T, Cydia pomonella (Grapholitini) (all except R:
H o r a k & B r o w n 1991: figs. 1.2.1, 1.2.3). Scale b a r = 2 m m .

and parallel at base or often stalked, but widely
separate in few unrelated taxa considered rela-
tively underived within their group; plesiomor-
phically all veins separate beyond discal cell, but
M3+ CuAi frequently stalked and rarely entirely
fused; CuP usually present at least distally (ab-
sent in Cochylini); 1 A + 2 A usually with a small
basal loop; 3 A usually present. Abdomen: Un-
spined dorsally; S2 apodemes tortricoid; segment
8 in males often with variously modified scaling
and/or coremata, infrequently elsewhere on ab-
domen; segment 7 in females sometimes with
The Tortricoidea
Fig. 12.2. Tortricid head structures: A, lateral view of
head of Cryptaspasma sordida, , denuded head with
mouthparts of Cydia pomonella (B: Horak 1991: fig.
1.1.5a).
Fig. 12.3. Tortricid antennae: A, tortricine antenna
with two rings of scales per segment (Epiphyas postvit-
tana, B, olethreutine antenna with one ring of scales
per segment and basal notch (Eucosmini), C and D,
antenna of Trymalitis sp., Sulawesi (Chlidanotini),
basal region (C) and distal region (D).
dense, deciduous scaling (corethrogyne) (Atteri-
ini, few Archipini); pairs of dorsal pits sometimes
present on tergum 2 and sometimes also on ter-
gum 3; female pheromone gland a dorsal invagi-
nation between segments 8 and 9. Male genitalia
(Fig. 12.5): Base of tegumen (pedunculi) some-
times with medially projecting attachment for
tergal flexors of valva (nu); uncus varying from
well-developed, often with a ventral brush below
apex (Archipini, Epitymbiini, Ceracini), to ab-
sent; socii large hairy or scaled lobes in unde-
rived condition, often small or absent; underived
gnathos with arms distally fused, but frequently
201
reduced or absent; hami (additional paired pro-
cesses from tegumen) rarely present (some
Schoenotenini, Chlidanotini and Hilarograph-
ini); anal tube sometimes sclerotized (as subs-
caphium) and fused with gnathos remnants; sac-
cus rarely developed (Phricanthini, few Chlida-
notini); underived valva simple and probably
weakly sclerotized, with a complete set of
muscles including intravalval muscles in Chlida-
notinae only, but usually one or several pairs of
muscles lost and valva variously modified; juxta
usually a simple plate, in Olethreutinae folded
and fused with anellus into functional unit;
transtilla present in Tortricinae and Chlidanoti-
nae, absent in Olethreutinae; aedeagus usually
with coecum in Tortricinae and Chlidanotinae,
without in Olethreutinae; vesica frequently with
fixed or deciduous cornuti. Female genitalia (Fig.
12.5): Ovipositor typically not telescopic, with
broad, flat, hairy ovipositor lobes and relatively
short apophyses, modified in most Cnephasiini
and some Tortricini (floricomous ovipositor);
distal part of oviduct (vagina) eversible from ovi-
pore as a complex structure (Fig. 12.4E) of un-
known function; ostium bursae usually on S8,
surrounding membrane often sclerotized, form-
ing so-called sterigma (often subdivided into la-
mella antevaginalis and lamella postvaginalis),
which in most Tortricinae, Chlidanotinae and
Microcorsini is connected to apophyses anteri-
ores (possibly underived condition), more rarely
fused with S7 (many Olethreutinae, some Epi-
tymbiini); bursa copulatrix frequently differenti-
ated into ductus bursae and corpus bursae, the
latter usually with one (most Tortricinae) or two
(most Olethreutinae) signa, rarely with numer-
ous scattered spines and/or small irregular scle-
rites (Cochylini, Euliini); ductus bursae fre-
quently with a sclerotized ring (colliculum) below
ostium and sometimes with a sclerotized ribbon
(cestum); ductus seminalis usually with a diver-
ticulum (bulla seminalis), more rarely corpus
bursae with a variously developed diverticulum
(accessory bursa).
Egg\ Of 'flat' type, with long axis horizontal
and micropyle at one end; roundish to oval,
dome-shaped, usually so flattened as to appear
scale-like; globose lemon-shaped egg of Phrican-
thini unusual for family and possibly plesiomor-
phic condition; chorion smooth, irregularly ru-
gose, or reticulate with pronounced ridges.
Larva\ Typical for generalized ditrysian larva,
with only primary and subprimary setae, three
pairs of segmented legs on thorax, ventral pro-
legs on (abdominal) segments A3 6, usually
slightly tapering towards apex, and anal prolegs
on segment AIO; crochets uniserial, on ventral
prolegs in a complete circle and on anal prolegs
in a continuous band; head usually semiprogna-
thous, shape of ocellar area reflecting feeding
mode (MacKay 1963); coronal suture nearly al-
ways short and adfrontals extending to epicra-
202 Marianne Horak

Fig. 12.4. Tortricid structures: A, typically spiny mid tibia (Epiphyas postvittana), B, hind tibial pencil of male
Olethreutini, C, forewing costal fold of Strepsicrates semicanella (Eucosmini), D, detail of sex scales in costal fold
of Crvptoptila sp. (Archipini), E, ovipositor lobes with everted distal portion of vagina of E. postvittana (A, C:
Horaic 1991: figs. 1.1.8b, 1.1.10a).

Fig. 12.5. Genitalia structures of Tortricinae (A, B), Chlidanotinae (D, E, F) and Olethreutinae (C, G): A, Acropol-
itis xuthobapta, male (Archipini), B, A. xuthobapta, female, C, Oxysemaphora sp., female (Olethreutini), D, Trymal-
itis climacias, female (Chlidanotini), E and F, Polylopha epidesma, male genitalia (E) and segment 8 with hair
pencils (F) (Polyorthini), G, Crocidosema plebejana, male (Eucosmini) (D: Common 1965: fig. 26 B).

nial notch or nearly so; six stemmata on each
side (absent in Thaumatographa), L-group on
prothorax (TI) with 3 setae (2 setae in Thauma-
tographa); SV-group on T l , 2 and 3 usually
2 : 1 : 1 ; LI and L2 adjacent on A 1 - A 8 (*); SV-
group nearly always trisetose on A3 6; SD on
A8 usually anterior, anterodorsal or antero-
ventral to the spiracle (*); A9 with both D2 on
common middorsal pinaculum and closer to
each other than to their own D1 (except Cera-
cini, some Cochylini and Tracholena) (*); A9
with D1 usually closer to SD than to D2, and
D1 and SD frequently on the same pinaculum
(*). If an anal fork is present, its shape (straight
prongs) is diagnostic for a tortricid larva and
probably a synapomorphy, albeit frequently lost,
The Tortricoidea
Fig. 12.6. Wing venation of representatives of Tortricinae (A), Olethreutinae (B) and Chlidanotinae (C), males:
A, Capnoptycha ipnitis (Epitymbiini), B, Sorolopha cyclotoma (Olethreutini), C, Polylopha epidesma (Polyorthini)
(Horak & Brown 1991: figs. 1.2.2d, 1.2.4a, b).
for at least a large subset of the family. Other-
wise, there is not one single character to identify
a tortricid larva, but most can be recognized by
a combination of four setal characters, marked
with (*) above (R. L. Brown 1987). Pupa: Adec-
ticous, obtect; abdominal segments A710 fused
in female, A 8 - 1 0 fused in male; head often with
large thorn on frons; maxillary palpi usually visi-
ble, alar furrows rarely present; two rows of dor-
sal spines present on abdominal segments A 3 - 7
(except Ceracini; few unrelated groups with cau-
dal row modified into band of scattered small
spines), with variable development on A2 and
A8-10; transverse dorsal groove or paired 'dor-
sal pits' may be present between anterior dorsal
row of spines and anterior margin of segments
A2 and A3; one or two hooked setae usually pre-
sent on each side of anus (anal rise) on AIO;
cauda usually with eight hooked setae and in the
more generalized condition with either dorsal
spines or derivations thereof, such as warts or
thorns (Cochylini, Cnephasiini, Hilarographini,
Eucosmini, Grapholitini), or produced into a
cremaster in numerous groups.
Biology. The flat tortricid ovipositor lobes, and
possibly also the eversible structure from the ovi-
pore, presumably are adaptations for a tortricid
egg-laying mode. Except for a few plesiomorphic
groups (Phricanthini, Arotrophora group, some
Cochylini) tortricid eggs are strongly flattened
and appear scale-like, and usually are deposited
exposed on the surface of the substrate. The
modified ovipositor complex could aid in flatten-
ing the eggs, though typically tortricid ovipositor
lobes are found also in Phricanthini. Eggs are
deposited singly, in small clusters or large imbri-
cated masses, to some degree reflecting tribal re-
lationships (Powell 1964, Powell & Common
1985). They are sometimes covered or fenced
with corethrogyne scales (Atteriini) or with de-
bris (Cnephasiini).
203
In accordance with the internal feeding modes
of primitive Lepidoptera, it was long assumed
that internal feeding was the ancestral feeding
mode also in tortricids (MacKay 1963, Powell
1964), corroborated by a similar biology in the
allegedly closely related cossids. However, within
the tortricids internal feeding is predominant in
some more derived tribes whilst several unrelated
groups of generalized Australian Tortricinae are
leaf-litter feeders, and a reassessment of the cos-
sid/tortricid relationship so far has merely re-
vealed symplesiomorphies. The presence of an
anal fork in the larva of some members of all
three tortricid subfamilies is a strong argument
for a non-boring ancestral larva, as most tortri-
cid borers lack this structure used to eject faeces
from the shelter. Whilst the Tortricidae as a fam-
ily use a vast array of plant families and fail to
specialize markedly on any one (Powell 1980),
there is some host plant correlation at the tribal
level. Two of the more generalized tortricine
groups show an exclusive association each with
a Gondwanan plant family: Phricanthini with
Dilleniaceae and the Australian Arotrophora
group with Proteaceae (Common 1963). The
Bactrini may be restricted to monocotyledons.
Leaf rolling larvae often pupate in a silk tied
shelter on the food plant whilst many boring lar-
vae pupate in the ground. Apart from Ceracini,
tortricid pupae have rows of dorsal spines and
are extruded from their cocoon before emer-
gence.
Adults are usually active in the early evening
and at night, but there are several brightly col-
oured day-flying groups (Ceracini, Zacorisca in
the Archipini, and possible the mimetic complex
including Atteria, comprising three genera from
three tribes (Obraztsov 1966)). Also many
Grapholitini are active during the day.
Information sources. Befitting an economically
important group a vast literature is available on
204
tortricids. Tortricid Pests (van der Geest & Even-
huis 1991) provides reviews of many aspects of
tortricid research, including extensive treatments
of tortricid morphology (Horak 1991) and tax-
onomy and systematics (Horak & Brown 1991),
with primary references to information on adult,
larval and pupal morphology. Comprehensive
studies of local faunas include: Nearctic region:
Heinrich 1923, 1926; Freeman 1958; Powell
1964; Miller 1987; Neotropical region: Powell
1986; J. W. Brown & Powell 1991; Razowski
1994; Palaearctic region: Obraztsov 1954-1968;
Hannemann 1961, 1964; Bentinck and Diako-
noff 1968; Kuznetsov 1978; Razowski 1987 a,
1989; Japan: Yasuda 1972, 1975; Kawabe 1982;
Oriental region: Diakonoff 1939, 1968, 1973; Mi-
cronesia: Clarke 1976; Hawaii: Zimmermann
1978: Phylogenetical aspects are addressed by
Powell (1964) from a biological perspective, by
Kuznetsov and Stekolnikov (1973, 1977, 1984)
and by Razowski (1976) mainly on the basis of
male genitalia musculature and by Horak
(1984 a) with an attempt at establishing a tortri-
cid groundplan. Egg morphology is described by
Peterson (1965) and, together with oviposition
pattern, by Powell (1964) for Nearctic and Pow-
ell and Common (1985) for Australian Tortrici-
nae. Swatschek (1958) and MacKay (1959, 1962,
1963) provide comprehensive treatments of tor-
tricid larval morphology of the Palaearctic and
Nearctic faunas respectively, and a more recent
summary is given by R. L. Brown (1987).
Mosher (1916) included tortricids in her compar-
ative study of lepidopterous pupae, and Patocka
(1980, 1982, 1983) describes in detail the pupae
of numerous Palaearctic olethreutines.
Classification. Though Tortricidae are easily rec-
ognizable by a combination of characters, the
only demonstrated autapomorphy for the family
is the large flat ovipositor lobes of the female
(Horak & Brown 1991; Dugdale 1988), with the
few exceptions obviously due to adaptive modifi-
cations (cnephasiine floricomous ovipositor for
covering eggs with debris). Upwardly directed
scales on the lower half of the frons (Zimmerman
1978) and the tortricid anal fork with straight
prongs may be further autapomorphies for the
family, but the anal fork is frequently absent (in
many groups with boring larvae) and has not
been found in the Phricanthini, possibly the
overall most plesiomorphic tribe. After a period
of very divergent subfamily and tribal classifica-
tions a consensus is now being reached (Razow-
ski 1987 a, 1989; Horak & Brown 1991). There
are strong arguments for the monophyly of two
of the three subfamilies, the Olethreutinae and
Chlidanotinae, whilst the Tortricinae, as pres-
ently delimited, are assumed to be paraphyletic,
comprising ancestral as well as derived groups.
The sister group of neither the Chlidanotinae nor
the Olethreutinae has been identified, but the
Marianne Horak
two groups share costal strigulae and the aedea-
gus in Olethreutinae and Polyorthini is encircled
by a ring of sclerotization in the diaphragm.
Some Tortricini, on the other hand, possess a
vertically folded juxta firmly linked with the ae-
deagus, recalling the olethreutine juxta/caulis
complex.
Key to tortricid subfamilies
1 a Antennal segments with two rings of scales
(very rarely with one ring but then always
strongly ciliate); juxta and aedeagus not
fused; sterigma usually connected with
ventral arm of anterior apophyses; cubital
pecten very rarely present 2
1 b Antennal segments with one ring of scales
and nearly always very short cilia; juxta-cau-
lis-aedeagus complex fused; sterigma usually
not connected with ventral arm of anterior
apophyses; cubital pecten usually present. .
Olethreutinae
2 a Glyphipterigoid wing pattern; very large
ocelli; short, slender, upcurved labial palpi;
antenna strongly ciliate in male
Hilarographini ( Chlidanotinae)
2 b Not with this character combination . . . . 3
3 a Antenna stout and apically flattened, cov-
ered with short, appressed scales, cilia ex-
tremely short also in male; Oriental/Austra-
lian taxa partly white with pronounced cos-
tal strigulae and with ocelli reduced
Chlidanotini (Chlidanotinae)
3 b Antenna not with these characters 4
4 a Labial palpi long, porrect; fore wing usually
with raised scale tufts and Rs4 to costa or
apex; valva very large, membranous, dorso-
longitudinally invaginated; anellus sclero-
tized dorsally to aedeagus
Polyorthini (Chlidanotinae)
4 b Not with this character combination
Tortricinae (includes Cochylini)
Tortricinae. Antenna nearly always with two
rings of scales per flagellar segment (Fig. 12.3 A),
often ciliate; forewing sometimes with costal
fold; hindwing only exceptionally with cubital
pecten; male genitalia variable, but juxta and ae-
deagus nearly always articulated (Fig. 12.5 A),
not strongly fused; aedeagus with coecum usu-
ally developed; sterigma connected with ventral
arms of anterior apophyses (Fig. 12.5 B) (con-
nection reduced in Cnephasiini, absent in few
Tortricini); usually only one signum, only excep-
tionally two or four; majority of pheromones
based on 14-carbon chain. The Tortricinae, as
presently delineated, constitute a paraphyletic
group, but there are no unequivocal synapomor-
phies to either associate particular tribes more
closely within the subfamily or refer them else-
where. Groupings into supertribes (Razowski
1981a; Kuznetsov & Stekolnikov 1984), based
The Tortricoidea
mainly on the loss of different muscles in the
male genitalia, have been largely abandoned (Ra-
zowski 1987 a). There are reasonable to convinc-
ing arguments for the monophyly of the majority
of the 11 tribes here included in the Tortricinae.
Phricanthini (Fig. 12.1 A). Adult'. Forewing
usually with tufts of raised scales, without costal
fold; pattern based on parallel transverse fasciae
and costal singulation; chorda and M not pre-
sent within cell; male abdomen with series of in-
wardly directed, curved, hair- or horn-like struc-
tures on anterior margin of third and/or fourth
terga; uncus absent; socii sclerotized and devel-
oped into a pair of hairy, beak-like posterior pro-
cesses of the tegumen; saccus usually large; sig-
num, if present, a curved dagger-shaped sclerite.
Larva: Without anal fork; SD and SD2 together
with spiracle on same enlarged pinaculum on
Al8; D l and SD on same pinaculum on A9;
SV group on A l , 2, 7, 8 and 9 is 3 : 3 : 2 : 2 : 2 .
Pupa: Alar furrow well defined, with granular
surface structure; Al anteriorly with longitudinal
median ridge; A8 with one row of dorsal spines,
A9 at most with few minute spine remnants;
cremaster irregularly subrectangular with serrate
apex, with eight weak, hooked setae; two hooked
setae on each side of anal rise. The Phricanthini
(Scolioplecta, Phricanthes) comprise 25 species in
4 genera and are centred on Australia, extending
into southeast Asia and to Madagascar. Known
larvae feed on Dilleniaceae, either in a shelter on
foliage or on living bark (Horak & Brown 1991).
The phricanthine synapomorphies (sclerotized
socii, series of inwardly projecting modified setae
on male abdomen, larva with SD, SD2 and the
spiracle on same pinaculum on Al8) were dis-
cussed by Common (1965). Diakonoff (1981)
raised the group to tribal rank and suggested
that it originated from ancestral tortricid stock
prior to its bifurcation into Tortricinae and Chli-
danotinae. The well-developed phricanthine sac-
cus, a forewing with Mi often distant from Rs,
the lemon shaped egg (Powell & Common 1985)
and the exclusive association with the archaic
Dilleniaceae are all arguments for an early deri-
vation. The so-called phricanthine socii will have
to be closely examined with a view to the genita-
lia of Denaeantha nivigera (Diakonoff 1981)
which seem to suggest a bifid hairy uncus with
lateral undifferentiated socii.
Tortricini (Fig. 12.1 B). Adult: Forewing often
with raised scales, without costal fold; chorda
only rarely present (Tortrix), M absent in cell;
M3 + CuAi often stalked; tergal flexor muscles
of the valvae (nu) absent; uncus often atrophied;
typical tortricine gnathos usually absent, but
ventral part of anal tube usually sclerotized as
subscaphium; valva usually with brachiola; juxta
usually a large, vertically folded plate; signum, if
present, usually a dentate plate, often star-
shaped. Larva: D1 and SD on A9 on separate
or on common pinacula; SD2 on separate pina-
205
culum on most or all A l - 7 ; L group on A9 tri-
setose; SV on A l , 2, 7, 8 and 9 is 3 : 3 : 2 : 2 : 2 , if
less highly unstable. Pupa: Maxillary palpi some-
times absent; spines on A2, at least cephalic row,
often absent; A8 and A9 with one row of dorsal
spines; cremaster short, truncate and curved
ventrally, multilobed or with two large ventrally
projected lateral thorns, with eight hooked setae;
usually two hooked setae on each side of anal
rise. Tortricini (Acleris, Aleimma, Tortrix), with
over 300 species in about 38 genera, are present
nearly worldwide except New Zealand and
southern South America, but are concentrated in
the Holarctic and Oriental regions (Razowski
1966, 1984, 1986). They oviposit singly and their
larvae are mostly leafrollers, mostly specialists
within species on a wide range of host plants.
The monophyly of the tribe is well demonstrated
(Obraztsov 1955), with the brachiola, the modi-
fied juxta and anal tube the most obvious apo-
morphies. Though long considered closest to the
Cnepasiini due to several symplesiomorphies no
convincing synapomorphy is known to provide
a link with any of the other tortricine tribes.
Schoenotenini (Fig. 12.1 C). Adult: Ocelli usu-
ally rudimentary; male often with hairpencil
from base of fore femur; both wings often
sparsely scaled and veins fringed with scales; cos-
tal fold only very rarely present; forewing often
with raised scales; discal cell narrow, often
curved and usually narrowed distally; chorda ab-
sent; M present at least as vestigial vein in cell,
terminating between bases of Mi and M2; uncus
variable but not with archipine-type brush of
hair beneath apex; hami sometimes present;
valva often with patches of flattened, pectini-
form bristles; sterigma usually intricate; signum,
when present, an often thorn- to sac-shaped scle-
rotized invagination, occasionally paired. Larva:
Anal fork often present; apical dorsal tarsal setae
often flattened; D1 and SD on A9 on same pi-
naculum; SD2 usually on same pinaculum as
SD on A 1 - A 7 ; SV group on A l , 2, 7, 8 and 9
from 3 : 3 : 3 : 2 : 2 to 1 : 2 : 1 : 1 : 1 ; D l , D2 and
SD on A9 often all on a common transverse
pinaculum. Pupa: With raised median ridge be-
tween antennal bases, in boring species with
large ventral thorn on frons; A8 with one or two
rows and A9 usually with only four dorsal
spines; cremaster very short with lateral angles
produced into ventrally curved thorns or cremas-
ter absent, with only two lateral thorns on AIO;
hooked caudal setae weak. The Schoenotenini
(Proselena, Tracholena, Palaeotoma, Schoeno-
tenes, Rhabdotenes, Barygnathella) comprise
about 200 species in over 20 genera and are most
diverse in the moss forests of the Papuan region,
with a small, presumably older group widely dis-
tributed from India to New Zealand. The feeding
habits are very diverse (Common 1965; Dugdale
1966), including leaf rolling, boring into various
plant tissues and tunnelling in bark. The unique
206
development of M within the fore wing cell
(Common 1965; Dugdale 1966; Horak 1984 a) is
a strong argument for the monophyly of the
Schoenotenini. After initially being accorded
family rank the group was later included in the
Chlidanotinae, based on the presence of hami in
both groups. However, hami are absent in the
more generalized Schoenotenini. J. W. Brown
(1990 c) found an expandable hairpencil from the
base of the fore leg femur in males of roughly
half the schoenotenine and euliine genera exam-
ined, and in a slight modification in nearly all
atteriine genera. As the structure is found in gen-
eralized as well as derived Euliini and in widely
divergent groups of Schoenotenini (Tracholena,
Schoenotenes group) it is a possible synapomor-
phy between these groups. J. W. Brown (1989)
provisionally transferred the Neotropical Ortho-
comotis and Paracomotis to the Schoenotenini on
the basis of shared medially extended and con-
nected chaetosemata. However, within the
Schoenotenini, this character appears to be an
apomorphy of the Schoenotenes group. It is not
present in the more generalized Schoenotenini,
and in the Australian fauna the chaetosemata are
unmodified in Epitrichosma whilst they are nar-
rowly triangular with a long medial extension,
albeit not medially connected into a band, in the
closely related Cornuticlava. Orthocomotis and
Paracomotis share none of the other apomor-
phies of the highly derived Schoenotenes group
(Ri, Rsi, Rs2 and RS3 nearly equally spaced,
veins fringed with denser scaling), nor do they
possess the characteristic schoenotenine fore
wing venation. The four schoenotenine genera
described from Madagascar have been removed
from this group (Diakonoff 1972) and the tribe
must until further evidence be considered to be
of Oriental and Australian distribution, centred
on New Guinea.
Cochylini (Fig. 12.1 D). Adult: Forewing with
termen oblique, pattern usually based on two in-
wardly oblique transverse fasciae; costal fold
sometimes present; CuP absent or strongly re-
duced, CUA2 from 2/3 cell or more distally; un-
cus often atrophied, gnathos reduced or atro-
phied; aedeagus usually very large, often with
numerous cornuti; bursa copulatrix rarely dif-
ferentiated into ductus and corpus, usually with
accessory bursa; antrum often heavily sclero-
tized; bursa with numerous spines and/or various
sclerites, never with well-defined signum. Larva:
With anal fork often present; D1 and SD on
same pinaculum on A9, but D2 setae sometimes
on two separate pinacula; SD2 at least some-
times on separate pinacula on A l - 7 ; L group
usually bisetose on A9; SV group on A l , 2, 7, 8
and 9 very variable, from 3 : 3 : 3 : 2 : 2 to
2 : 2 : 2 : 1 : 1 ; VI on A9 further apart than on A8.
Pupa: Frons sometimes with large ventral thorn;
alar furrow a shallow depression; AIO always
with several spines, often nearly encircling anal
Marianne Horak
rise; no cremaster, but two spines often enlarged
into two subdorsal thorns, and smaller thorns
often present on each side of anal rise; AIO with
weak hooked setae on apex and close to lateral
thorns, but no distinct setae on anal rise. The
Cochylini (Aethes, Cochylis, Eupoecilia, Phaloni-
dia) with over 800 species in over 55 genera are
present in all faunal regions, with high diversity
in the Palaearctic and Neotropical regions and
only two native species known from Australia.
Trachysmia, the most plesiomorphic genus, is
virtually restricted to the Holarctic. The mostly
oligophagous Cochylini usually feed internally in
flower heads, seed capsules, stalks and roots,
rarely in rolled leaves (Razowski 1970). Numer-
ous apomorphies (forewing pattern and vena-
tion, male genitalia) illustrate the monophyly of
the Cochylini, and the group has long been ac-
corded family rank. Except for the inwardly
oblique transverse fascia of the cochyline fore-
wing with its distal origin of CuA 2 all the charac-
ters used to justify family status are also found in
other Tortricidae (Horak 1984 a). The Cochylini
have been variously associated with other tortri-
cid groups on the strength of symplesiomorphies
(Horak & Brown 1991), but none of these associ-
ations is supported by a unique synapomorphy.
Cnephasiini. Adult: Maxillary palpi often four-
segmented; forewing narrow, distally expanded,
termen strongly oblique; without costal fold;
chorda present or vestigial, R in cell at most ves-
tigial, all veins separate; uncus nearly always spi-
nulose laterally; socii and gnathos well-devel-
oped; ovipositor nearly always floricomous;
ventral arms of anterior apophyses not devel-
oped, not connected with sterigma; signum, if
present, a dentate band, rarely stellate; known
pheromones 12-carbon compounds. Larva: D2
usually on separate pinaculum on Al7; SD
and D l on separate or common pinacula on A9;
L group bi- or trisetose on A9; SV group on A l ,
2, 7, 8 and 9 is 3 : 3 : 3 : 2 : 2 to 3 ( 2 ) : 3 : 2 : 1 : 1 .
Pupa: A2 usually with two, A 3 - 7 with two and
A8 and A9 with at most one row of spines; crem-
aster short with two dorsal thorns or cauda with
two large, usually dorsally curved thorns only;
smaller thorn sometimes present lateral of anal
rise; setae on AIO straight or hooked, but no
prominent setae on anal rise. The basically Hol-
arctic Cnephasiini (Cnephasia, Eana, Exapate,
Neosphaleroptera) comprise about 300 species in
about 19 genera. The Australian Arotrophora
group (Common 1963), restricted to Proteaceae,
shares only symplesiomorphies with Cnephasia
and has to be excluded from the Cnephasiini. It
might be found to deserve tribal rank once the
more plesiomorphic Archipini have been thor-
oughly studied. Female Cnephasiini collect detri-
tus with the modified ovipositor lobes, and
spread it onto the newly deposited eggs (Powell
1964). The larvae feed in spun leaves or flowers,
sometimes mining in the first instar. Relatively
The Tortricoidea
generalized members of other tribes poorly rep-
resented in the Holarctic region have long been
included in the Cnephasiini s. lat. (Horak &
Brown 1991), but in its restricted sense the tribe
is characterized by several apomorphies (spined
uncus, floricomous ovipositor, band-shaped den-
tate signum, 12-carbon pheromone) (Powell
1986).
Archipini (Figs. 12.1 E, 12.5 A, B). Adult: An-
tenna in male variably ciliate, often dentate and
exceptionally bipectinate; forewing often with
costal fold; chorda rarely present; uncus with
brush of hair below apex (Fig. 12.5 A); transtilla
often bipartite and spined above; valva usually
with pulvinus, nearly always with a sclerotized
sacculus, disc and dorsal margin often membra-
nous and variously plicate; ductus bursae often
with split colliculum, sometimes with cestum;
signum often dagger-shaped with a capitulum
(Fig. 12.5 B). Larva: Anal fork usually present;
D1 and SD on A9 on separate pinacula (except
Capua), SD2 on pinaculum of SD on Al 8; L
group trisetose on A9; SV group on A l , 2, 7, 8
and 9 is usually 3 : 3 : 3 : 2 : 2 . Pupa: A8 with one
or two rows, and A9 only rarely with a well-
developed row of dorsal spines; 'dorsal pits'
sometimes present; cremaster well-developed,
often cylindrical and longer than wide, with eight
strong hooked setae. The Archipini (Adoxophyes,
Choristoneura, Ctenopseustis, Epichoristodes,
Pandemis, Ptycholoma) with well over 500 spe-
cies in about 150 genera have a worldwide distri-
bution but are poorly represented in the Neo-
tropical region. Archipine eggs are usually de-
posited in masses of imbricate rows and the lar-
vae are mostly polyphagous leafrollers. The Ar-
chipini are presently a polyphyletic grouping,
comprising several derived lineages as well as
plesiomorphic residual taxa. The Archipini s. str.,
i. e. the genera characterized by a dagger-shaped
signum with a capitulum, consist of at least 3
distinctive lineages (Horak 1984 a): (1) Archips
and its relatives, with the most plesiomorphic
members in Africa/Madagascar, and just reach-
ing northern Australia (Homona) across the Ori-
ental region; (2) Epiphyas, Clepsis and other
members with strongly plicate valva, centred on
Australia and New Guinea; and (3) a much less
derived group with a relatively simple valva, typ-
ified by the New Zealand Planotortrix. A close
relationship has recently been recognized be-
tween Pseudargyrotoza and an Oriental/Austra-
lian group containing Drachmobola and Taeni-
archis (Yasuda & Razowski 1991), but as these
genera lack euliine or cnephasiine apomorphies
it seems best to retain them among the assem-
blage of generalized Archipini rather than ob-
scure otherwise monophyletic groupings.
Epitymbiini (Figs. 12.1 F, 12.6 A). Adult: Head
and wings often sexually dimorphic; forewing
often with costal fold in male, frequently of Ho-
mo/ja-shape in female, with strong tendency to
207
stalking of Rs2, Rs3 + Rs4 and M3 + CuAi (Fig.
12.6 A); hindwing often with Rs + Mi and M 3
+ CuAi stalked; male genitalia strongly sclero-
tized; pedunculi strong, their base often club-
shaped; uncus with brush below apex; transtilla
well developed, not spined; valva often sclero-
tized along base and dorsal margin, never with
pulvinus; sacculus, if present, centred on ventral
part of valva base, often extending deep into
disc; female genitalia often with strongly sclero-
tized, asymmetrical sterigma, never with a split
colliculum; signum diverse, sometimes paired,
never a dagger with a capitulum; known phero-
mones 14 carbon compounds with the double
bond other than in the eleventh position. Larva:
Anal fork present; pinaculum with D1 and D2
on T1 elongate, caudally extended; SD2 on same
pinaculum as SD on Al8; Dl and SD on
same pinaculum on A9; SV group on A l , 2, 7, 8
and 9 is 3 : 3 : 3 : 2 : 2 . Pupa: A8 with one or two
rows and A9 with few spines only; cremaster
flattened, rectangular to apically trilobed, with
eight strong hooked setae. The Epitymbiini (Epi-
tymbia, Meritastis, Asthenoptycha, Rhomboceros)
with well over 100 species in about 13 genera are
primarily Australian (Horak & Common 1985),
with only a small group extending through New
Guinea to Thailand. The Australian Epitymbiini
appear all to feed on dead leaves, nearly always
eucalypt leaf litter. A recognizably monophyletic
group (genitalia, pheromones) including Epitym-
bia is at the core of the tribe, but the delineation
between Epitymbiini and some other leaf lit-
terfeeding generalized Australian tortricines has
yet to be established.
Sparganothini (Fig. 12.1 G). Adult: Labial
palpi porrect, usually long to extremely long; an-
tenna ciliate, fasciculate to pectinate, scaling
sometimes only one ring per flagellar segment;
ocellus rarely absent; male often with very large,
hood-like frontal tuft; chorda always absent; M
in cell, if present, usually to between Mi and M2;
cubital pecten usually present; (Se + Ri) + Rs
often stalked; uncus usually long and very slen-
der, socii usually scaled, with free broad caudal
lobe (except Niasoma) and free apically slender
ventral lobe; gnathos arms usually atrophied;
valva with pulvinus (except Niasoma), sterigma
a finely scobinate flat pocket with a cup-shaped
depression in lamella postvaginalis, unless
strongly derived; collicum present; signum very
variable, often a sclerotized crease. Larva: Anal
fork usually present; head with PI closer to P2
than to Adf2; D1 on separate pinaculum on A9;
SD2 on Al8 on same pinaculum as SD; L
group trisetose on A9; SV group on Al, 2, 7, 8
and 9 is 3 : 3 : 3 : 2 : 2 . Pupa: Often with median
ridge between antennal bases; labial palpi usually
long; A8 with one or two rows and A9 and AIO
without spines; transverse groove on A2 and A3
between segment margin and anterior row of
spines sometimes with 'dorsal pits'; cremaster
208
well-developed, usually long and with eight
hooked setae. The Sparganothini (Amorbia,
Platynota, Sparganothis, Sparganothoides, Spar-
ganopseustis) with about 130 species in 14 genera
are nearly all confined to the New World (Lam-
bert 1950, Powell 1986), with only few species of
Sparganothis from the Palaearctic region and the
monotypic Lambertiodes from India. Sparga-
nothini deposit their eggs in imbricated groups,
and the majority are polyphagous leafrollers.
The monophyly of the core group is well demon-
strated, but the delineation and affinities of the
tribe are still controversial. The modified socii/
gnathos complex seems to consist of a reduction
of the gnathos and parallel modification of the
socii and not of a primary fusion of the two parts
(Horak 1984 a), and it is now doubtful whether
Heterochorista from New Guinea should be in-
cluded in the tribe (Horak 1984 b). Powell (1985)
demonstrates that Niasoma and some related
genera are merely very specialized Sparganoth-
ini. Association of the Sparganothini with either
Atteriini or Cochylini are based on plesiomor-
phies and not supported by derived characters
(Horak & Brown 1991).
Atteriini (Fig. 12.1 H). Adult: Labial palpi usu-
ally short to moderate, rather slender, ascending
but weakly curved, terminal segment very small;
antenna in male usually strongly ciliate, fascicu-
late to bipectinate, scaling sometimes only one
ring per flagellar segment; ocellus sometimes re-
duced; male fore leg usually with hairpencil from
base of femur; forewing in females of some gen-
era of Homona-shape; chorda and M present or
residual in cell; cubital pecten absent; M2 and M3
closely approximated at base; sterna of segments
6 and 7 in female with corethrogyne scaling: S6
with dense, fine scales, S7 with bundles of broad,
distally bent scales; uncus usually apically di-
lated; socii large, not produced posteriorly be-
yond basal attachment; gnathos well-sclerotized
and arms apically joined; valva with sclerotized
dorsal margin and sacculus, nearly always with-
out pulvinus; sterigma a membranous, finely
scobinate flat pocket; signum a flat horn or hol-
low keel, often associated with a membranous,
capitulum-like structure. Larva: Anal fork pre-
sent; D1 and SD on separate pinacula on A9;
SD2 on same pinaculum as SD on Al8; SV
group on A l , 2, 7, 8 and 9 is 3 : 3 : 3 : 2 : 2 ; VI
setae much further apart on A9 than on A8 or
A7. Pupw. With wide median keel between anten-
nal bases; with a deep transverse dorsal groove
on A2 and A3 between segment margin and an-
terior row of spines; A9 with an indistinct row
of spines; cremaster long, cylindrical, apically
rounded, with eight strong, hooked setae. The
Atteriini (Atteria, Anacrusis, Templemania, Ti-
nacrucis, Sisurcana), comprising about 40 species
in 8 genera, are largely restricted to the Neotrop-
ical region, reaching Arizona (Powell 1986). Pru-
nus is the only host recorded, but several genera
Marianne Horak
have been reared on synthetic diet, suggesting
polyphagy (J. A. Powell, pers. comm., 1986).
Many atteriine features are plesiomorphies, but
the modified abdominal scales used by females
to erect a fence around their large, imbricated
egg masses is an obvious apomorphy (Powell
1976, 1986). The delineation of the group has
long been obscured by the aberrant wing pattern
of Atteria, linking an assemblage of unrelated
but presumably mimetic tortricids from three
tribes (Obraztsov 1966). Atteriini and Sparga-
nothini have been closely associated by several
authors, but venation and genitalia provide
strong arguments against monophyly of the two
groups (Horak 1984 a, b). The expandable hair
pencil from the base of the male femur in Atteri-
ini is very similar to that found in numerous
schoenotenine and euliine genera, suggesting a
possible synapomorphy (J. W. Brown 1990 c).
Euliini (Fig. 12.1). Adult. Labial palpi moder-
ate, porrect or ascending; male fore femur fre-
quently with expandable hair pencil from base;
forewing without costal fold; M absent or re-
duced to trace within cell, chorda sometimes pre-
sent; all veins separate; cubital pecten exception-
ally present (some Orthocomotis)\ uncus well de-
veloped, without ventral brush of hair; gnathos
well developed, arms joined apically at least by
membranous band; valva without differentiated
pulvinus; cornuti usually non-deciduous; bursa
copulatrix often not differentiated into ductus
and corpus, rarely with a well-defined signum,
bursa usually lined with spiculae. Larva: D1 and
SD on separate pinacula on A9; SV group on
A l , 2, 7, 8 and 9 is 3 : 3 : 3 : 2 : 2 ; VI setae on A9
not further apart than on A8. Pupa: Without
spines or thorns on AIO, with long, smooth
cremaster with eight hooked setae. The Euliini
(.Eulia, Proeulia, Anopina), comprising 230 spe-
cies in about 47 genera, are nearly restricted to
the Neotropical region (Powell 1986, J. W.
Brown & Powell 1991), with only a few Nearctic
species and the single Holarctic Eulia ministrana.
Eggs are deposited singly or in small groups. The
larvae of E. ministrana and Proeulia are polypha-
gous leafrollers and those of Anopina are be-
lieved to be leaf litter feeders (J. W. Brown &
Powell 1991). Presence of an expandable hair
pencil from the base of the male femur in 48%
of 42 euliine genera (J. W. Brown 1990 c), in gen-
eralized as well as highly derived groups, indi-
cates that the Euliini are indeed a monophyletic
group. This character is shared with the Schoe-
notenini, and, in a modified version, with the At-
teriini, suggesting possible synapomorphies. The
inclusion of a group including Pseudargyrotoza
and Drachmobola in the Euliini (Yasuda & Ra-
zowski 1991) seems questionable, given the dif-
ferences in larval morphology between Pseudarg-
yrotoza and Eulia (Swatschek 1958). No hair
pencil is present on the fore leg femur in males
of Australian species of Drachmobola and the
The Tortricoidea
closely related genera Taeniarchis and Protop-
terna.
Ceracini (Fig. 12.1 J). Adult: Large (up to
60 mm), brightly coloured, day-flying; head
small, smoothly scaled; labial palpi short, subas-
cending; antenna fasciculate-ciliate in male; hind
tarsal segments with apical row of bristles; fore-
wing elongate, often with notch at apex, without
costal fold; red-brown to blackish, dotted with
numerous usually yellow spots; chorda usually
present, M present (rarely forked) or absent in
cell; all veins separate; hindwing with dark mac-
ulation; M2 distant from M3, Rs and Mi and M3
and CuAi at most connate. Uncus with brush of
hair below apex; socii large; valva without pulvi-
nus; transtilla band-like or absent; ostium fun-
nel-shaped, sclerotized and fused with S7, only
lamella postvaginalis connected with ventral
arms of anterior apophyses; signum, if present, a
serrate plate. Larva: Anal fork present; D setae
and SD setae arranged more or less vertically on
T2 and T3; SD2 on Al8 either on separate pi-
naculum or on SD pinaculum; DI, SD and
each of the D2 setae on separate pinacula on A9;
SV group on A l , 2, 7, 8 and 9 is 3 : 3 : 3 : 2 : 2 .
Pupa: Without dorsal spines and not extruded
at emergence; cremaster long, with knife-shaped
setae. The Ceracini (Bathypluta, Cerace, Eury-
doxa), comprising about 30 species in 4 genera,
are Himalaya-centred, reaching Japan and Su-
lawesi. They deposit egg masses and their larvae
are leaf rollers. Several unique apomorphies
(pupa without dorsal spines and not extruded,
arrangement of tarsal setae, ostium not con-
nected with anterior apophyses) demonstrate the
monophyly of the Ceracini. The dense hair cover
of the ceracine valva, absent from its more mem-
branous basal part, is reminiscent of a general-
ized eucosmine valva. Further comparative study
is needed to decide whether this is only a superfi-
cial similarity. The reduced connection between
lamella antevaginalis and anterior apophyses
and suggestions of a forked M within the fore-
wing cell of some Ceracini (Horak 1984 a) are
also of interest in this context.
Chlidanotinae. Forewing without costal fold;
hindwing without cubital pecten, with M2 distant
from M3 at base (Fig. 12.6 C); vinculum wide
and firmly fused to tegumen; valva with intraval-
val muscles present and at least in some taxa of
each tribe with a dorso-longitudinal invagi-
nation, containing hair-pencil arising from
eighth segment (Figs. 12.5 E, F); female in at
least some taxa of each tribe with an accessory
bursa arising from near a signum shaped like a
bunch of spines (Fig. 12.5 D). The monophyly of
the Chlidanotinae is well demonstrated by
unique synapomorphies in both male and female
genitalia (Diakonoff 1977 b; Tuck 1981). The Po-
lyorthini seem the overall most generalized chli-
danotine tribe (venation, well-developed
gnathos, female genitalia of Neotropical taxa), in
209
accordance with their apparent southern-dis-
junct distribution. Chlidanotini and Hilaro-
graphini share pronounced costal singulation,
presence of hami and very similar female geni-
talia.
Polyorthini (Figs. 12.1 K, 12.5E, F, 12.6C).
Adult: Labial palpi usually porrect, long. Fore-
wing without costal fold, usually with raised
tufts of scales (AclerisAi)-, chorda and M pre-
sent or absent in cell; R.S4 nearly always to costa
or apex; hindwing often with Rs and Mi distant
at base, with M2 widely separate and parallel to
M3 (Fig. 12.6C); valva large and membranous,
dorso-longitudinally invaginated, containing
hairpencil arising from eighth segment (Fig.
12.5 E, F); anellus sclerotized dorsally to aedea-
gus, joined with aedeagus and laterally with
valva. Larva: Anal fork present; SD and SD2
on Al 7 on same or separate pinacula; D1 and
SD on A9 on separate pinacula or close to-
gether and with D2 setae on one large transverse
pinaculum; L group trisetose on A9. Pupa: Last
three segments lengthened in male; cremaster
short, with straight apex and hooked caudal se-
tae. The Polyorthini (Polyortha), comprising
about 120 species in 18 genera, have two distri-
bution centres, the Neotropical and Oriental/
Australian region, which are linked by closely re-
lated genera in South America (Lypothora) and
Australia (Polylopha). Two isolated genera occur
in the Palaearctic region (Isotrias, Olindia).
Rearing records are scarce and feeding modes di-
verse, including leaf rolling and boring (Horak &
Brown 1991). The monophyly of the Polyorthini
is well demonstrated (valva and anellus struc-
ture), and apart from the brightly coloured
Pseudatteria its members are characteristic by
their Acleris-like appearance (Common 1963;
Dugdale 1966; Diakonoff 1974; Razowski 1979,
1981 b). The female signum ist very diverse; pos-
sibly structures like the cestum and the signum
found in the plesiomorphic Pseudatteria each
gave rise to the signum of different polyorthine
groups.
Chlidanotini (Figs. 12.1 L, 12.3 C, D, 12.5 D).
Adult: Antenna stout and apically flattened, en-
tirely covered with short appressed scales (Figs.
12.3 C, D), cilia extremely short even in male;
labial palpi long and porrect in Neotropical gen-
era, shorter and ascending in Oriental/Australian
taxa, second segment with loose scales beneath;
ocelli often reduced or absent; chaetosemata
small; forewing without costal fold, with costal
strigulation; chorda absent, M present or absent
within cell; CuP often vestigial; hindwing with
Rs + Mi stalked and M2 and M3 widely separate
at base; male genitalia usually with hami which
may be fused with socii; transtilla well devel-
oped; valva sometimes with dorsal invagination
and corresponding hairpencil; female genitalia
with accessory bursa usually from near signum
(Fig. 12.5 D); signum a bunch of spines, some-
210
times strongly reduced. Larva and pupa un-
known. The Chlidanotini (Trymalitis, Pseu-
docomotis), comprising about 60 species in over
20 genera, are known from the Neotropical and
Oriental/Australian regions, with an endemic ge-
nus from New Caledonia. There is evidence that
the larvae are internal feeders (Horak & Brown
1991). The recent discovery of overall general-
ized Chlidanotini in the Neotropical region (Ra-
zowski 1987 b, J. W. Brown 1990 a) changes the
concept of the tribe which had been based on the
highly derived Oriental/Australian genera with
strongly stalked or even fused wing veins (Com-
mon 1965, Tuck 1981). Auratonota (Razowski
1987 b, J. W. Brown 1990 b) combines a hilaro-
graphine wing pattern with antennae and palpi
typical for a Chlidanotini. The genitalia in both
sexes of the more plesiomorphic Neotropical
taxa are virtually identical between the two
tribes, leaving only the modified, distally com-
pressed antenna as a possible autapomorphy for
the Chlidanotini. This suggests that the Hilaro-
graphini could be a subordinate group within the
Chlidanotini.
Hilarographini (Fig. 12.1 M). Adult: Antenna
with long cilia in male; labial palpi short to mod-
erate, slender, curved and ascending; ocellus very
large; forewing apically dilated, brightly col-
oured with metallic markings and long costal
strigulae, termen usually with whitish scales be-
low apex, thus seemingly notched; chorda and
trace of M present in cell, usually all veins sepa-
rate beyond cell; hindwing with M2 distant from
and parallel to M3 at base; male genitalia usually
with hami and with dorso-longitudinally invagi-
nated valva; female genitalia typically with an
accessory sac arising from near usually stellate
signum. Larva: Essentially tortricid, but aberrant
in some features; stemmata and anal fork absent;
pinacula weakly sclerotized; on Al 7 SD2 sepa-
rate and dorsal to SD; Dl and SD on same
pinaculum on A9; L group bisetose on T l , with
L2 anterodorsal of LI, and trisetose on A9; SV
group multisetose (59 bristles) on Al 7, biset-
ose on A8 and A9; VI on coxa on Tl and T2.
Pupa: Dorsal spines very small, an anterior band
of 5 6 irregular rows on each segment and a
posterior, sometimes partly interrupted, single
row; no cremaster, but AIO with four small
thorns, two subapical and one each laterally,
with small caudal setae; two setae on each side
of anal rise (Diakonoff & Arita 1981 for imma-
tures). The Hilarographini (Hilarographa, Thau-
matographa), comprising about 60 species in 5
genera, are a largely pantropical group (Heppner
1982). All known larvae are borers in various
plant parts (Horak & Brown 1991), some in cam-
bium of Pinus sp. With their glyphipterigid wing
pattern the Hilarographini have only recently
been recognized as tortricids (Diakonoff 1977 a,
b), with some superficially similar but unrelated
groups included among them (Horak & Brown
Marianne Horak
1991). Synapomorphies for the tribe are the slen-
der, smooth, upcurved palpi, ciliate antenna in
male and large ocelli, apart from the characteris-
tic wing pattern.
Olethreutinae. Antenna with one ring of scales
per segment (Fig. 12.3 B); forewing with costal
strigulae and often with costal fold; hindwing
usually with cubital pecten (Fig. 12.6 B), male
genitalia with transtilla absent, gnathos reduced,
aedeagus fused with anellus and juxta (Fig.
12.5 G); valva with costal hook and with inner
face strongly sclerotized except for basal excava-
tion; female genitalia with sterigma usually not
connected with anterior apopyhses (Fig. 12.5 C);
signa often two. Pheromones usually based on
12C-chains, except for Microcorsini, Gates-
clarkeanini and Bactrini (Horak & Brown 1991).
The monophyly of the Olethreutinae is demon-
strated by several apomorphies, with the fusion
of aedeagus, juxta and anellus into a functional
unit and the modified flagellum with only one
ring of scapes per segment both diagnostic. De-
spite their highly derived genitalia, the Olethreu-
tinae have usually retained M and often the
chorda in their forewing cell, in contrast to most
Tortricinae. Of the seven tribes here recognized
the four smaller ones are considered to be mono-
phyletic (pending the final position of Endo-
thenia), whereas the present Olethreutini and Eu-
cosmini are probably paraphyletic, though each
with a monophyletic core, and the Grapholitini
polyphyletic.
Microcorsini (Fig. 12.1 N). Adult: Costal fold
and cubital pecten sometimes present; tibial pen-
cil absent (the tibial pencil of an Olethreutini in
Horak 1991: figs. 1.1.8(d) and (e) is erroneously
ascribed to Cryptaspasma sp.); hindwing with M2
and M3 separate and in male often with dorsal
fold on anal margin; male usually with sex scales
on S8; aedeagus with coecum; valva usually bul-
bous, with small basal excavation and large basal
pulvinus; funnel-shaped sterigma connected to
apophyses anteriores; ductus bursae with long
colliculum; two large blade- or horn-like signa.
Larva: Without anal fork; D2 on A8 on same
pinaculum and closer together than DI sea te;
D l , D2 and SD on A9 all on same large pinacu-
lum; SD2 on same pinaculum as SD on Al8;
L group on A9 bisetose. Pupa: Alar furrow with
raised margins; cremaster a low transverse ridge
with lateral angles as ventrally projecting thorns.
The Microcorsini (Diakonoff 1959; Kuznetsov
1970), including over 30 species in 5 subgenera
of Cryptaspasma, are found on all southern
hemisphere continents and in the Oriental re-
gion, extending to Japan, a distribution probably
reflecting their early derivation. Their larvae are
borers in nuts and kernels (Horak & Brown
1991). The monophyly of the group is demon-
strated by the funnel-shaped sterigma, the two
horn-shaped signa close to the 'neck' of the cor-
pus bursae and the structure of the valva. In ac-
The Tortricoidea
cordance with their numerous plesiomorphies
(sterigma connected with apophyses anteriores,
large coecum, alar furrow in pupa) the Microcor-
sini are here treated as the most plesiomorphic
olethreutines. Recent studies (Horak in prep.) re-
sulted in the finding of a small basal excavation
in Gatesclarkeana, thus removing the main
reason for treating the Gatesclarkeanini as the
basal group (Horak & Brown 1991).
Gatesclarkeanini (Fig. 12.1 O). Adult: Costal
fold absent; hindwing with M2 and M3 sepa-
rated; male often with anal area of hindwing and
hind tibia modified, and with pouches with sex
scales on abdomen; tarsal spines reduced; vincu-
lum broad; valva often asymmetrical, with small
basal excavation and sacculus with spined basal
pulvinus; sterigma often asymmetrical, inserted
in membranous pocket; one or two signa (second
usually smaller), a flat sclerite with dentate or
star-shaped projections. Larva and Pupa un-
known. The Gatesclarkeanini, about 23 species
in 4 genera, are an Oriental group, extending
into the eastern Palaearctic and Australian re-
gions (Diakonoff 1973; Razowski 1989). Larvae
of Gatesclarkeana are leaf rollers. According to
Kuznetsov & Stekolnikov (1984) the Gates-
clarkeanini lack tergal extensors of the valvae
(012) and the sternal extensors (3) are used for
closing the valvae, whereas in Endothenia and
other Olethreutinae 3 are reduced, and the val-
vae are closed by contraction of the retractors of
the aedeagus (me). Gatesclarkeanini were re-
ferred to the base of the Olethreutinae based on
the absence of a basal excavation in Ukamenia
sapporensis (as Aphiaris mirana), but it is present
in Gatesclarkeana, though hidden by the pulvi-
nus lobe. Horak and Brown (1991) included En-
dothenia in the Gatesclarkeanini, but Dang
(1990) found m2 to be present in the Endothenia
species he examined and refers the genus to the
Bactrini. Obvious similarities in the genitalia and
shared reduction of tarsal setae suggest a close
relationship between Gatesclarkeanini, Endo-
thenia and Bactrini.
Bactrini (Fig. 12.1 P). Adult : Costal fold or any
other scent organs absent; hindwing with M2, M3
and CuAi approximated; tarsal setae reduced;
valva with small basal excavation and greatly en-
larged sacculus, usually with heavy spines or
large hairpencils; sterigma anteriorly invagi-
nated; signum a small scobinate invaginated
plate. Larva: Anal fork absent; legs with dorsal
setae nearly twice length of claws; A9 with D1
and SD on same pinaculum; SV bisetose on
A79. Pupa: AIO at most with two small me-
dian spines; cremaster absent, cauda with two
small simple setae; anal rise with one small seta
on each side. The Bactrini, comprising about 105
species in 7 genera, are present worldwide, but
concentrated in the Oriental/Australian, African
and Palaearctic regions (Diakonoff 1956, 1962,
1963, 1973). The larvae of Bactra bore in stems
211
of Cyperaceae, Juncaceae and Gramineae. Mo-
nophyly of the Bactrini is demonstrated by the
characteristically modified sacculus and by the
anteriorly invaginated sterigma. A transtilla has
been erroneously reported for Protobactra\ the
sclerite in question is the displaced vinculum.
Dang (1990) includes Endothenia in the Bactrini
on the basis of uncus, tegumen and valva base
structures.
Olethreutini (Figs. 12.1 Q, 12.4B, 12.5C,
12.6 B). Adult: Costal fold absent; hindwing with
M2, M3 and CuAi approximated or connate at
base (Fig. 12.6 B); males often with sex scales in
roll on inner margin of hindwing, with hair pen-
cil (Fig. 12.4 B) or brush on hind tibia and/or
abdominal pouches or brushes; vinculum nar-
row, coecum absent; valva with large basal exca-
vation, costal hook usually two-branched, saccu-
lus with one or two spine clusters, cucullus nar-
row; cornuti fixed or deciduous; sterigma derived
from spinulose membrane around ostium, not
fused with S7 (Fig. 12.5 C); usually two signa,
plate-like, scobinate or cornuate. Larva: Anal
fork often present; T1 with LI usually closer to
L2 than to L3; on A l - 7 L2 usually not much
smaller than LI; SV group on A 7 - 9 is usually
3 : 2 : 2 ; A9 with D1 and SD usually on separate
pinacula. Pupa: A8 with one or two rows, A9
with one row and on AIO spines random or ab-
sent; cremaster present, bearing eight hooked se-
tae, anal rise with two hooked setae on each side.
The Olethreutini (Apotomis, Cymolomia, Dud.ua,
Endopiza, Hedya, Lobesia, Olethreutes), include
about 1400 species in over 144 genera and occur
worldwide, with their greatest diversity in south-
east Asia (Diakonoff 1973) and the Holarctic re-
gion (Heinrich 1926; Miller 1987; Razowski
1989). Their larvae are mainly leaf rollers. The
concept of the Olethreutini (hindwing with M3
and CuAi separate or connate at base, sacculus
with two clusters of spines) was based on the Pa-
laearctic fauna, and no apomorphy has been
found to define the group. The group probably
is paraphyletic in terms of the other olethreutine
tribes, but there is a large core group of related
taxa with similarly derived signa, sterigma, valva
and male tibial pencil (the latter also present in
some Enarmoniini), and with a cremaster in the
pupa. Present work (Horak in prep.) on the Aus-
tralian fauna largely confirms Diakonoff's (1973)
groupings within the tribe, once the numerous
Enarmoniini are removed. Signum development
apparently follows a similar course as in Enarm-
oniini: a gradual reduction from cornuate struc-
tures to derivations of the scobinations at their
base.
Enarmoniini (Fig. 12.1 R). Adult: Labial palpi
mostly sinuate-porrect and with a yellowish
oblique streak or small spot subapically from
dorsal margin of second segment, often sur-
rounded by dark scales with bluish iridescence;
forewing with strongly falcate apex to nearly ree-
212
tangular, but always with sinuate termen; usually
with lines of silvery scales curving from costa be-
fore apex to termen below apex, preceded by a
series of silvery strigulae along distal part of
costa; hindwing with M3 and CuAi separate,
stalked or fused and often with modified anal
margin with edge thickened and/or rolled dor-
sally, and with axillary tuft developed into long
pencil (some Ancylis, Oriodryas) concealed
within anal roll; rarely hind tibia with pencil
(Metaselena); uncus usually absent (present in
Ancylis) and socii often fused to form bristled
apex of a weakly sclerotized tegumen; pedunculi
often with a medially projecting lobe (attach-
ment points of the tergal flexors of the valva,
nu); valva usually with very large basal excava-
tion and often with distal thorn; caulis very long
with the aedeagus protractor (ms) attached to its
base; with numerous deciduous lanceolate cor-
nuti; often with a tuft of large scales from mem-
brane between tegumen and valva; sterigma with
at least lamella postvaginalis spinulose; signa
two or one, in more generalized taxa two hollow
blade-to-horn-shaped structures (Ancylis, Neo-
anathamna), often with a granular base, and one
or both lost or reduced to a granular patch in
more derived genera (Metaselena); sometimes
additional sclerites at entrance to bursa. Larva:
Anal fork present or absent; D1 and SD on A9
on same (Enarmonia, Eucosmomorpha) or sepa-
rate (Ancylis) pinacula; SV group on A 7 - 9 is
usually 3 : 2 : 2 ; VI very close to or on coxa on
T2. Pupa: A8 with one or two rows of dorsal
spines, A9 with one or without, AIO with dorsal
row (Eucosmomorpha, some Ancylis), with scat-
tered spines (some Ancylis) or without spines
(.Enarmonia), but always with cremaster; cremas-
ter with 8 hooked terminal setae, two setae (An-
cylis) or less on each side of anal rise. The En-
armoniini, comprising over 150 species in over
25 genera are found nearly worldwide, with their
greatest diversity in the Oriental/Australian re-
gion and with numerous species of Ancylis in the
Holarctic region (Razowski 1989). Their larvae
are either leaf rollers (Ancylis) or borers (Enarm-
onia). An ongoing study of the Australian
Olethreutinae (Horak in prep.) has shown the
Enarmoniini to be much more diverse and nu-
merous than anticipated, and after long delibera-
tion (Horak 1981) tribal rank, as suggested by
Kuznetsov and Stekolnikov (1973), is here ac-
cepted for the group for mainly pragmatic
reasons. With its pupa with a cremaster and a
spinulose sterigma the group certainly is not sub-
ordinate within the Eucosmini, and its removal
from the Olethreutini will help to recognize mo-
nophyletic Olethreutini s. str. Possible synapo-
morphies are the characteristically marked se-
cond segment of the labial palpi, forewing shape,
the hindwing roll with a well-defined axillary
pencil, pedunculi with attachment lobes for nu,
socii fused with each other or/and with tegumen,
Marianne Horak
basal excavation large, ms attached to base of
long caulis, numerous deciduous lanceolate cor-
nuti and pupa with a cremaster. Several of these
characters are absent in Ancylis, and probably
very few are found in all Enarmoniini. Reduction
of the signa from initially two cornuate or
wedge-shaped hollow blades to a single, scobi-
nate patch (Metaselena) can be traced through
the intermediate steps and no doubt hindered re-
cognition of the group. Ancylis may well be the
sister group to the rest of the tribe.
Eucosmini (Figs. 12.1 S, 12.3 B, 12.4C, 12.5 G).
Adult : Male antenna frequently with basal notch
(Fig. 12.3 B); costal fold often present (Fig.
12.4 C); hindwing with M3 and CuAi stalked,
sometimes with anal roll containing hair pencil
from base of wing; attachment points of muscles
m4 not prominently projecting from pedunculi;
valva with larga basal excavation, costal hook
single- or two-branched, cucullus well-developed
and usually delimited by neck (Fig. 12.5 G); ae-
deagus with deciduous cornuti; sterigma derived
from a smooth sclerotized area around ostium,
in deep emargination of S7 or fused with S7 in
derived genera; ductus bursae often with sclero-
tized ring near ductus seminalis; signa often cor-
nuate or reduced and entire bursa finely spined.
Larva: Anal fork often present, T1 with LI usu-
ally equidistant from L2 and L3; on Al 7 L2 at
most half as long as LI; SV group on A 7 - 9 usu-
ally bisetose or unisetose; A9 with D1 and SD
on same pinaculum; A8 with SD anterior or
dorso-anterior to spiracle. Pupa: A9 and AIO
with one row of dorsal spines; without cremas-
ter; cauda with 48 flat hooked setae; anal rise
with 1 2, rarely 3 setae on each side. The Eucos-
mini (Crocidosema, Epiblema, Spilonota, Strep-
sicrates, Zeiraphera), with about 1000 species in
over 100 genera are found worldwide but are
predominantly Holarctic (Heinrich 1923;
Obraztsov 1964-68; Miller 1987; Razowski
1989). The larvae of the more generalized Eucos-
mini are usually leaf rollers or webbers (Epi-
notia), and those of the more derived taxa are
borers in roots, stems and fruit. The tribe, as
presently delineated (base of M2 bent towards
base of stalked M3 and CuAi) is probably pa-
raphyletic (relative to the Grapholitini) as well
as polyphyletic (relative to the Olethreutini),
though the eucosmine/grapholitine sterigma that
is apparently derived from a smooth sclerite as-
sociated with the ostium is probably apomorphic
relative to the olethreutine/enarmoniine sterigma
that is derived from a spinulose membrane. Also,
with the Enarmoniini removed, the Eucosmini
and Olethreutini seem to be characterized by dif-
ferent secondary sexual structures (if present),
the Eucosmini by a costal fold and the Olethreu-
tini by a tibial pencil.
Grapholitini (Fig. 12.1 T). Adult: Forewing
often with distinct speculum, rarely with costal
fold (Dichrorampha); hindwing with bases of Rs
The Tortricoidea
and Mi approximated or distant (Dichroram-
pha), base of M2 parallel to and distant from M3;
uncus and socii reduced or absent; valva with
costal hook single or reduced; aedeagus often
with spiraled apex, projections or fixed cornuti;
sterigma derived from smooth sclerite, usually in
deep emargination of S7; ductus bursae often
with sclerotized pockets near ostium; usually
with two cornuate signa. Larva: Anal fork often
absent; spinneret simple; A9 with D1 and SD
on same pinaculum; A8 with spiracle distinctly
close to caudal margin, with SD ventral ante-
rior or dorsal posterior to spiracle; SV group on
A 7 - 8 bisetose or unisetose, on A9 unisetose.
Pupa: A8 with one to two rows of dorsal spines,
A9 with one row, rarely absent or irregular, AIO
with one row, rarely irregular; 10th segment usu-
ally short; cremaster absent; cauda usually with
four recurved setae; anal rise with two setae on
each side, rarely one or absent. The Grapholitini
(Collicularia, Pammene, Selanla), comprising
some 600 species in over 40 genera are repre-
sented in all regions of the world, but over two-
thirds of the described species of Cydia and
related genera are Holarctic (Heinrich 1926;
Obraztsov 1958-61; Miller 1987; Razowski
1989). Most species are monophagous borers in
fruits, roots, and apical shoots. The Grapholitini
seem to be a polyphyletic assemblage (Horak &
Brown 1991), historically characterized by a
hindwing with M2 and M3 widely separated and
parallel at base and male genitalia with a
strongly reduced dorsal complex. A presumably
monophyletic core group around Cydia and
Grapholita is probably subordinate within the
Eucosmini.
Acknowledgments
I wish to thank the following colleagues for gen-
erously providing information and valuable com-
ments on the manuscript: Dr. I. F. B. Common,
Toowoomba; Dr. J. W. Brown, Chula Vista; Dr.
R. L. Brown, Mississippi State University, Mis-
sissippi State; Dr. J. S. Dugdale, Landcare Re-
search, New Zealand; Mr. E. D. Edwards and
Dr. E. S. Nielsen, CSIRO, Canberra; Dr. F. Ko-
mai, Osaka University of Arts, Osaka; Dr. J. A.
Powell, University of California, Berkeley; Mr.
K. R. Tuck, The Natural History Museum, Lon-
don; Dr. J. Razowski, Polish Academy of Sci-
ences, Krakow. Permission for republishing Figs.
12.1, 12.2 B, 12.4A, C and 12.6 was given by
Elsevier Science Publishers, and for Fig. 12.5 D
by the editor of the Australian Journal of Zool-
ogy, as indicated in the figure captions. The re-
maining illustrations were provided by Mr. J. P.
Green (photography), Mrs. E. Alyta (printing),
Mrs. H. Geier (SEM) and Mr. C. Davies and
the graphics section (production of plates), all
CSIRO Division of Entomology, Canberra.
213
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Addendum
Important references published after conclusion
of the manuscript include a field guide to the
microlepidoptera of South-East Asia (Robison et
al. 1994), and the tortricid chapters for the Neo-
tropical (Powell et al. 1995), Australian (Horak
et al. 1996) and European (Razowski 1996)
checklists.
Recent studies by F. Komai on the Australian
Grapholitini suggest that Cryptophlebia, with a
bifid uncus in a plesiomorphic Australian species
and with cornuate signa and a smooth, sclero-
tized sterigma, belongs to the Grapholitini
(Horak et al. 1996).
Horak, M., Common, I. F. B. & Komai, F. (1996):
Tortricidae. Pp. 123-136, 349-351. In: Nielsen,
E. S., Edwards, E. D. and Rangsi, T. V. (eds.).
Checklist of the Lepidoptera of Australia. Mono-
graphs on Australian Lepidoptera 4. CSIRO Pub-
lishing, Collingwood.
Powell, J. ., Razowski, J., Brown, J. W. and Brown,
R. L. (1995): Tortricidae. Pp. 138-157, 177. In:
Heppner, J. B. (ed.). Atlas of Neotropical Lepido-
ptera. Checklist 2. Association of Tropical Lepido-
ptera, Gainsville.
Razowski, J. (1996): Tortricidae. Pp. 130-157,
313-318. In: Karsholt, O. & Razowski, J. (eds).
The Lepidoptera of Europe. A distributional check-
list. 380 pp. Apollo Books, Steenstrup.
 13. The Smaller Microlepidoptera-Grade Superfamilies
John S. Dugdale, Niels P. Kristensen, Gaden S. Robinson & Malcolm J. Scoble
As outlined in 12 and 13 the demand that
taxa must be monophyletic in the strict cladistic
sense has led to a proliferation of superfamilies,
which are either monofamilial or comprise a very
small number of families. Several taxa treated in
this chapter have until comparatively recently
been treated as subordinate in some of the larger
micro-moth superfamilies, but in no case is there
strong evidence for such arrangements.
Ditrysia Incertae Sedis
GALATICOIDEA superfam. nov.
In recent decades a group of small genera
centred on Galactica has been ranked as a sub-
family in the Plutellidae (Friese 1966; Heppner
1987), Glyphipterigidae (Clarke 1968, Moriuti
1977), Urodidae (Heppner 1991), or as a distinct
family in the Tineoidea (e. g. Common 1990, Mi-
net 1991). Minet (1986) suggested a close rela-
tionship between galacticids and psychids to be
indicated by the larval pronotal shield including
the L setae, and by the anteriorly directed spines
on the pupal abdomen. However, the apomor-
phic nature of the former trait is debatable, and
the galacticids are here ( 1 - 7 ) excluded from the
Tineoidea. The sternum II structure provides
ambiguous clues to the affinities of the group,
but a placement in the Apoditrysia cannot be
ruled out: while the apodemes are slender; they
are moderately short, have broad bases and are
not continued into venulae; the anterior sternal
corners are produced laterally, but not as much
as is usual in the Apoditrysia. Since the super-
family category is consistently used in current
Lepidoptera classifications (see 13) a (informa-
tion-wise redundant) superfamily is here erected
for the Galacticidae, pending better insights into
their affinities.
Galacticidae. Family autapomorphies remain to
be worked out in detail, but postabdominal mod-
ifications in both sexes will probably prove valid
as such: male tergum VIII forming a hood over
genitalia; female ostium bursae a minute orifice
on curved, sclerotized tube projecting from
membrane behind segment VII.
Small moths, forewing length 48 mm. Head
vestiture smooth, mostly of broad scales; ocelli
and chaetosemata absent; antennal scape usually
with pecten, flagellum filiform or bipectinate.
Maxillary palps short, 12 segmented. Labial
palps rather short, porrect or lightly upcurved.
Epiphysis present; spurs 0-2-4. Fore wing with
chorda present and M-stem usually well defined;
Rs configuration variable; CuP in both wing
pairs distinct near margin.
Larvae gregarious in webbings on Legumino-
sae-Mimosaceae. Prothoracic L trisetose, in pro-
notal shield; LI and 2 close together on same
pinaculum on IVI, farther part (on separate pi-
nacula) on VIIVIII; proleg crochets in irregu-
larly biordinal circle (uniordinal half-circle on
anal prolegs). Pupa with variably developed
bands of posteriorly directed dorsal spines/ridges
on abdomen, at least in some taxa also with
whorl of anteriorly directed spines near hind
border of IVVII; cremaster with hooked setae,
or absent; cocoon dense, whitish.
An Old World family with 17 named species,
ranging from North Africa through Asia to Aus-
tralia; there are undescribed species from New
Caledonia and the Afrotropics. Most species (8)
are in Homadaula; H. anisocentra ('mimosa web-
worm', Fig. 13.2 A), a pest on ornamentals, has
been introduced to Eastern N. America.
Moriuti (1963), Friese (1966), Common
(1990).
SIMAETHISTOIDEA superfam. nov.
Minet (1991) assigned the small micro-moth ge-
nus Simaethistis to a family of its own, because
no good support could be found for its previous
placements in the Tineoidea or Crambidae-Pyra-
ustinae. At present it still seems necessary to
leave the question of simaethistid affinities open;
the taxon has even not been unambiously as-
signed to the Obtectomera or the non-obtectom-
eran grade. As with the Galacticidae and Whal-
leyanidae, a new superfamily for the Simaethisti-
dae is here erected for formal classificatory
reasons. The following is based on Minet's ac-
count.
Simaethistidae (Fig. 13.2B) are perhaps superfi-
cially most similar to some Crambidae-Pyrausti-
nae; they are contrast-rich, broad-winged moths
spanning about 1219 mm. The family is largely
characterized by the absence of apomorphies
diagnostic of other ditrysian superfamilies. Ocelli
large; chaetosemata absent; cephalic vestiture of
very narrow (piliform-looking) scales with
deeply scalloped apices. Antenna filiform, with
distinct setae in both sexes. Maxillary palps at
least 2-segmented, small; proboscis unsealed. La-
bial palps porrect, segment 1 rather long, seg-
ment 3 slender. Epiphysis present; spurs 0-2-4;
218 John S. Dugdale, Niels P. Kristensen, Gaden S. Robinson & Malcolm J. Scoble

Fig. 13.1 Choreutidae: - B ; Schreckensteiniidae: CD; Epermeniidae: EG; Alucitidae: H I; Pterophoridae:

JO. A, Choreutis head; B, same, venation: C, Schreckensteinia larva; D, same, adult's hind tibia: E, Epermenia
hind tibia; F, Thambotricha venter II; G, Epermenia venation; H, Alucita venation; I, same, sternum II (ap:
apdeme; ve: 'venula'; si. ts: 'latero-tergal sclerite'; tg. 1: tergum I); J, hindwing underside with specialized scales
(DS); K, pterophorine (Emmelina) male genitalia with asymmetrical valves; L, Agdistis sternum II (ve: 'venula');
M, Pterophorus venation; , 'platyptiline' hind wing venation; O, pterophorine larva (right: abdominal segment
III); P, Leioptilus pupa. A after Robinson et al. (1994); after Heppner & Duckworth (198); C after Heppner
(1987); D E original (B. Rubaek del.); F redrawn after Dugdale (1987); G - H after Nielsen & Common (1991);
I, L after Minet (1983); J - K after Hannemann (1977); M - N after Wasserthal (1974): O after Neunzig (1987);
after Yano (1963).
The Smaller Microlepidoptera-Grade Superfamilies
pulvillus unspecialized. Meso-basisternum and
metascutellum not exhibiting specializations
common to alucitoids and pterophoroids. Fore-
wing chorda vestigial (a fold); spinarea present;
all Rs branches separate. Hindwing SC + R ap-
priximated to Rs beyond discal cell. CuP absent
in both wing pairs. Some wing scales with promi-
nent median ridge. Tergum I without postspira-
cular lobes characteristic of thyridoids; sternum
II without V-shaped venula pattern characteristic
of alucitoids. In male tergum VIII narrowed;
juxta with long lateral lobes; gnathos spinose.
While the loss of CuP is compatible with sub-
ordination in the Alucitoidea, the thoracic and
sternum II structure is not. This loss is also com-
patible with a subordination in the Thyridoidea,
and this placement could receive additional sup-
port from the male's tergum VIII-, juxta- and
gnathos-specializations, which all have counter-
parts within the Thyridoidea. On the other hand
a subordination within the Thyridoidea is con-
tradicted by the large ocelli (very small in thyri-
doids, when at all present), the presence of a spi-
narea, and the absence of lateral tergum I lobes.
Similarly a placement within the Pyraloidea is
decisively contradicted by the unsealed probos-
cis, separate forewing Rs2 and Rs3, and lack of
tympanic organs. Also a placement as the closest
relative of either of the two lastmentioned super-
families is difficult to accept because of the un-
specialized Simaethistis pulvillus. It is true, how-
ever, that the ridged wing scales is a specializa-
tion which is recurrent in 'lower Obtectomera'
(Hyblaeoidea, many Pyraloidea and Thyri-
doidea, a few Immoidea and Copromorphoidea).
Simaethistis, with 2 species, occurs in China
and Northern India. The immatures are un-
known. The Australian Metaprotus (2 species) is
also included here (M. Shaffer pers. comm.).
Non-Obtectomeran Apoditrysia
CHOREUTOIDEA
The taxon centred on Choreutis was long in-
cluded in the non-apoditrysian Glyphipterigidae
(now in the Yponomeutoidea), and subsequently
transferred to the Sesioidea (Brock 1971,
Heppner 1977, Heppner & Duckworth 1981).
Minet (1986, 1991), however, refuted alleged sy-
napomorphies with the sesioid assemblage and
placed the choreutids in a superfamily of their
own.
Choreutidae. Probable autapomorphies (Heppner
1982, Minet 1991): proboscis scaled (Fig. 13.1 A,
parallelism with Gelechioidea in the non-apodi-
trysian grade and Pyraloidea in the Obtectom-
era); maxillary palp minute, at most 2-seg-
mented; egg upright. The arrangement of the lar-
val proleg crochets in a circle/penellipse rather
than a transverse row is likely to be plesiomor-
219
phic rather than an additional choreutid ground-
plan autapomorphy as believed by Heppner; the
same may be true of the presence of a forewing
pterostigma.
Small (wingspan some 5 - 2 0 mm) diurnal
moths with large ocelli but without chaetosem-
ata. Face (Fig. 13.1 A) smooth-scaled, vertex
with loosely appressed scales. Antenna filiform,
in male usually with long ventral setae; some-
times strongly scaled dorsally. Labial palp up-
curved, often with ventral scale tuft on segment
2; vom Rath's organ terminal, cup-shaped.
Epiphysis present; spurs 0-2-4; scale tufts present
near spurs. Wings usually broad, forewing often
with metallic markings, chorda present or ab-
sent, M-stem at most just discernible, all veins
'sessile' (Fig. 13.1 B). Genitalia quite diverse in
both sexes.
Larval structure and biology variable; proleg
crochets uniordinal (except for a single millieri-
ine genus); LI and 2 on IVIII widely separated.
Pupa protruded before eclosion; segment X with
several hook-tipped setae.
Two subfamilies are here recognized; they can-
not be separated on the basis of adult characters.
Millieriinae. Probable autapomorphy: Larval
prothoracic L-group bisetose (may be related to
endophytic life style). The larvae also differ from
those of choreutines in having D2's on VIII as
close or closer than Dl's, and short prolegs; the
pupae have two spine rows (or, in Phormoestes,
a spine field and a row) on several abdominal
segments. These traits occur elsewhere among
lower Apoditrysia and are here considered plesi-
omorphic.
3 monobasic genera (Heppner 1982 b). Mil-
lieria (W. Palaearctic, Fig. 13.2D) larvae are leaf
miners (a life-style otherwise unknown in the
family) on Aristolochia\ Phormoestes (Florida)
larvae are borers in the inflorescens of Sabal
palms; they are unique in the family in having
biordinal proleg crochets; Nyx (Chile).
Choreutinae. Probable autapomorphies: Lar-
val prolegs slender, elongate (reminiscent of
those of some Yponomeutoidea); on VIII Dl's
closer than D2's. Pupal abdominal segments with
single spine row.
402 species worldwide (Heppner 1991). A sep-
arate subfamily Brenthiinae has been made for
Brenthia (Fig. 13.2 E; 6 5 + species, all regions,
but mainly Oriental) and Litobrenthia which are
characterized by rounded forewings and a suite
of other specializations (Heppner & Duckworth
1981); it seems, however, that the remaining
choreutines are thereby rendered a paraphyletic
assemblage. Choreutis (85 4- species, cosmopoli-
tan): C. pariana skeletonizes apple leaves in
Europe; many other species feed on Moraceae,
C. tigroides being a pest on jackfruit (Artocarpus)
in Java. Other main genera include Prochoreutis,
Tebenna, Asterivora (Australia, New Zealand, on
Asteraceae), Saptha (Fig. 13.2 C).
220 John S. Dugdale, Niels P. Kristensen, Gaden S. Robinson & Malcolm J. Scoble
Dugdale (1979), Arita (1987 a, b), Diakonoff
(1986), Heppner & Duckworth (1981), Kuznet-
zov & Stekolnikov (1981).
URODOIDEA
The Urodus group of genera was excluded from
the Yponomeutoidea by Kyrki (1984, 1988).
While these moths seem appropriately placed in
the Apoditrysia, their affinities are at present
unresolved. Derived characters shared with
Schreckensteinioidea (elongate larval prolegs,
reduced pupal spinosity, large-meshed cocoon)
are obviously homoplasious, hence ambiguous
(Kyrki 1988). The Galactica group of genera, by
Heppner (1991) considered a urodid subfamily,
are here treated as a separate family (see above).
Urodidae. Probable autapomorphies (Kyrki
1988): Male antennae lamellate; male hindwing
with costal hair pencil (occasionally reduced).
Larval pro thorax without seta MXD1; segment
VIII with L3 ventroanteriad from LI and L2,
SVI almost as high as L3; prolegs long and slen-
der, constricted at midlength, crochets in mesose-
ries.
Kyrki's statement that the presence of a single
metathoracic seta M D is among the "good diag-
nostic characters for the family" must be a lapsus
calami; the metathoracic M D is bisetose only in
the Noctuoidea.
Small to medium-sized micro-moths, wing-
span 1137 mm. Cephalic vestiture more or less
appressed, ocelli and chaetosemata absent.
Maxillary palps minute; proboscis unsealed, with
prominent sensilla. Labial palps medium-sized,
variably upcurved. Epiphysis present; spurs 0-2-
4. Forewing with distinct chorda and long ptero-
stigma; Rs4 to costa or apex; CuP distinct near
margin only. Sternum II with prominent antero-
lateral processes; 'sternal rods' (or 'venulae') pre-
sent or not. Female apophyses very long.
The few known larvae free-living on broad-
leaved trees. Pupal abdominal segments with few
spines, IIIVII movable in males, IIIVI in fe-
males; in meshed cocoon, protruded before eclo-
sion.
3 genera with about 60 species. Urodus (Fig.
13.2 H) and Spiladarcha are Neotropical (the for-
mer with a single N. American species), Wockia
is Eurasian.
Kyrki (1988).
SCHRECKENSTEINIOIDEA
Previous assignments of Schreckensteinia to the
Yponomeutoidea or Copromorphoidea were
shown ill-founded by Minet (1983) and Kyrki
(1984), and while the taxon can be placed among
the non-obtectomeran Apoditrysia no better sub-
stantiated inclusion in another superfamily has
been proposed. Characters supporting a close re-
lationship with urodoids (see above) are homo-
plasious, as is the hindtibial spinosity which
might support a close relationship with epermen-
ioids.
Schreckensteinidae. A very small family charac-
terized by the presence of stiff dorsal spines on
the hind tibia (Fig. 13.1 D) and a single-bristle
female frenulum.
Small narrow-winged moths. Head smooth-
scaled; ocelli and chaetosemata absent; scape
without pecten. Maxillary palps minute; probos-
cis unsealed. Labial palps medium-sized, porrect-
upcurved. Epiphysis present; spurs 0-2-4, spurs
in each pair of subequal length; hind legs ob-
liquely elevated at rest. Forewing without pteros-
tigma; chorda and M-stem vestigial; spinarea
present; all veins sessile in both wings; forewing
Rs4 to termen; CuP in both wings vestigial (basal
portion may be well defined in forewing).
Egg upright. Larvae (Fig. 13.1 C) external
feeders; prolegs unusually slender, with very few
crochets (Fig. 13); prothoracic prespiracular L-
group trisetose; LI and 2 on IVIII on same
pinaculum; D setae (one or both) on mesotho-
rax-IX strikingly modified, with expanded apex.
Pupa with considerable abdominal mobility and
strong, paired dorsal spines on several abdomi-
nal segments; cremaster with hooked setae. Pu-
pation in large-meshed cocoon; pupa protruded
prior to eclosion.
Schreckensteinia (Holarctic and Neotropical)
comprises 5 species. Recorded host plants in-
clude Anacardiaceae, Scrophulariaceae and Ro-
saceae. The widespread S. festaliella (Fig. 13.2 G)
has been introduced to Hawaii to control foreign
Rubus. Corsocasis (3 species) is Oriental.
Zimmermann (1978), Kuznetzov & Stekolni-
kov (1984).
EPERMENIOIDEA
Like the preceding taxon this one has been
placed in the Yponomeutoidea as well as the
Copromorphoidea, and there are intriguing in-
dications of affinities with the alucitoid/ptero-
phoroid assemblage; see 12.
Epermeniidae. Probable autapomorphies: hind
tibia with stiff bristles (Fig. 13.1 E; not confined
to dorsal surface as in Schreckensteinioidea).
Forewing fringe with groups of lamellar scales
(as in Alucitoidea and Pterophoroidea, Minet
1991). Larval submentum with posterior protu-
berance (as in Alucitoidea and some Copromor-
phoidea, Heppner 1987); prothoracic prespiracu-
lar L group bisetose (as in Alucitoidea and most
Obtectomera). Pupa unspined, abdominal seg-
ments IIV immovable (as in Yponomeutoidea,
Gelechioidea, Alucitoidea and Obtectomera);
segment IX with characteristic paired lateral pits
(Common 1990).
The Smaller Microlepidoptera-Grade Superfamilies 221

Fig. 13.2 Galacticidae: A; Simaethistidae: B; Choreutidae: CE; Epermeniidae: F; Schreckensteiniidae: G; Urodi-

dae: H; Tineodidae: I, MO; Alucitidae: J - L ; Pterophoridae: P - S . A, Homadaula anisocentra; B, Simaethis
tricolor, C, Saptha smaragditis; D, Millieria dolosana; E, Brenthia albimaculana; F, Epermenia chaerophylella; G,
Schreckensteinia festaliella; H, Urodus venatella, I, Cenoloba obliteralis; J, Alucita hexadactyla; K, Triscaedecia
dactyloptera; L, Hexeretmis argo, M, Tanycnema anomala, , Tineodes pterophoralis; O, Oxychirota paradoxa; P,
Agdistis bennettiv, Q, Ochyrotica concursa; R, Deuterocopus planeta; S, Oidaematophorus rogenhoferi. Uniform
magnification, scale (millimetres) in A.

Small narrow-winged moths. Head smooth-
scaled; ocelli and chaetosemata absent; scape
with pecten of piliform scales or, (in Thambot-
richa) an awning of broader scales. Maxillary
palps 3-segmented, folded over proboscis base;
proboscis unsealed. Labial palps medium-sized,
ascending. Epiphysis present; spurs 0-2-4, inner
spurs markedly longer than outer. Forewing
(Fig. 13.1 G) Rs4 to termen, sessile or stalked
with Rs3; CuP discernible near termen or altoge-
222 John S. Dugdale, Niels P. Kristensen, Gaden S. Robinson & Malcolm J. Scoble
ther absent. Hindwing without CuP and only
one, very short, A. Female frenulum with two
bristles. Abdominal terga with or without spine
patches. Male pleuron II sometimes with eversi-
ble or permanently protruded setose process
(Fig. 13.1 F). In female genitalia anterior apoph-
ysis at base continued into dorsal and ventral
costae (as in Schreckensteinioidea, Yponomeu-
toidea, Tineoidea and Gracillarioidea, but not
Copromorphoidea, Dugdale 1987).
Eggs flat. Larvae endophytic, or external leaf/
seed feeders; several species change from the for-
mer life-style to the latter after the first instars.
LI and 2 on I - V I I I on separate pinacula; proleg
crochets uniordinal, in complete circle. Recorded
food plants belong to several, partly distantly re-
lated dicot families. Pupa not protruded from co-
coon prior to eclosion.
83 species, represented in all regions (Heppner
1991). The current division into two subfamilies
Epermeniinae and Ochromolopinae is not
adopted here, since the latter (characterized by
the loss of one of the apical forewing veins, and
widening of ventral costa at base of anterior
apophysis, Gaedike 1978) is likely to be cladisti-
cally subordinate in the former. The monobasic
New Zealand Thambotricha may be the sister-
group of all other members of the family; its im-
matures are unknown. Main genera: Epermenia,
E. chaerophyllella (Fig. 13.2 F) being a minor
pest on cultivated umbellifers in Europe; Ochro-
molopis; Gnathifera.
Gaedike (1966, 1970, 1977, 1978, 1979), Minet
(1983), Kuznetzov & Stekolnikov (1984), Dug-
dale (1987), Heppner (1987).
ALUCITOIDEA
This superfamily is probably the sister group of
the Pterophoroidea, but the evidence is ambigu-
ous since some traits indicate closer affinities
which copromorphoids and/or epermenioids;
see 12.
Probable autapomorphies (Minet 1991): pupal
segments IIV immovable (as in Yponomeu-
toidea, Gelechioidea, Epermenioidea and Obtec-
tomera), loss of CuP in both wings, and sternum
II with V-shaped venula configuration (Fig.
13.11). Common (1990) mentioned the presence
of anterior spinose bands on abdominal terga
IIVII as another characteristic of the super-
family; a similar spinosity is also present in the
basic epermenioid Thambotricha, but it is debat-
able whether this trait is apomorphic at the
apoditrysian level. The larval submental process
(recurring in the obtectomeran Copromor-
phoidea), as well as the bisetose prothoracic L-
group and unspined pupal abdomen are further
apomorphies at this level; all traits also occur in
Epermenioidea among the non-obtectomeran
Apoditrysia.
Prominent divisions of the wings occur within
the Alucitoidea and Pterophoroidea and no-
where else in Lepidoptera; however this trait
cannot well be attributed to the ground plan of
either superfamily, and hence it is at most an 'un-
derlying synapomorphy' sensu S aether. Aero-
dynamic aspects of these specialized wing mor-
phologies are discussed by Norberg (1972).
Alucitoids are delicate micro-moths with vari-
able head vestiture and ocellus development.
Maxillary palp in ground plan with as many as
5 segments (Philpott 1927), an apparently plesio-
morphic condition otherwise unknown from
Lepidoptera above the tineoid grade (and here
probably derived from subdivision of one or
more segments from a groundplan state of fewer
than five); proboscis unsealed. Labial palps mod-
erately long. Epiphysis present; spurs 0-2 (0 in a
specialized tineodid genus-4; hind tibial vestiture
variable. Pterostigma absent; spinarea absent
(except in some Alucitidae).
Larvae semiprognathous concealed feeders;
LI and 2 on IVIII close; proleg crochets unior-
dinal.
Tineodidae (including Oxychirotidae auct.). This
taxon comprises an assemblage of alucitoids with
entire or bilobed wings; autapomorphies have
not been worked out, so it is not excluded that
it will prove to be paraphyletic with respect to
the Alucitidae.
Facies variable (Figs. 13.21, M - O ) . Head
smooth-scaled, or with projecting facial tuft;
ocelli present or absent. Maxillary palp report-
edly with up to 4 segments; proboscis unsealed.
Labial palp porrect. Epiphysis present; spurs 0-
2-4, occasionally 0-2-2; hind tibia smooth-scaled.
Spinarea absent; female frenulum with 1 3 bris-
tles.
Larvae known for only a few genera; these
feed in joined leaves of Oleaceae and Euphor-
biaceae or (Cenoloba) bore in seeds of Avicennia
(Verbenaceae), eating the developing cotyledons.
LI and 2 on IVIII on single pinaculum; proleg
crochets in mesal penelipse. Pupa in cocoon; ab-
dominal segments VVI movable; apex with
hooked setae.
18 named species in 10 genera, confined to
Oriental and Australasian regions (with the ex-
ception of the monobasic Tephroniopsis from
Iran, provisionally placed here). Some genera,
including Tineodes (monobasic, with large ocelli
and unusually long labial palps, Fig. 13.2N) are
crambid-like, while Tanycnema (monobasic, Fig.
13.2 M) resembles a primitive pterophorid, and
Cenoloba (4 species, Fig. 13.21) with their bi-
lobed wings superficially resemble 'typical' pter-
ophorids (all of which, however, have fnlobed
hindwings); Oxychirota (3 species, Fig. 13.20)
have almost linear wings.
Alucitidae. A conspicuous apomorphy shared by
most members is the multiple deep division of
The Smaller Microlepidoptera-Grade Superfamilies
both wings (Figs. 13.1 H, 13.2 J - L ) : between Rs
and M, between the M branches (only 2 present
in hindwing), between M3 and CuA, between the
CuA branches, and, in the hindwing, behind
CuA; hindwings may be 6- or 7-'plumed', in 6-
plumed taxa Sc + R anastomoses with Rs,
whereas in 7-plumed taxa Sc + R supports a
plume of its own. Only in a few taxa are the divi-
sions (primarily?) little pronounced.
Head smooth-scaled, except for raised dorso-
lateral tufts; ocelli present or absent. Maxillary
palp 5-3 segmented or absent. Labial palps fairly
long, more or less strongly upcurved, sometimes
with apical segment porrect, segment 2 usually
with projecting scales beneath. Hind tibia
smooth-scaled, or above with raised long and
stiff piliform scales, which may be scattered or
set in a group near midlength. Spinarea small,
but distinct in some species (Fig. 13.1 H).
Larvae borers in buds, flowers, fruits and
shoots, or gall makers; proleg crochets in circle,
sometimes with small lateral gap. Pupae in larval
shelter, or in a cocoon away from the latter; in
both sexes only abdominal segments VVI mov-
able.
130 + species worldwide, few in Nearctic
(Heppner 1991). Principal genus Alucita (= Or-
neodes, Fig. 13.2 J. The host spectrum comprises
at least eight dicot families including Caprifolia-
ceae, Bignoniaceae and Rubiaceae; the Afrotrop-
ical A. coffeina is a coffee pest. In the large moths
belonging to Hexeretmis (2 species, Neotropical)
and Triscaedecia (2 species; Sri Lanka, Trobriand
Isles; Fig. 13.2 K) the wings are divided for a
tenth their length or less; H. argo (Fig. 13.2 L)
has wings that just appear deeply scalloped.
Meyrick (1910), Zimmermann (1978), Kuznet-
zov & Stekolnikov (1979 b), Scholz & Jckh
(1994).
PTEROPHOROIDEA
The true plume moths may be most closely re-
lated to the Alucitoidea, but the evidence bearing
on the issue is ambiguous; see 1 - 2 . Minet (1991)
and Gielis (1993) are in agreement that Agdis-
topis is the sister group of all other pteropho-
roids, but differ in their ranking of the former.
We here adopt Minet's inclusion of all pteropho-
roids in a single family, since its monophyly is
apparently well supported even in this broad
sense. Moreover, we subjectively consider the
phenetic discontinuity between Agdistopis and
the remaining pterophoroids insufficient to war-
rant a family-level separation, particularly in
view of the phenetic heterogeneity of the latter
entity.
Pterophoridae ('Plume moths'). Probable autapo-
morphies (Minet 1991): loss of ocelli and spi-
narea; reduction or loss of 'fenestra media' be-
hind metascutellum; metepimeron forming an
223
acute angle near the posterior extremity of the
subalare; hind leg (at least in ground plan) with
tibia more than twice as long as femur; in I 'ter-
gal rim' very narrow (in posterior view); apo-
demes of sternum II very reduced (Fig. 13.1 L);
abdominal terga II and III elongated. The ar-
rangement of the proleg crochets in a mesoseries/
mesopenellipse can also be considered an auta-
pomorphy (note: the proleg structure of Agdis-
topis is undescribed); it is an unusual condition
in the non-macrolepidopteran grade, although
parallelled in the Urodoidea, in several Zygae-
noidea and in the Immoidea.
Pterophorids are medium-sized micro-moths
with very characteristic facies (Figs. 13.2 PS):
the long and slender bodies, legs and wings, and
the resting posture with the latter held out-
stretched, are parallelled only in the Alucitoidea-
Tineodidae. Head smooth-scaled, occasionally
with a conical tuft on the frons. Ocelli and chae-
tosemata absent. Labial palps very variable in
shape and vestiture. Epiphysis present; spurs 0-
2-4; hind tibia usually smooth-scaled, sometimes
with long piliform scales above. CuP present (in
ground plan) in both wing pairs. Pterostigma
sometimes present; spinarea absent; female fren-
ulum with 12 bristles. Abdominal terga un-
spined. Sternum II (Fig. 13.1 L) with venulae
long, sometimes joined by transverse bar. Asym-
metry of the genital apparatus is frequent in both
sexes (male example in Fig. 13.1 K) and has ap-
parently been evolved on several occasions. The
same is true for the absence of anterior apophy-
ses in the female.
The deep indentations in the fringed wing
margins (Fig. 13.1 M - N , 13.2 R - S ) , from which
the family derives its colloquial name, are a con-
spicuous specialization of the great majority of
pterophorids, but they are not a groundplan trait
of the family. Wasserthal (1974) related wing di-
vision to the moths' complex longitudinal wing
folding/rolling and noted a relation between hab-
itat preferences and degree of wing folding. He
further suggested that wing division has evolved
independently in Pterophorinae and 'Platyptili-
nae', but the evidence (non-derivability of one
extant venation pattern from another extant one)
is not compelling, and the scheme is difficult to
reconcile with Gielis' (1993) finding that 'Platy-
ptilinae' are paraphyletic in terms of Pteropho-
rinae.
Egg flat. Larval spiracles sometimes slightly
protruding; prothoracic L-group trisetose; LI
and 2 on IVIII approximated; secondary setae
often numerous, they may arise on distinct Ver-
rucae; prolegs long and slender (Fig. 13.10). Lar-
vae may be endophytic, boring in roots, stems
etc., or they may be external feeders. In the for-
mer case they are mostly short-haired, while ex-
ternal feeders often have long setae which may
be forked, clubbed and glandular, secreting a
sticky fluid, which is probably a protective de-
224 John S. Dugdale, Niels P. Kristensen, Gaden S. Robinson & Malcolm J. Scoble
vice. Pupae more or less strongly setose (Fig.
13.1 P); abdomen with several movable segments
and dorsal spinosity; apex with two groups of
hooked setae (on IX and X respectively) attach-
ing the fully exposed pupa to a silk pad on the
host plant or in leaf litter.
The following subfamily division (Macropira-
tinae ranking excepted) and recorded species
numbers are from Gielis (1993).
Key to subfamilies
1 Wings undivided 2 phorid subfamily. It now includes the genera pre-
Wings devided 4 viously assigned to a subfamily 'Platyptilinae',
2(1) Hindwing underside with specialized, erect
scales along Cu A (Fig. 13.1 J) 3 13.1 N), in which M3 is retained, and CuP termi-
Hindwing underside without such scales . .
Macropiratinae
3 (2) Fore wing with sparsely scaled field along
termen, abdomen unicolorous. . Agdistinae
Forewing without sparsely scaled field, ab-
domen with alternating white and coloured
segments Ochyroticinae
4(1) Fore wing with single cleft. . Pterophorinae
Fore wing with 2 or 3 clefts
Deuterocopinae
Macropiratinae. Monogeneric. Agdistopis (=
Macropiratis, Indo-Australian) with 2 species
lacks the maxillary, metasubalar and scale spe-
cializations shared by all other pterophorids. The
moths are superficially similar to agdistines (Fig.
13.2 P). One species is known to feed in the fruit
of Secamone (Asclepiadaceae) (Common 1990).
The remaining subfamilies constitute a mono-
phylum characterized by loss of the maxillary
palp, by having the metathoracic subalare mark-
edly narrowed and by the presence of specialized
scales (believed to be scent scales) along M3/
CuA on the hindwing underside (Fig. 13.1 J).
Agdistinae. Monogeneric. Agdistis (Holarctic/
Oriental, (Fig. 13.2P) with 80 species comprises
the bulk of the 'non-plumed plume moths', char-
acterized by a sparsely scaled triangular area ex-
tending inwards from forewing termen. Male
genitalia asymmetrical as in many pterophori-
nae; this is believed to be a homoplasy (Gielis
1993).
Larvae leaf feeding, recorded from several di-
cot families.
Ochyroticinae. Monogeneric. The pantropical
Ochyrotica with 22 species are 'non-plumed'
pterophorids. In male genitalia sacculus with
spine directed distad, and cucullus (apparently
secondarily) undifferentiated. In many species
forewing and abdomen strongly patterned (more
so than in O. concursa illustrated in Fig. 13.2 Q).
Recorded hosts are Convolvulaceae, Vitaceae
and Leguminosae.
Deuterocopinae together with the following
subfamily probably constitute a monophylum
characterized by cleft wings and double teeth (in
the ground plan, reduction to one is common-
place within both subfamilies) of enlarged scales
on the third hindwing lobe. The deuterocopine
autapomorphy is the presence of two, or even
three, forewing clefts. Male genitalia symmet-
rical.
Pantropical, with 30 species in 4 genera. Deut-
erocopus (19 species, Fig. 13.2R) is mainly Indo-
Australian, Leptodeuterocopus (8 species) mainly
neotropical. Recorded hosts are Leeaceae and
Vitaceae.
Pterophorinae. This is by far the largest ptero-
characterized by the hindwing venation (Fig.
nates in the bottom of the second cleft instead
of extending onto the hindmost wing lobe (Fig.
13.1 M); this assemblage was found in Gielis'
analysis to be paraphyletic. A groundplan auta-
pomorphy of the Pterophorinae as presently de-
limited is the double signum in the bursa copu-
latrix of the female.
852 species in 66 genera, largest among which
are the cosmopolitan Hellinsia (= Leioptilus; 197
species) and Platyptilia (97 species), the similarly
cosmopolitan Stenoptilia (67 species) and the Old
World Pterophorus (58 species). While Asteraceae
are the principal hosts, the total host spectrum
of the subfamily comprises some twenty system-
atically scattered dicot families. The majority of
pterophorines have herbaceous hosts, but some
do feed on woody plants; thus recorded hosts of
the remarkably widespread tropical/subtropical
Megalorhipida defectalis include woody Fabaceae
(Acacia, Mimosa) in addition to Amaranthaceae
and Nyctaginaceae. The known larvae in the
small genus Buckleria are remarkable for feeding
on sundews (Droseraceae). A few species are mi-
nor horticultural pests, such as the N. American
artichoke-feeding Platyptilia carduidactyla.
Lange (1950), Yano (1963), Wasserthal (1970),
Kuznetzov & Stekolnikov (1979 b), Gielis (1991,
1993); the extensive literature on W. Palaearctic
pterophorids is being summarized in a manual
by Gielis (1996).
Superfamilies Assigned to Apoditrysia-
Obtectomera
Members of this clade are characterized by un-
spined pupae which have abdominal segments
I - I V (at least) immobile, and are not protruded
from the pupal shelter prior to eclosion (excep-
tions are encountered in a few Macrolepidoptera,
but are tentatively interpreted as character rever-
sals). Moreover, the adult moths have specialized
pretarsal pulvilli. The pupal characteristics are
encountered also in a number of ditrysian lin-
eages, which are excluded from the Obtectomera,
The Smaller Microlepidoptera-Grade Superfamilies 225

Fig. 13.3. Copromorphidae: A; Carposinidae: A - ; Hyblaeidae: F; Thyrididae: G - H . A, Copromorpha venation;

B, Carposina venation; C, Heterogymna head; D, "Bondia" larval head; E, Carposina sternum II; F, Hyblaea
venation; G, Striglina venation; H: Thyris abdominal base lateral (arrow: postspiracular lateral lobe). AB, FG
after Nielsen & Common (1991); C after Robinson et al. (1994); D after Heppner (1987): E redrawn after Kyrki
(1983); H original (B. Rubaek del.).

since they do not have the pulvillus apomorphy.
See 1 - 2 for further details.
COPROMORPHOIDEA
The content of a superfamily centred on the
Copromorphidae has varied considerably. In
particular, copromorphoids have often been
merged with alucitoids and epermenioids with
which they share specializations in larval and pu-
pal structure. Minet (1991) considered these sim-
ilarities homoplasious and excluded the lastmen-
tioned taxa from the clade Obtectomera, to
which the Copromorphoidea were assigned. See
1-2.
The superfamily as here delimited comprises
only the Copromophidae and Carposinidae, and
the distinction between the two families is not
well supported.
Probable autapomorphies according to Minet
(1986): forewing with raised scale-tufts and
hindwing with cubital pecten (basal fringe of pili-
form scales); in some taxa the latter is developed
in the female only, and occasionally it is absent
in both sexes. Anterolateral process on sternum
II usually long and curved (Fig. 13.3 E; Kyrki
1983); near-absence in some genera probably sec-
ondary. Phallus with coecum. Larva usually with
spiracles slightly protruding and those on VIII
largest and most dorsal (Common 1990); these
traits may also be groundplan autapomorphies
of the superfamily.
Small to medium-sized micro-moths, wing-
span 10 (exceptionally) 50 mm. Facial scales
smooth, all downwards pointing; lateral scale
tufts on vertex more or less raised; ocelli present
or absent; chaetosemata absent; antennae vari-
able, scape without pecten. Maxillary palp 1 4
segmented; proboscis unsealed. Labial palp
prominent, upturned or porrect. Epiphysis pre-
sent, spurs 0-2-4; hind tibia smooth or with long
roughened piliform scales above. Forewing pter-
ostigma variably developed; chorda and M-stem
at most faintly indicated; spinarea present; fe-
male frenulum of 23 bristles. Forewing with all
veins separate, or with Rs2/Rs3, Rs3/Rs4 or M3/
CuAl stalked; CuP distinct near margin (and
sometimes discernible to base), or greatly re-
226 John S. Dugdale, Niels P. Kristensen, Gaden S. Robinson & Malcolm J. Scoble
duced. Hindwing M3 separate, connate or
stalked with CuAl; CuP present. Male with
paired coremata near posterior end of abdomen,
sometimes also in anterior part. Copromor-
phoids are relatively broad-winged moths with
some superficial resemblance to tortricoids
(which, in addition to the lack of corpromor-
phoid autapomorphies, differ in having the lower
facial scales upwards pointing and in the more
proximal original of the forewing CuA2) and im-
moids.
Larva may have well developed processes on
postlabium or stipes (Fig. 13.3 D). Prothoracic
L-group bisetose. LI and 2 on same pinaculum
in carposinids and some 'copromorphids', but on
separate pinacula in other 'copromorphids'
(Heppner 1987, Common 1990). Prolegs crochets
in uniserial (or, in some 'copromorphids') partly
biserial circle or mesal penellipse. Pupal integu-
ment pallid, flimsy; head and thorax angular in
outline; segments VVII (male) and V - V I (fe-
male) movable; tip with clusters of hooked setae.
Adults nocturnal. Larvae mostly concealed
feeders, borers or living between joined leaves or
within/amongst fruits. Pupation in larval gallery
or away from host plant, in a cocoon usually
covered with debris.
Note. The implications of the aberrant New
Zealand Isonomeutis for the copromorphoid
ground plan are uncertain. This genus, hitherto
placed in the 'Copromorphidae', does have the
peculiar 'flimsy' pupal cuticle characteristic of
the Copromorphoidea, but its sternum II has
both anterior and posterior tergosternal bars
arising from the anterior (lateral) arm and en-
closing spiracular area; the phallus devoid of a
coecum, the unmodified larval postlabium and
the trisetose prothoracic L-group do not accord
with those of other Copromorphidae. The larva
is predatory, feeding on Coccoidea-Margarodi-
dae (Hemiptera). Isonomeutis may be erron-
eously placed in the copromorphoid lineage, or
it could be the sister group of all other members
of the superfamily.
Heppner (1987), Common (1990).
Copromorphidae. Those copromorphoid genera
which lack specializations characterizing the
Carposinidae. We here retain the family distinc-
tion with express reservations, pending a more
detailed inquiry.
Most, but not all, 'Copromorphidae' are cryp-
tically coloured moths with more or less rounded
forewings. Antennae sometimes simple, mostly
thickened or unipectinate (sometimes even bipec-
tinate) in males. Maxillary palp primitively 4-
segmented, folded over proboscis base; may be
reduced to single segment. Labial palp upturned
or porrect; segment 2 longest. Hindwing with all
three M branches (Fig. 13.3 A).
40 + species from all regions except the Pa-
laearctic (Heppner 1991) are currently placed in
this family, more than half belonging to the
Indo-Australian Copromorpha (Fig. 13.4B). Lar-
val biology diverse. Copromorpha and Phyco-
morpha comprise a number of species which are
twig/inflorescence borers, or fruit or leaf eaters
(between joined leaves) on Moraceae (fig family);
other recorded host families include Ericaceae
and Berberidaceae. Sisyroxena from Madagascar
is remarkable for its autapomorphic venation,
with minute discal cells, and prominent, aligned
scale sockets.
Carposinidae. Probably cladistically subordinate
in preceding family. Autapomorphies: Hindwing
M2 (often also M l ) absent (Fig. 13.3 B). Antenna
(sometimes unipectinate) densely set with long
setae in male, setae shorter in female. Male with
conspicuously elongated phallic coecum (Com-
mon 1991).
Forewings less rounded than in most 'copro-
morphids'. Ocelli usually absent, if present then
reduced. Maxillary palp 1 - 3 segmented (when
1-segmented, occasionally with a tiny terminal
vestige of a second segment). Labial palp in
males usually somewhat upcurved, with segment
3 porrect, in female porrect and particularly
long; both sexes have enlarged scale covering on
segment 2 (Fig. 13.3 C). Male wings may bear
patches of modified (presumably scent) scales on
upper and underside.
Larvae modified for endophytic life, in flow-
ers, fruits, buds, shoots, or in galls or other
wound-reaction tissue on living plants; 20 + host
plant families have been recorded. Pupation in
the larval shelter or in a cocoon on the ground.
273 species in 20 + genera (Heppner 1987,
1991); worldwide, but not in NW Palaearctic. A
very large proportion is included in Carposina
(Fig. 13.4C), which is predominantly Indo-Aus-
tralian, but includes a couple of garden and or-
chard pests in the Holarctic. Other important
genera are Bondia (Australian; B. ' comonana is
a N. American Prunus pest), Heterocrossa, Meri-
darchis, Peragrarchis, Paramorpha, Coscinop-
tycha. Host plants have been recorded from Po-
docarpaceae as well as several disparate dicot
families including, i. a., Fagaceae, Ericaceae, Ro-
saceae, Myrtaceae, Proteaceae, Rutaceae, Cam-
panulaceae and Asteraceae.
Davis (1969), Kuznetzov & Stekolnikov
(1979 b), Diakonoff (1989).
IMMOIDEA
The taxonomic history of the Imma group is sim-
ilar to that of Choreutoidea in that it was origi-
nally included in the Glyphipterigidae, later
transferred as a separate family, Immidae,
(Heppner 1977) to the Sesioidea and finally given
status as a separate superfamily Immoidea
(Common 1979).
The Smaller Microlepidoptera-Grade Superfamilies
Immidae. Family autapomorphies of adults have
not so far been worked out, but the following
are potentially significant and should be checked
for consistency in the generic groups recognized
by Heppner (1982): ovipositor lobes fused dor-
sally into a single, bent disc, with strong setae
confined to the sclerotized rim; male segment
VIII with tergum and sternum anteriorly fused,
forming a slender, looped structure; the few
known larvae are unique among non-macrolepi-
dopteran 'Obtectomera' in having the proleg
crochets (which are uniordinal) in a mesoseries.
Adults are small to medium-sized (wing-span
ca. 1245 mm), usually round winged moths.
Head with smooth, mesally directed scale tufts
above, face with long rough scaling pointing up-
wards, or short smooth scaling pointing down-
wards (sometimes pointing upwards on lower
face area); ocelli or absent; chaetosemata usually
present. Antenna filiform, usually short-ciliate in
male; scape without pecten. Maxillary palp min-
ute, 1 - 2 segmented, ventrally directed; probos-
cis unsealed. Labial palp with segment 2 ascend-
ing, with scaling more or less erect, and ending
on a level with dorsal margin of compound eye;
segment 3 either short, smooth-scaled, or re-
duced to a minute remnant hidden by the apical
scaling of segment 2, where not reduced it has
the cup-shaped vom Rath's organ at half-length;
males may have an apical or subapical process
(lobe, hook) on segment 2. Epiphysis present;
spurs 0-2-4; hind tibiae smooth scaled, with tufts
around spurs. Wings smooth-scaled; forewing
pterostigma, when present, bounded by Sc;
chorda vestigial; M-stem absent; spinarea pre-
sent; Rs3 and 4 often stalked and both postapi-
cal; CuA2 arising near end of cell; CuP distinct
near margin. Hind wing sometimes with hyaline
patch; CuP present near margin; 3 A present.
Male sometimes with coremata near abdominal
end; genitalia quite variable. Female ovipositor
lobes sometimes floricomous; antrum floor usu-
ally scaled; ductus bursae often multicoiled. Im-
mid moths are most likely to be confused with
tortricids (which have the apical labial palp seg-
ment shorter and blunter, forewing Rs3 preapical
and CuA2 arising from before 4/5 discal cell
length), copromorphoids (which usually have
raised scale tufts on forewing, a scale pecten on
the hindwing CuA and the forewing Rs3 preapi-
cal), or small noctuoids (which have a metathor-
acic tympanum and no hindwing CuP).
Known larvae feed externally on leaves of co-
nifers and a wide spectrum of dicot families; usu-
ally cryptic, sometimes exposed and aposematic
{Birthana species on podocarps). Some dorsal
head and trunk setae barbed; prothoracic L
group trisetose (a rare condition within Obtec-
tomera); LI and 2 on segments IVIII on sepa-
rate pinacula, more or less horizontally aligned
below spiracles; prolegs slender. Pupa with (at
least) prothoracic spiracle protruding, with sharp
227
process; segments V - V I movable; abdominal
apex with up to 16 hooked setae; pupation in
loose, sometimes net-like cocoon. When dis-
turbed larvae of some Imma species 'catapult'
frass in the general direction of the disturbance.
246 species, pantropical (Heppner 1991). The
bulk of the species are in just two genera, Moca
(ca. 50) and Imma (150 +, Fig. 13.4E). The Indo-
Australian Bursadeila and Birthana contain
large, robust and conspicuously coloured species,
some of which (Fig. 13.4D) may mimic Noctui-
dae-Aganainae.
Common (1979), Heppner (1982 a).
HYBLAEOIDEA
As noted by Minet (1983) no previous superfam-
ily assignment of the Hyblaeidae is known to be
well supported, so a placement in a superfamily
of their own is appropriate at present. Hyblaeid
larvae have a capacity of regurgitating gut
contents in squirts of several centimetres upon
disturbance. Since defence by regurgitation is
also known from some members of another fam-
ily among the 'basal Obtectomera', viz., the
Thyrididae, a search for a possibly synapomor-
phic gut specialization is very desirable.
Hyblaeidae. Probable autapomorphies: male
hindleg modified, with large concave coxal pro-
cess into which a tibial hair-pencil fits, tarsi
strongly spined beneath. Metathoracic secondary
furcal arms strongly enlarged. Forewing A veins
not anastomosed, CuP absent.
Large, robust and broad-winged micro-moths
(Fig. 13.4 A), which have some superficial sim-
ilarity to small catocaline noctuids. Head
smooth-scaled; ocelli present; chaetosemata ab-
sent; antennae filiform. Maxillary palp porrect,
34 segmented and densely scaled; proboscis un-
sealed. Labial palp porrect. Epiphysis present;
spurs 0-2-0 (male)/4 (female). Forewing (Fig.
13.3 F) without pterostigma, chorda and M-
stem; no stalked veins in either wing pair; fore-
wing Rs4 postapical; hindwing Sc/R/Rs configu-
ration variable, CuP vestigial near base. Sternum
II with apodemes prominent and clubbed; no
venulae or sternotergal sclerites.
Egg flat. Larvae leaf feeding on Verbenaceae,
Rhizophoraceae, Myrtaceae and Bignoniceae,
sheltering in tied leaves. Prothoracic L bisetose;
proleg crochets in triordinal (or partly tri-, partly
biordinal) circle. Pupa in tough cocoon, with
segments VVI movable in both sexes; cremas-
ter with cluster of setae with bulbous hooked
tips.
18 species in 2 genera. Hyblaea (17 described
species) ranges through the Old World tropics;
H. puera is a serious defoliator of Tectona, Vitex,
Catalpa and other trees in the Oriental region,
and it has been introduced into the New World.
Torone (monobasic) is Neotropical.
228 John S. Dugdale, Niels P. Kristensen, Gaden S. Robinson & Malcolm J. Scoble

w
G H I

Fig. 13.4. Hyblaeidae: A; Copromorphidae: B; Carposinidae: C; Immidae: D - : Thyrididae: F - H ; Whalleyani-

dae: I. A, Hyblaeaavifasciata; , Copromorpha eforescens; C, Peragrarchis sp.; D, Birthana saturata; E, Imma
albofasciala\ F, Arniocera cyanoxantha; G, Dysodia rajah; H, Siculodes triumphans; I, WJwlleyana vroni. Uniform
magnification, scale (mm) in A.

Koning & Roepke (1949), Singh (1956), Naka-
mura & Iwata (1969), Common (1990).
THYRIDOIDEA
The Thyrididae have often been included in Pyr-
aloidea, but no synpomorphies have been iden-
tified to support this assignment (Minet 1983).
Unlike Pyraloidea, Thyridoidea lack abdominal
tympanal organs and the proboscis is without
basal scaling. The group may be related to
Hyblaeidae (see above), but this suggestion re-
quires further investigation.
Thyrididae. Probable autapomorphies (Minet
1991) include: tergum I with very large latero-
ventral lobes immediately behind the spiracle
(Fig. 13.3 H); resting posture with mid legs not
touching substrate; spinarea absent; hindwing
with CuP vestigial. Adults rest in a distinctive
position with the body steeply raised and the
wings held outstretched. Larva with only two L
setae occurring on either side of all thoracic seg-
ments (non-trisetose condition on T2 and T3 is
exceptional in Lepidoptera, Hasenfuss 1980).
Medium to large microlepidopterans exhibit-
ing considerable phenetic heterogeneity. Wing-
span 1272 mm. External ocelli rarely present,
when occurring small. Chaetosemata absent.
Frons occasionally extended into prominent pro-
cess. Cephalic vestiture appressed. Compound
eye sometimes bearing interfacetal hairs. An-
tenna typically minutely ciliate, but may be cili-
ate, dentate, or pectinate. Maxillary palpi min-
ute; proboscis often reduced or even vestigial
but, when present, lacking scales. Labial palpi
usually 3-segmented, occasionally 2-segmented.
Epiphysis rarely absent. Tibial spur formula 0-2-
2, 0-2-3 or 0-2-4. Hair pencil sometimes occur-
ring on hind tibia of male. Wings in 'typical thyr-
idids' (Charideinae excepted) broad; pattern usu-
ally similar on both forewing and hindwing;
often reticulate both on upper surface and under
surface, and with hyaline spots or patches; often
brown to red-brown; many species mimic dead
leaves. Venation (Fig. 13.3G) little reduced.
Forewing Rsl Rs4 usually arising indepen-
dently from the cell, sometimes Rsl and Rs2 or
Rs2 and Rs3 stalked; lower branch of anal loop
variably developed, sometimes very short.
Hindwing: Se + RI usually approaching Rs part
way along cell, but occasionally veins fusing for
a short distance. Spinarea generally absent, but
apparently (Kuijten 1974) present in some spe-
cies of Thyris. Frenulum very rarely present. Ovi-
positor short. Male genital musculature: Kuznet-
zov & Stekolnikov 1979 a.
Eggs upright, ribbed. Larva semiprognathous
to hypognathous. Secondary setae present (re-
corded from Thyris by Hasenfuss) or absent.
The Smaller Microlepidoptera-Grade Superfamilies
Proleg crochets uni- to biordinal, on IIIVI ar-
ranged in circles or broad ellipses. Concealed
feeders living within stems or in rolled or tied
leaves; some inducing formation of galls (Hasen-
fuss 1979, 1980; Neunzig 1987; Common 1990).
Recorded host plants belong to 30 families,
prominent among which are Euphorbiaceae, Le-
guminosae, Sapotaceae, Myrtaceae, Dipterocar-
paceae, Capparidaceae and Combretaceae.
Thyrididae are mainly lowland forest-dwellers
in the tropics and subtropics (Robinson et al.
1994). There are about 760 described species, and
a further 4 5 0 + species and subspecies are un-
named in the collections of the Natural History
Museum, London (M. Shaffer, pers. comm.).
Key to subfamilies ( modified from Whalley 1971 )
1 Tarsi with apical pairs or rows of spines 2
Tarsi without spines . . Siculodinae (part)
2(1) Tarsi with apical pairs of spines only . . 3
Tarsi with rows of spines on each segment
4 Alberti (1954).
3 (2) Male genitalia with small process on tegu-
men in front of uncus; valves simple. Fe-
male with secondary sac on bursa. Stout-
bodied moths; hind wing often with large
translucent areas Thyridinae
Male genitalia without process on tegu-
men, often complex, with bifid or other-
wise modified uncus; valves variously mod-
ified. Females usually without secondary
bursal sac. Hind wing usually without
large translucent areas Striglininae (part)
4 (2) Narrow-winged moths with Zygaena-like
facies, colours mostly aposematic, metallic
Charideinae
Moths not pronouncedly narrow-winged,
wing pattern cryptic, shades of brown . . 5
5 (4) Hind wing often with Se + R and Rs anas-
tomosing for a short distance. Male genita-
lia complex, with bifid or otherwise modi-
fied uncus Striglininae (part)
Hind wing with Sc + R and Rs free. Male
genitalia relatively simple, uncus always
single Siculodinae (part)
Striglininae are pantropical. Unlike in the
following subfamilies, the metathoracic meral
sclerotization is not produced posterodorsally
(Minet 1991). About 200 species; major genera
include Striglina, Aglaopus and Banisia (Whal-
ley 1976).
Thyridinae (= Pachytyrinae). Mostly stout-
bodied. Pantropical with a few genera such as
the small diurnal Thyris extending into the Hol-
arctic. About 115 species; principal genus Dy-
sodia (Fig. 13.4G). The oriental Glanycus are
aposematic, and probably distasteful; the larva
of G. tricolor stridulates when disturbed, appar-
ently using its thoracic legs (Li in Robinson
et al. 1994).
229
Siculodinae including the tribes Siculodini
(Fig. 13.4H), Argyrotypini and Rhodoneurini,
are also widely distributed, but the subfamily
may not be monophyletic. At least some siculod-
ines have a modified forewing base which may
be an ear (Minet 1983, 1991, but see the caveat
by Yack & Fullard 1994). 420+ species; major
genera include Hypolamprus, Collinsa and Rho-
doneura.
Principal references on 'typical thyridids' are
Whalley (1971, 1976).
Charideinae. Diurnal narrow-winged moths
with little resemblance to 'typical thyridids'.
With their metallic, aposematic colours they
have a marked superficial resemblance to some
Zygaenidae, and they were long included in this
family from which they were transferred by Mi-
net (1991). Immature stages undescribed, but an
Arniocera larva is recorded from Grewia (Tilia-
ceae) (Sevastopulo, MS information in Nat. Hist.
Mus. moth host database). Afrotropical, 20 spe-
cies. Principal genera Arniocera, Marmax.
WHALLEYANOIDEA s u p e r f a m . nov.
Minet (1991) assigned the small moth genus
Whalleyana to a family of its own, since no good
reasons for its previous placement in the Thyridi-
dae could be found. While the placement of the
family in the Obtectomera seems reasonably
founded it remains uncertain whether its closest
relationships are with some 'micro-moth-grade'
lineage or with the Macrolepidoptera. As with
the Galacticidae and Simaethistidae (above) a
new superfamily for the Whalleyanidae is here
erected for formal classificatory reasons.
Whalleyanidae. Whalleyana comprises 2 species.
The somewhat arctiid-like moths (Fig. 13.41)
have greatly reduced ocelli, but large contiguous
chaetosemata (with short scales and long setae)
in the posterior vertex. Antennae in both sexes
scaled dorsally, setose ventrally. Flagellum in dis-
tal section curved (as in Callidulidae). Maxillary
palp short, 3-segmented; proboscis unsealed. La-
bial palps with segment 1 long and arched, 2
short and 3 thick, rather short. Epiphysis pre-
sent; hind spur complement different in the spe-
cies: 0-2-4/2; pulvillus bilobed. Forewing base in
one of the two known species with first axillary
caudally elongated towards median notai wing
process; in the other species this specialization is
not evident. Spinarea absent; female frenulum
with several bristles. Forewing with all Rs
branches stalked; Rs3 and Rs4 postapical; lower
branch of anal loop very slender. CuP absent in
both wings. Hindwing Se + R approximated to
Rs beyond cell; 3 A slender, but discernible. Ster-
num II with apodemes reduced; complete prespi-
racular bar present between tergum I and ante-
230 John S. Dugdale, Niels P. Kristensen, Gaden S. Robinson & Malcolm J. Scoble
rior corner of sternum II; tergum I with slightly
produced lateral lobe far behind spiracle I.
While the loss o f the spinarea and the hind-
wing C u P could support the original placement
o f Whalleyana in the Thyrididae, it is true that
these simple, regressive features are very h o m o -
plasious within the Lepidoptera and carry little
weight. The presence of well developed chaeto-
semata (with scale mixing, as in some Macrolepi-
doptera) is incompatible with a subordination in
the Thyrididae, and the small posterolateral
lobes o f tergum I (similar to formations which
are recurrent in the Ditrysia) are very different
from the large lobes, which are the m o s t impres-
sive thyridid autapomorphy. A placement as the
closest relatives of the Thyridoidea or Pyraloidea
remains a real possibility, and is compatible with
the hindwing Sc + R / R s configuration and
ridged scales. On the other hand the posterior
elongation o f the first axillary sclerite in one o f
the species might be taken as evidence for a
placement within the Macrolepidoptera, but the
absence of this trait in the other species renders
its phylogenetic significance problematical:
which o f the states pertains to the whalleyanid
ground plan?
The genus is confined to Madagascar, and the
immatures are unknown.
Viette (1977), Minet (1991).
Acknowledgments
The section o n Galacticidae was based originally
on a draft kindly provided by Dr. D . R. Davis.
We are indebted to Mr. O. Karsholt for useful
c o m m e n t s o n the manuscript.
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moths of the genus Platyptilia in California. -
Hilgardia 19: 561-668.
Meyrick, E. (1910): Lepidoptera Heterocera. Fam. Or-
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Minet, J. (1983): tude morphologique et phylogn-
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231
tions of its larva and pupa. Trans. Lep. Soc. Jap.
XIV, 3: 5 2 - 5 9 .
- (1977): Fauna Japnica. Yponomeutidae s. lai. (In-
serta: Lepidoptera). 327 pp., 95 pis., Tokyo.
Nakamura, M. & Iwata, K. (1969): Larva and pupa
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Heterocerists' J. 55: 619-622.
Neunzig, H. H. (1987): Hyblaeidae (Pyraloidea). Thyr-
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Norberg, R. . (1972): Flight characteristics of two
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Scoble, M . J . (1992): The Lepidoptera. Form, function
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- Indian Forest Ree. (. S., Ent.) 9: 1 - 1 6 .
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Morph. Tiere 77: 127-155.
Whalley, P. E. S. (1971): The Thyrididae (Lepidoptera)
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Suppl. 17: 1 - 1 9 8 , 68 pis.
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Yack, J. E. & Fullard, J. H. (1993): What is an insect
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Addendum
T h e g e n u s Tanaoclena f r o m A u s t r a l i a , N e w Z e a -
l a n d a n d R a p a is i n c l u d e d in t h e G a l a c t i c i d a e
in Nielsen et al. (1996). T h e larvae, e n d o p h y t i c
R u b i a c e a e feeders, h a v e the p r o t h o r a c i c L biset-
ose a n d s e p a r a t e f r o m the p r o n o t a l shield; seta
D 2 is e n l a r g e d a n d d a r k e n e d o n III, V, V I I - I X .
232 John S. Dugdale, Niels P. Kristensen, G a d e n S. Robinson & Malcolm J. Scoble
The Palaearctic Pterophoridae-Agdistinae,
-Ochyroticinae and -'Pterophorinae' (as pre-
viously delimited) were monographed by Aren-
berger (1996).
A placement of the Simaethistoidea in the
Apoditrysia is indicated by the structure of the
adult's sternum II (Minet pers. comm.).
Arenberger, E. (1996): Pterophoridae [1]. Microlepi-
doptera palaearctica 8: 258 pp. + Plates volume
(153 pis.).
Nielsen, E. S., Edwards, E. D. and Rangsi, T. V. (eds.)
(1996): Checklist of the Lepidoptera of Australia.
- Monographs on Australian Lepidoptera 4, xiv +
529 pp. C S I R O Publishing, Collingwood.
 14. The Pyraloidea
Eugene Munroe and M. Alma Solis
The Pyraloidea are one of the larger superfami-
lies of Lepidoptera. Their two families, Pyralidae
and Crambidae, comprise about 16000 named
species worldwide; probably at least as many
more remain to be described. Linnaeus (1758)
recognized the Pyraloidea as a subgenus, Pyralis,
of his comprehensive moth genus Phalaena;
however, he included a number of pyraloid spe-
cies in the subgenus Geometra, and members of
narrow-winged subfamilies such as Crambinae
and Phycitinae in the subgenus Tinea. Subse-
quent authors such as Fabricius, Cramer, Stoll,
Denis & Schiffermller, Hbner, Curtis, Ha-
worth, Treitschke, and Stephens described many
additional genera and species, and the group
achieved the rank of family or group of families,
though with little agreement or discussion as to
limits or subdivisions. Herrich-Schffer (1843
56) showed that deltoids and nolids belonged
with noctuids rather than with pyralids, and he
associated crambines and phycitines with the lat-
ter. Lederer (1863) confirmed these findings, and
reviewed the work of previous authors, with em-
phasis on that of Herrich-Schffer (op. cit.),
Guene (1845, 1854), and Walker (1859). Mey-
rick (1882 et seq.), Ragonot (1891-92, 1893),
and Ragonot and Hampson (1901) built on Led-
erer's foundation, and Hampson (1895, 1896a,
1896 b, 1899 a, 1899 b, 1917) developed the ideas
of these three authors into a comprehensive clas-
sification that stood virtually unchallenged until
the period following World War II, and that until
very recently has been adopted in almost all sys-
tematic lists and collections. Forbes (1920, 1923,
1926) made interesting suggestions on classifica-
tion. Brner (1925) was the first to point out the
significance of the tympanal organs in dividing
the Pyraloidea into two major series, though
presence or absence of the praecinctorium had
already been used in a haphazard way by
Guene (1854) in the definition of groups.
The definition of Pyraloidea has narrowed
considerably over the past few decades; classi-
cally the Pyraloidea included Pterophoridae,
Alucitidae, Thyrididae, Hyblaeidae, Oxychiroti-
dae, and Tineodidae (reviews in Fletcher & Nye
1984; Minet 1986; Nielsen 1989; Common 1990).
Minet (1986) also included the Dudgeoneidae,
but later he withdrew this placement (Minet
1991). At present the Pyraloidea are defined as
ditrysian moths with abdominal tympanal or-
gans consisting of paired tympanal chambers on
the ventral part of segment two, supporting a
tympanum and conjunctiva, the former with a
sensory scoloparium attached (Figs. 14.2, G - J ) ;
in addition, the maxillary palpus is usually pre-
sent (Fig. 14.1 D), the proboscis, if present, is
scaled at its base (Fig. 14.1 E), R3* and R4 of the
forewing are stalked or fused (Fig. 14.2 A), and
Se + R| and Rs of the hindwing are anasto-
mosed or approximated for some distance distad
of the discal cell (Fig. 14.2 B). This definition ex-
cludes all the above-named families, although a
few pyraloids have tympanal structures reduced
or absent (Clarke 1971; Minet 1991; Solis in
prep.)
The Thyrididae resemble some Pyraloidea in
general appearance and in having only two setae
on the prespiracular tubercle of the larval pro-
thorax. For these reasons authors such as Forbes
(1923), Whalley (1971, 1976), and Fletcher & Nyl
(1984) placed them in the superfamily. However,
the Thyrididae differ in fundamental character
states of high fidelity, such as the absence of ab-
dominal tympanal organs and of scaling on the
base of the proboscis, as well as in having the
maxillary palpi greatly reduced, not distinctly de-
veloped as in most true pyraloids; also the thyri-
dids have a reticulate wing pattern of a primitive
type, whereas only vestiges of this are seen in
pyraloids. Accordingly both Whalley (op. cit.)
and Munroe (1978) expressed reservations as to
the placement of the the Thyrididae in the Pyra-
loidea. Similarly Forbes (1933) associated the
Hyblaeidae with the Pyraloidea, largely on tho-
racic and pupal characters. However, these char-
acters were contested by de Koning and Roepke
(1949) and Singh (1955). Traditionally hyblaeids
had been placed in the Noctuidae, but hyblaeids
have well-developed maxillary palpi like those of
pyraloids, whereas maxillary palpi are absent in
true noctuids. However, hyblaeids lack noctuoid-
like thoracic or pyraloidlike abdominal tympanal
organs, and the proboscis lacks basal scaling.
Minet (1981) also separated them as a superfam-
ily. In Forbes' classification the Pterophoridae
and Alucitidae were also placed in the Pyra-
loidea. The latter have little resemblance to the
pyraloids, but the Pterophoridae often have Sc
+ Ri of the hindwing anastomosing with Rs as
in the Pyraloidea, and in the larva they often
have two prespiracular setae, as in Pyraloidea
and Macrolepidoptera; however, the proboscis is
naked at the base, and the maxillary palpi are
reduced. Common (1970) recognized the Ptero-
phoroidea as a distinct superfamily, to which
Munroe (1982) transferred the Tineodidae and
* The vein notations R 2 - R 5 in this chapter equal
R s l - R s 4 used elsewhere in this volume. Editor
234 Eugene Munroe and M. Alma Solis

Oxychirotidae. The latter two families had been Table 1: Summary of families and subfamilies included
placed in the Pyraloidea by Common (1970). Mi- in the Pyraloidea.
net transferred the Oxychirotidae (Minet 1986) PYRALIDAE
and the Tineodidae (Minet 1991) to the Aluci- Galleriinae Zeller
toidea. Finally, Clarke (1971) proposed the fam- Chrysauginae Lederer
ily Lathrotelidae, on the basis of a single female Pyralinae Latreille
from Rapa in French Polynesia. This specimen Epipaschiinae Meyrick
had pyraloid characters except for the absence of Phycitinae Zeller
the tympanal organs. There was some question CRAMBIDAE
as to whether the Lathrotelidae of Clarke (1971) Scopariinae Guene
are Pyraloidea (Nielsen 1989; Common 1990), Heliothelinae Amsel
but Minet (1991) dissected a male, which proved New Synonymy
to have reduced tympanal organs of nymphuline Crambinae Latreille
type. Accordingly, he sank Lathrotelidae as a Schoenobiinae Duponchel
synonym of Nymphulinae. Cybalomiinae Lederer
Linostinae Amsel
On the basis of morphology of the abdominal Midilinae Munroe
tympanal organ, the Pyralidae (sensu lato) were Musotiminae Meyrick
divided by Munroe (1972) into three groups: Nymphulinae Duponchel 1
Pyraliformes, Crambiformes, and Midiliformes. Noordinae Minet
However, Minet (1983), in a broad study of lepi- Odontiinae Guene
dopteran tympanal organs, showed that those of Wurthiinae Roepke
Midilinae are of the crambiform type, though re- Evergestinae Marion
duced. Accordingly, he separated Pyralidae Glaphyriinae Forbes
Dichogaminae Amsel
(sensu lato) into two sister families, Pyralidae
New Synonym
(sensu stricto), corresponding to Pyraliformes,
Alatuncusiinae Amsel
and Crambidae, comprising Crambiformes plus New Synonym
Midiliformes of Munroe (1972). We follow this Cathariinae Minet
division (Table 1) and the terminology of tympa- Pyraustinae Schrank
nal organs used by Minet & Surlykke (2-11). Spilomelinae Guene Revised
We note that some lepidopterists do not recog- Synonymy
nize the two monophyletic groups as families. We
1
consider that the morphological and cladistic An application has been made by M. Alma Solis (in
evidence for the monophyly of the Pyralidae and press) to the International Commission on Zoological
Crambidae and their sister group relationship is Nomenclature to conserve the name Nymphulinae in
preference to Acentropinae.
clear enough to warrant their separation at the
family level.
Authors have differed somewhat in their clas-
sifications of the subfamilies of the Pyraloidea. made in the study of pyraloid relationships and
Though efforts were made by Herrich-Schffer classification. Among the more important works
(1849), Guene (1845, 1854) and Lederer (1863) in this period are: Sylvn (1947); Heinrich (1956);
to develop natural classifications, these were not Marion (1952, 1957a, 1957b, 1953-1976):
based on evolutionary concepts. The first such Munroe (1958 b, 1959, 1961,1964 a, 1964 b, 1970,
was that of Meyrick (1879, 1882 a, 1882 b, 1972-1976); Common (1990); Roesler (1973,
1884 a, 1884b, 1885 a, 1885b, 1885 c, 1890,1895). 1978); Gaskin (1975 a, see other references under
Initially Meyrick used characters of the male Crambinae); Kuznetzov & Stekolnikov (1979);
genitalia as well as external characters, but un- Minet (1981, 1983, 1985); Solis & Mitter (1992);
fortunately he did not continue this practice in Maes (1985, 1995 b); Landry (1995); Munroe
his later work. Ragonot (1891a, 1891b, 1892, (1995); Shaffer, M. & Solis (1995); Solis, Becker,
1893), Ragonot & Hampson (1901), and Hamp- Munroe (1995); Solis (1995); J. C. Shaffer (1995);
son (1895, 1896a, 1896b, 1897a, 1897b, 1899a, ShafTer, M., Nielsen & Horak (1996). Roesler
1899 b, 1917) developed and elaborated Mey- (1973) presented a phylogeny based on adult and
rick's classification. Hampson in particular used immature characters. Kuznetzov & Stekolnikov
what purported to be phylogenetic principles (1979) proposed one based on functional mor-
throughout this work, though injudicious selec- phology of the male genitalia. Minet (1981, 1985)
tion of characters and defects of methodology suggested subfamily autapomorphies of the ab-
led him into many mistakes. Nonetheless, no ac- dominal tympanal organs. Yoshiyasu (1985) pre-
ceptable substitute for this classification emerged sented his concept of relationships among Japan-
for nearly 100 years, and it still is the arrange- ese crambid subfamilies on the basis of charac-
ment of most museum collections in the world. ters of adults and immatures. Roesler (1973) and
It is only since World War II, with the wide- Minet (1981, 1985) attempted to reconstruct re-
spread use of genital and latterly of tympanal lationships in the Pyraloidea based on Hennigian
characters, that further major progress has been phylogenetics. The best defended sister group re-
The Pyraloidea
lationship is the Nymphulinae-Schoenobiinae
clade. Passoa (1988) summarized synapomor-
phies for this sister group relationship, with em-
phasis on immatures. Solis & Mitter (1992) con-
ducted a cladistic analysis of the pyralid subfam-
ilies. J. Shaffer (1968, 1976) redefined the peori-
ines based on characters of the female and male
genitalia of North American species. The peori-
ines are closely related to the Phycitinae and we
prefer to consider them a tribe within that sub-
family (see Solis & Mitter 1992). The monobasic
Hypotiinae have been included in the Pyralinae
(Solis & Mitter 1992). The Endotrichini, found
in Europe, Africa and Asia, were placed in the
Pyralinae by Whalley (1961), but Minet (1985)
proposed that the endotrichines be returned to
subfamily level; we consider this step premature.
The arrangement used in this work (Table 1) rep-
resents a synthesis and partial revision of cur-
rent views.
Work on immature stages is sparse and scat-
tered, and keys are regional or include only a
selection of pest species. Scientific study of the
immature stages began with Dyar (1900, 1901,
1903), Forbes (1910, 1923), Fracker (1915) and
Mosher (1916), but with little impact on overall
classification. The important papers of Gerasi-
mov (1939, 1947, 1949) had little direct influence
in western countries because of limited availabil-
ity and the language barrier. Modern work on
pyraloid larvae begins with that of Bollmann
(1955) and Hasenfuss (1960). Recent major work
on last instar larvae in subgroups of the Pyra-
loidea includes Allyson (1977 a, 1977 b, 1981,
1984), Neunzig (1986, 1987), Passoa (1985), and
Yoshiyasu (1985). The standard work on pyra-
loid pupae is still Mosher (1916). Recent work
on eggs is limited to SEM morphological work
on stored product pests (Arbogast Van Byrd,
1981); earlier Peterson (1963) published pho-
tographs and descriptions of eggs of common
pyralid species in the U. S.
Larval stages of some primitive Pyralidae are
associated with fungi and/or dried or decaying
plant or animal matter. Among Crambidae, lar-
vae of a number of primitive forms feed on cryp-
togams: Diptychophorini and some Crambini in
the Crambinae as well as many Scopariinae on
mosses, the terrestrial nymphuline genus Nym-
phicula on liverworts (Yoshiyasu 1980), some
Glaphyriinae on lichens, species of Phenacodes
in the Scopariinae, and all known larvae of Mu-
sotiminae on ferns, and the larvae of many
stream-living Nymphulinae on algae (Lange
1956 a, 1956 b; Yoshiyasu 1985). The wide associ-
ation of primitive pyraloids with primitive plants
may indicate that the family originated before
flowering plants achieved their present domi-
nance. Whether or not this is true, the great ma-
jority of present-day pyraloid larvae feed on the
tissues of vascular plants, either gymnosperms or
angiosperms. However, in addition to plant-feed-
235
ers and to the scavengers already mentioned,
there are some species predaceous on Ho-
moptera and other insects, or that are inquilines
or parasites in the nests of social insects. Pyra-
loid larvae are characteristically concealed feed-
ers: folding, rolling, webbing or tying leaves;
making sand, silk or frass tunnels or tubes; bor-
ing in stems, roots, shoots, buds, fruits, or galls;
mining leaves; making cases of various kinds; or
sheltering in nests of ants, bees, wasps and ter-
mites. Larvae of pyraloids cause major damage
to crops worldwide (see Clausen, 1978 for litera-
ture review). Species of several subfamilies may
defoliate shrubs and forest, shade, and fruit
trees. Species directly benefitting humans are
few, but include some used in biological control
of weeds.
Pyraloidea occur on all continents except Ant-
arctica, and on most islands, including oceanic
ones such as Hawaii, Rapa, the Azores, St. Hel-
ena, and Amsterdam Island. Though best repre-
sented at low and middle elevations in the trop-
ics, they range from the high arctic of Ellesmere
Island to Tierra del Fuego and from equatorial
lowlands to alpine snowline. Their patterns of
distribution indicate repeated dispersal at many
evolutionary levels. Some nearly cosmopolitan
species must have been spread by human agency.
However, at least some close-knit species groups,
such as the Nomophila noctuella complex
(Munroe 1973 b), have wide natural distribu-
tions. Overseas dispersal has been relatively easy
in some genera, such as Udea (Pyraustinae) and
Eudonia (Scopariinae). These have endemic spe-
cies or species clusters on remote islands in all
oceans, as well as wide continental ranges. In
contrast, Scoparia, while also widely distributed
on continents, is poorly represented on islands
(Munroe 1958 a). Other genera are more re-
stricted in range: Synclera (Pyraustinae) is
pantropical; Dioryctria (Phycitinae) is holarctic;
many genera are restricted to single faunal re-
gions or parts of faunal regions; some genera,
such as Mestolobes (Crambinae) in Hawaii and
Idioblasta (Crambinae) in the Marquesas, are
confined to oceanic islands, where some of them
have evolved species clusters (Clarke 1986;
Munroe 1989, Zimmerman 1958). Continental
islands also may have endemic genera, e. g., Ba-
songa in Jamaica. On the continents, too, some
genera have very restricted ranges (Munroe
1975). Ranges of some genera or groups of gen-
era are strikingly disjunct: Choristostigma (Pyra-
ustinae) has several species in North America
and one in Central Asia; the tropical American
Cliniodes (Odontiinae) is represented by the vic-
ariant Metrea in northeastern United States and
southeastern Canada. Siga (Pyraustinae) of trop-
ical America is represented in tropical Africa by
Eporidia.
Several subfamilies, including Phycitinae, Epi-
paschiinae, Galleriinae, Crambinae, Scopariinae,
236
Musotiminae, Nymphulinae, Schoenobiinae, Ev-
ergestinae, Odontiinae and Pyraustinae, are
widely distributed, with complex patterns of in-
terregional vicariance. In Pyraustinae, for exam-
ple, many genera or groups of genera are shared
between the tropics of America and the Old
World, some showing evidence of multiple inter-
changes. The possibility of past faunal links via
the north is clear, as many mainly tropical as-
semblages extend at present into subtropical and
even temperate North America and Asia. South
circumpolar links are less obvious. Though there
are well-developed cool-temperate pyraloid fau-
nas in America, Africa, Australia and New
Zealand, they have few components in common,
and even related forms so far appear to be inde-
terminate or negative indicators. The genus Sco-
parla, for example, has numerous species in Chile
and Patagonia, though Eudonia is absent there
(Munroe 1958 a). Both genera are rich in species
in New Zealand, but Chilean Scoparla species do
not appear to be more closely related to New
Zealand ones than to those of any other conti-
nent. Australian and New Zealand Scopariinae
show some vicariance, but the few South African
species appear distinct. Crambini are numerous
in temperate South America, Africa, Australia,
and New Zealand, but the genera appear to dif-
fer from continent to continent. Not all the ma-
jor groups are globally distributed. Midilinae
and Linostinae are Neotropical, the last edging
into the North American subtropics; Cacothera-
piini in the Galleriinae and Argyriini in the
Crambinae occur in both temperate and tropical
America. Glaphyriinae and Chrysauginae are
mainly Neotropical but are represented signifi-
cantly in the Nearctic and sparingly in the Old
World as well. The following tribes and subfami-
lies are confined to the Old World: Hypotiini
(Palaearctic) (Pyralinae) and probably Endot-
richini (general) (Pyralinae); Wurthiinae (Indo-
Australian); Eclipsiodini (Indo-Australian) (Sco-
pariinae), Heliothelini (general) (Scopariinae),
and Hoploscopini (Indo-Australian) (Scoparii-
nae). No subfamily or tribe is known to be re-
stricted to the Afrotropical region. The Ethio-
pian and Neotropical faunas have a substantially
smaller proportion of the world fauna in Pyra-
loidea than in Lepidoptera as a whole. The other
four regions have a larger proportion of pyraloid
species, the relative excess being fairly evenly di-
vided among them, though larger in the Oriental
and Australian. In numbers of species, the Neo-
tropical and Oriental pyraloid faunas are almost
the same and the Australian about three-fourths
as large, whereas in total Lepidoptera the Neo-
tropical fauna is larger than the Oriental and
Australian put together. The smallest pyraloid
fauna is the Nearctic; the Ethiopian is not much
larger, probably with many more species to be
described; the Palaearctic is almost as large as
Eugene Munroe and M. Alma Solis
the two together, and somewhat larger than the
Australian.
Munroe 1994; Solis 1996 a, 1996 b.
PYRALOIDEA
Diagnosis: Maxillary palpus usually present,
small to prominent, scaled, generally three- or
four-segmented, not folded. Proboscis when pre-
sent scaled at base, sometimes vestigial. Foretibia
with epiphysis. Tibial spurs normally 0-2-4. Tho-
rax without primary tympanal organs. Abdomen
with paired tympanal chambers on ventral part
of base, facing more or less forward into space
between thorax and abdomen. Hindwing with Sc
+ Ri approximated to or anastomosed with Rs
for some distance beyond discal cell. Eggs of flat
type, ellipsoidal or lenticular, with thin cuticle
and inconspicuous sculpture. Larvae with two
prespiracular setae on prothorax. Primary setae
and often well-developed pinacula present, but
secondary setae few or absent. Abdominal pro-
legs generally with crochets in a multiordinal
penellipse or complete circle, but in at least some
Chrysauginae arranged in a mesoseries. Pupae
with epicranial suture present, except in Epi-
paschiinae, and lobes indicating the presence of
pilifers, except in Galleriinae (Mosher 1916).
Adult very small to large and heavy, but typi-
cally midsized or smaller, ditrysian Lepidoptera.
Length of forewing 5 to 75 mm, but mostly un-
der 30 mm. Build slender and delicate to thick
and robust.
Head: (Figs. 14.1 D - ) frons smoothly or
roughly scaled; rounded, flat and oblique, or,
less commonly, prominent or with variously
shaped spines, ridges or processes. Vertex with
erect scaling in tufts. Labial palpus normally
prominent and 3-segmented, articulations with
limited movement, palpus rarely reduced, some-
times excavated dorsally to receive maxillary pal-
pus; normal position decurved, porrect, ob-
liquely ascending, or upturned against frons or
even over vertex; scaling well developed, various,
sometimes including androconia. Maxillary pal-
pus reduced, 1- to 4-segmented, sometimes ab-
sent; scaling compressed, distally dilated, plu-
mose, or forming an aigrettelike tuft. Proboscis
primitively well developed, coiled when not in
use, exposed anterior surface of basal coil
densely scaled; proboscis sometimes reduced or
absent; even very rudimentary probscides often
retain basal scaling. Eye never with macroscopic
setae, usually large, globular, but sometimes re-
duced, especially in diurnal species. Ocellus most
often present, situated near dorsal margin of eye,
reduced or absent in some species and genera.
Chaetosema often present, but in reduced form,
as a somewhat radiating group of short fine setae
near posteroventral angle of vertex; a raised boss
or specialized setal bases rarely visible on the de-
nuded vertex. Antenna various, filiform, annu-
The Pyraloidea 237

Fig. 14.1. Structural Features of Pyraloidea. A, Ostrinia nubilalis, foreleg; B, Ostrinia nubilalis, midleg; C, Ostrinia
nubilalis, hindleg; D, Ostrinia nubilalis, frontal view of descaled head; E, Mecyna submedialis, lateral view of head
with scales (modified from Munroe, 1950); F, Mediavia discalis, female genitalia (from Solis, 1993); G, Deuterollyta
conspicualis, male genitalia (from Solis, 1993); H, Tancoa califas, aedoeagus (from Solis, 1993).

late, prismatic, laminate, or less commonly uni-
or bi-pectinate; often sexually dimorphic.
Thorax without primary tympanal organs; in
males often with variously disposed androconia.
Thoracic structure variable, but so far not com-
paratively studied. Legs (Figs. 14.1 AC) slender
to thick and robust, smoothly scaled or with
thick vestiture of normal or modified scales, in
males often with prominent androconia, some-
times in retractile tufts or with accompanying
distortions of the leg; foretibia with epiphysis;
tibial spurs 0-2-4. Wings (Figs. 14.2AB) vari-
able in shape; forewing wide to narrow, hindwing
wide, with fringe much narrower than mem-
brane; costa usually straight or arched, but
sometimes concave, sinuous or distorted in either
forewing or hindwing, especially in males; ter-
men usually convex or sinuate, tending to be
more strongly curved in cell M3. Forewing primi-
tively with Sc, 5-branched R, R3 and R4 stalked,
no areole, 3-branched M, M2 closer to M3 than
to Mi, stem of M absent or faintly indicated, dis-
cal cell usually closed, CuA 2-branched, CuP
represented by a vein, fold, a weak tubular ves-
tige adjacent to termen, or absent. 1 A strong and
tubular, 2 A shorter and weak, often joined to
238
1A by a crossvein; both sexes with a retinaculum
of stiff scales on underside in cubital area, males
primitively with a sclerotized frenulum hook ex-
tending posterodistad from behind costa, but the
frenulum hook lost in some groups. Hindwing
with Se + Ri approaching or anastomosed with
Rs for some distance beyond end of discal cell,
then diverging again, Rs not branched, basally
often short-stalked with Mi from anterior angle
of cell, but sometimes connate with Mi or the
latter arising separately from discocellular, M 3-
branched, M2 closer to M3 than to Mi, as on
forewing, CuA 2-branched, CuP usually well de-
veloped, 1 A + 2 A always and 3 A usually pre-
sent. Discal cell open in many Crambinae. Stem
of CuA in some groups with pecten of fine setae.
Frenulum unisetose in male, multi- or unisetose
in female.
Male genitalia (Figs. 14.1 G - H ) diversified.
Tegumen and vinculum forming a complete ring,
often with complex lateral folding; vinculum
usually produced anteriad into a midventral sac-
cus. Uncus various, almost always present.
Gnathos primitively consisting of a pair of lat-
eral arms arising from junction of tegumen and
uncus, medially fusing to form a posteriorly di-
rected median process. Basal articulations of
gnathos either movable, hinging at tegumino-un-
cal suture, or fixed, in the latter case often joined
to distolateral margins of tegumen. Transtilla a
transverse sclerotized band between costae of
valves, often medially narrowed or absent, rarely
with special armature. Aedoeagus a variously
sclerotized tube, sometimes distorted or ornate,
projecting through a membranous or spinulose
sleeve, the manica, or sometimes a sclerotized
troughlike or tubular anellus. Inside aedoeagus a
membranous eversible vesica, generally minutely
spinulose, often also with single or grouped
larger spines, the cornuti; the latter sometimes
deciduous, capable of being shed into the female
bursa during copulation. Juxta variable in shape
and size, ventral to manica or anellus, connect-
ing ventrolaterally with bases of valves. Valve ex-
tremely various in form, primitively with subpar-
allel costal and ventral margins, each somewhat
inflated, the latter inflation known as the saccu-
lus, and with rounded or oblique terminal mar-
gin; costa articulating basally with transtilla;
most of base hinged with vinculum; ventral an-
gles articulating with juxta; mesal surface gen-
erally with variously grouped setae and some-
times spines, often also with variously disposed
ridges and processes; in certain groups the costa
and/or sacculus produced distally as a free pro-
cess.
Female genitalia (Fig. 14.1 F) primitively with
a pair of membranous setose ovipositor lobes,
approximated or fused dorsally, that of each side
supported by a usually T-shaped posterior
apophysis. Ostium base simple or variously
armed. Corpus bursae variable in shape and ar-
Eugene Munroe and M. Alma Solis
mature; ductus bursae usually differentiated
from corpus bursae; sometimes an accessory sac
arising from corpus bursae; ductus seminalis
usually arising from ductus bursae, but some-
times from corpus bursae; ductus bursae often
with a collarlike or complex sclerite at or just
posterior to opening of ductus seminalis.
Common 1990; Falkovich 1986; Falkovich &
Zagulyaev 1986; Goater 1986; Hanneman 1964;
Inoue et al. 1982; Martin 1986 a, b, c; Munroe
1972, 1973, 1976; Neunzig 1986, 1990, 1997;
Palm 1986; Shaffer, Nielsen & Horak 1996; Rob-
inson, Tuck & Shaffer 1994; Sinev 1986; Wang
1980; Zimmerman 1958.
Key to the Families and Subfamilies of the
Pyraloidea
1 Praecinctorium present; tympanal case
(= caisse tympanique "open" ante-
romedially; tympanum meeting con-
junctiva at a distinct angle
Crambidae, 1
Praecinctorium absent; tympanal case
"closed" medially and open anteriorly
only; tympanum and conjunctiva in the
same plane Pyralidae, 2
2(1) Secondary venulae present 3
Secondary venulae absent 5
3 (2) Female frenulum with one bristle; male
genitalia with uncus arms extending at
an angle of 110 degrees or more from
midline . . Phycitinae (Peoriini, in part)
Female frenulum with three bristles;
male genitalia with uncus arms extend-
ing at an angle of 90 degrees from mid-
line 4
4 (3) Gnathos absent; males with simple
wing-locking mechanism and forewing
without costal folds and/or resonating
chamber Galleriinae
Gnathos usually present; males of most
species with wing-locking mechanism
modified and forewing with costal folds
and/or resonating chamber
Chrysauginae
5 (2) Female frenulum with one bristle; duc-
tus seminalis usually originating from
corpus bursae Phycitinae
Female frenulum with two bristles; duc-
tus seminalis originating from ductus
bursae 6
6 (5) Phallobase of male aedoeagus curved
ventrad; male genitalia with uncus arms
extending at an angle of 110 degrees or
more from midline; third labial palpus
segment of both sexes upturned and
pointed at apex Epipaschiinae
Phallobase of male aedoeagus not
curved ventrad to tip; male genitalia
with uncus arms extending at an angle
The Pyraloidea
of 90 degrees from midline; third labial
palpus segment of both sexes upturned,
but blunt at apex Pyralinae
7 (1) Chaetosema present 8 process but, except in Heliothela, with-
Chaetosema absent 13
8 (7) Forewing with distal part of CuP devel-
oped as a tubular vein 9 ferns, and roots of seed-bearing vascu-
Forewing with CuP absent, not devel-
oped as a tubular vein 10
9 (8) R.2 of forewing stalked with R3+4; pro-
boscis normally developed; M2 of
hindwing absent; termen of hindwing
usually with a series of black and met-
allic spots; vinculum of male genitalia
without shell-like accessory sclerites;
larvae aquatic, in silk webs on rocks or
feeding on submerged plant tissues . . .
Nymphulinae (part)
R.2 of forewing most often separate
from R3+4; proboscis reduced; M2 of
hindwing present; termen of hindwing
without a series of black and metallic
spots; vinculum of male genitalia usu-
ally with a pair of shell-like accessory
sclerites; larvae borers, generally in
semiaquatic graminaceous plants . . . .
Schoenobiinae
10 (8) Proboscis present, but reduced; tympa-
nal cases reduced and widely separated;
praecinctorium reduced; known larvae
on monocotyledonous plants, mostly
Araceae Midilinae
Proboscis usually and tympanal organs
almost always normally developed . 11
11 (10) R.2 of forewing at least closely apposed
to and usually stalked with R3+4; labial
palpus usually upturned, basal segment
often longer than second segment;
wings mostly with conspicuous pattern
of transverse bands on a pale ground;
larvae aquatic, rarely in damp ter-
restrial habitats . . Nymphulinae (part)
R2 of forewing well separated from
R3+4; labial palpus often porrect, basal
segment much shorter than second seg-
ment; wings usually without conspicu-
ous pattern of transverse bands on pale
ground; larvae usually terrestrial, some-
times stem borers in aquatic gramina-
ceous plants 12
12(11) Forewing usually with weakly raised
patches of black scales; cubitus of
hindwing usually not pectinated with
hairlike scales; lateral arms of tegumen
of male genitalia about as long as un-
cus, little tapered ventrally; uncus of
moderate length, pyriform, hood-
shaped or bilobed, not obviously de-
curved from base to tip; gnathos with
median element spikelike, sword
shaped, or digitate, straight or de-
curved, rarely reduced; uncus and
239
gnathos not forming a jawlike structure,
and not widely separated dorsad from
valves; valve sometimes with a ventral
out strong costal or medial armature;
known larvae on mosses, lycopods,
lar plants Scopariinae
Forewing without raised patches of
black scales; cubitus of hindwing usu-
ally pectinated with hairlike scales, lat-
eral arms of tegumen of male genitalia
much longer than uncus or narrowed
ventrally, usually both; uncus usually
long, acuminate, and more or less de-
curved from base to tip; gnathos with
lateral arms articulating at base of un-
cus, medial element various in shape,
often rodlike or forming a dorsally di-
rected hook; uncus and gnathos form-
ing a jawlike structure, widely separated
dorsad from valves; valve often with
strong costal process or medial arma-
ture; larvae feeding at bases or on roots
of grasses, mosses, or other plants, or
stem borers in grasses . . . . Crambinae
13 (7) Valve of male genitalia with costal pro-
cess; simple uncus and well-developed
gnathos widely separated from valve by
deep lateral elements of tegumen . . . .
Cybalomiinae
Valve of male genitalia without costal
process, or uncus bilobed and laterally
decurved and setose, or gnathos absent
14
14(13) Uncus of male genitalia bilobed;
gnathos well developed 15
Uncus of male genitalia simple, or
gnathos reduced or absent 16
15(14) Uncus laterally setose; gnathos with
oblique lateral arms, median element
usually distinct, spikelike; moth not li-
macodiform; distal part of CuP absent;
larvae leaf miners, leaf tiers, or seed
feeders, not myrmecophilous
Odontiinae
Uncus not laterally setose; gnathos
short, with two prongs curving medially
and then laterally, median element not
present; moth limacodiform, with
rounded wings, pale ground, and little
or no maculation; distal part of CuP de-
veloped and tubular; larvae in leaf nests
of Oecophylla tree ants . . . Wurthiinae
16 (14) Male genitalia with gnathos well devel-
oped, slender, dorsally toothed near
apex 17
Male genitalia with gnathos, if well de-
veloped, not dorsally toothed near apex
19
17 (16) Male genitalia with uncus narrow, digi-
tate or distally pointed; gnathos slender,
240
with dorsal row of spinules; valve usu-
ally oblong or rounded, if narrow and
elongate not tapering 18
Male genitalia with uncus hood-shaped;
gnathos with strongly sclerotized, verti-
cally compressed median element end-
ing posteriorly in a sharp spike; valve
narrow and elongate, irregularly taper-
ing, narrowly rounded at tip Linostinae
18(17) Forewing subtriangular; maculation,
when distinct, of narrow, often oblique,
transverse lines; hindwing almost im-
maculate; termina of forewing and
hindwing regular or weakly angulate;
Mi of forewing from near anterior an-
gle of discal cell; Mi of hindwing usu-
ally short-stalked with Rs; tympanal
chamber not enlarged; male genitalia
with valve somewhat rhomboidal, sim-
ple, or with a clasper on mesal face; ae-
doeagus bent at middle; manica spinu-
lose; female genitalia with ductus bur-
sae of moderate length; corpus bursae
globular, with paired spinulose signa;
ductus seminalis arising from ductus
bursae; larvae leaf webbers, mostly on
Cruciferae and Capparidaceae
Evergestinae
Forewing short and wide, with arched
costa; maculation of forewing and
hindwing similar, usually of sinuated
transverse bands; termina of forewing
and hindwing sinuate or excavated; Mi
of forewing from behind anterior angle;
Mi of hindwing not stalked with Rs;
tympanal chamber enlarged to posterior
border of first abdominal segment; male
genitalia with valve short and rounded,
often with a process from sacculus; ae-
doeagus not bent at middle; manica not
spinulose; female genitalia with ductus
bursae shorter than corpus bursae, the
latter oval, not with paired spinulose
signa; ductus seminalis arising from
corpus bursae; larvae feeding on foliage
of ferns Musotiminae
19 (16) Male genitalia with gnathos well devel-
oped, not toothed dorsally; uncus trian-
gular Pyraustinae (part)
Male genitalia with gnathos reduced;
uncus various 20
20 (19) Hindwing with base of CuA pectinated
with hairlike scales; uncus davate or
gladiate; gnathos reduced; tympanum in
same plane as metathoracic coxal mem-
brane; tympanal chamber atrophied,
imbedded in metathorax; tympanal
ridge long and spatulate . . Noordinae
Hindwing with base of CuP not pecti-
nate; uncus various; gnathos reduced or
absent; tympanal organs not as de-
scribed for Noordinae 21
Eugene Munroe and M. Alma Solis
21 (20) Praecinctorium simple 22
Praecinctorium bilobed
Pyraustinae (most)
22 (21) Male genitalia with gnathos absent; un-
cus wide, distally rounded, with close-
set vestiture of fine anteriorly directed
setae on apical and lateral areas; valve
not fanlike, but narrow, elongate, dis-
tally rounded, unarmed; aedoeagus bent
behind middle with long, anterior coe-
cum, with conspicuous cornutus; female
genitalia with bursa copulatrix un-
armed, with ductus seminalis arising
from ductus bursae; tympanal organs
sexually dimorphic, fenestra media ab-
sent; coxal countertympanum very
large, internal Cathariinae
Male genitalia with gnathos reduced or
absent; uncus various, but not wide, dis-
tally rounded and apically spinose,
valve fanlike, often with costal arma-
ture; juxta ringlike or with high lateral
processes; aedoeagus generally short
and cylindrical or flask shaped, with
conspicuous cornutus/i; female genitalia
with heavily armed, distorted bursa,
with ductus seminalis arising from a di-
verticulum of corpus bursae; tympanal
organs not sexually dimorphic and not
as indicated for Catharinnae
Glaphyriinae
Pyralidae. Figs. 14.3 AJ. Diagnosis: Tympanal
organs with tympanal case almost completely
closed; conjunctiva and tympanum in the same
plane; praecinctorium absent; vein R5 of fore-
wing stalked or fused with R3 + R4; segment A 8
of larvae almost always with a sclerotized ring
around base of seta SD; male genitalia with un-
cus arms, a pair of processes arising laterally
from the base of the uncus (Hasenfuss 1960; Mi-
net 1981, 1985; Solis & Mitter 1992).
Description: Head with frons rounded; scaling
of frons usually smooth. Labial palpus almost
always 3-segmented, porrect, obliquely ascend-
ing, or upturned in front of face. Maxillary pal-
pus usually 3-segmented, sometimes minute or
absent, usually shorter than labial palpus, highly
modified in some groups. Proboscis usually well
developed, but in various groups reduced or ab-
sent. Eye normally large and gobular, without
macroscopic setae.
Wing venation as described for the superfam-
ily. Forewing with R2 closely apposed to, not
usually stalked with, R3 and R4; R3, R4, Rs
sometimes reduced to two veins or one vein. Mi
from near anterior angle of cell. M2, M3 and
CuAi from posterior angle of cell or near it, M2
and M3 sometimes stalked. CuP well developed,
or incomplete with only its distal part well devel-
oped, or absent, reduced to a fold. 1 A strongly
developed; 2 A distally free or connected by a
The Pyraloidea 241

Fig. 14.2. Structural Features of Pyraloidea. A, Cacozelia basiochrealis, forewing venation (from Solis, 1993); B,
Cacozelia basiochrealis, hindwing venation (from Solis, 1993); C, Azamora sororia, ventral view of pupa; D, Aza-
mora sororia, lateral view of pupa; E, Eoparargyractis plevie, lateral view of last instar larva (modified from
Fiance & Moeller, 1977); F, Eoparargyractis plevie, dorsal view of terminal abdominal segments (modified from
Fiance & Moeller, 1977); G - J : Tympanal organs of the Pyraloidea: G & I, Oneida luniferella, Pyralidae, praecinct-
orium absent; H & J, Desmia vulcanalis, Crambidae, praecinctorium present; G, Pyralidae, tympanum and con-
junctiva in the same plane (line horizontal only); H, Crambidae, tympanum and conjunctiva meeting at a distinct
angle (lines horizontal and vertical); I, Pyralidae, tympanal case "closed" medially and with a small, anterior
aperture (bold dotted lines); J, Crambidae, tympanal case "open" anteromedially (bold dotted lines).

c r o s s v e i n t o 1 A t o f o r m a closed cell, a n d s o m e - sent. F r e n u l u m single in m a l e s , m o s t l y m u l t i p l e

t i m e s w i t h a f r e e p o r t i o n e x t e n d i n g b e y o n d this
cell. I n t h e h i n d w i n g Sc + R i a n d R s m a y b e
anastomosed or separate. M2 a n d M3 usually
s e p a r a t e , b u t s o m e t i m e s f u s e d , t h e a m o u n t of f u -
sion sometimes variable within taxa. C u A i a n d
C U A 2 u s u a l l y a r i s i n g s e p a r a t e l y f r o m cell; C u A i
r a r e l y f u s e d w i t h M3. C u P a n d 1 A + 2 A p r e -
in f e m a l e s , b u t single in t h o s e o f P h y c i t i n a e . L e g s
varying f r o m long a n d slender to short a n d
s t o u t , o f t e n w i t h m o d i f i e d scaling. P r a e c i n c t o r -
i u m a b s e n t ; t y m p a n a l cases " c l o s e d " , i. e., w i t h a
r e s t r i c t e d a n t e r i o r a p e r t u r e ( = t y m p a n a l areole);
c o n j u n c t i v a a n d t y m p a n u m in t h e s a m e p l a n e ;
secondary venulae present or absent.
242
In male genitalia, uncus usually well devel-
oped, but reduced in certain genera in all sub-
families. Uncus with ventro-anteriorly elongate
arms articulating with base of gnathos, extend-
ing laterally in most chrysaugines and pyralines
at 90 degrees to midline and in all epipaschiines
and most phycitines at 110 degrees or more; in
some phycitines these arms very short or absent.
Uncus with many setae caudally, sometimes in
a regular pattern. Tegumen in epipaschiines and
phycitines abutting on only a very small portion
of base of uncus, in chrysaugines and pyralines
abutting on at least its middle half, and in galleri-
ines extending entire width of base. Tegumen
complete, or in some Epipaschiinae with a sepa-
rate sclerite articulating with gnathos. Gnathos
usually present, absent in galleriines, reduced or
absent in some chrysaugines and pyralines. Dis-
tal end of gnathos either a simple hook or
curved, s-shaped or often with variously modi-
fied apex as in many Phycitinae. Valve most
often simple, but in some groups with modified
lobes or setae. Transtilla well developed only in
some phycitines and endotrichines. Juxta plate-
like, U-shaped, or variously modified. Aedoea-
gus usually a short cylinder, with or without cae-
cum; cornuti often absent, but present in many
phycitines, Old World epipaschiines, endotrich-
ines and the pyraline genus Vitessa. Ovipositor
lobes membranous, with many, usually unmodi-
fied setae. Lamella antevaginalis membranous,
but sclerotized in some species. Ductus bursae
usually membranous, but sometimes with sclero-
tized or scobinated areas. Corpus bursae mem-
branous or spinulose, globular, oval, or variously
wrinkled and contorted, without special scleroti-
zation, or with one or two elongate-conical, spi-
nelike projections, or one or two flat, scobinate
patches, or with various, often bizarre spining or
sclerotization. Ductus seminalis usually originat-
ing from corpus bursae in Phycitinae but from
ductus bursae in all other subfamilies.
Galleriinae. Fig. 14.3 A. Small to very large
moths, mostly with narrow wings and stout
body. Ocellus absent in Tirathabini and Galleri-
ini, present in Megarthridiini. Chaetosema ab-
sent. Frenulum of females with three bristles, of
males with one. Tympanal organs with curved
line between conjunctiva and tympanum, tympa-
nal areole absent or weak, secondary venulae
well developed. Uncus arms absent; tegumen ex-
tending entire length of base of uncus; gnathos
absent, vestigial in a few genera (e. g., Perine-
toides). In tribes Tirathabini and Gallerini a su-
pra-transtilla sac present ventral to anal tube and
attached to a membrane on dorsal side of
transtilla. Ductus seminalis originating on duc-
tus bursae. Most larvae with sclerotized ring
around SD of Al; pupae with prominent me-
dian ridge on dorsum of thorax and abdomen.
(Forbes 1923; Hasenfuss 1960; Kuznetzov &
Eugene Munroe and M. Alma Solis
Stekolnikov 1979; Minet 1985; Munroe 1972;
Solis & Mitter 1992).
Larvae of some Galleriinae such as Corcyra
cephalonica feed on dry vegetable matter such as
rice, while a few species, for example Galleria
mellonella, feed on combs of hymenopteran
nests. Tirathaba spp. are pests of coconut and
other palms. Spangler (1988) studied sound pro-
duction by tegulae of adult male Galleriinae for
purposes of monitoring and control of pest spe-
cies.
261 species (Heppner 1991). Principal genera
include: Achroia, Aphomia, Eldana, Paralipsa.
Cosmopolitan in distribution, some species ac-
companying man.
Hampson 1917; Martin 1986a; Munroe 1995;
Ragonot & Hampson 1901; Whalley 1964.
Chrysauginae. Figs. 14.3 B - C . Small to very
large moths, often with simple pattern of two
transverse lines on a green, brown, gray, yellow
or reddish ground, but sometimes with bizarre
or mimetic patterns, or with bright red, yellow,
or green areas. Ocellus present. Chaetosema pre-
sent or absent. Forewing often sexually dimor-
phic: females unmodified, males with costa,
shape of discal cell, stalking of radial veins, and
retinaculum altered. Frenulum of females usually
with three bristles, of males with one, sometimes
thickened, bristle. Retinaculum in the latter case
hoop shaped. Secondary venulae present. Uncus
arms arising laterally from uncus and extending
at about 90 degrees: tegumen abutting on at least
half length of base of uncus; gnathos usually pre-
sent, but absent in Satole and Clydonopteron.
Ductus seminalis originating from posterior end
of ductus bursae. Larvae usually with a sclero-
tized ring at base of SD on metathorax, but this
ring missing in some genera such as Azamora
(Cashatt 1968; Hasenfuss 1960; Minet 1985;
Solis & Mitter 1992).
No major pests are known. The larvae have a
variety of habits, including seed, fruit, stem, and
root boring, and leaf rolling and tying; some are
myrmecophilous. Adults of several genera
{Cryptoses, Bradypodicola, Bradypophila) live in
fur of sloths (Dyar 1908; Waage & Montgomery
1976; Bradley 1982), and their immature stages
develop in sloth dung. Larvae of other species
have been found feeding on ant and wasp nests
(Hampson 1905; Pastrana 1953) and on spines
of caterpillars of Automeris spp. (Saturniidae)
(Jordan 1926).
400 species (Solis, Becker & Munroe 1995).
Principal genera include: Azamora, Clydonopt-
eron, Pachypodistes, Galasa. Primarily Neotropi-
cal, but a few genera in North America and the
tropics of Asia and Australia.
Cashatt 1968; Hampson 1897 a; Martin
1986 b.
Pyralinae. Figs. 14.3 DF. Small to medium-
sized, mostly dark red or brown colored moths,
but a few groups dull yellow or orange, brightly
The Pyraloidea
colored, or mimetic. Ocellus present or absent.
Chaetosema present. Frenulum with one bristle
in males and two in females. Tympanal organs
with secondary venulae usually absent, but pre-
sent in a few taxa. Uncus arms arising laterally
from base of uncus, extending at about 90 de-
grees from longitudinal axis of uncus; tegumen
abutting on at least half width of base of uncus;
gnathos usually a hook, but in Endo tricha a flat
plate articulating with the uncus; sometimes re-
duced or absent, but gnathos arms always pre-
sent. Corpus bursae membranous and barely ex-
tending cephalad of segment 7; ductus seminalis
originating from ductus bursae (Forbes 1923;
Minet 1985; Munroe & Shaffer 1980; Solis &
Mitter 1992; Whalley 1963).
The pyraline larvae with well-known biolo-
gies, such as Pyralis farinalis, are cosmopolitan
stored products pests. One of these Pyralis mani-
hotalis, has been reared from bat guano (V. O.
Becker, pers. comm.). A number of species feed
on dead vegetation; some species are leaf feeders.
Over 900 species (Shaffer & Salis 1995 and un-
published). Principal genera include Aglossa,
Dolichomia, Herculia. The pyralines are more di-
verse in Asia and especially in Africa than in the
Western Hemisphere.
Hampson 1896 a; Martin 1986 b; Whalley
1961.
Epipaschiinae. Figs. 14.3 GH. Small to large,
but mostly medium-sized moths. Third segment
of labial palpus always upturned and pointed at
apex. Ocellus and chaetosema present. Frenulum
of males with one bristle, of females with two.
Tympanal organs without secondary venulae.
Uncus arms elongate, arising laterally from base
of uncus and extending at an angle of 110 de-
grees or more; tegumen abutting on only a very
small portion of base of uncus; a weakness in
sclerotization of base of tegumen defining a sep-
arate sclerite. Phallobase of males usually ex-
tending beyond ductus ejaculatorius and curved
ventrad; ductus seminalis originating from duc-
tus bursae (Solis & Mitter 1992; Solis 1993).
Larvae are known to be leaf rollers, leaf tiers,
or leafminers, but their morphology is not well
known (Allyson 1977 a; Passoa 1985). A few spe-
cies are minor pests of mahoganies, avocado,
and corn (Zea mays) in the Neotropics.
572 species (Solis 1992, 1993). Principal genera
include Pococera, Phidotricha, Macalla, Epi-
paschia, Stericta. Tropical and temperate re-
gions, excluding Europe.
Hampson 1896a; Hulst 1889; Janse 1931;
Martin 1986 b; Solis 1993, 1995.
Phycitinae. Figs. 14.31J. Mostly small
moths, with long, narrow forewings, some with
raised scales. Ocellus present or absent. Chaeto-
sema present. Frenulum with one bristle in both
sexes. Uncus arms of males extending at 110 de-
grees or more from longitudinal axis; tegumen
abutting a small portion of base of uncus. Duc-
243
tus seminalis of females generally originating
from corpus bursae, rarely from ductus bursae.
Tympanal organs with a circular sclerotization
around distal insertion of scoloparium; second-
ary venulae usually absent, but present in most
genera of the Peoriini. Larvae with a sclerotized
area encircling base of seta SD of mesothorax
(Hasenfuss 1960; Minet 1985; Solis & Mitter
1992).
The larvae are mostly leaf rollers, some in a
tube of silk mixed with frass. Others are borers
in buds, shoots, cones, fruits, galls, cankers and
cambium; leafminers in early instars; inquilines
in galls and in nests of Hymenoptera; predators
on Homoptera (Laetilia sp.); seed feeders (in-
cluding stored products pests: Plodia interpunc-
tella, Anagasta kuehniella, Cadra cautella, Ecto-
myelois ceratoniae, Etiella zinckenella; in part).
A number of species have been used to control
noxious weeds. The best example is Cactoblastis
cactorum, which has helped to reduce prickly
pears (Opuntia spp.) in Hawaii, Australia, South
Africa, West Indies, and Mauritius (Huffaker &
Messenger 1976; Clausen, et al. 1978).
Solis and Mitter (1992) estimated close to 4000
species. Principal genera include Acrobasis, Cryp-
toblabes, Homoeosoma, Nephopterix, Dioryctria,
Paramyelois, Hulstia, Peoria. Distributed
throughout the world except in the southern ex-
tremes.
Janse 1941, 1942, 1944, 1946; Heinrich 1956;
Neunzig 1986, 1990; Ragonot 1893; Ragonot &
Hampson 1901; Shaffer 1968, 1995; Sinev 1986;
Roesler 1965, 1973, 1983; Whalley 1970.
Crambidae. Figs. 14.3 K - T ; Figs. 14.4 A - T . Di-
agnosis: Tympanal organs with tympanal cases
"open", i. e. with a wide anteromedial aperture;
conjunctiva and tympanum not in the same
plane, meeting at a distinct angle; praecinctor-
ium present, usually well developed; vein R5 of
forewing most often not stalked with R3+4; seg-
ment A8 of larvae without sclerotized ring
around base of SD; male genitalia without un-
cus arms.
Head with frons various in shape - rounded,
flat, conical, cylindrical, carnate, thorned, or in-
flated - scaling usually smooth. Vertex with
tufts of erect scales, especially a pair extending
anteriad between antennae, and a larger postero-
lateral pair. Labial palpus almost always 3-seg-
mented: porrect, obliquely ascending, or up-
turned in front of face. Maxillary palpus usually
shorter than labial, sometimes minute or absent,
typically 3- or 4-segmented; often with distal
scaling forming a flattened tuft, connecting pro-
file of frons with that of dorsal scaling of labial
palpus. Proboscis usually well developed, but in
various groups reduced. Eye without macro-
scopic setae; normally large and globular, in di-
urnal forms sometimes reduced, then often bor-
dered by a scaleless zone. Legs most often long
244
and slender; in males often with androconial
tufts and sometimes with structural distortions.
Praecinctorium simple or bifid; tympanal cases
"open", i. e., with a wide anteromedial aperture;
tympanal ridge present; spinula typically present;
scolopal swellings enlarged; in species so far
studied, two of the four scolopal bodies dif-
ferentiated into squamiform structures (Minet
1985); secondary venulae present.
Wing venation normal for the superfamily.
Forewing with R2 often closely apposed to or
stalked with R3+4 beyond cell; R5 often curved
and approximated to R3+4, stalked with it in
many Crambinae and Schoenobiinae, and spo-
radically in other groups. Mi from near anterior
angle of cell. M2, M3 and CuAi from posterior
angle of cell or near it, M2 and M3 sometimes
stalked. CuP reduced to a fold or with its distal
part developed as a weak tubular vein. 1A + 2 A
strongly developed, usually ending on termen
near tornus. 2 A distally free or connected by a
crossvein to 1 A to form a closed cell, but hardly
ever with a free portion extending beyond this
cell as sometimes occurs in Pyralidae. Hindwing
with M2, M3, and CuAi from posterior angle of
cell or near it, often basally approximated, some-
times M2 and M3 stalked or even fused. CuP, 1 A
+ 2 A, and 3 A present, simple. Frenulum single
in males, usually multiple in females.
Uncus primitively tapering, distally rounded
or weakly bilobed, and with dorsal setae tending
to be separated in lateral groups; in derived
forms uncus variously spined, in shape long and
rodlike, beaklike or distally dilated, or alterna-
tively strongly bifid, abbreviated, or virtually
aborted. Gnathos often with posteriorly directed
median element, this frequently dorsally denticu-
late; in some groups median element missing and
lateral arms separate or the whole gnathos rudi-
mentary or absent. Base of genitalia with weak
to large and bushy tufts of variously shaped an-
droconia, often forming a prominent anal tuft or
anal brush; these tufts arising from various parts
of the genitalia proper, especially from processes
on the vinculum, but also from intersegmental
membrane V i l i - I X , and often from the sternite
VIII, which may have special support sclerotiza-
tions, the culcitae. The muscles of the male geni-
talia appear to have taxonomic importance, but
they have been looked at in only a few represen-
tative forms (Kuznetsov & Stekolnikov 1979).
Ovipositor lobes as described for the super-
family or sometimes reduced or with reduced se-
tae, or heavily spinose, or elongate and strongly
sclerotized or even bladelike. Such modifications
often coupled with lengthening or hypertrophy
of the apophyses, presumably reflecting differ-
ences in oviposition habits.
Scopariinae. Figs. 14.3 KL. Heliothelinae,
new synonym. Forewing usually with tufts of
raised scales, pattern primitively noctuidlike.
Uncus triangular and hood-shaped, bilobed, or
Eugene Munroe and M. Alma Solis
pyriform; gnathos usually with median element
long, sharp, and with dorsal denticles, in most
genera articulated at junction of tegumen and
uncus; transtilla incomplete; valve simple or with
terminal process from sacculus, rarely with cos-
tal process; aedoeagus simple, with or without
cornuti.
Larvae tunnelling in roots and stems of
mosses and lycopods (Eudonia), feeding on roots
of seed-bearing vascular plants (Scoparla), or
webbing fronds of large ferns (Phenacodes).
479 species (Heppner 1991). Principal genera
include Eudonia, Scoparla, and Micraglossa. Al-
most universal, including many oceanic islands
(Munroe 1958 a).
Klima 1937; Leraut 1983, 1989; Martin 1986 c;
Munroe 1958 a, 1972, 1995.
Crambinae. Figs. 14.3 M - N . Labial palpus
long, porrect. Maxillary palpus with distally ex-
panded scaling. Forewing generally long, nar-
row, with grass-matching pattern, wider in tribe
Diptychophorini and a few insular or montane
groups; termen of forewing often notched or
with a break in pattern; wings longitudinally
folded or rolled at rest. Hindwing with discocel-
lulars usually incomplete; stem of CuA mostly
with pecten of erect setae. Uncus and gnathos
far removed from vinculum, juxta, and valves;
gnathos with lateral arms articulating with junc-
tion of tegumen and uncus; valve usually with
sclerotized and process-bearing costal region and
sacculus. Ovipositor lobes reduced; ostium with
taxonomically important sclerites; signa one or
two inconspicuous groups of spicules, or absent.
Larvae of two general habits: (a) root-feeders
or ground-living leaf feeders, on grasses or
mosses, rarely on other plants, and (b) stem-bor-
ers in grasses, sedges, and rushes. Both types
cause economic damage, the former as turf and
sod pests, also attacking roots of maize and
other field crops, the latter in grain crops, includ-
ing sugar cane, maize, and rice. The giant Mye-
lobia smerintha bores in stems of bamboo. Cram-
bine webworms pass the day in silk-lined tunnels
in the ground, and emerge to feed at night on
sod grasses, maize and other plants.
1877 species (Heppner 1991). Principal genera
include Glaucocharis, Mestolobes, Argyria, Cala-
motropha, Charltona, Euchromius, Mesolia, Prio-
napteryx, Surattha, and Talis. The stem-boring
Chilo, Diatraea, Bissetia, Girdharia, and Eoreuma
include pests of sugar cane; the first two of these
genera also contain borers in rice. Other major
stem-boring genera, though of less agricultural
importance, are Eschata, Calamotropha, Erupa,
Haimbachia, and Xubida. Root and sod-feeding
genera include especially Crambus and its rela-
tives such as Agriphila, Catoptria, Culladia, Fer-
nandocrambus, Fissicrambus, Hednota, Micro-
crambus, Pediasia, and Thaumatopsis; many spe-
cies in this group of genera regularly or occasion-
ally injure lawns and turf. Arctic to Subantarctic
The Pyraloidea 245

Fig. 14.3. Pyraloidea. AJ, Pyralidae: A, Cacolherapia leucocope [10 mm*]; B, Parachma ochracealis [9 mm]; C,
Casuario catocalis [11 mm]; D, Vitessa splendida [20 mm]; E, Cledeobia moldvico [13 mm]; F, Pyralis farinalis
[10 mm]; G, Epipaschia superatalis [9 mm]; H, Carthara abrupta [13 mm]; 1, Peoria roseotinctella [8 mm]; J, Dior-
yctria rossi [13 mm]; K - T , Crambidae: K. Eudonia torniplagalis [8 mm]; L, Gesneria centuriella [11 mm]; M, Euch-
romius ocelleus [10 mm]; N, Agriphila vulgivagella [11 mm]; O, Rupela tinctella [15 mm]; P, Cybalomia pentadalis
[7 mm]; Q, Linosta sinceralis [17 mm]; R, Midila daphne [26 mm]; S, Neurophyseta completalis [8 mm]; T, Musotima
acclaraiis [7 mm]. [* = length of forewing]

zones, also throughout Polynesia and on some
other oceanic islands. Root-feeding forms best
developed in north and south temperate zones,
stem borers in the tropics and subtropics.
Bleszynski 1963, 1965, 1969, 1970; Bles-
zynki & Collins 1962; Box 1931; Falkovitch
1986; Gaskin 1971, 1973, 1974 a, b, 1975 a, b,
1985, 1986 a, b; Klots 1970; Landry 1992, 1995;
Munroe 1995; Schouten 1992; Wang, et al. 1988.
Schoenobiinae. Fig. 14.3 O. Wings usually long
and narrow, with white or brown ground color
and reduced maculation. Proboscis reduced.
Chaetosema present. Forewing with radial vena-
tion variable and usually with reduced macula-
tion; CuP with terminal portion preserved as a
tubular vein in most genera. Hindwing with CuA
not pectinate. Frenulum hook present in males;
frenulum single in males, single or double in fe-
246
males. Males in all but four primitive genera with
a rounded membranous area on sternite VIII,
covered by a posteriorly directed brush of stiff
scales from posterior margin of sternite VII
(Common 1960), and associated with presence of
platelike, scale-bearing coremata flanking vincu-
lum (Lewvanich 1981). Uncus short, with wide
base and acuminate tip; gnathos usually similar
in shape, but sometimes modified (Heinrich
1937); lateral margin of tegumen on each side
with a conspicuous subteguminal process, its
shape different in different taxa; valve simple or
with modified sacculus. Ovipositor with high and
narrow or elongate, setose, membranous lobes;
apophyses slender; ostium variously sclerotized;
ductus bursae short, with sclerotized collar; duc-
tus seminalis simple and narrow; corpus bursae
short and oval, finely spinulose, usually without
signa.
Larvae borers in marsh-living Gramineae; spe-
cies in Scirpophaga and Rupela important pests
of rice (Khan, et al. 1990).
169 species (Heppner 1991). Principal genera
include: Donacaula, Patissa, and Leptosteges.
Temperate and tropical regions of both hemi-
spheres.
Common 1960; Heinrich 1937; Lewvanich
1981; Martin 1986c; Munroe 1995.
Cybalomiinae. Fig. 14.3 P. Proboscis and
maxillary palpus well developed. Chaetosema
absent. In forewing of males a fovea between
R.3+4 and R5. Hindwing with Mi from anterior
angle of cell, not stalked with Rs; CuA not pecti-
nate. Praecinctorium unilobed, its apex trans-
versely rounded. Gnathos with short lateral
arms, articulating at junction of uncus and tegu-
men, distal part triangular to spatulate, strongly
sclerotized, as long as uncus and held parallel to
it; tegumen high and narrowly arched, as in
Crambinae.
Larvae of most known species on Cruciferae
(Luquet & Minet 1982) and Capparidaceae.
About 55 species. Principal genera include:
Cybalomia, Schacontia, Stiphrometasia, Styphlo-
lepis. Southern Europe, Africa, Asia, Australia,
western North America, Central and South
America, Antilles, especially in dry regions.
Luquet & Minet 1982; Munroe 1995.
Linostinae. Fig. 14.3 Q. Robust, midsized,
broad-winged moths, with white wings, and re-
stricted black markings. Labial palpus short,
porrect. Maxillary palpus, proboscis, ocellus,
and chaetosema absent. Forewing with R2
stalked with R3+4; R5 stalked with Mi; in males
a fovea between R3+4 and R5 + Mi; CuP absent;
2 A short, free. Se + Ri of hindwing anasto-
mosed with Rs distad of cell. Frenulum hook
present in males. Praecinctorium unilobed. Un-
cus hood-shaped; gnathos with lateral arms
strong, almost transverse, widely attached to
margins of tegumen, median element shorter and
much narrower than uncus, ending in a heavily
Eugene Munroe and M. Alma Solis
sclerotized, thornlike process; valve narrow, ir-
regularly tapering, narrowly rounded at tip, un-
armed.
Early stages unknown.
3 species. Sole genus Linosta. Southern Mex-
ico to Bolivia and southern Brazil.
Munroe 1959, 1962, 1995.
Midilinae. Fig. 14.3 R. Moths often very large
and robust; wings broad, mostly with prominent
hyaline discal spots and angulate or sinuate ter-
minal margins. Maxillary palpus prominent, al-
most as long as labial palpus, its distal scaling
usually dilated. Proboscis greatly reduced in
most genera. Chaetosema well developed, often
differentiated from surrounding cranium. An-
tenna short and thick, prismatic, laminate or un-
ipectinate. Forewing with CuP absent; 2 A free.
Hindwing with Sc + Ri approximated to or
anastomosed with Rs for some distance beyond
cell; Rs and Mi not stalked. Tympanal organs
reduced. Uncus long, hardly setose; gnathos ar-
ticulating at junction of uncus and tegumen; me-
dian element usually with dorsal spinules; valve
mostly unarmed. Ovipositor short, wide; lobes
not fused dorsally; ostium unarmed; bursa with-
out signum, or at most with diffuse patches of
spinules.
Known larvae borers in Araceae: Cacographis
osteolalis from roots of Colocasia sp. (Munroe
1970) and Caladium sp. (USNM larval collec-
tion), and Eupastranaia fenestrata from stems of
Philodendron corcovadense (Pimentel, et al.
1991).
47 specis. Principal general include: Hositea,
Cacographis, Dismidila, Midila. Mexico to Boli-
via and Argentina; absent from Antilles.
Munroe 1970, 1995.
Musotiminae. Figs. 14.3 ST. Broad-winged
moths, usually small and delicate. Ocellus pre-
sent or absent. Chaetosema absent, but a radiat-
ing scale-tuft in its usual occipital position. Fore-
wing with CuP absent; 2 A short, free. Hindwing
with Sc + Ri anastomosed with Rs beyond cell;
Rs not stalked with Mi, often separating from it
well based of end of cell. Praecinctorium uni-
lobed. Uncus long, sparsely setose; gnathos usu-
ally nearly reaching tip of uncus, rarely shorter
or divided; articulating with junction of uncus
and tegumen and bearing a ventrally directed
process.
Larvae on ferns, sometimes destructive to or-
namentals.
166 species (Heppner 1991). Principal genera
include: Musotima, Ambia, Cymoriza, Undulam-
bia, Neurophyseta. Mexico to Bolivia and Argen-
tina; Old World Tropics to South Africa, Siberia,
Japan, New Zealand, Micronesia, Samoa.
Munroe 1972, 1995; Phillips & Solis 1996;
Yoshiyasu 1985.
Nymphulinae. Figs. 14.4 AD. Mainly with
slender build, long legs and long narrow wings,
usually with banded maculation, often with ca-
The Pyraloidea
taclystiform metallic spots on posterior margin
of hindwing. Forewing usually with subcostal
and radial venation crowded; R2 often stalked
with R3+4; 2 A short and free or absent. Hind-
wing with Sc + Ri anastomosed with Rs beyond
discall cell; Rs and Mi usually not stalked; discal
cell often open. Frenulum hook present in males.
Praecinctorium unilobed. Uncus long; gnathos
basally articulated with uncus and tegumen, its
median process long, slender, usually dorsally
denticulate; a sclerite connecting uncus and
gnathos with costa of valve; valve elongate, gen-
erally simple, often with hypertrophied setae or
scales on mesal surface; aedoeagus with ductus
ejaculatorius entering near middle. Ovipositor
long, extensible; lobes elongate, dorsally fused,
with reduced setae.
Early stages almost always aquatic, living
either in standing water on vascular plants, often
in cases and in some species swimming from
plant to plant, or in rapid streams, usually under
webs on rocks and feeding on algae; exception-
ally the larvae of Nymphicula are terrestrial case-
makers, feeding on mosses and liverworts in
moist environments (Yoshiyasu 1980). In Acen-
tria the larvae are standing-water casemakers
and most females are aquatic, wingless and Par-
thenogenese; winged males and winged sexually
reproducing females emerge in a brief swarm late
in the season and disperse the species (references
in Batra 1977; Speidel 1984). Larvae may be air-
breathers with open spiracles and hydrofuge in-
tegument, living in air-filled cases or alterna-
tively may have closed spiracles and breathe dis-
solved oxygen through tracheal gills. Gilled lar-
vae may live in water-filled cases in various habi-
tats or under silken nets in rapid streams. Lange
(1956 a) supposed that the latter mode of life was
correlated with certain genital and venational
characters and used it to separate the tribes
Nymphulini and Argyractini, but it is now
known that this correlation breaks down (Fi-
ance & Moeller 1978; Herlong 1979; Yoshiyasu
1985). Yoshiyasu (op. cit.) has published a tabu-
lation of larval habits showing that the actual
relationships are more complex. Pupae breathe
air through the spiracles, and are formed in
cases, in chambers in the host plant, or in water-
proof but gas-permeable cocoons. Nymphuline
larvae disfigure waterlilies and other aquatic or-
namentals, and are minor pests in greenhouses.
They have been investigated for biological con-
trol of aquatic weeds (e. g. Parapoynx diminu-
talis), but their effectiveness remains to be de-
monstrated.
716 species (Heppner 1991). Principal genera
include: Nymphula, Nymphicula, Chrysendeton,
Parapoynx, Elophila, Aulacodes, Eoophyla, Pet-
rophila. Temperate and tropical regions of conti-
nents and continental islands; Micronesia; Poly-
nesia to Fiji; Cocos; introduced in Hawaii.
247
Klima 1937; Lange 1956a; Martin 1986c;
Munroe 1972, 1995; Speidel 1984; Yoshiyasu
1985.
Noordinae. Fig. 14.4 E. Moths small, robust.
Labial palpus porrect; maxillary palpus dilated
with scales distally. Ocellus present. Chaetosema
absent. Forewing with CuP absent; 2 A forming
a large loop with 1 A. Hindwing with Sc + Ri
anastomosed with Rs for a considerable distance
beyond cell; Mi and Rs short-stalked; CuA bas-
ally with pecten of hairlike scales. Males with
frenulum hook present; frenulum single in both
sexes. Tympanal structures partly imbedded in
thorax; praecinctorium reduced, unilobed. Un-
cus long and slender, acute tipped and with fine
lateral setae; gnathos with very short median ele-
ment arising from a transverse bridge; transtilla
rudimentary; aedoeagus with ductus ejacula-
torius entering at middle; valve broadly rounded;
tergum VIII and sternum VIII with complex
sclerotization bearing androconia.
Larvae living on buds or leaves of Moringa
(Mathew & Menon 1984; Tarns 1938).
Six species. Sole genus: Noorda. Near East; In-
dian subregion and islands of the Indian Ocean;
Madagascar; Australia (introduced?).
Amsel 1965; Minet 1980.
Odontiinae. Figs. 14.4 FH. Chaetosema ab-
sent. Posterior margin of forewing often with a
prominence and sometimes with a tooth of
scales; 2 A of forewing free or forming a large
pyraustine-like loop with 1 A. Uncus bilobed,
semimembranous, with anteriorly directed lat-
eral setae, sides tapering or curving to a posterior
emargination; gnathos basally fused to postero-
lateral margins of tegumen, median element
spikelike; vinculum shallow, in tribe Eurrhypini
with a corema-bearing process at each ventrolat-
eral angle; valve more or less rounded at apex,
with radially arranged recurved setae, and often
with an array of radially arranged striations;
sternite VIII with specialized androconia-bear-
ing sclerites.
Larval habits diverse: in tribe Odontiini, gen-
erally leafminers and in Eurrhypini, leaf folders,
flower and bud feeders, fruit and stem borers.
Host plants diverse, but mostly dicotyledons.
367 species (Heppner 1991). Principal genera
include: Cynaeda, Frechinia, Titanio, Tegostoma,
Noctueliopsis, Cliniodes, and the Indo-Australian
aganaid-mimetic Heortia. Arctic to south tem-
perate zones, and from equatorial rainforests to
polar and alpine tundra; numerous and diverse
in eremic habitats; often diurnal in dry or cold
environments; found on all continents and most
continental islands, including Madagascar but
not New Zealand; in the Pacific area extending
to Micronesia and Samoa.
Leraut & Luquet 1982; Martin 1986 c; Munroe
1964 a, 1972, 1977, 1995.
Wurthiinae. Fig. 14.41. Small moths with
broad, rounded, white or light-colored wings and
248 Eugene Munroe and M. Alma Solis

Fig. 14.4. Pyraloidea. AT, Crambidae: A, Parapoynx seminealis [11 mm*]; B, Petrophila jaliscalis [11 mm]; C,
Acentria ephemerella male, [6 mm]; D, Acentria ephemerella female, wingless form, [1.5 mm]; E, Noorda blitealis
[9 mm]; F, Pseudoschinia elautalis [11 mm]; G, Cliniodes vinacea [15 mm]; H, Sobanga rutilalis [12 mm]; I, Niphopyr-
alis suffidalis [9 mm]; J, Evergestis limbata [9 mm]; K, Chalcoela pegasalis [9 mm]; L, Abegesta reluctalis [8 mm];
M, Catharia pyrenaealis [8 mm]; N, Pyrausta tyralis [6 mm]; O, Pyrausta unifascialis [11 mm]; P, Uresiphila rever-
satis [12 mm]; Q, Syngamia florella [8 mm]; R, Pantographa acoetesalis [14 mm]; S, Liopasia andrealis [28 mm]; T,
Glyphodes extorris [13 mm]. [*= length of forewing]

bombycoid or eucleoid facies. Head small. La-
bial palpus short, upturned, somewhat divergent.
Maxillary palpus, proboscis, ocellus, and chaeto-
sema absent. Antenna bipectinate in males, ser-
rate in females. Body short and stout. Forewing
with R3 and R4 short-stalked, other radiais well
separated; cell open or hardly closed; CuP pre-
sent at termen; 2 A short, weak, free. Hindwing
with frenulum single in both sexes; Sc + Ri anas-
tomosed with Rs; cell open; base of CuA without
pecten. Praecinctorium transversely flattened, bi-
lobed; tympanal chamber large, transversely
oval, extending posteriad beyond sternite II;
spinula apparently absent. Male genitalia com-
pact, highly modified, uncus short, deeply di-
vided; gnathos short, bilobed; subteguminal pro-
cess prominent; vinculum massive; valve reduced
and thornlike; aedoeagus long and decurved. Fe-
male genitalia with ovipositor lobes fitting into
the broad tergite VIII; apophyses reduced; due-
The Pyraloidea
tus bursae short; corpus bursae elongate with
longitudinal folds except posteriorly; signa ab-
sent; ductus seminalis from right posterior part
of corpus bursae.
Larvae and pupae distinctive in ecology and
structure, living in silken cases in the arboreal
nests of Oecophylla and Polyrhachis ants. Larvae
with head small, body fusiform, constricted be-
tween segments, pinacula absent, shields weak or
absent; thoracic legs normal, abdominal legs nor-
mal or reduced in size, crochets in a uniordinal
circle; chaetotaxy and other features illustrated
and biology discussed by Kemner (1923) and
Roepke (1916).
7 described species plus a considerable number
undescribed. Sole genus: Niphopyralis [= Wur-
thia]. The Indo-Malayan region. We are indebted
to Michael Shaffer of the Natural History Mu-
seum, London, for information on this sub-
family.
Evergestinae. Fig. 14.4 J. Resembling Pyrausti-
nae in most respects. First segment of porrect or
obliquely ascending labial palpus usually at least
as long as second, third segment short. Ocellus
present. Chaetosema absent. Forewing with CuP
absent; 2 A short and free. Hindwing with Sc +
Ri anastomosed with Rs for some distance be-
yond cell; Mi short-stalked with Rs. Males with
subcostal frenulum hook. Praecinctorium uni-
lobed or weakly bifid. Uncus long; gnathos long,
slender, articulated with sides of tegumen, spi-
nose dorsally; transtilla complete; valve usually
unarmed; aedoeagus generally angled behind
middle, with ductus ejaculatorius entering at that
point. Corpus bursae with a pair of signa, usu-
ally round and spinulose, occasionally fused.
Larvae with morphology similar to that of
Pyraustinae and diagnostic characters uncertain;
feeding mainly on Cruciferae and Capparidaceae
occasionally on other plants.
137 species (Heppner 1991). Principal genera
include: Evergestis, Crocidolomia, both contain-
ing crucifer pests, Orenaia, notable for reduced
eyes and other high-altitude adaptations. All
continents and most continental islands, from
arctic and alpine to tropical and south temperate
zones, in the Pacific ranging to Samoa and Mi-
cronesia; species most numerous in Holarctic re-
gion, but genera more diverse in tropical Amer-
ica and Asia.
Martin 1986 c; Munroe 1972, 1995.
Glaphyriinae. Figs. 14.4 - L . Dichogaminae,
new synonym; Alatuncusiinae, new synonym.
Small to midsized moths. Labial palpus with
basal segment long, second and third segments
often progressively diminishing. Ocellus usually
present. Chaetosema absent. Forewing with CuP
absent, 2 A short, diverging from 1 A, distally
free. Hindwing with Sc + Ri anastomosed with
Rs beyond end of discal cell; Rs connate or
short-stalked with Mi from anterior angle of cell;
CuP present; base of CuA pectinate with fine
249
scales or a basal patch of such scales present be-
hind CuA, grading into an area or patch of
wider, spatulate scales toward medial area. Fren-
ulum hook strong in males. Frenulum single in
males, multiple in females. Praecinctorium sim-
ple. Male genitalia with uncus well developed;
gnathos much reduced or absent; tegumen short,
forming an arch much higher than vinculum;
juxta U-shaped or annular, with high, narrow,
often spinose, lateral elements; penis short and
cylindrical, usually with short, strong cornuti.
Short, high ovipositor, with membranous,
strongly setose ovipositor lobes and slender,
moderately short apophyses; ductus bursae
short, with sclerotized collar and sometimes ad-
ditional armature; ductus seminalis joining cor-
pus bursae posteriorly on right side, often wid-
ened and distorted or coiled near junction, often
with sclerotized armature, tending to extend
onto corpus bursae.
Larval habits various: leaf folders, often on
Cruciferae or Capparidaceae; on Opuntia stems;
borers in heads of Typha spp.; makers of flat-
tened cryptic cases on lichens; parasites of
psychid caterpillars; and inquilines in nests of
vespid wasps.
About 200 species. Principal genera include
Glaphyria, Hellula (pests of crucifers), Lipo-
cosma, Dicymolomia, Dichogama. Nearctic and
Neotropical, a few species of Hellula in the Old
World.
Munroe 1964 b, 1972, 1995; Solis & Adamski
in press.
Cathariinae. Fig. 14.4 M. Moths small, dark
gray to blackish with rounded wings, resembling
other high-altitude holarctic pyraloids such as
Metaxmeste (Odontiinae) by convergence. Labial
palpus with bushy scaling. Ocellus present.
Chaetosema absent. Forewing of males with
frenulum hook. Frenulum of males single, of fe-
males multiple. Male genitalia with gnathos ab-
sent; uncus wide, distally rounded, with scattered
spines dorsally; valve not fanlike, but narrow,
elongate, distally rounded, unarmed; aedoeagus
bent behind middle with long, anterior coecum
and one prominent cornutus. Female genitalia
with bursa copulatrix unarmed. Tympanal or-
gans sexually dimorphic; reduced in females;
praecinctorium simple, with long scales; fenestra
media absent; coxal countertympanum very
large, internal; fornix (= cadre tympanique) pro-
truding, not angled; tympanum without spinula;
tympanal ridge lateral in males, almost dorsal in
females; tympanal sac replaced by wide depres-
sions.
Larvae and biology undescribed.
One species. Sole genus: Catharia. Europe: at
high altitudes in the Pyrenees and Alps.
Marion 1962; Minet 1980.
Pyraustinae. Figs. 14.4N-T. Chaetosema ab-
sent. Forewing with CuP absent or represented
by a fold; 2 A often forming a large closed loop
250
with 1A. Hindwing with Mi usually short-
stalked with Rs, base of CuA not pectinate. Sub-
costal frenulum hook in males present or absent.
Praecinctorium bilobed, in tribe Spilomelini
strongly so. Gnathos almost always rudimentary
or absent; a straplike subscaphium fused to
ventral wall of anal tube; vinculum usually with
a pair of rods descending from ventrolateral an-
gles and bearing androconia. Female genitalia di-
verse.
Adults of pyraustine webworms (Loxostege
sp.) may migrate in swarms and immatures can
defoliate fields and forage areas. Stem-borers in-
clude the European corn borer and relatives
(Ostrinia). Species of several genera bore in
stems and fruits or cucurbits, sweetpotato, egg-
plants and tomatoes, and in shoots of mahogany,
coffee, and other tropical trees. Species of Ategu-
mia have been utilized as biological control
agents against weedy Melastomaceae in Hawaii
and elsewhere in the Old World and Salbia haem-
orrhoidalis has been introduced to combat Lan-
terna spp. in Hawaii. The aquatic Samea multipli-
calis and Niphograpta albiguttalis, have been con-
sidered for control of aquatic weeds in tropical
and subtropical countries. The colors of adult
pyraustines may be cryptic, sometimes in a
bright, flower-matching mode, and mimicry of
aposematic Lepidoptera and of Hymenoptera is
common, especially in the tropics. Black-and-
white dazzle patterns occur in such genera as An-
ania and Desmia.
About 7400 species (Heppner 1991). Principal
genera include (tribe Pyraustini): Pyrausta; Lox-
ostege and Achyra, containing pests of pastures
and field crops; Hahncappsia; Paratalanta; Ec-
pyrrhorrhoe; Thliptoceras; Placosaris; (tribe
Spilomelini): Phaedropsis; Lygropia; host ria;
Syllepte; Herpetogramma; Pilocrocis; Lineodes,
and Leucinodes, Sceliodes, Daraba, and their al-
lies, borers in solanaceous fruits; Udea which in-
cludes the greenhouse leaf tier and relatives;
Lamprosema; Blepharomastix; Nacoleia; Bradina;
Desmia, including leaf rollers on grape; Hymenia
and Spoladea, pests of beets and relatives; Pilet-
ocera, with extremely varied androconia-bearing
modifications of antennae, legs, wings and geni-
talia, widely distributed in Indo-Australian and
Pacific regions; Anarmodia; Diaphania, with
pests of cucurbits; Glyphodes; Parotis; Stemorr-
hages, containing large, Old World species of
silky green color; Omiodes, including pests of
beans, palms and bananas; Maruca, tropical
bean pests; Palpita; Polygrammodes; Pachynoa;
Conogethes, including pests of peaches and coni-
fers in the Palaeotropics; Cirrhochrista; Azochis,
shoot borers in coffee and other tropical trees;
Omphisa, Megastes, and relatives, borers in
sweet-potato stems; Siga, large and massive
pyraustines, with wingspans exceeding 100 mm;
Agathodes and Terastia, borers and webbers of
Erythrina shoots and flowers; Nomophila, whose
Eugene Munroe and M. Alma Solis
type species, N. noctuella, formerly considered a
paradigmatic cosmopolitan insect, is now known
to be an Old World member of a complex of 12
geographically restricted species (Munroe
1973 b); Psara; Pycnarmon; Marasmia; Salbia;
Dichocrocis; and the mimetic Hyalea and
Trichaea, with dioptid and hymenopterous mod-
els, respectively. Cosmopolitan, except in Ant-
arctica.
Clavijo A. 1990; Hampson 1899 a, b; Klima
1937, 1939; Maes 1987, 1994, 1995 a, b; Martin
1986c; Mathew & Menon 1988; Munroe 1973b,
1976, 1995; Munroe & Mutuura 1968, 1969,
1970, 1971; Mutuura & Munroe 1970; Popescu-
Gorj & Constantinescu 1977.
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 15. The Axioidea and Calliduloidea
Jol Minet
Although they probably do not represent sister
groups, these two monotypic superfamilies ap-
parently belong to a monophyletic group also in-
cluding the Hedyloidea, Hesperioidea, Papilio-
noidea, Drepanoidea and Geometroidea (Minet
1991). These seven superfamilies share a few sy-
napomorphies, notably - in the adult - a more
or less pronounced reduction of the fenestrae lat-
erales (a pair of metathoracic "windows" adja-
cent to, and mesad of, the supraphragmal scle-
rites - see Minet 1982), and a distinct modifica-
tion of the secondary arms of the metafurca: the
mesal edges of their internal laminae are indeed
closely parallel (e.g. Brock 1971: Fig. 46a) or
even posteriorly convergent.
Axioidea and Calliduloidea share at least the
closeness of hindwing veins Sc + R and Rs, but
this apomorphy is also observable in several
other ditrysian groups: Pyraloidea, Thyridoidea,
Drepanidae, and so on. Contrary to those of the
Axioidea, the mature larvae of the Calliduloidea
lack, on their abdominal segments 17, a
secondary seta (or group of setae) which is
present - cephalad of L3 - in the Drepanoidea,
most Geometroidea, and at least some members
of the butterflies (e. g. Parnassius).
Axiidae. The only member of the Axioidea. This
family has at least two imaginai autapomor-
phies, viz.: (a) hindwing with Se + R approxi-
mated to Rs beyond the apex of the discal cell;
(b) A7 with a pair of pocket-like organs, asso-
ciated with the spiracles. Apomorphy (b) occurs
nowhere else in the Lepidoptera.
Adult medium-sized, broad-winged and often
brightly coloured (Figs. 15.2 A, B). Antennae
scaled dorsally, bipectinate in both sexes (to or
almost to the apex). Eyes without interommati-
dial setae. Ocelli present, usually rather small.
Chaetosemata absent. Pilifers either weak or well
developed. Proboscis unsealed, varying from
short to rather long. Maxillary palpi very small
(unsegmented in Axia margarita). Labial palpi
porrect or slightly ascending; segment 3 short.
Tegula: ventral process never sharply pointed.
Mesepimeron and meron with a short, or rather
short, line of junction. Metascutum extremely
narrow mesally. Tibiae occasionally with numer-
ous loose hair-scales; epiphysis present; spurs
(rather long): 0-2-4. Tarsomeres spined only dis-
tally (ventrally). Arolium and pulvilli present;
claws often with a ventral tooth (on their inner
surface). Frenulum well developed in both sexes,
multiple in the female; male retinaculum: a long
subcostal hook. Spinarea present (Fig. 15.1 A: sp
- definition: Minet 1990). Forewing venation
(same figure): R free, arising far from apex of
discal cell (beyond the latter, Rs2 touches Rs3 or
fuses with it for a distance, thus forming an are-
ole); M2 arising about midway between Ml and
M3; no distinct M stem in the cell; M3 and
CuAl separate, CuAl and CuA2 being still more
widely separated; CuP absent. Hindwing: Se + R
approximated to Rs for some distance, especially
beyond the apex of the discal cell; M2 arising
midway between Ml and M3, or slightly closer
to Ml; CuP absent; 3 A long. No antero-abdomi-
nal tympanal organs; Al dorsum with a fairly
broad median sclerite; sternum 2 with short an-
terolateral processes, short apodemes and, usu-
ally, distinct venulae; A7 with a pair of lateral
pockets (tympanal organs?) which open caudad
and are associated with the spiracles (Forbes
1936; Bourgogne 1951; Minet 1983). Male ster-
num 8 plate-like; saccus present (e. g. Forbes
1936, Rungs 1947); valvae undivided; uncus
hooklike; gnathos reduced to a pair of thin lat-
eral arms which run along the tegumen edges;
phallus not fused to the juxta. Female genitalia:
corpus bursae unarmed, elongate, with one or
two constrictions; venter 8 membranous (ostium
bursae located on it); apophyses all present, of
moderate length.
Descriptions of the early stages are available
for only two species, viz. Axia margarita (Chr-
tien 1916) and Axia napoleona (Reisser 1933; Mi-
net in press). Egg of the flat type, more or less
ovoid, with a few broad longitudinal grooves
(very shallow in Axia napoleona). Last instar
larva: cervix with a small, fleshy, middorsal pro-
tuberance; forecoxae close together, but not
fused; most segments divided into annulets; on
thorax and abdomen, most setae short and
clubbed; secondary setae scarce, one lying cepha-
lad of L3 on A 1 - A 7 ; LI and L2 remote from
each other on A 1 - A 8 ; ventral prolegs short,
anal prolegs stronger; proleg crochets in a
curved, roughly triordinal, mesoseries. Pupa: an-
tennae long; labial palpi, maxillary palpi and
fore femora concealed; pronotum broad; abdo-
men punctate, with two moveable segments (A5
and A6); AIO more or less conical (at least poste-
riorly), with 3 to 4 pairs of pointed teeth. The
earthy cocoon lies in the soil, near the surface.
The adults come to light at night. At rest, their
forewings form a steep ridged roof concealing
hindwings and abdomen. The known larvae feed
externally on certain species of Euphorbia (Eu-
phorbiaceae). The family inhabits southern
Europe, North Africa and the Middle East (War-
258
Tetragonus lycaenoides. Scales = 5 mm. sp: spinarea (G. Hodebert).
ren 1912, Freina & Witt 1987, Freina 1994). It
contains six species, traditionally placed in two
genera: Epicimelia (Fig. 15.2 A) and Axia (Fig.
15.2B).
Callidulidae. This family is the only member of
the Calliduloidea. It is characterized by at least
seven apomorphic traits (Minet 1991), among
which three seem to be especially significant: (a)
foreleg with an apical pair of stronger spines on
tarsomere 4, but without well developed spines
on tarsomere 5; (b) presence of a sclerotized
bridge uniting the valvae ventrad of the juxta; (c)
ovipositor flat and quadrilobate.
Medium-sized to fairly large moths, with
broad wings and rather slender bodies (Figs.
15.2C I). Antennae filiform or slightly clubbed,
but never pectinate (dried specimens with the
subapical part of the antenna somewhat bent,
the apex being turned up); flagellum scaled dor-
sally (its proximal section may also be scaled
ventrally); sensilla trichodea short. Compound
eyes: interommatidial setae minute. Ocelli pre-
sent, except in Pterothysaninae. Chaetosemata
well developed (very wide in Callidulinae), with
small scales between their setae. Pilifers and pro-
boscis always present; proboscis unsealed.
Maxillary palpi minute. Labial palpi: segment 2
porrect or more or less ascending; segment 3
often porrect, showing at the apex a well defined
vom Rath's organ. Ventral process of tegula
never pointed, frequently rather short. Meso-
thoracic anepisterna well developed. Metascu-
tum either fairly broad mesally (Pterothysani-
nae), or practically divided (Griveaudiinae, Calli-
dulinae). Rear part of the metascutellum often
high, flat and vertical. Tibiae sometimes spined
(hindtibiae occasionally with androconial tufts);
epiphysis short to rather long; spurs 0-2-4
(hindleg medial spurs often shorter than apical
spurs), 0-2-3, or 0-2-2. Foretarsus: tarsomere 4
Jol Minet
ventrally provided with a pair of strong apical
spines; distitarsus without spines or - rarely -
with a few minute spines. Arolium well devel-
oped; pulvilli cleft; claws simple (i. e. without
teeth). Male without subcostal retinaculum; fren-
ulum present (short) in both sexes, even when
humeral angle is expanded. Spinarea often ab-
sent, but present in at least two genera: Griveau-
dia and Tetragonus (contrary to previous state-
ments concerning the Callidulinae: e. g. Com-
mon 1969 and Minet 1990). Wings occasionally
with a number of loose spatulate scales; in males,
fringes of long hair-scales may occur along the
hindwing inner margin (Pterothysanus: Fig.
15.21) or beneath the forewing dorsum (Corn-
elia)'. Forewing venation (Figs. 15.1 B, C): R free;
areole rarely present; Rs2 and Rs3 always
stalked; Rsl either free or stalked with other Rs
branches; Rs4 usually free (not so in Griveaudia);
M and CuA veins free, M2 arising distinctly
closer to M3 than to M l ; CuP replaced by a fold;
1 A + 2 A without basal fork. Hindwing: Se + R
shows a recurrent humeral spur, sometimes very
short, and it approaches or touches Rs for a dis-
tance, sometimes before the end of the discal cell;
this cell frequently open; M2 closer to M3 than
to M l ; no tubular CuP; 3 A either reduced or
well developed. Abdomen: no tympanal organs;
lateral bars of dorsum 1 anteriorly connected to
sternum 2 through complete tergosternal scle-
rites; sternum 2 often elongate (its apodemes
short); terga 3 - 6 usually wide, but narrowed in
Pterothysaninae. In the male: tergum 8 consider-
ably lengthened in Griveaudia, sternum 8 reduced
to a pair of rods in Callidulinae; at least one pair
of androconial pencils on intersegmental mem-
branes 7 8 or 89; vinculum sometimes with a
saccus; valvae joined mesally, ventrad of the
juxta; no complete gnathos; uncus narrow dis-
tally, more or less hook-like; phallus with a coe-
cum, sometimes very short. Female genitalia:
The Axioidea and Calliduloidea
bursa copulatrix unarmed or variously orna-
mented; ostium bursae touching or emarginating
the anterior edge of the arched 8th sternum;
apophyses rather well developed; ovipositor flat-
tened, quadrilobate.
Eggs of the fiate type, ovoid in Helicomitra
(Minet 1990), elliptical and flattened in Griveau-
diinae (D. C. Lees, pers. comm.) and Calliduli-
nae (e.g. Nagano 1916, Holloway et al. 1987,
Nishio 1987). Only known in Callidulinae (e. g.
Chistyakov & Belyaev 1987 and Chistyakov
et al. 1994), the last instar larva has conspicuous
primary setae, but no secondary setae; pronotal
shield wide, with 5 pairs of setae; L group bise-
tose on TI; Al with a pair of lateral glands; LI
and L2 remote from each other on Al - A 7 , but
close together on A8; prolegs short (five pairs;
crochets arranged in biordinal circles). Pupa:
maxillary palpi sometimes exposed; labial palpi
largely concealed; proboscis either short {Heli-
comitra) or long {Tetragonus, Pterodecta); fore-
femora concealed; midlegs extending further,
caudad, than the antennae; abdomen with two
moveable segments in Callidulinae (A5; A6), and
possibly fewer in Pterothysaninae (Minet 1987);
cremaster present in Callidulinae, absent in Pter-
othysaninae. Pupation in a flimsy silken cocoon
(spun between foliar lobes) in Callidulinae; pupa
probably subterranean in Pterothysaninae.
The adults may be nocturnal or diurnal. Early
stages are still poorly known. The larvae of Calli-
dulinae feed on various ferns, rolling or tying fo-
liar lobes to shelter (e. g. Holloway et al. 1987,
Nishio 1987, Kobes 1990, Wang 1993, Chistya-
kov et al. 1994).
Callidulidae contain eight genera and approxi-
mately 60 species. Until recently, most authors
have regarded "Pterothysanidae" and Calliduli-
dae as separate families, but these taxa were
placed in a single family by Minet (1986 and
1990), who also assigned to it a genus previously
referred to the Noctuidae {Griveaudia). Calliduli-
dae comprise three subfamilies, which are easily
distinguishable at the imaginai stage:
Key to the subfamilies
1 Forewing crossvein rm short and rather
thick; in the hindwing: discal cell closed,
Se + R and Rs separate beyond its upper
angle, M3 and CuAl arising separately, 3 A
very short (Fig. 15.1 B); male 8th abdominal
sternum well developed 2
- Forewing crossvein r - m thin and compara-
tively long; hindwing with the discal cell
open between Ml and M2, Se + R approxi-
mated to or fused with veins Rs beyond
Ml point of origin, M3 and CuAl connate
or stalked, 3 A usually well developed (Fig.
15.1 C); male 8th abdominal sternum re-
duced to a pair of rods Callidulinae
259
2 Ocelli present; forewing with falcate apex
and Rsl to Rs4 on a common stem (Fig.
15.1 B); foretibia without strong apical
hook; abdomen: terga large to very large . .
Griveaudiinae
- Ocelli absent; forewing not falcate, with Rs4
separated from the other branches of Rs;
foretibia short, producing a strong apical
hook; abdominal terga 3 - 6 distinctly nar-
rowed Pterothysaninae
Pterothysaninae have one genus in Madagas-
car, Helicomitra, and another, Pterothysanus
(Fig. 15.21), in the oriental region. Adults are
essentially nocturnal in the former, but diurnal
or both diurnal and nocturnal in the latter. Lar-
vae and their hostplants remain unknown.
Griveaudiinae contain a single genus, Griveau-
dia, and are restricted to Madagascar. Adults
show a pronounced sexual dimorphism (Figs.
15.2 F - H ) , and sometimes fly during the day
(D. C. Lees, pers. comm.). At rest, they hold
their wings in a roof-like arrangement. Their
early stages remain to be studied/discovered.
Callidulinae are mainly oriental, with a few
species that reach palaearctic Asia. Adults usu-
ally hold their wings vertically like the advanced
butterflies (e. g. Tweedie 1988); they are nor-
mally diurnal, but some of them occasionally
come to light at dusk. This subfamily is com-
posed of five genera: Tetragonus (Fig. 15.2E),
Cornelia, Pterodecta (Fig. 15.2C), Callidula and
Cleis (Fig. 15.2 D).
Acknowledgements
I am extremely grateful to the reviewers of the
manuscript, Dr D. C. Lees (NHM, London) and
Dr M. J. Scoble (NHM, London). The former
provided valuable, unpublished information on
the biology of Madagascan Callidulidae. Useful
information on this family was also received
from Dr M. A. Solis (USDA, Washington,
D. C.), Dr P. Viette (Bar-sur-Aube) and Dr J. D.
Weintraub (Philadelphia). I thank Dr C. Rungs
(Ajaccio) and my wife, Christiane, for helping
me to find the early stages of Axia napoleona
(Axiidae). Finally, I am indebted to Mr G. Hode-
bert (MNHN, Paris) for illustrating wing vena-
tions.
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Fig. 15.2. Adults of Axiidae (A, B) and Callidulidae ( C - I ) : A, Epicimelia theresiae; B, Axia nesiota; C, Pterodecta
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 16. The Butterflies: Hedyloidea, Hesperioidea and Papilionoidea
Phillip R. Ackery, Rienk de Jong & Richard I. Vane-Wright
Butterflies (Rhopalocera), comprising the ditry-
sian superfamilies Hesperioidea (skippers), Papi-
lionoidea (true butterflies) and, according to
some authors, the Hedyloidea, are amongst the
most conspicuous, aesthetically pleasing and
therefore popular of all groups of organisms.
The wealth of taxonomic studies available, as in
most groups, tends either to be faunistic, cover-
ing all butterflies of a given region, or systematic,
treating entire groups of butterflies on a global
basis. The emphasis of this chapter is on the lat-
ter type of approach, but the value of faunistic
studies on butterflies cannot be overlooked.
Ackery (1984) gave an overview of faunistic
studies related to butterflies. We restrict our-
selves here to noting principal works published
subsequently. These include: Saudi Arabia
(Larsen, 1984), Egypt (Larsen, 1990), western
Palaearctic (Weidemann, 1986, 1988), north-
western E u r o p e (Bink, 1992), Switzerland (Gei-
ger, 1987), Caucasus (Nekrutenko, 1990), Afro-
tropical region (Ackery et al., 1995), Tanzania
(Kielland, 1990), Kenya (Larsen, 1991), Sikkim
(Haribal, 1992), Thailand (Pinratana, 1977-88),
Malay Peninsula (Corbet & Pendlebury, 1992),
Laos ( M o t o n o & Negishi, 1989), Borneo (Ot-
suka, 1988, 1991), the South East Asian islands
(Tsukada, 1982), P a p u a New Guinea (Parsons,
1991), N o r t h America (Scott, 1986), Hispaniola
(Schwartz, 1989), and Costa Rica (DeVries,
1987). In addition, D'Abrera's ambitious series
of volumes is now nearing completion (D'-
Abrera, 1971 - 1 9 9 5 ) . These works, together with
the extensive older literature, contain a wealth of
distributional, biological and phenological data
for butterflies, f r o m global down to local levels.
Lepidoptera biogeography, including many ex-
amples based on butterflies, is discussed by Hol-
loway & Nielsen (121).
In a c o m p a n i o n publication (de Jong, Vane-
Wright & Ackery, 1996) we have addressed vari-
ous problems associated with the classification of
butterflies, specifically the relationship of Papili-
onoidea to Hesperioidea, their inter-relation-
ships with the Hedyloidea and other m o t h
groups, and the associations of the convention-
ally recognized butterfly families and subfamil-
ies. For this earlier study we examined 103 char-
acters across 59 species representing all recently
recognized butterfly subfamilies, together with
exemplars of 15 m o t h groups. We introduce our
review here with an identification key to subfam-
ilies, before going on to discuss the available
characters widely cited in support of both the
traditional concept of 'butterflies' and the gen-
erally recognized butterfly sub-groups. Through-
out, we relate this accepted classification to the
results of our preliminary phylogenetic analysis
(de Jong et al., 1996), and to general butterfly
biology.
Key to families and subfamilies of Hedyloidea,
Hesperioidea and Papilionoidea
1 Antennae at least twice the width of the
scape apart; eyes with a complete mar-
ginal ring of reduced ommatidial facets;
forewing radial veins (Fig. 16.9 A) all
arising directly f r o m the discal cell
(Hesperioidea) Hesperiidae 2
Antennae at most 1.5 times the width
of scape apart; eyes usually without 'eye
ring' (some Papilionidae possess a par-
tial eye ring b u t it lacks distinct o m m a -
tidial facets); forewing with at least one
radial vein stalked, either with another
radial (Fig. 16.9B, C) or with M l . . 7
2(1) Antennal base lacking a lateral scale
tuft extending over the eye ('eyelash');
third segment of labial palp long, slen-
der, awl-like, perpendicular to the erect
second segment; head with one pair of
chaetosemata; Palaeotropical
Coeliadinae
'Eyelash' present; head with two pairs
of chaetosemata; third segment of labial
palp variable but never as above . . . 3
3 (2) Forewing cell always very long, longer
than posterior margin and more than
two-thirds as long as costa; forewing
vein M 2 arising nearer to vein M 3 t h a n
to vein M l ; antennal club stout, bent
back at its base; male secondary sexual
characters absent or, at most, as tibial
brushes; heavily built, large, wingspan
often more than 40 mm; Neotropical
(one species just reaching U S A )
Pyrrhopyginae
Forewing cell generally shorter; anten-
nal club variable, bending in middle or
beyond (if bent at base then Palaeotrop-
ical); generally not heavily built, some-
times slender and delicate 4
4 (3) Forewing vein M 2 arises mid-way be-
tween M l and M3; mid tibiae unspined
(one exception); male secondary sexual
characters, if present, never as a brand
or stigma on forewing, often as a costal
fold and/or tibial brushes or hairpencils;
264 Phillip R. Ackery, Rienk de Jong & Richard I. Vane-Wright

Fig. 16.1. Papilionidae ( A - F ) and Pieridae ( G - M ) (natural size). A, Troides helena helena. , Parnassius apollo
apollo. C, Bhutanitis lidderdalii lidderdalii. D, Baronia brevicornis brevicornis. E, Graphium sarpedon choredon. F,
Papilio machaon gorganus. G, Delias aglaia aglaia. H, Pieris brassicae brassicae. 1, Euchloe ausonia simplonia. J,
Colias hyale. K., Eurema daira. L, Dismorphia lata laia. M, Pseudoponlia paradoxa.

larvae on dicotyledons (with rare excep-
tions); cosmopolitan Pyrginae
Forewing vein M2 arises mid-way be-
tween Ml and M3, or is closer to M3;
midtibiae spined or unspined; male sec-
ondary sexual characters never as a cos-
tal fold or tibial brushes, often as a
stigma or brands on the forewing; lar-
vae on monocotyledons (with rare ex-
ceptions) 5
5 (4) Forewing vein M2 straight and about
midway between veins Ml and M3 bas-
The Butterflies: Hedyloidea, Hesperioidea and Papilionoidea
ally; hindwing discal cell longer than 11 (8)
half length of wing; foretibial epiphysis
fully developed, reduced or absent; no
male secondary characters; slender-bod-
ied; basking with all wings fully spread;
cosmopolitan except Indo-Australian
area Heteropterinae
- Forewing vein M2 basally curving to
M 3 (mostly well-marked but indistinct 12(11)
in a number of genera); hindwing cell as
long as or shorter than half length of
wing; foretibial epiphysis always fully
developed; generally with peculiar bask-
ing position (forewings partially spread,
hindwings fully spread) 6 Forewing vein M2 arising from end
6 (5) Median nervure within hindwing cell
forked, the lower branch terminating at
the origin of vein M3; discocellular
veins of hindwing angled in a line that
crosses the posterior margin near the 13 (12)
tornus; midtibiae unspined; Papuan-
Australian Trapezitinae
- Median nervure within hindwing cell, if
present, unforked or, if forked, lower
branch terminating well above the
origin of vein M3; discocellular veins
of hindwing angled in a line that
crosses the outer margin near the tor- 14(13)
nus; midtibiae spined or unspined; fore-
wing vein M2 basally often markedly
curved to vein M3; cosmopolitan . . . .
Hesperiinae
7(1) Antennae usually filiform but some-
times bipectinate; males with fully func-
tional frenulo-retinacular system (He-
dyloidea Hedylidae 15 (11)
- Antennal flagellum usually distinctly
thickened or davate distally, never pec-
tinate; males lacking frenulo-retinacular
system (Papilionoidea) 8 Antennal base usually displaced from
8 (7) Vein 2 A of forewing runs to, or almost
to, the posterior margin (Fig. 16.9 B);
3 A of hindwing absent (Fig. 16.9 B, ex- 16(15)
cept Baronia), foretibial epiphysis pre-
sent Papilionidae 9 fied, less than half the length of the
- Vein 2 A of forewing either loops with
1A (Fig. 16.9 A), is very short, or is ab-
sent altogether (Fig. 16.9 C); 3 A of
hindwing invariably present; foretibial
epiphysis absent 11
9 (8) Pretarsal claws strongly curved; 3 A of
hindwing present; monobasic, a single
species from Mexico Baroniinae 17(16)
- Pretarsal claws long and almost straight
(at least in female); 3 A of hindwing ab-
sent 10
10 (9) Forewing with a long 'basal spur' or cu-
bital cross-vein (Fig. 16.9 B); pretarsal
claws symmetrical Papilioninae 18(17)
- Forewing 'basal spur' short or, more
commonly, absent; pretarsal claws of
male asymmetrical Parnassiinae
265
Foreleg fully developed, bearing dis-
tinctly bifid claws (Fig. 16.9 E) Pieridae
12
Number of foreleg tarsal segments usu-
ally reduced at least in males; if tarsi of
males are not reduced in number, then
the pretarsal claws are simple or only
bear a basal endodont 15
Forewing vein M2 stalked with Ml +
Rs2 + 3 + 4; hindwing vein Sc + R
secondarily fused with Rs before middle
of the wing; hindwing vein M2 stalked
with Ml; monobasic, a single African
species Pseudopontiinae
of discal cell; hindwing vein Se + R
ot secondarily fused with Rs; hind-
wing vein M2 arising from discal cell
13
Forewing with five radial veins present,
usually all arising from a common stalk;
forewing cubitus 'quadrifid'
Dismorphiinae
Forewing with three, four or five radial
veins present, at least one arising inde-
pendently from the cell; forewing cubi-
tus 'trifid' 14
Hindwing humeral vein usually long;
patagia membranous; tegumen longer
than uncus; "whites" Pierinae
Hindwing humeral vein usually greatly
reduced or absent; patagia strongly scle-
rotized; tegumen usually considerably
shorter than uncus; "yellows or sul-
phurs" Coliadinae
Antennal base adjacent to the margin of
the compound eye, and usually indent-
ing it; antennae lacking carinae Lycae-
nidae 16
margin of compound eye; antennae tri-
carinate (Fig. 16.91) Nymphalidae. . 21
Hindwing humeral vein usually present;
male prothoracic leg considerably modi-
pterothoracic legs; male foreleg coxa
usually strikingly extended (Fig.
16.9 G) Riodininae
Hindwing humeral vein seldom present;
male prothoracic leg not greatly modi-
fied; male foreleg coxa at most only
slightly extended 17
Hindwing usually 'tailed' on vein
CuA2; foreleg coxa not arched upward
distally Lycaeninae
Hindwing lacking tails on vein CuA2;
foreleg coxa arched upward at distal
end 18
Pterothoracic tibial spurs present; eyes
invariably 'hairy' Curetinae
Pterothoracic tibial spurs absent; eyes
usually 'bare' 19
266 Phillip R. Ackery, Rienk de Jong & Richard I. Vane-Wright

Fig. 16.2. Nymphalidae (natural size). A, Libythea geoffroy philippina. B, Argynnis paphia paphia. C, Acraea horta.
D, Heliconius charitonia simulator. E, Vanessa polychloros. F, Melitaea cinxia. G, Megalura coresia. H, Colobura
dirce. I, Biblis hyperia. J, Limenitis populi. K, Apatura iris. L, Charaxes jasius jasius.

19(18) Forewing with only four radial veins; 20 (19) Male foretarsus segmented and clawed
forewing veins Sc and R do not anasto- Miletinae (Liphyrini)
mose Miletinae (Miletini) Male foreleg tarsomeres reduced to a
Forewing usually with five radial veins single stubby-tipped segment Poritiinae
(if only four, then R anastomoses with 21 (15) Terminal structure of female foreleg
Sc) 20 with equal and unreduced c l a w s . . . 22
The Butterflies: Hedyloidea, Hesperioidea and Papilionoidea
Terminal structure of female foreleg
usually lacking claws (labial palp sel-
dom elongate, thorax lacking dense
golden, orange or red pubescence) . 23
22(21) Labial palp very long; thoracic pubes-
cence sparse Libytheinae
Labial palp unexceptional in length;
thorax clothed in dense golden, orange
or red pubescence Calinaginae
23(21) Forewing vein 2 A present, looped to
vein 1A (and hindwing discall cell al-
ways closed by a tubular v e i n ) . . . . 24
Forewing vein 2 A usually present only
as a basal spur or completely absent
(when rarely present, hindwing discal
cell is open) 25
24 (23) Forewing with veins greatly swollen at
the base Satyrinae (Haeterini)
Forewing with veins not greatly swollen
at base Danainae
25 (23) Forewing usually with at least one vein
greatly swollen at the base (Fig. 16.9 C);
hindwing discal cell usually distinctly
closed Satyrinae (in part)
Forewing with veins usually not greatly
swollen at base; hindwing discal cell
either open or closed 26
26 (25) Antennal flagellum usually bare dor-
sally, except at extreme base; frequently
large, 'owl' or 'blue' butterflies
Morphinae
Antennal flagellum scaled dorsally (but
sometimes sparsely) for most of length
27
[The remaining nymphalid subfamilies are really only
separable on characters of the early stages (see Harvey,
1991 ) - supplementary features based on adult mor-
phology are non-exclusive and can only be used as a
guide. ]
27 (26) Forewing often narrow; hindwing cell
may be closed by a tubular vein; ptero-
thoracic pretarsal claws simple or bifid;
larva with lateral scoli arising between
segments T1 and T2 and between seg-
ments T2 and T3; larval hosts fre-
quently Passifloraceae, Flacourtiaceae
or Violaceae Heliconiinae
Forewing usually broader; hindwing cell
never closed by a tubular vein (Fig.
16.9 D); pterothoracic claws simple; lar-
val lateral scoli arising either at mid-
points of segments T2 and T3, or ab-
sent; larval hosts various 28
28 (27) Body of robust build; aedeagus strik-
ingly long; larva often 'smooth', with
bifid tail 29
More slightly built butterflies; aedeagus
unexceptional in length; larva usually
spinose, lacking a bifid tail 30
29 (28) Forewing veins Rs3 and Rs4 shortly
stalked; parapatagia with at least some
267
trace of sclerotization; saccus unexcep-
tional in length; forewing costa fre-
quently strikingly serrated; larval hosts
diverse Charaxinae
Forewing veins Rs3 and Rs4 with a
longer common stalk; parapatagia un-
sclerotized; saccus long; forewing costa
smooth; larval hosts commonly Ulma-
ceae Apaturinae
30 (28) Larvae spinose, covered with rows of
scoli; larval hosts various, principally
Asteraceae, Urticaceae, Acanthaceae,
Lamiaceae, Scrophulariaceae; eggs
never 'spiny'; males always lacking hy-
pandrium; forewing cubitus lacks a
basal spur Nymphalinae
Larval form variable may have long
pectinate lateral spines, large median
dorsal spines only, incomplete but
prominent dorso-lateral rows of spines;
larval hosts principally Euphorbiaceae,
Sapindaceae, Caprifoliaceae; eggs may
be 'spiny'; hypandrium may be present;
spur may be present on forewing cubi-
tus Limenitinae
Relationships of Hedyloidea,
Hesperioidea and Papilionoidea
Six characters are identified by de Jong et al.
(1996) which might support monophyly of
Hesperioidea plus Papilionoidea: brain with
large optic lobes, small protocerebral lobe and
small deuterocerebrum (Kristensen, 1976, after
Ehnbohm, 1948); oblique dorsal longitudinal
muscle of mesothorax twisted (Kristensen, 1976,
after Ehrlich & Ehrlich, 1963); mesothoracic an-
episternum varying from less than one third of
episternum to virtually absent; second median
plate in forewing base partly or entirely beneath
base of vein 1A + 2 A; splitting back of vein
Rs3 + 4 in pupal forewing (after Spuler, 1892;
Headlee, 1907; Tillyard, 1919); tenth segment of
pupal abdomen ventrally with pair of anterior or
anterolateral tubercles (Minet, 1991). Of these,
only two (reduced mesothoracic anepisternum
and second median plate of forewing base be-
neath vein 1 A + 2 A) seem entirely reliable. The
other four require further investigation.
With respect to grouping the Hedyloidea,
Hesperioidea and Papilionoidea as a clade, first
suggested by Scoble (1986), de Jong et al. (1996)
discuss ten characters that relate to this problem:
mesothoracic aorta with horizontal chamber
(Kristensen, 1976, after Hessel, 1966, 1969); lat-
eral arm (including fused elements from the 1st
and 2nd tergites) present between the 1st and 2nd
abdominal segments (Minet, 1991, after Brock,
1971); mesoscutum with secondary line of weak-
ness near median notai wing process (Minet,
1991); metascutellum dorsally very narrow (Mi-
268 Phillip R. Ackery, Rienk de J o n g & Richard I. Vane-Wright

Fig. 16.3. N y m p h a l i d a e (natural size). A, Morpho achilles achilles. B, Amathusia phidippus niasana. C, Haetera
piera. D, Satyrus actaea castiliana. E, Danaus plexippus plexippus. F, Brassolis sophorae sophorae. G , Calinaga
buddha buddha.

net, 1991); dorsal laminae of secondary meta-
thoracic furcal arms fused mesially (Kristensen,
1976, after Brock, 1971); apophysis of metatho-
racic furca sagittate (Scoble, 1986, after Ehrlich,
1958 ; Brock, 1971); tergum of first abdominal
segment strongly pouched (Scoble 1986); eggs
upright (Minet 1991, after Common, 1975); lar-
val anal comb present (Scoble, 1986); pupa gir-
dled (Minet, 1991). Not one of these characters
is universal and unique for (Hedyloidea +
Hesperioidea + Papilionoidea). In our parsi-
mony analyses (de Jong et al., 1996) we only pre-
sent information for six of these suggested char-
acters. The position of Macrosoma (the represen-
tative genus of Hedyloidea) varies according to
the sub-set analysed but is usually closely associ-
ated with either Hesperioidea or Papilionoidea
(see also section on Hedyloidea).
The Butterflies: Hedyloidea, Hesperioidea and Papilionoidea
HESPERIOIDEA
Hesperioidea are generally treated as comprising
a single family, Hesperiidae. Separation of the
giant skippers as a distinct family (e. g. Freeman,
1969) is not tenable (see Hesperiinae, below).
Hesperiidae (skippers). Most Hesperiidae are
easily recognized as such, in set specimens by
their wide head and general appearance, or in
the field by their characteristic flight. Although
there is some disagreement about relative rank
of the higher taxa, monophyly of Hesperiidae
has not been seriously challenged. The family
emerged as a monophyletic group in all our par-
simony analyses (de Jong et al., 1996). Five of
the following seven features are unique and uni-
versal for Hesperiidae.
Antennae at least twice the width of the scape
apart. In all other Lepidoptera, including Papili-
onoidea, the antennae are at most 1.5 times the
width of the scape apart, and often considerably
less. Eyes with a complete marginal ring of re-
duced ommatidial facets (Ehrlich, 1960). A par-
tial, glossy eye ring lacking distinct ommatidial
facets occurs in many Papilionidae. It is not cer-
tain if this is a transformation of the same char-
acter. Third axillary sclerite of forewing base very
wide and indented anteriorly, narrow and curved
posteriorly, forming a very irregular Y-shaped
structure. Remarkably constant in Hesperiidae
(Fig. 16.8 F) and not found in other Lepidoptera
(e. g. Fig. 16.8 D, E). In addition, there is a liga-
ment from the middle of the posterior end of this
sclerite to the base of 2 A. Vein R of hindwing
apparently simple, touching Sc at a short distance
from base to form a narrow cell; R completely
fused with Sc, indiscernible as a separate vein. In
other Lepidoptera with a humeral cell, the base
of R is free, closing the humeral cell, with R be-
coming indiscernible only if the bases of Sc and
R are fused into a common stem that forks into
Sc and Rs. Area of very small, specialized scales
on upperside of hindwing, covering basal part of
R (and Sc if both veins are proximate). Special-
ized scales are found in the humeral area (be-
tween Sc and costa) on the upperside of the
hindwing in all Papilionoidea and Hesperioidea,
as well as in a number of ditrysian moths (Kuy-
ten, 1974). The Hesperiidae, however, possess
this unique additional area of specialized scales.
Mesoscutellum overhanging metanotum. This is
due to a strong development of the mesoscutum
and mesoscutellum. The only other Lepidoptera
with a comparable condition are Libytheinae
(Nymphalidae), but this does not appear to be
an homologous development as in this case the
metathorax is drawn under the mesoscutellum
and the latter "topples over" so that its dorsal
side takes on a caudal position. Larvae with a
"neck". Found in all Hesperiidae for which lar-
vae are known, except the Megathymus group;
not known in any other Lepidoptera.
269
Many Hesperiidae have hyaline or semi-hya-
line spots, especially on the forewings. The col-
ourless or pallid scales of these spots stand per-
pendicularly to the wing membrane. This feature
seems unique for Hesperiidae, but it is absent in
many species, or sometimes present in females
only. Kristensen (1976) and Nielsen & Common
(1991) record the antennal thickening in Hesperi-
oidea as subapical, as against apical in Papilio-
noidea. In many Hesperiidae, however, the thick-
ening is apical and ends bluntly. Philpott (1924)
mentioned an aperture in the foretibial epiphysis
of Hesperiidae. Robbins (1990) has suggested
that this "aperture", which he observed as a
groove devoid of acanthae (microtrichia), is an
autapomorphy for the family, an interesting pos-
sibility deserving a more extensive study over a
wide array of species of skippers, butterflies and
moths.
There are about 3500 species of Hesperiidae,
the family having a worldwide distribution (al-
though absent from New Zealand), with greatest
diversity in the tropics, especially the Neotropi-
cal region. Well over 500 genera are recognized,
most of which are restricted to either the Old
World or the New World. Some, however, have
a wider distribution, occurring in the Holarctic
(e. g., Hesperia, Ochlodes, Carterocephalus), the
Holarctic plus Neotropical regions (Erynnis, Pyr-
gus), or show a pantropical distribution (Celaen-
orrhinus). Most skippers are small, inconspicu-
ous and dull-coloured, but a number of tropical
species are of medium size, and a few show bril-
liant colours. The vernacular name relates to the
typically very fast and darting flight. Most are
generally day-active, but a number, such as Cel-
aenorrhinus fritzgaertneri (DeVries et al., 1987)
and various species belonging to Erionota and
related genera, are crepuscular, or may even be
partially nocturnal (e. g., Hasora on Sulawesi -
pers. obs.; Chaetocneme in Papua New Guinea
Parsons, 1991). Many of these species have red
eyes instead of the usual dull brown (de Jong,
1983).
There are some remarkable examples of
pattern convergence affecting genera of different
subfamilies, particularly in the Neotropical re-
gion. For example, the structurally uniform pyr-
gine genus Phocides includes species that in ap-
pearance differ widely amongst themselves, but
resemble closely species of pyrrhopygine genera
Pyrrhopyge, Jemadia and Mimoniades. These
cases of putative mimicry have not been studied
and it remains to be discovered, for example, if
particular groups can be regarded as models or
mimics. Numerous secondary sexual characters
occur in skippers, mainly in males in the form of
presumed scent-distributing organs (andro-
conia). These occur most obviously on the wings
(Sellier, 1972) and legs (Watson, 1893), but in
some cases they are located on other parts of the
body (e. g. the metathorax). Frequent occurrence
270 Phillip R. Ackery, Rienk de Jong & Richard I. Vane-Wright

Fig. 16.4. Lycaenidae (1.5 natural size). A, Styx infernalis. B, Riodina lysippus. C, Hamearis lucina. D, Corrachia
leucoplaga. E, Euselasia gelon. F, Emesis tenedia. G, Poritia hewitsoni. H, Liptena undularis. I, Miletus symethus.
J, Curetis thetis. K, Lucia limbaria. L, Polyommatus icarus. M, Lycaena phlaeas eleus. N, Anthene (Lycaenesthes)
emolus. O, Thecla betulae.

of such organs in Lepidoptera prompted Jordan
(1923) to remark that "insects possibly have less
sense than the higher mammals, but probably
more senses".
Hesperiid eggs are dome-shaped, with a nearly
smooth to strongly ribbed or reticulate chorion.
Larvae, generally elongate, with a large head and
narrow neck, are usually greenish, yellowish or
whitish, but in some cases brightly coloured
(e. g., Coeliadinae: Igarashi, 1992). They usually
live in shelters made of leaves spun together, or
just folded over, although a few, such as Mega-
The Butterflies: Hedyloidea, Hesperioidea and Papilionoidea 271

Fig. 16.5. Lycaenidae (A, B), Hesperiidae (CL), Hedylidae (M) (natural size). A, Eumaeus minijas peruviana. B,
Liphyra brassolis major. C, Pyrrhopyge phidias bixae. D, Bibasis sena sena. E, Pyrgus malvae malvoides. F, Urbanus
proteusproteus. G, Celaenorrhinus eligiuspunctiger. H, Euschemon rafflesia. I, Heteropterus morpheus. J, Trapezites
symmomus symmomus. K, Hesperia comma benuncas. L, Megathymus yuccae yuccae. M, Macrosoma tipulata.

thymus and related genera which feed in the cau-
dex of Agavaceae (Freeman, 1969), are borers.
Pupation usually takes place in the shelter. Clark
(in Pennington, 1978) depicted early stages of a
number of African species in some detail. About
70 families of flowering plants are utilised as
hosts. The larvae of a few species may be eco-
nomically important, such as those of Erionota
thrax which feed on b a n a n a leaves, Hidari irava
on palms, Nyctelius species on sugar cane, and
Parnara species on rice (Ralshoven, 1981). Lar-
vae of Metardaris cosinga (Pyrrhopyginae) are
eaten by people. Larvae of Megathymus species
are the " w o r m s " in bottles of mescal.
Subfamily divisions and relationships. The clas-
sification of Hesperiidae is mainly based on Wat-
son (1893) and Evans (1937, 1949, 1951-55).
Evans' classification is only worldwide at sub-
family level. Within subfamilies, Evans proposed
separate generic groupings for Africa, the rest of
the Old World, and for America. Other world-
wide classifications, such as those of Mabille
( 1 9 0 3 - 0 4 ) and Voss (1952), do not change this
overall picture. Scott (1985) and Scott & Wright
(1990) introduced larval characters and sug-
gested a phylogeny, but they did not document
their samples and used characters in an ambigu-
ous way. Six subfamilies are recognized here:
272 Phillip R. Ackery, Rienk de Jong & Richard 1. Vane-Wright
Coeliadinae, Pyrrhopyginae, Pyrginae, Heterop-
terinae, Trapezitinae and Hesperiinae. Some are
characterized by unique features, but Pyrginae
and Hesperiinae are possibly paraphyletic assem-
blages, since they are heterogeneous (the
Hesperiinae less so) and their unique features are
absent in many members. They are only distin-
guished here for the sake of convention.
In our parsimony analyses (de Jong et al.,
1996) not a single grouping of the representatives
of these six subfamilies appears consistently, with
the exception of Celaenorrhinus + Euschemon,
and all skippers other than the Coeliadinae. Thus
the well-characterized Coeliadinae are probably
the sister group of the rest of Hesperiidae, and
this can be supported by two synapomorphies
shared by Pyrrhopyginae, Pyrginae, Heteropteri-
nae, Trapezitinae and Hesperiinae: Head with
two pairs of chaetosemata (Jordan, 1923). Not
known elsewhere in Lepidoptera. Coeliadinae,
Papilionoidea and a variety of moths have a sin-
gle pair, while some moths appear to lack chae-
tosemata altogether. Antennal base with a lateral
scale tuft extending over the eye (eye lash). This
feature does not occur in any other Lepidoptera,
including Coeliadinae. However, its occurrence
within Pyrrhopyginae is sporadic, being present
in Metardaris and Oxynetra, but absent in a
larger number of genera, including Pyrrhopyge.
Non-coeliadine Hesperiidae could be divided
into two groups, but these are not supported in
our parsimony analyses. Pyrrhopyginae + Pyrgi-
nae share the following features: Discal cell of
forewing longer than posterior margin. Universal
for Pyrrhopyginae, but many Pyrginae have a
shorter discal cell. A long cell also occurs in Coe-
liadinae. Male secondary sexual characters con-
sist of a costal fold on the forewing and tibial
brushes or hair pencils (sometimes associated
with metathoracic structures). No brands occur
on the forewing. However, many species lack
male secondary sexual characters, and the only
secondary sexual characters in Pyrrhopyginae
are tibial brushes. Tibial brushes are also found
in some Coeliadinae. At first sight there is no
apparent reason why Pyrrhopyginae could not
be a subordinate taxon of Pyrginae. However, if
the various types of tibial brushes are all homol-
ogous, which is not at all certain, and the Coelia-
dinae are sister to the rest of the Hesperiidae,
then the grouping (Pyrrhopyginae + Pyrginae)
would be based on symplesiomorphies. If, on the
other hand, the tibial brushes are not all homolo-
gous, then there is no reason for uniting Pyrrho-
pyginae and Pyrginae.
Since Pyrrhopyginae are well-defined, they are
kept separate here. The grouping (Heteropteri-
nae 4- Trapezitinae + Hesperiinae) is little better
characterized: Male secondary sexual characters
consist of brands or stigmata on the forewings.
There is never a costal fold or tibial hairpencil.
However, many species, and even the whole sub-
family Heteropterinae, lack secondary sexual
characters. Brands and stigmata are also found
in Coeliadinae, so this could be a symplesiomor-
phy and thus not a sound basis for uniting Tra-
pezitinae and Hesperiinae. The far from univer-
sal occurrence of a peculiar basking posture in
Trapezitinae and Hesperiinae (see below) could
be an indication of close relationship between the
two subfamilies, but it remains to be seen if unit-
ing the two while omitting the Heteropterinae
would not create a paraphyletic group. Larval
foodplants monocotyledons (with very few excep-
tions). Almost all other Hesperiidae feed on di-
cotyledons. This appears to be a synapomorphy,
if we accept Coeliadinae as sister group of the
rest of Hesperiidae. Possibly related to a diet of
grasses and other tough monocotyledons is a re-
duction and eventual loss of larval mandibular
teeth (Scott & Wright, 1990). Larval mandibles
of grass-feeding Satyrinae and Morphinae
(Nymphalidae) also lack teeth.
Coeliadinae. The subfamily is defined by a sin-
gle, unique and universal feature: Third segment
of labial palp long, slender, awl-like, perpendicular
to the erect second segment. There is much varia-
tion in the palps of other Hesperiidae, but the
condition found in Coeliadinae is never ap-
proached. Secondary sexual characters in this
subfamily comprise brands and stigmata on the
wings, otherwise characteristic for Hesperiinae
and Trapezitinae, as well as tibial hair tufts as
found in Pyrginae. Species of Choaspes are re-
markable for their strongly hairy eyes; most
Hesperiidae have bare eyes. Evans (19515) be-
lieved Coeliadinae to be Old World equivalents
of New World Pyrrhopyginae. There is little con-
vincing morphological evidence for such a sister-
group relationship, although Bibasis and Pyrrho-
pyge did combine in some cladograms produced
in our parsimony analyses.
This small subfamily of about 75 medium-
sized species is confined to the Old World trop-
ics, and consists of 8 genera (including Coeliades,
Bibasis and Hasora). Athough most species can
be found by day, many are mainly active (or at
least observed) in early morning and early eve-
ning, and sometimes individuals come to light.
The common African species Coeliades forestan
has been observed about 5 km off the coast of
Senegal, coming to the lights of a ship (R. de
Jong, pers. obs.). The Indo-Australian species
Badamia exclamationis is strongly migratory, at
least in part of its distributional range (Com-
mon & Waterhouse, 1981). Larvae feed almost
exclusively on dicotyledons. About 20 plant fam-
ilies have been recorded (most of which are also
utilised by Pyrginae), including Asclepiadaceae,
Combretaceae, Myristicaceae and Solanaceae.
Pyrrhopyginae. Monophyly of this subfamily
is supported by two characters: Abdominal ter-
gum I compressed between thorax and tergum II,
vertical, more or less scale-like. In other Hesperii-
The Butterflies: Hedyloidea, Hesperioidea and Papilionoidea
dae tergum I is never so compressed, being more
or less inflated, oblique to almost horizontal.
Club of antenna stout, spindle-shaped, bent at its
commencement. The vernacular name Golf-Club
Skipper for the North American Pyrrhopyge ar-
axes reflects this character. Similarly-shaped an-
tennae, but of much lighter build, occur in a few
pyrgine genera (e. g., Caprona, Netrobalane, Ab-
antis).
The Pyrrhopyginae comprise almost 150 me-
dium-sized, strongly built and sometimes bril-
liantly coloured species, in 20 genera including
Pyrrhopyge, Sarbia, Mysoria, Myscelus and Pas-
sava. The group is Neotropical, with a single spe-
cies just extending into North America. Al-
though well represented in Central America, the
subfamily is absent from the Caribbean islands.
At rest, individuals sit with their wings spread
flat (like many Pyrginae). In addition to being
mimicked by (or mimicking) species belonging to
other hesperiid subfamilies, many pyrrhopygines
mimic each other. For example, Elbella includes
butterflies similar to species of Pyrrhopyge, Mi-
moniades and Jemadia. Larvae are only known
to feed on dicotyledons, of which 11 families (all
utilised by the Pyrginae) have been recorded, in-
cluding Fagaceae, Anacardiaceae and Sterculia-
ceae.
Pyrginae. This subfamily, which never ap-
peared as a monophyletic group in our parsi-
mony analyses, is weakly characterized by sec-
ondary sexual characters that, although appar-
ently unique, are absent in many species: Male
forewing with costal fold. The fold encloses an-
droconial scales (de Jong, 1972). Female with an
abdominal pheromone gland at tergum VII (de
Jong, 1975). The distribution of this character
needs further study. We follow Evans (1949) in
including Euschemon rafflesia in this subfamily.
Curiously, this Australian species has a frenulum
and retinaculum in the male. In our parsimony
analyses it always formed a monophyletic group
with Celaenorrhinus.
Comprising about 1000 species in 160 genera,
including Spialia, Astraptes, Euschemon, Chae-
tocneme, Coladenia, Tagiades and Erynnis, the
Pyrginae form the second-largest subfamily of
Hesperiidae. Distribution is worldwide, but over
600 species occur in the Neotropical region only.
Celaenorrhinus, with about 75 species, is amongst
the largest genera of Hesperiidae, and the only
one with a pan tropical range (de Jong, 1982).
The widespread Pyrgus is remarkable for the
large number of apparently incomplete or only
recently completed speciation events (de Jong,
1987). Many pyrgines rest on the underside of
leaves, with their wings spread out flat. Larvae
feed almost exclusively on dicotyledons, of which
about 45 families have been recorded, including
Acanthaceae, Fabaceae, Rosaceae, Malvaceae,
Tiliaceae, Annonaceae, Bignoniaceae and Lecy-
thidaceae. A few species, including certain mem-
273
bers of Urbanus, have switched from dicotyle-
dons to grasses or other monocotyledons.
Heteropterinae. Two weak features support
monophyly: Second segment of labial palp porrect
and hairy. The palpi are very similar to those
found in Pyrgus and relatives (Pyrginae). Epiphy-
sis of fore tibia reduced or absent. Evans ( 1951 - 5)
suggests that absence is due to the epiphysis be-
ing deciduous in many species. There are also
species in this subfamily with a fully developed
epiphysis, as in other Hesperiidae. Evans (1949)
based his Heteropterus group, comparable with
the present subfamily, mainly on the long discal
cell of the hindwing, reaching beyond the middle
of the wing. However, a long hindwing cell is
also found in some other Hesperiidae, some Pap-
ilionoidea, as well as other Lepidoptera.
With about 150 species in 16 genera, the Het-
eropterinae are a small subfamily. Distribution is
remarkable, the group being well represented in
the Afro tropics (including Metisella) and the Pa-
laearctic, very weakly represented in the Nearc-
tic, and having about 100 species in the Neotrop-
ics. One genus, Carterocephalus, is Holarctic.
The Neotropical Dalla is one of the largest gen-
era of Hesperiidae (about 80 species). Heteropte-
rine skippers are generally small, slender-bodied
and broad-winged. Larvae have only been found
on grasses (Poaceae). Contrary to the next two
subfamilies, species belonging to this group lack
secondary sexual characters.
Trapezitinae. A single character supports mo-
nophyly of this subfamily: Discocellular veins on
hindwing directed toward wing apex (Fig. 16.9 A;
Evans, 1949). In other Hesperiidae, the discocel-
lular veins are either vertical or point toward the
base of the wing.
This small subfamily of about 60 species in 16
genera is mainly restricted to Australia, with a
northward extension through New Guinea to the
Moluccas. The genera include Trapezites, Neo-
hesperilla and Anisynta. Individuals of some spe-
cies bask with forewings spread partially and
hindwings spread fully, a posture otherwise
found only in Hesperiinae. Three groups of gen-
era can be distinguished based on larval food
preferences: (a) feeding on Xanthorrhoeaceae
and Poaceae (e. g., Trapezites, Toxidia), (b) on
Cyperaceae (e. g., Hesperilla), and (c) on Irida-
ceae (Mesodina). These groupings are supported
by morphological differences in larvae and pu-
pae (Common & Waterhouse, 1981).
Hesperiinae. This large and diverse group is
loosely held together by a single, unique but non-
universal venational character: Vein M2 of fore-
wing basally curving to M3. Well-marked in most
genera, but indistinct in others. Hesperiinae can
be divided into a number of well-defined sub-
groups, some of which may deserve formal re-
cognition as tribes, but a number of genera are
difficult to place. We include within the subfam-
ily the peculiar "Megathymidae" of authors (e. g.
274
Lindsey et al., 1931; Freeman, 1969), even
though in our parsimony analyses the genera
Hesperia and Megathymus did not always group
together. The peculiarities of the "Megathymi-
dae" (24 genera; sometimes even divided into
two tribes, e. g. Scott, 1986), particularly the nar-
row head of the adult and larva, and the absence
of the typical hesperiid neck in the latter, appear
to relate to the unique larval habit of boring in
stems and roots of Yucca and Agave. The epiphy-
sis of Megathymus and relatives is shorter than
in other Hesperiinae.
The Hesperiinae are by far the largest subfam-
ily of skippers, including more than 2000 species
in 325 genera. Distribution is worldwide, but
about half of the species occur in the Neotropics.
Hesperia and Ochlodes are Holarctic genera.
Other genera include Polites, Amblyscirtes, Ked-
estes, Telicota and Halpe. As remarked under the
previous subfamily, Hesperiinae exhibit a pecu-
liar basking posture, with the hindwings fully
spread horizontally, while the forewings are kept
only half-opened. Larvae feed almost exclusively
on monocotyledons, of which about 10 families
are utilised, including Araceae, Arecaceae, Musa-
ceae, Poaceae and Liliaceae.
PAPILIONOIDEA
Papilionoidea, the true butterflies, comprise four
families, Papilionidae, Pieridae, Lycaenidae and
Nymphalidae. Riodinidae and Libytheidae
{sensu Harvey, 1987; Ehrlich, 1958 b) are in-
cluded here within Lycaenidae and Nymphali-
dae, respectively. Despite such disagreements
about the relative rank of higher taxa, the mono-
phyly of Papilionoidea has been widely accepted,
albeit on rather scant evidence. In our parsimony
analyses (de Jong et al., 1996), the Papilionoidea
consistently form a monophyletic group. Five
thoracic characters support this arrangement:
Anepisternum of mesothorax present as a tiny
sclerite, hardly discernible, or absent (Jordan,
1902; Ehrlich, 1958 b). Elsewhere in Lepidop-
tera, the mesothoracic anepisternum is distinct,
either well-developed (most moths) or small
(Hesperiidae) (Fig. 16.8 B). If present in Papilio-
noidea, the anepisternum is a very small sclerite
at the dorsal end of the katepisternum (Fig.
16.8 A, C). Reduction is due to a dorsal shift of
the anapleural cleft. The anepisternum has seem-
ingly disappeared in a few other Lepidoptera,
e. g. Urania. In these cases, however, the an-
apleural cleft has not moved but simply disap-
peared, and so the anepisternum, although indis-
cernible as a separate sclerite, is still present.
Parepisternal suture running in a straight or
smoothly curved line from dorsal end to base of
sternum (Fig. 16.8 A, C). In non-papilionoid lepi-
dopterans, the parepisternal suture is strongly
curved before reaching the margin of the ba-
sisternum or, if not reaching the margin, is co-
Phillip R. Ackery, Rienk de Jong & Richard I. Vane-Wright
linear with it (Fig. 16.8 B). Mesophragma with
dorsal processes (Ehrlich, 1958 ). A universal
character in Papilionoidea, often very pro-
nounced, but sometimes reduced (particularly in
Lycaenidae). In other Lepidoptera there are no
traces of such processes, but in Hesperiidae in-
distinct ridges or small dorsal plates, not pro-
cesses, occur. Ventral edge of tegula attached by
membrane to the mesonotum (Jordan, 1928; He-
ring, 1958). In Hesperiidae and other lepidopter-
ans, the attaching membrane is set some distance
from the ventral edge of the tegula. Secondary
sclerite behind metascutellum (Brock, 1971).
Originally thought to be restricted to Pieridae +
Lycaenidae + Nymphalidae, and used to support
the monophyly of this grouping (Kristensen,
1976). However, it also occurs in Papilionidae,
sometimes in rudimentary form, but it can be
very pronounced (e. g., Graphium).
Brock (1971) discussed two more characters
that, according to him, supported monophyly of
Papilionoidea: fusion of precoxal and margi-
nopleural sutures in the mesothorax, and pres-
ence of an anterior sclerite at abdominal sternite
2. The first character is not only non-universal in
Papilionoidea, but also widespread in Hesperii-
dae. The second character is often so obscure,
especially in Papilionidae and Pieridae, that it is
not very useful. Elsewhere, Kristensen (1976)
suggested two autapomorphies, neither of which
appears convincing: wing-coupling mechanism
amplexiform (as it is in all Hesperioidea, except
male Euschemon) and antennae with apical clubs
(certainly poorly developed in many species, and
occurring in a number of Hesperiidae as well).
Amplexiform wing-coupling occurs in a variety
of moths (e. g. Urania, Callidula, Poecilocampa,
Thaumetopoea), and the same is true for clubbed
antennae (e. g. Zygaena, Castnia, etc.). Absence
of medial spurs on the hind tibia, proposed by
Minet (1991) and others, is certainly not unique;
a comparable condition occurs in Megathymus
and Euschemon (Hesperiidae), and Thaumeto-
poea, Mustilia and Poecilocampa. In addition,
Minet (1991) re-iterates his earlier character (Mi-
net, 1986) "dorsal enlargement, in lateral view,
of the metathoracic mera". Of the seemingly im-
pressive array of further characters presented by
Scott & Wright (1990), most require further in-
vestigation.
There are about 14000 species of Papilio-
noidea. The group has a world-wide distribution,
with greatest diversity in the tropics.
Family Divisions and Relationships. The out-
standing question centres around two conflicting
hypotheses of relationships: ((Pieridae + Papilio-
nidae) + (Nymphalidae + Lycaenidae)) (see
Scott & Wright, 1990), and (Papilionidae +
(Pieridae + (Nymphalidae + Lycaenidae))) (see
Kristensen, 1976).
The Butterflies: Hedyloidea, Hesperioidea and Papilionoidea
A number of the characters presented by
Scott & Wright (1990) to support the first hy-
pothesis, notably those based on adult muscula-
ture and eyes, and pupal structure, have for the
most part only been verified for a limited range
of taxa. Nevertheless, at least one such character
may prove to be of particular significance (R. K.
Robbins, pers. comm.): a peculiar arrangement
of proboscis extensor muscles reported by
Schmitt (1938) for Papilionidae and Pieridae, but
not found in other butterflies (which proved very
similar to each other and to a variety of moths).
Some of the characters cited by Scott &
Wright (1990) in support of Papilionidae + Pier-
idae are certainly not universal: presence of cover
flap over ostium bursae; postdiscrimen repre-
sented internally by a small anterior prong or la-
mella; forewing vein M2 close to M3 (presuma-
bly equates to 'quadrifid' condition of cubitus
see below); profurcal arms with secondary ante-
rior prong or lamella; and adult tarsi with min-
ute 'spines' (alpha-type trichoid sensilla: Ma &
Schoonhoven, 1973) on the upperside. Even the
five remaining characters are equivocal: Spitia-
sternum heavily sclerotized between the two points
of attachment to the remainder of the mesothorax.
This seems to be an extrapolation from Ehrlich's
(19586) statement "in the Pieridae the spinaster-
num is widened into a small oval or diamond-
shaped plate between the furcasternum and the
spina. This is faintly reminiscent of the lateral
expansion of the papilionids". Forewing basal ra-
dial plates hardened posteriorly. Introduced by
Scott (1985). He neither elaborates further, nor
discusses the condition found in other lepidop-
terans. We could not confirm this character. Pro-
thoracic presternum indistinguishable (after Ehr-
lich, 19586). Of doubtful polarity, Kristensen
(1976) employed presence of the presternum to
characterize Nymphalidae + Lycaenidae (see be-
low). Metathoracic scutum small, only slightly vis-
ible from the rear. Scott (1985) delimits this char-
acter thus: scutum somewhat visible from the
rear (Hesperiidae), very visible from the rear
(Nymphalidae + Lycaenidae) and scarcely visi-
ble from the rear (Papilionidae + Pieridae).
Brock (1990) suggests that the latter condition is
in fact ancestral to the Papilionoidea. According
to our findings, however, there is too much vari-
ation within the families to render this character
useful. It cannot even be stated as a generality
that the metascutum is more readily visible from
the rear in Nymphalidae and Lycaenidae than in
Papilionidae and Pieridae. Mesothoracic second-
ary sternopleural sulcus absent (after Brock,
1971). Both Brock and Kristensen (1976) suggest
that presence of this structure (in Nymphalidae
and Lycaenidae; Fig. 16.8 C) is the derived state
(hence the adjective secondary), which view we
endorse since, among Lepidopterans, the sulcus
is unique to Nymphalidae and Lycaenidae.
Scott & Wright (1990) unjustifiably employ ab-
275
sence of the sulcus as a synapomorphy of Papili-
onidae and Pieridae, together with its presence
as a synapomorphy of Nymphalidae and Lycae-
nidae.
Another potential character is the upright
pupa supported by a girdle which, where it
crosses the abdomen, is received by a thickened
groove. Poulton (1908), who investigated this in
Pieris in some detail, speculated that the pupal
girdle of the Papilionidae and Pieridae represents
the "persistent trace of a vanished cocoon". If
so, then the complete absence of a girdle in the
suspended pupae of the Nymphalidae and some
Lycaenidae might represent the final stage of this
transformation, and this could argue against the
girdle of the Pieridae and Papilionidae as a link-
ing synapomorphy. The girdle crosses different
body parts in the two families, and so even the
thickened groove may not be homologous. Fi-
nally, we must consider the unique 'quadrifid'
forewing cubitus (Clench, 1955), found in all
Papilionidae and in the Dismorphiinae amongst
the Pieridae. Generally, the Dismorphiinae have
been considered internal to the Pieridae, but they
may prove to be the sister group of all other
whites and sulphurs (see below). If so, the quad-
rifid cubitus might be a convincing synapomor-
phy for Papilionidae + Pieridae, even though
this would require interpretation of the trifid
condition in all higher Pieridae as homoplasious.
Kristensen (1976) presents four characters in
support of the alternative grouping of Pieridae
+ Nymphalidae + Lycaenidae. Two are loss-
apomorphies: absence of a fore-tibial epiphysis
(otherwise present throughout the Papilionidae,
almost all Hesperiidae and most moths), and loss
of the prospinasterno-procoxal muscle (after
Ehrlich & Ehrlich, 1963, and confirmed for only
a limited number of taxa). The third character,
maxillary palp unsegmented or absent, is also
true of most Papilionidae except Baronia. The
sole remaining character is also ambiguous:
Secondary sclerite present behind metascutellum
(after Brock, 1971). Originally scored as present
in Pieridae, Nymphalidae and Lycaenidae but
subsequently found, at least in rudimentary
form, within Papilionidae. Our investigations
suggest that the secondary sclerite is most dis-
tinct in Lycaenidae and Nymphalidae, a condi-
tion apomorphic with respect to Papilionidae
and Pieridae.
On this evidence, the case for a sister-group
relationship between the Papilionidae and Pier-
idae looks more plausible. In contrast, the results
from our parsimony analyses (de Jong et al.,
1996) tend to support Kristensen's view when
taking all butterflies and one moth at a time.
Monophyly of Papilionidae + Pieridae is only
supported when analysing all butterflies and
moths together. We conclude that the position of
the Pieridae needs further attention.
276 Phillip R. Ackery, Rienk de Jong & Richard I. Vane-Wright
Lycaenidae + Nymphalidae is a widely ac-
cepted grouping (e. g. see Kristensen, 1976;
Scott & Wright, 1990), monophyly of which is
based on the following characters: Conspicuous
differentiation of secondary sclerite behind metas-
cutellum (Brock, 1971, 1990). Apparently the
most derived state of a transformation series (see
also above). Our own findings suggest that the
metascutellar apex is unmodified in Hesperiidae,
but bears a weakly differentiated secondary scle-
rite in Papilionidae and Pieridae, a structure that
is strongly differentiated in Nymphalidae and
Lycaenidae. Secondary sternopleural suture in
mesothorax present (Brock, 1971). Universal and
unique in Lycaenidae and Nymphalidae (see also
above). Adult prothoracic presternum present
(Kristensen, 1976, after Ehrlich, 1958 6). Gen-
erally present throughout Lycaenidae and
Nymphalidae, although sometimes only indis-
tinctly separated from the propleura. Muscle
connecting fronto-clypeus and proboscis absent
(Scott & Wright, 1990, after Ehrlich & Ehrlich,
1962). This is "character 3" of Ehrlich & Ehrlich,
a loss-apomorphy only verified for a limited
number of taxa. Foreleg strongly modified, at
least in males. Modification in foreleg tarsal
structure is a universal feature of the Nymphali-
dae (Fig. 16.9 H) that recurs in males of most
Lycaenidae (Fig. 16.9 G). The mode of antennal
cleaning (by means of the midleg) must be
strongly correlated with this character (see Rob-
bins, 1990, for review of antennal grooming by
means of leg structures).
The following additional features have been
employed in attempts to further justify this
grouping (others see Scott & Wright, 1990
need verification across a far wider range of
taxa): Tarsal 'spines' confined to the under surface
(Borner, 1939). Alpha-type trichoid sensilla
(Ma & Schoonhoven, 1973), often referred to er-
roneously as 'spines', occur on the upper surface
of the tarsi in all Papilionidae, most Pieridae,
many Nymphalidae, some Hesperiidae and occa-
sionally in moths; they are never found in Lycae-
nidae. The character is clearly non-unique and
non-universal for Lycaenidae + Nymphalidae.
Adult eyes hairy (Scott & Wright, 1990). Wide-
spread but not universal within either Nymphali-
dae or Lycaenidae. A comparable condition is
found in certain moths and even in some Hesper-
iidae, e. g. Choaspes. Male foreleg coxa extending
beyond articulation with the trochanter (Fig.
16.9 G; Scott & Wright, 1990). True of many ly-
caenids but we have not identified a comparable
condition in any nymphalid. In our parsimony
analyses of all butterflies plus one moth at a time,
Nymphalidae + Lycaenidae always emerged as a
monophyletic group, but not so when taking all
butterflies and moths together (de Jong et al.,
1996).
Papilionidae (swallowtails). The phylogeny of
Papilionidae is probably the best documented of
any lepidopteran family (e. g. Munroe & Ehrlich,
1960; Munroe, 1961; Hancock, 1983; Igarashi,
1984; Miller, 1987 6). Despite this, many prob-
lems remain (Huser, 1993 ). Monophyly of the
family is, however, well supported by a number
of autapomorphies that are unique and uni-
versal:
Vein 2 A of forewing runs freely to, or almost to
posterior margin (Miller, 1987 6). In other Lepi-
doptera, forewing 2 A either anastomoses with
1 A, is very short, or may be absent altogether.
Although such a condition might appear plesio-
morphic, this feature is diagnostic for Papilioni-
dae. Third axillary sclerite of forewing base
strongly extended anteriorly, more or less reaching
anterior angle of base of 1A + 2 A. In other Lepi-
doptera it hardly reaches the midline (see Fig.
16.8 D, F). At the same time the second axillary
is more or less overhung by the raised radial
plate (Fig. 16.8 E), and the first median plate de-
flects downward. The second median plate is not
visible in dorsal view, being covered by the base
of 1 A + 2 A (Fig. 16.8 E). The latter condition
is, in varying degree, also found in other Papilio-
noidea and Hesperioidea (e. g. Fig. 16.8 F). Ap-
parently the changes are not all interrelated, and
we have treated them separately in our character
matrix (see de Jong et al., 1996). Cervical scle-
rites joined ventromedially (Miller, 19876, after
Kristensen, 1976). According to Kristensen, close
proximity of the laterocervicalia is probably an
ancestral state of Lepidoptera, but this condition
is otherwise only found in certain primitive
groups (e. g. 'Zeugloptera', 'Dacnonypha'). Cu-
bitus quadrifid (Fig. 16.9 B; Clench, 1955; Ehr-
lich, 1958 6). Within Papilionoidea this condition
is otherwise only found in Dismorphiinae. Larva
with osmeterium on prothorax (Igarashi, 1984;
Miller, 19876). This character is very distinctive,
and no swallowtail for which the early stages are
known lacks it.
A number of other characters recorded in the
literature either need confirmation, or appear
non-unique for Papilionidae, or are not universal
within the family. Such characters include ana-
tomical features of muscles (Ehrlich & Ehrlich,
1963) and heart (Kristensen, 1976), and bio-
chemical characters (presence of papiliochrome
in the wing scales; Umebachi, 1977, 1978; Umeb-
achi & Aburano, 1978; Descimon, 1983). R. K.
Robbins (pers. comm.) has pointed out to us that
the absence of an endodont on the pretarsal
claws of the Papilionidae may be a further (loss)
apomorphy for the family, especially if the Papi-
lionidae are eventually resolved as sister group
to the Pieridae (in which a long endodont is uni-
versal; see below). In our parsimony analyses (de
Jong et al., 1996), Papilionidae always emerged
as a monophyletic group.
There are about 600 species of swallowtail
butterflies, with a worldwide distribution (see
Scriber, 1984; Collins & Morris, 1985). Most spe-
The Butterflies: Hedyloidea, Hesperioidea and Papilionoidea 277

Fig. 16.6. Larvae of Papilionidae (A, B) and Nymphalidae ( C - F ) . A, Ornithoptera alexandrae (after Igarashi,
1979). B, Graphium agamemnon (after Igarashi, 1979). C, Danaus plexippus (after Stehr, 1987). D, Adelpha ixia
(after Aiello, 1991). E, Marpesia petreus (after Stehr, 1987). F, Caligo beltrao (after Whalley, 1988).

cies are spectacular, and the family includes the
birdwings (Troides and Ornithoptera), usually
considered to include the largest of all butterflies
(e. g. Ornithoptera alexandrae, O. goliath), the
true swallowtails (Papilio), kite swallowtails
(Graphium), and apollo butterflies (Parnassius).
The family affords many examples of mimicry,
including models (e. g. Pachliopta and Battus
species) and mimics, many of which are poly-
morphic and have been the subject of detailed
genetic investigations (e. g. Papilio dardanus, P.
memnon, P. polytes, P. glaucus). Much of the
classical mimicry literature is surveyed by
Turner (1984).
Papilionid eggs are circular and dome-shaped,
globular or bi-concave. Larvae have been char-
acterized as either "red-tuberculate" (Fig.
16.6 A) or "smooth green" (Fig. 16.6 B) (see
Munroe, 1961), while pupae are girdled or some-
times terrestrial (Parnassius, Baronia). Igarashi
(1979) illustrates the early stages of many Papili-
onidae in great detail.
278 Phillip R . Ackery, Rienk de J o n g & R i c h a r d I. Vane-Wright
Subfamily divisions and relationships. Three
subfamilies have generally been recognised: Bar-
oniinae, Parnassiinae and Papilioninae. The last
two can be considered sister groups based on the
following characters: Third anal vein of hindwing
absent (Fig. 16.9B; Miller, 1987 ft). This loss apo-
morphy is unique to Papilionidae among both
Hesperioidea and Papilionoidea, but occurs
widely among moths. Pretarsal claws long and
almost straight (at least in female). This condi-
tion, which recurs in some Danainae (Nymphali-
dae), needs to be checked in a wider range of
Papilioninae and Parnassiinae. The claws in Bar-
onia are strongly curved, as in most butterflies
and moths. Potential synapomorphies linking
these two subfamilies that need further study in-
clude a ventral sclerite in the cervical membrane
(Miller, 19876, after Ehrlich, 1958 b), and a lat-
eral production of the prothoracic spinasternum
at the spina (Ehrlich, 1958 ft).
Our parsimony analyses (de Jong et al., 1996)
do not support a sister group relationship be-
tween Baroniinae and the rest of the Papilioni-
dae. Instead, in most cases, a sister group rela-
tionship between Parnassius and the rest of the
family, including Baronia, is apparent. Huser
(1993) has raised a number of serious doubts
regarding the current classification of the Papili-
onidae, and it is clear that further work is still
needed.
Baroniinae. The single species included, Baro-
nia brevicornis, is well characterized by several
autapomorphies: Patagia with small, elongate
sclerotization (Ehrlich, 19586). There is some
variation in the degree of sclerotization through-
out the butterflies, but the condition encountered
in Baronia is apparently unique. Base of radial
vein in forewing upperside thickened and with a
conspicuous sclerite. Unknown in other butter-
flies. Male valve with a membranous lateral lobe.
Recorded by Miller (1987 ft) to be unique in Pap-
ilionidae, and not recorded from other butter-
flies. Two venational characters (loss of Rs3 and
fusion of R and Sc in forewing) are homoplasi-
ous within Papilionidae (Miller, 1987 ft).
B. brevicornis (which is restricted to central
Mexico) adults are both polymorphic and poly-
typic; the larvae feed on Fabaceae (Vasquez &
Perez, 1961, 1967).
Parnassiinae. Monophyly of this subfamily is
very weakly based on the following characters:
Pretarsal claws asymmetrical (Ehrlich, 1958 ft).
Not universal (e. g. does not occur in Hyper-
mnestra) but otherwise the usual condition in
males, and occurs in females of some species. A
comparable condition is found in the pterothora-
cic legs of Acraea (Nymphalidae), and even in
some Papilioninae (e. g. Parides DeVries,
1987; Huser, 1993 ). Aedeagus narrow and
heavily sclerotized. Although Miller (1987ft) rec-
ognized this as a synapomorphy for Parnassii-
nae, he recorded a similar condition in some
members of the Troidini (e. g., Pachliopta\ to
which we can add subgenus Danaus (Nymphali-
dae)), while Huser notes that this feature does
not occur in Hypermnestra. Ostial region of fe-
male abdomen heavily sclerotized. According to
Miller (1987 ft), this is a unique configuration,
but Huser (1993) again notes absence from
Hypermnestra. Possibly correlated with previous
character, as a comparable condition also recurs
in subgenus Danaus. Miller (1987 ft) and Huser
(1993 ) discuss other characters recorded in the
literature, but point out that these also exhibit
homoplasy (e. g. elongate labial palpi, incurved
m l - m 2 in forewing) or, like the sphragis, are
neither unique nor universal. In our current par-
simony analyses (de Jong et al., 1996) the two
exemplar genera (Parnassius and Bhutanitis) rar-
ely group together. For future analyses, inclusion
of the enigmatic Hypermnestra, which Hiura
(1980) and Huser (1993) have both suggested
should be placed in a separate group, will be
essential.
The subfamily as presently recognised (54-76
species in 8 genera, including Parnassius, Zeryn-
thia, Bhutanitis and Luehdorfia, together with the
doubtfully placed Hypermnestra, Huser, 1993 ft)
is largely Holarctic. Larvae mainly feed on Aris-
tolochiaceae, Crassulaceae, Fumariaceae and
Saxifrageaceae. Many Parnassius species (e. g.
P. apollo, P. mnemosyne), which are mostly mon-
tane, show spectacular but confusing geographi-
cal variation, and have been divided into innu-
merable subspecies (e. g. Bryk, 1935).
Papilioninae. Several features have been cited
to support monophyly of the subfamily, of which
perhaps only the first is convincing: Metathorax
with a distinct meral suture, including internal la-
mella (Miller, 1987 ft, after Ehrlich, 1958 ft). Ap-
parently universal in this subfamily and absent
in all other butterflies, including Baroniinae and
Parnassiinae. Forewing with a long cubital cross-
vein or 'basal spur' (Fig. 16.9 B; Miller, 1987 ft).
This character is obvious in all Papilioninae ex-
cept Teinopalpus, in which the spur is short.
Some Parnassiinae (e. g. Sericinus, Bhutanitis)
also have a short spur, and a long spur occurs in
a number of nymphalid butterflies (e. g., IIeli-
co nius, Charaxes, etc.) and in some moths (Ura-
nia). Often considered to be a vestige of CuP,
Huser (1993) questions the validity of this
character for defining the Papilioninae. Male ter-
gile VIII with pseuduncus (Miller, 1987 ft). Found
in many Papilioninae (absent from a number of
kite swallowtails; Huser, 1993 ), but not else-
where in Papilionidae. However, a structure of
similar appearance occurs in the pierid genus
Colias (Miller, 1987 ft), the nymphalid Calinaga
(Ehrlich, 1958ft) and some other butterflies.
Miller (1987 ft) and Huser (1993 ) discuss a
number of additional characters (larval abdomi-
nal saddle, ventral anal brushes along vein 2 A of
male hindwing, relative length of forewing disco-
The Butterflies: Hedyloidea, Hesperioidea and Papilionoidea
cellular veins, etc.) which, although distinctive,
are variable in occurrence. In the great majority
of our parsimony analyses (de Jong et al., 1996)
the three exemplar genera (Papilio, Graphium
and Troides) did not form a monophyletic group.
Papilioninae comprise about 550 species in 17
genera, including Teinopalpus, Lamproptera,
Graphium (kites), Parides, Pachliopta, Atropha-
neura, Troides (birdwings) and Papilio (true swal-
lowtails). A cosmopolitan group, but diversity is
greatest in the Old World tropics (Collins &
Morris, 1985). The familiar European swallow-
tail (Papilio machaon) belongs to a complex of
species found throughout the Holarctic. Adults
of many species are attracted to flowers, mud-
puddles, salt-licks and carrion, from which they
often feed while continuing to beat their wings
in a steady, characteristic fashion. With respect
to larval hosts, Troidini offer one of the best ex-
amples of oligophagy in butterflies, being exclu-
sive to a small variety of Aristolochiaceae. Other
Papilioninae use a wider variety of foodplants,
notably Annonaceae, Apiaceae, Canellaceae,
Hernandiaceae, Lauraceae, Rutaceae, Wintera-
ceae, etc. (see Ehrlich & Raven, 1965; Miller,
1987 a). Igarashi (1987) has recently described
the early stages of Teinopalpus from Magno-
liaceae, also utilized by other Papilioninae.
Pieridae (whites & sulphurs). The classification
of Pieridae is largely based on Klots (1933) and
Ehrlich (1958 ). Characters supporting mono-
phyly include:
Lateral plates of pronotum not fused medially.
These plates are fused in all other butterflies ex-
amined (own observation, after Ehrlich, 19586;
listed as "presence of a membranous cleft medio-
posteriorly in the pronotum" by Kristensen,
1976). Fore tarsus with distinctly bifid claws (Fig.
16.9E). Butterfly pretarsal claws occur with or
without an inner tooth (endodont: Clench,
1955). In Pieridae the endodont is subequal to
the outer tooth (somewhat shorter in Neophasia;
R. K. Robbins, pers. comm.), and thus the claws
are truly bifid. Ehrlich (19586) also called the
claws in the nymphalid tribe Acraeini (Heliconii-
nae) bifid, but of 25 species checked none had
the endodont longer than half the length of the
outer tooth and most had it much shorter; Pierre
(1985) notes other anatomical differences.
Amongst the Lycaenidae, the Aphnaeini have a
well-developed endodont (Eliot, 1973). Samanta
and Mana tar ia (Satyrinae; Miller, 1968) also
have a strong endodont, and genera of the satyr-
ine tribe Melanitini have fully bifid claws (Miller,
1968 - but this is restricted to the pterothoracic
or even hind legs only: Common & Waterhouse,
1981). With the exception of the Papilionidae, all
butterfly families include many species possess-
ing an endodont, but the equal 'bifid' condition
affecting all legs is confined to Pieridae. Outer
edge of forewing third axillary with tooth. In all
279
Papilionoidea examined (de Jong et al, 1996) the
third axillary is an elongate, straight or slightly
curved structure, with a smooth outer edge, ex-
cept in Pieridae which have a distinct, larger or
smaller blunt tooth, whereby the sclerite hinges
with the base of vein 1A + 2 A (Fig. 16.8 D).
Median plate 1 of forewing basal sclerites fused
with third axillary to form a single structure (de
Jong et al., 1996). As a result there is no separate
sclerite between the third axillary and the radial
plate (Fig. 16.8 D). This character has not been
found elsewhere in butterflies, or in other Lepi-
doptera studied so far. Pterin pigments in wing
scales. Pterins are thought to be of universal oc-
currence in wing scales of Pieridae, occurring in
six major forms: pterin, leucopterin, isoxan-
thopterin, xanthopterin, sepiapterin, and eryth-
ropterin. The last-named has also been found in
one (possibly two) Heliconius species (Nymphali-
dae) (Baust, 1967). According to Descimon
(1983) pterins have also been found in a few Rio-
dininae (Lycaenidae) and some moths, but he did
not state whether they were found in wing scales
or in the body. Pterins occur in a wide array of
animals, including Amphibia and Hymenoptera,
and even in plants. Possibly related to the pres-
ence of pterins in pierid wing scales is absence of
-alanine, a substance commonly found in scales
of other Papilionoidea (Umebachi & Aburano,
1978).
A number of anatomical or other characters
recorded in the literature also need further inves-
tigation. These include uni- or tri-sulcate anten-
nae (Jordan, 1898), loss of sclerotization of the
labial sclerite behind the palpal sockets, reduc-
tion or absence of prespiracular bar at base of
abdomen, absence of a muscle in prothorax run-
ning from furca to cervicle, a more posterior at-
tachment to the scutum of a muscle on the meso-
thoracic postcoxal sclerite, and frequent use of
ultraviolet coloration for specific signalling. In
the great majority of our parsimony analyses (de
Jong et ai, 1996) the Pieridae emerge as a mono-
phyletic group.
The family includes about 1000 species and is
almost cosmopolitan (New Zealand lacks native
species). Of the 75 genera, Colotis, Delias, Colias,
Catasticta, Eurema and Belenois are amongst the
most speciose. Many pierids are very conspicu-
ous, of medium size, and characteristically white,
orange or yellow in colour (striking exceptions
include the flame-red Appias nero, tiger-pat-
terned Dismorphia, and the curious jezebels, De-
lias species). The South American Dismorphiinae
include the classical mimics described by Bates
(1862), and the family provides many other cases
of mimicry, including poorly understood intra-
familial examples (e. g. Mylothris and Belenois in
Africa). Pieridae are remarkable amongst butter-
flies for the number of migrant species (Williams,
1930), seasonal variation (Shapiro, 1984), and
pest status, including the cabbage butterflies
280 Phillip R. Ackery, Rienk de Jong & Richard I. Vane-Wright
(.Pieris) and clouded-yellows or sulphurs (Colias,
on alfalfa). For access to much information on
pierid ecology in general, see Courtney (1986)
and, for Pieris in particular, Feltwell (1982).
Pierid eggs are spindle-shaped and ribbed, and
often white, yellow, orange or red. The cylindri-
cal larvae have no obvious protuberances and
are cryptic, usually green or brown, patterned
with longitudinal stripes; pupae are girdled. One
Neotropical pierid, Eucheira socialis, forms re-
markable silk nests (described and figured by
Westwood, 1834) in which the larvae feed gregar-
iously. Adult Eucheira socialis socialis, although
fully winged, fly weakly, if at all (Sattler, 1991).
Larvae are ranched for human consumption in
southern Mexico (and Aztecs evidently used the
nests to make paper), and man is now apparently
the principal agent for dispersal of this curious
butterfly (Geiger et al., 1989).
Subfamily divisions and relationships. Four
subfamilies are currently recognized: Pseudopon-
tiinae, Dismorphiinae, Pierinae (whites includ-
ing Anthocharinae and Euchloeini) and Coliadi-
nae (sulphurs). The last two subfamilies, how-
ever, are not convincingly characterized, and the
relationships between the four subfamilies are
uncertain. Two male genitalia characters suggest
a relationship between Pseudopontiinae and Dis-
morphiinae (uncus bilobate, and valvae fused
along ventral and distal margins), but these fea-
tures are not convincingly homologous in the
two groups. Klots (1933) considered the Pseudo-
pontiinae + Dismorphiinae to form a group sep-
arate from all other Pierinae.
Scott (1985) and Scott & Wright (1990) sought
to associate Pierinae and Coliadinae on the basis
of a "bump" between the base of the forewing
cubitus and the radial plate, but this was not evi-
dent in material studied by us. These two groups
do, however, share a distinctive mate refusal sig-
nal (females settle with wings spread and abdo-
men raised vertically) that does not occur in the
Palaearctic dismorphiine Leptidea (Wiklund,
1977). In a preliminary and unpublished cladistic
analysis by the late Adrienne Venables (1993)
based on morphological data (including charac-
ters studied by Klots, 1933), all eight genera of
sulphurs in her sample formed a paraphyletic
series of four or more clades at the base of the
Pierinae. The latter, however, did emerge as a
monophyletic group (which Venables classified
as the Pierini, the major subordinate taxon
within the Pierinae redefined to include the Coli-
adinae). In contrast, the membranous patagia of
the adult prothorax suggest that (Pseudopontii-
nae + Dismorphiinae + Pierinae) could form a
natural group, to the exclusion of Coliadinae.
Sulphur butterflies have strongly sclerotized pa-
tagia, considered by Ehrlich (1958 b) to be a
primitive trait. However, this could well be a re-
versal.
In conclusion, limited evidence might suggest
that the classification of pierid subfamilies
should be ((Pseudopontiinae + Dismorphiinae)
(Pierinae + Coliadinae)), as proposed by Ehrlich
(1958 6). Such a grouping, however, is not ap-
parent from our parsimony analyses (de Jong
et al., 1996). The only "traditional" grouping
found in a majority of analyses is Colias + Eu-
rema (Coliadinae), while the grouping (Dismor-
phia + rest of Pieridae) also appeared in most
results. In this context it is particularly interest-
ing to note that Venables (1993), who studied
both Leptidea and Dismorphia in addition to
Pseudopontia and a wide variety of Pierinae and
Coliadinae, concluded that the cladistic relation-
ships of the major groups within Pieridae would
probably prove to be (Dismorphiinae (Pseudo-
pontiinae (Pierinae [= paraphyletic Coliadinae
+ Pierini]))). This seems to reflect the earlier con-
clusion of Clench (1955), who recognized only
three pierid subfamilies, with Pseudopontia "an-
nectant" between Dismorphiinae and Pierinae.
However, in our parsimony analyses Pseudopon-
tia appears to be a subordinate taxon of the Pier-
inae.
Dismorphiinae. Although several characters
have been recorded as supporting monophyly of
this subfamily, only one seems reliable: Antenna
trisulcate. This condition, first described by Jor-
dan (1898), is most strongly developed at the tip
of the flagellum. Other potential characters in-
clude the short tegumen (much shorter than un-
cus); bilobed uncus; fused valvae (ventral and
lower distal margins); single corpus bursae
(double in at least a number of other Pieridae);
and presence of flavone pigments in the wings
(flavones are considered to be largely absent
from Pierinae, Coliadinae and Pseudopontia
Ford, 1941). According to Ehrlich (1958 b, after
Klots) all forewing radial veins are stalked in
Dismorphiinae; in the Dismorphia species exam-
ined by us (de Jong et al., 1996), however, vein
R arises separately from the discal cell.
This subfamily, including about 100 species (in
6 genera, including Dismorphia and Enantia
see Llorente-Bousquets, 1983), is predominantly
Neotropical with the exception of a single dis-
junct genus, Leptidea (wood-whites), in the Pa-
laearctic. Most species are fragile creatures, with
narrow bodies and wings. Larvae feed on Faba-
ceae.
Pseudopontiinae. The single species included
here, P. paradoxa, has several well-defined auta-
pomorphies: Forewing veins Ml and M2 stalked
with Rs2 + 3 + 4. This condition appears to be
unique to Pseudopontia. Hindwing vein Se + R
secondarily fused with Rs before middle of the
wing. Although most unusual for a butterfly, this
configuration recurs in a number of moths, in-
cluding Lymantria, Crambus and Thaumetopoea.
Hindwing vein M2 stalked with Ml. Appears to
be a unique feature. Lamella of metadiscrimen
The Butterflies: Hedyloidea, Hesperioidea and Papilionoidea
runs straight into the furca. Elsewhere in Papilio-
noidea the lamella of the discrimen either curves
downward to the base of the furca, or joins the
furca above its base (see Ehrlich, 1958 6).
Pseudopontia paradoxa, a fragile, whitish in-
sect with rounded, almost hyaline wings, is re-
stricted to African rainforests. In the past this
curious species has been placed in a variety of
butterfly families or elsewhere in Lepidoptera, or
even outside Lepidoptera altogether. Our parsi-
mony analyses (de Jong et al., 1996) consistently
support the idea that Pseudopontia is a pierid,
even suggesting that it may be highly "internal".
Venables (1993; see above) considered Pseu-
dopontia to be the sister group of the Pierinae
(including the Coliadinae). Heath (1977) re-
corded larvae from Opiliaceae.
Pierinae. Several characters have been pro-
posed to support monophyly of Pierinae, partic-
ularly by Scott & Wright (1990), including: uncus
hinging on tegumen so that downward flexion is
possible; antennae more clubbed; fimbriate an-
droconial scales scattered over the wings; and
first stage larva without proprioceptor seta
MXD1. However, all of these appear homoplasi-
ous, or require more investigation. Adult Pieri-
nae can be separated from all other butterflies by
the combination of bifid pretarsal claws, unisul-
cate antennae, membranous patagia, hindwing
Se + RI running freely to margin, fully devel-
oped forelegs lacking tibial epiphyses, and hind-
tibia with spurs. In our parsimony analyses, the
three exemplar genera rarely if ever group to-
gether, and the possibility that Pierinae as cur-
rently defined are a paraphyletic or even poly-
phyletic assemblage must be considered. Ven-
ables (1993), however, appears to have found a
number of synapomorphies for the group.
This subfamily comprises approximately 700
species, in about 55 genera, including Mylothris,
Delias, Colotis, Belenois, Catasticta, Euchloe, An-
thocharis, Leptosia, Ascia, Appias, Ixias, Cepora
and Tatochila. Although cosmopolitan, the Pieri-
nae have greatest diversity in the tropics. Com-
pared with other Pieridae, the larvae feed on a
different suite of plant families (mainly Brassica-
ceae, Capparidaceae, Loranthaceae, and Santa-
laceae); the American genus Neophasia feeds on
pines. The cabbage butterflies, Pieris brassicae,
P. rapae and close relatives (the latter sometimes
placed in a separate genus or subgenus, Arto-
geia) are amongst the most familiar of all insects.
Sexual signalling in Pieris, and probably many
other Pierinae, can involve hidden UV pattern
differences between males and females, produced
by the far more abundant and UV-absorbent
pterin pigments found in wing scales of males;
females are usually diffusely UV-reflectant
(Obara, 1970).
Coliadinae. As with Pierinae, there are no ob-
vious autapomorphies for Coliadinae. Dubious
characters include reduction or absence of hu-
281
meral vein in hindwing; reduction of third seg-
ment of palpus; 'raised line' on forewing (Klots,
1933); and loss of VT3 (= V3) on first stage lar-
val head. Within butterflies, Coliadinae can be
recognized by the same combination of features
as Pierinae, except the strongly sclerotized pata-
gia (which will distinguish the former); as already
noted, Venables (1993) considered the group to
be paraphyletic.
The Coliadinae, as recognized by Ehrlich
(1958 ), contains approximately 250 species in
12 genera; cosmopolitan, but with greatest diver-
sity in the tropics. Larvae feed mainly on Astera-
ceae, Fabaceae, Rhamnaceae and Zygophylla-
ceae. Sulphur butterflies include such common
and familiar genera as Colias, Eurema, Phoebis,
Nathalis, Catopsilia and Gonepteryx. Specific and
sexual signalling in sulphur butterflies often in-
volves iridescent UV reflectance patterns pro-
duced by the structure of special scales on the
upperside of the wings (Silberglied & Taylor,
1978), most notably in the males.
Lycaenidae (blues, coppers & metalmarks). Pa-
pers by Stichel (e.g. 1910-11), Clench (1955),
Shirzu & Yamomoto (1956, 1957), Stempffer
(1967), Eliot (1973, 1990; see also Fiedler, 1991),
Harvey (1987) and Robbins (1988, b) have con-
tributed to development of a workable lycaenid
classification. As considered here, Lycaenidae in-
clude Riodinidae (metalmarks) and Curetidae of
authors, but the position of the Riodinidae re-
mains debatable. Conventionally, the lycaenids
and riodinids are perceived as intimately related,
although on scant evidence. Harvey (1987) iden-
tified five characters, all based on early stages,
that favour co-ordinate sister-relationship be-
tween the two groups. In contrast, Robbins
(1988 a, 1990) presented several features of male
foreleg structure and function suggestive of a
close relationship between Riodinidae and
Nymphalidae. Both suites of characters have the
same weakness the several individual charac-
ters could be merged into a single composite (the
inevitable result of myrmecophily on the one
hand, and the non-ambulatory, sensory function
of the forelegs on the other). Martin & Pashley
(1992) have recently suggested, based on ribo-
somal RNA sequence data, that metalmarks
may form a clade with Pieridae and Nymphali-
dae but such a grouping has never been seri-
ously proposed on morphological criteria. Kris-
tensen (1976) was first to claim that, if riodinids
are excluded, Lycaenidae are rendered paraphy-
letic. Our parsimony analyses (de Jong et al.,
1996) for all butterflies support this view, the 17
exemplars nearly always forming a single group,
with the 6 riodinids consistently forming a single,
highly internal subgroup. Monophyly of the Ly-
caenidae is only upheld if the metalmarks are in-
cluded. The following characters offer some jus-
tification for recognizing Lycaenidae sensu lato:
282 Phillip R. Ackery, Rienk de Jong & Richard I. Vane-Wright

Fig. 16.7. Larvae of Nymphalidae (A, B) and Lycaenidae (C, D, F); pupae of Papilionidae (G), Nymphalidae (H)
and Lycaenidae (E). A, Hamanumida daedalus (after Van Son, 1979). B, Morpho amathonte (after De Vries, 1987).
C, Curetis brunnea (after Fukuda et al., 1987). D, Deloneura sheppardi (after Clark & Dickson, 1971). E, Liphyra
brassolis (empty puparium, after Common & Waterhouse, 1981). F, Liphyra brassolis (larva, after Common &
Waterhouse, 1981). G, Papilio ulysses (after Common & Waterhouse, 1981). H, Doleschallia bisaltide (after Com-
mon & Waterhouse, 1981).

Antennal base adjacent to margin of compound
eye, and usually indenting it (Ehrlich, 1958 ft). A
traditional character that is universal in Lycaeni-
dae, but a similar condition does occur in other
butterflies (e. g. Eurema, Biblis), and in a variety
of moths (e. g. Macrosoma, Castnia, Urania). Re-
duction of the metathoracic wing case of the pupa.
According to Harvey (1987), in all butterflies ex-
cept Lycaenidae the metathoracic wing case ex-
tends at least to abdominal segment 3, but in Ly-
caenidae it extends no further than segment 2, or
even segment 1. Spatulate lobe on larval abdomi-
nal prolegs which interrupts the mesoseries of
crochets (Borner, 1939). Unknown from other
butterflies, it is also absent in a few Riodininae
(which Harvey, 1987, interprets as secondary
loss). Absence of larval eversible prothoracic
gland. According to Harvey (1987) this gland is
present in Hesperiidae, Nymphalidae and Pier-
idae, and has been lost independently in Papilio-
The Butterflies: Hedyloidea, Hesperioidea and Papilionoidea
nidae (interpreted in this case to be correlated
with the origin of a dorsal eversible prothoracic
gland, the osmeterium) and in Lycaenidae. This
character would be more informative if we had
comparable information about moth larvae.
Other characters recorded in the literature, in-
cluding several larval features noted by Scott &
Wright (1990), need further investigation or are
also found outside Lycaenidae. The latter is also
true for the membranous thoracic patagia used
by Kristensen (1976, after Ehrlich, 1958 ) to
link Lycaenidae with riodinids. Renewed work
on functional modifications associated with the
biotic relationships between lycaenids and ants
(DeVries e tal., 1986), sometimes considered to
be a ground-plan feature of Lycaenidae + Rio-
dinidae (e. g. Vane-Wright, 1978), would seem
particularly valuable. Fiedler (1991), based on
the idea that the myrmecophilous organs of rio-
dinids are not homologous with those of true Ly-
caenidae (Harvey, 1987; DeVries, 1991), pro-
posed that the two groups should be maintained
as separate families in which myrmecophily has
evolved independently. If this is so, then it raises
a number of critical questions about parallelisms
in the two groups, such as larval shape (sug-
gested to be an adaptive response to myrme-
cophily; Malicky, 1969, 1970), or the fleshy pads
found on the larval prolegs and compact pupae
in both groups (R. K. Robbins, pers. comm.). In
practice, adult Lycaenidae s. 1. can be separated
from other butterflies by the combination of
modified male forelegs, non-tricarinate antenna,
and indentation of the eye next to the antennal
base.
This is the largest and biologically most di-
verse of all butterfly families, with more than
6000 species (Robbins, 1982), divided by Eliot
(1990) into five subfamilies (Riodininae, Poritii-
nae, Miletinae, Lycaeninae and Curetinae).
Overall, the family has a cosmopolitan distribu-
tion (including species endemic to New Zealand,
Hawaii and many other Pacific islands), but with
greatest diversity in the tropics. Although many
subgroups (e. g. blues, coppers, hairstreaks) ex-
hibit characteristic colour patterns, some (e. g.
Riodininae) resemble an almost bewildering
variety of other butterflies, and even certain
moths; the significance of this mimicry or
pseudo-mimicry is not understood. The vast ma-
jority of lycaenid butterflies are small, with rare
exceptions such as the peculiar, moth-like Li-
phyra. Some Lycaeninae, and in particular cer-
tain Theclini, are remarkable for their false head
patterns, including fluttering hindwing tails
which simulate antennae, and special behaviours
to enhance this protective illusion (Robbins,
1980).
Lycaenid eggs are frequently dome-shaped,
often with a dorsal depression surrounding the
micropyle, and usually have a densely reticulate
or deeply sculptured surface. In larval and pupal
283
stages, many lycaenids are associated with ants
(Fiedler, 1991), some larvae are carnivorous on
ant-brood or Homoptera (Cottrell, 1984), while
others feed on lichens (Henning, 1983 ). Ant as-
sociation involves many adaptations, including
special myrmecophilous organs that mediate in-
teractions with ants, notably by chemical (Hen-
ning, 1983 b) and auditory (DeVries, 1990)
means. The chemical interactions have been
shown to include, in certain cases at least, release
of amino acids as rewards or bribes for ants
(Pierce et al., 1991). This peculiarity may relate
to the use made by many phytophagous lycae-
nids of nitrogen-rich plant families (Pierce & El-
gar, 1985) or plant parts. Many specialize on
flowers or seeds, and it is possible that this spe-
cial nutritional need may have had a stronger
influence on lycaenid hostplant choice than
either plant chemistry or co-evolution (Pierce,
1984 - but see also Fiedler, 1991). These factors
could be responsible for occasional pest status
of some species, such as Lampides boeticus on
legumes. Lycaenid pupae may be girdled or un-
girdled.
Subfamily divisions and relationships. Eliot
(1973) divided Lycaenidae, excluding riodinids
and stygids, into eight subfamilies. Kristensen's
(1976) inclusion of riodinids within Lycaenidae
thus resulted in recognition of ten subfamilies by
Ackery (1984). Harvey (1987), in reinstating Rio-
dinidae to family status, distinguished several
subfamilies within the group. However, evidence
for inter-relationships between the increasing
number of subgroups remained scant.
Against this trend of splitting, Eliot (1990), in
a paper on Curetis, has remarked that the sub-
family divisions should be reduced to five: Rio-
dininae, Curetinae, Poritiinae, Miletinae and Ly-
caeninae. Although he did not give supporting
characters, this is simply formal recognition of
the five major branches shown in his earlier den-
dogram of Lycaenidae (Eliot, 1973: fig. 1), which
was based on "an intuitive evaluation of [19] se-
lected diagnostic characters", scored across 33
tribes {op. cit. : 454). Thus there may be some evi-
dence that the five subfamilies are monophyletic,
and we accept his simplified classification.
However, in our parsimony analyses (de Jong
et al., 1996) only Riodininae amongst these five
subfamilies emerge consistently as a monophy-
letic group (apart from Curetinae which com-
prise a single genus). In contrast, Martin &
Pashley (1992) were unable to demonstrate
monophyly for the two riodinids included in
their ribosomal R N A study. Another highly con-
sistent feature in all our analyses is the grouping
of riodinids with Liptena, Poritia and Miletus, in
that order. However, given many other prob-
lems, it would be premature to include Miletinae
and Poritiinae within an extended Riodininae.
A possible relationship between Poritiinae,
Miletinae and Curetinae could be supported by
284 Phillip R. Ackery, Rienk de Jong & Richard I. Vane-Wright
the following character. Foreleg coxa arched up-
wards distally, and sometimes extending beyond its
articulation with the trochanter (Robbins,
1988 a). This character, which affects both sexes
identically, seems to be the final stage of a devel-
opment starting with an unextended and un-
arched coxa, but homoplasy may be involved
here. This grouping is not supported by our par-
simony analyses. A relationship between Poritii-
nae and Miletinae is supported by the following
loss apomorphy: Pterothoracic tibial spurs absent
(Eliot, 1973; Harvey, 1987). The loss, however, is
not completely restricted to these two subfamil-
ies (e. g., it recurs in two out of six species in
Eumaeus; R. K. Robbins, pers. comm.). In our
parsimony analyses, Poritiinae and Miletinae
cluster as a paraphyletic assemblage in relation
to Riodininae + Liptena.
There are four general larval forms in Lycaeni-
dae (onisciform, carapaced, "lymantriine", tu-
berculate: Eliot, 1973). Although the taxonomic
distribution of these is not inconsistent with our
classification, we are unable to evaluate these
generalized larval phenotypes in a phylogenetic
sense. The classification adopted here must be
seen as highly provisional.
Riodininae. The six riodinid exemplars we
studied (including Styx and Corrachia) always
group in our parsimony analyses (de Jong et al.,
1996), although the grouping did not emerge in
Martin & Pashley's (1992) study. Support is of-
fered by the following three characters: Male pro-
thoracic leg less than one half the length of the
pterothoracic legs. This character needs quantifi-
cation (and recurs in Nymphalidae), but in other
Lycaenidae the male prothoracic legs are notice-
ably longer. Trichoid sensilla of female foretarsus
arranged in a single cluster (Harvey, 1987). In all
other Papilionoidea examined they are arranged
in paired clusters. Posterior apophyses of female
genitalia absent (Harvey, 1987). This loss apo-
morphy recurs in a few species of Poritiinae. Tra-
ditionally, riodinids have been recognized by the
striking extension of the male foreleg coxa
(Stichel, 1910-11). However, this feature is only
weakly developed in Styx (Robbins, 1988 a) and
Corrachia, while it is quite well-developed in var-
ious Poritiinae and Curetinae (Robbins, 1988 a).
Jordan (1898) considered the antennae of Rio-
dininae to be characteristic, having a single
ventral groove, and this was repeated by Scott &
Wright (1990). However, according to Robbins
and Harvey (pers. comm.), this may be just an
artifact caused by drying.
There are about 1250 described species, repre-
senting some 140 genera. Distribution is
worldwide but with greatest diversity in the Neo-
tropics. Major or better-known genera include
Hamearis, Dodona, Abisara, Praetaxila, Eusela-
sia, Mesosemia, Eurybia, Ancyluris, Melanis, Ca-
lephelis, Riodina, Mesene, Symmachia, Anteros,
Helicopis, Emesis, Lemonias, Audre, Theope,
Nymphidium and Stalachtis. The strange Styx in-
fernalis from Peru has been regarded as distinct
at family level. Corrachia leucoplaga represents a
monobasic tribe. Riodinine eggs are diverse
(Downey & Allyn, 1980), some being only
weakly sculptured, although most have notable
ornamentation like other lycaenids. Larvae, of
diverse general form, have been reported from
over 40 families of flowering plants (Harvey,
1987). Those larvae known to have biotic rela-
tionships with ants are usually smooth in general
appearance, while non-myrmecophilous species
are distinctly hairy (DeVries, 1991); at least one
species is recorded to feed on Homoptera (Har-
vey, 1987). Although pupae are usually girdled,
there is some variation in girdle position (Har-
vey, 1987).
Poritiinae. In our parsimony analyses, the two
genera studied (Poritia and Liptena) always
emerged as a paraphyletic group. However, the
following characters support monophyly of Pori-
tiinae sensu Eliot (1990), to include Liptenini:
Saccus in situ directed posteriorly (Eliot, 1973).
Elsewhere in Lycaenidae the saccus points ante-
riorly, as in other butterflies. However, in the
type species of Poritia the saccus is apparently
reduced to a vestige, and so cannot be assessed
in this case. Larval skin attached around pupal
cremaster (Eliot, 1973; Scott & Wright, 1990). It
is not yet certain that this condition is absent
from all other Lycaenidae; it is listed here tenta-
tively.
Two subgroups (tribes) are recognized. Lip-
tenini are African, with about 530 species (in 47
genera, including Alaena, Telipna, Durbania, Bal-
iochila, Liptena, Epitola), some of which mimic
other butterflies (e. g. Mimacraea). Larvae live
on lichens and microscopic algae, apparently a
unique habit amongst butterflies, and do not
have mutual interactions with ants (Henning,
1983 ; Fiedler, 1991). Poritiini occur in the Ori-
ental Region, with just over 50 species in 7 gen-
era (e. g. Poritia, Deramas). Larval hosts are Fa-
gaceae, and again mutual interactions with ants
have not been confirmed (Fiedler, 1991). Both
groups have similar, 'lymantriine'-like larvae,
with hairy tufts (Fig. 16.7 D), and the pupae
lack girdles.
Miletinae. In our parsimony analyses, the gen-
era Liphyra and Miletus never group together,
and we are not aware of any unambiguous syna-
pomorphy to unite them and their close relatives
as a well-founded subfamily. This may even ap-
ply to their unusual larval form and food. Larvae
with carapace under which the legs, head and anal
claspers are hidden. This is clearly the case for
Liphyrini (Fig. 16.7 F) but, although Miletini
were scored as 'carapaced' by Scott & Wright
(1990), Eliot (1973) described them as 'onisci-
form'. Larvae aphytophagous, living on homopter-
ans, homopteran secretions, ant regurgitations, or
ant brood (Cottrell, 1984). Although this is the
The Butterflies: Hedyloidea, Hesperioidea and Papilionoidea
only major group of butterflies that appears to
be entirely aphytophagous, such eating habits re-
cur in a variety of other lycaenids, as well as in
a few moths.
This subfamily contains approximately 150
species, in 18 genera and two tribes (Miletini, Li-
phyrini; Eliot, 1986, 1990). Although found
mainly in the Old World tropics (e. g. Spalgis,
Taraka, Miletus, Allotinus, Megalopalpus, Lach-
nocnema, Aslauga), a few representatives occur
in the Holarctic region. Larvae are wholly aphy-
tophagous. Pupae of Liphyrini and those of most
Miletini lack a girdle. Bethune-Baker (1925) il-
lustrates the remarkable armoured early stages
of Liphyra (see also Fig. 16.7 E, F). Pupae of
North American Feniseca species, and the Old-
World tropical genus Spalgis, are celebrated for
their curious skull-like appearance (Hinton,
1974).
Curetinae. This subfamily seems firmly based
on the following characters: Male with two inva-
ginateci tergal brushes on abdominal segment 2
(Chapman, 1915; Scott & Wright, 1990). Such a
condition is not known elsewhere in Lycaenidae.
Larvae with permanently exserted pair of cylindri-
cal tubercles on abdominal segment 11 (see
DeVries, 1984, and Fig. 16.7 C). This larval char-
acter appears to be unique.
Curetinae comprises a single, distinctive genus
(Curetis) with 18 species (Eliot, 1990), mostly
Oriental but a few just extend into the eastern
Palaearctic. The tuberculate larvae feed on Faba-
ceae, and may have facultative relationships with
ants (DeVries, 1984); pupae are weakly girdled.
Lycaeninae. Following Eliot (1990), the Poly-
ommatinae, Aphnaeinae, Theclinae and Lycaeni-
nae are all included here. In our parsimony
analyses for all butterfly groups (de Jong et al.,
1996), the six lycaenine genera studied never
clustered together as a single unit. Three charac-
ters offer some evidence for monophyly of this
subfamily: Hindwing 'tailed' on vein CuA2 (Eliot,
1973). Tails are essentially absent from other Ly-
caenidae. They are also absent from a number
of Lycaeninae, but such species are apparently
closely related to tailed species. According to
Robbins (1980; pers. comm.), in life the hindw-
ing tails of Lycaeninae are uniquely bent inwards
across the sagittal plane. Hindwings moved while
settled (Robbins, 1980). Characteristic "circular"
hindwing movements, which appear to enhance
the false-head-effect of the tails on CuA2, occur
throughout the group (including Lycaena\ pers.
obs.). The hindwing movements of certain Rio-
dininae (Robbins, 1986) and both old and new
world Charaxinae are considered to be non-ho-
mologous (R. K. Robbins, pers. comm.). Larva
with true dorsal nectary organ (DNO) (Cottrell,
1984). Considered by Fiedler (1991) to be a basic
synapomorphy for Lycaeninae, absence of the
DNO in Lycaenini and Theclini is interpreted as
independent secondary loss (Fiedler, 1991). A
285
group of polyommatine genera can be charac-
terized by a remarkable modification of the post-
abdomen, in which the ostium bursae occurs on
the end of a protrusible chitinous tube (Chap-
man, 1916).
This cosmopolitan subfamily (approximately
4000 species, in about 500 genera) includes spe-
cies endemic to New Zealand, Tahiti and Hawaii.
According to Eliot (1990), the group now in-
cludes all the familiar blues (Polyommatini: e. g.
Candalides, Anthene, Nacaduba, Theclinesthes,
Jamides, Lampides, Uranothauma, Leptotes, Ziz-
ina, Zizeeria, Zizula, Everes, Cupido, Celastrina,
Glaucopsyche, Maculinea, Lepidochrysops, Plebe-
jus, Polyommatus), Aphnaeini (e. g. Aphnaeus,
Spindasis, Phasis, Aloeides, Poecilmitis), coppers
(Lycaenini: e. g. Heliophorus, Lycaena), and
hairstreaks (Theclini: e. g. Lucia, Hypochrysops,
Ogyris, Jalmenus, Arhopala, Thecla, Chrysozeph-
yrus, Quercusia, Loxura, Iolaus, Hypolycaena,
Deudorix, Capys, Eumaeus, Satyrium, Cal-
lophrys, Strymon). The onisciform larvae feed on
a great variety of flowering plants. Many species
are aphytophagous for at least part of their lar-
val development, some feeding variously on Ho-
moptera; others have species-specific relation-
ships with ants (Thomas et al., 1989), feeding
either on ant brood (e. g. Maculinea arion) or ant
regurgitation (e. g. Maculinea alcon, M. rebeli),
while obligate or facultative symbiotic relation-
ships with ants are also widespread (Fiedler,
1991). The range of flowering plant families uti-
lised by the Lycaeninae as larval hosts is exten-
sive (Ehrlich & Raven, 1965; Vane-Wright,
1978), but most belong to subclass Rosidae
(Fiedler, 1991). A number of species are pests on
seeds, fruits or flowers. Lycaenine pupae are usu-
ally girdled.
Nymphalidae (browns, fritillaries, admirals and
monarchs). Classification of this very large
group has recently been reviewed by Ackery
(1988) and Harvey (1991). There is a single com-
plex character unique to all members of the fam-
ily: Ventromesial surface of antennae with three
longitudinal ridges (carinae), separating two con-
tinuous sulci or a pair of shallow depressions on
each segment (Fig. 16.91; Jordan, 1898). Carinae
occur in all species examined, including Libythei-
nae and Calinaginae, and have not been found
in any other Lepidoptera. Female abdomen with
von Siebold organ (Fig. 16.9 F; Huser, 1993 c).
Huser has demonstrated presence of this organ
within a range of Nymphalidae, including Liby-
thea, but did not find it in a large number of
other butterflies examined. This feature may also
prove to be universal for the group. Although
presence of antennal carinae is the only unique
and universal character for Nymphalidae en-
coded in our data matrix, in all our parsimony
analyses (de Jong et al., 1996) Nymphalidae
emerged as monophyletic.
286 Phillip R. Ackery, Rienk de Jong & Richard I. Vane-Wright

mm

Fig. 16.8. AC: ventral view of mesopleurosternum (left half). A, Papilionidae, Parnassius apollo. , Hesperiidae,
Euschemon rafflesia. C, Nymphalidae, Danaus plexippus. aep = anepisternum; bs = basisternum; ec = eucoxa II;
me = mesepimeron; mm = mesomeron; mp = marginopleural suture; pc = precoxal suture; pe = parepisternal
suture; sp = secondary sternopleural suture. D - F : dorsal view of basal sclerites of right forewing. D, Pieridae,
Pier is brassicae. E, Papilionidae, Papilio machaon. F, Hesperiidae, Hesperia comma, ax 1 - 3 = axillary 1 3; b l 2 =
base vein 1 A + 2 A; hm = hind margin forewing; mpl = median plate 1 ; mp2 = median plate 2; rp = radial plate.
The broken line indicates the right side of the thorax, slightly hanging over the basal structures.

According to Harvey (1991) nymphalid larvae
possess a filiform seta on A9 that is absent in
other papilionoid families, but it is not yet clear
how reliable this character is. All other suppos-
edly diagnostic features recorded in the literature
appear either non-unique or non-universal. They
include the following: female forelegs always re-
duced or modified (less so in Libythea, and re-
curs in some Lycaenidae), and usually clawless
(clawed in Libythea, Calinaga, some Satyrinae,
some Danainae); antero-lateral internal trichoid
sensilla on trochanter of male foreleg absent (a
comparable condition is found in Lycaenidae:
Riodininae); very distinct separation of upper
laterocervico-tentorial muscles into two bundles
(needs further investigation; not universal); pu-
pal head without cleavage line (recurs in Papilio-
nidae and Pieridae); pupa suspended by crem-
astral hooks only (or sometimes lying on the
ground), unattached (absence of a silken girdle is
universal in Nymphalidae, but this condition is
also found in species of Hesperiidae, Papilioni-
dae and Lycaenidae, and in the last family the
pupa may even be suspended by the cremaster
only).
With about 6000 species, Nymphalidae con-
tain approximately one-third of the entire butter-
fly fauna (Robbins, 1982). Currently divided into
about 350 genera (Harvey, 1991), ranging from
monobasic groups to the diverse Charaxes
(which includes more than 200 species). Nymph-
alids occur throughout the world, except for the
extreme Arctic or Antarctica. The family in-
cludes such widely studied groups as milkweeds
and glasswings (Danainae), passion-vine butter-
flies (Heliconiini), and checkerspots (Melitaeini).
Nymphalids range in size from small (e. g. Phyci-
odes) to very large (e. g. Caligo). Colour pattern
phenomena include sexual dimorphism (e. g. Ca-
tonephele, Zeuxidia), geographical variation
(e. g. Heliconius), seasonal variation (e. g. Ju-
nonia, Bicyclus), warning coloration (e. g.
Amauris, Acraea), polymorphic mimicry (e. g.
Hypolimnas, Pseudacraea), and crypsis (includ-
ing several genera of leaf butterflies, such as Kal-
lima, Doleschallia and Consul).
The usually ribbed and globular eggs are laid,
depending on species, in batches or singly, on or
off the foodplant. There are at least five general
forms of larvae: smooth with bifid tails (e. g. Sa-
The Butterflies: Hedyloidea, Hesperioidea and Papilionoidea
tyrinae), spinose (e. g. Nymphalinae), with large
dorsal spines (e. g. Limenitinae: Cyrestini), with
filaments (e. g. Danainae), and tufted (e. g. Mor-
phinae). Feeding may be diurnal, nocturnal, soli-
tary or gregarious. Hostplant relationships of
Nymphalidae are of outstanding interest (Ehr-
lich & Raven, 1965; Ackery, 1988). About 100
flowering plant families are utilised, including
Poaceae, Fabaceae, Acanthaceae, Passifloraceae,
Asteraceae, Scrophulariaceae and Euphor-
biaceae. In addition, particular species specialize
on cycads and club mosses. The pupae, some of
which have golden spots (responsible for the
term chrysalis), are almost always suspended by
the cremaster (Fig. 16.7 H) and never have a
girdle.
Subfamily divisions and relationships. Opinions
differ widely as to how the family should be sub-
divided and the many genera assigned. Here,
with the exception of Brassolinae (which we in-
clude within Morphinae) and Danainae (in
which we now include Ithomiinae and Tellervi-
nae), we follow Harvey (1991), and thus recog-
nize 10 subfamilies (Libytheinae, Heliconiinae,
Nymphalinae, Limenitinae, Charaxinae, Apatur-
inae, Morphinae, Satyrinae, Calinaginae, Danai-
nae). Although Harvey used some of these sub-
families in a more inclusive way than previous
authors (e. g. Ackery, 1988), most remain only
weakly supported by autapomorphies.
The adult forelegs of male Riodininae and the
forelegs of the vast majority of both male and
female Nymphalidae are useless for walking. In
consequence, the Nymphalidae were called Tet-
rpoda even before this name was applied to ver-
tebrates. In these butterflies the male forelegs are
not sensory, being almost devoid of trichoid sen-
silla, and are covered in hair-like scales and held
tucked-up against the body (so-called brush-
footed butterflies). The forelegs of female
Nymphalidae retain trichoid sensilla, and are
used for testing potential oviposition sites by
scraping or 'drumming'. A simple transforma-
tion series of the female foreleg may establish the
basal clades of the Nymphalidae. A sister group
relationship exists between Libytheinae (often
treated as a separate family; Ehrlich, 1958 ft) and
all other Nymphalidae, a relationship corrobo-
rated by Martin & Pashley (1992), using ribo-
somal RNA data. In Libytheinae, the female
forelegs are slightly modified, but still fully de-
veloped and can be used for walking. All other
Nymphalidae can then be grouped by the
following synapomorphy: Female foreleg modi-
fied, the tarsus often having less than five seg-
ments and usually lacking prelarsal claws; leg not
used for walking (Fig. 16.9H; Ehrlich, 1958 ). In
the monobasic Calinaginae the overall modifica-
tion of the female forelegs is less extreme, retain-
ing fully developed pretarsal claws. This might
suggest that Calinaga forms the sister group of
all Nymphalidae other than Libytheinae. How-
287
ever, as pointed out by Miller (1968), females of
some Danainae (Ithomiini), Satyrinae (Para-
lethe) and Morphinae (Antirrhea) have claws on
the foreleg, and Corbet & Pendlebury (1992)
note foreleg claws in female Amnosia (Nymph-
alinae) and Stibochiona (Nymphalidae, incertae
sedis - see Harvey, 1991). In our parsimony
analyses (de Jong et al., 1996), Libytheinae al-
ways appear in the expected position, but not so
Calinaginae.
Close relationship between Heliconiinae and
Nymphalinae may be suggested by the unique
presence of scoli on A9 of the larva. According
to Harvey (1991), however, the scoli are not ho-
mologous in the two subfamilies, occurring in a
different position. This is perhaps debatable and,
in our parsimony analyses four out of five exem-
plar genera of these two groups usually cluster
together. Based on characters of the female geni-
talia, Huser (1993 c) suggests possible relation-
ships between Heliconiinae, some Nymphalinae
(Melitaeini) and Danainae (including the ithomi-
ines).
As the larvae of many species of Morphinae
and Satyrinae feed on monocotyledons, close
relationship between the two groups has often
been supposed. However, most Morpho species
feed on legumes, and monocot-feeding has been
observed in various other nymphalids (e. g.
Charaxinae; Ackery, 1988). The mandibles of
first instar larvae lack teeth, a character usually
associated with eating grass (recurring, for ex-
ample, in grass-feeding Hesperiidae; Scott &
Wright, 1990).
Most relationships within and between the
various groups remain obscure. Even though
some of our parsimony analyses restricted to
nymphalid exemplars produced an apparently
well-resolved result, in general this was far from
typical, and continuing changes in higher classi-
fication must be anticipated, including alter-
ations to limits of many of the subfamilies them-
selves.
Libytheinae. This small subfamily can be rec-
ognized by at least four characters: Metatergum
overhung by mesotergum (Ehrlich, 1958 ft). The
whole metathorax is drawn under the mesoscu-
tellum, giving the dorsal side of the latter a cau-
dal position (unknown in other butterflies). Sec-
ondary pleural sulcus defining the mesothoracic
pre-epimeron forms a thickened band of cuticle
(Scott & Wright, 1990). Presence of the second-
ary pleural sulcus is regarded by Scott & Wright
as a "ground plan" feature of butterflies. We do
not concur with this view (see above, Papilio-
noidea). However, the form that it takes in Liby-
theinae is apparently unique. Sclerotized area of
patagia very small (Ehrlich, 1958 ft). Generally,
within Nymphalidae, the patagia are prominent
and uniformly well-sclerotized. Male genitalia
with bifid superuncus, and female genitalia with
a corresponding structure (Corbet & Pendlebury,
288 Phillip R. Ackery, Rienk de Jong & Richard I. Vane-Wright
1992). Although some other butterflies have a
male superuncus, including Calinaginae among
nymphalids, the combination of male and female
specializations in Libytheinae appears to be
unique.
The twelve included species (divided between
two genera Libythea, Libytheana) form an al-
most cosmopolitan group (Shields, 1985; but see
also Okano, 1987) which includes an endemic
species on the Marquesas, but lacks representa-
tion in New Zealand. Adult Libytheinae are re-
markable for their migratory behaviour (Shields,
1987). The great length of their labial palpi (the
origin of the popular names, beak or snout but-
terflies) is neither universal nor unique: the labial
palpi of male Libythea geoffroy are not particu-
larly long, while the longest are equalled or even
exceeded in a few other nymphalids (e. g., species
of Mycelia and Libythina cuvierii: Limenitinae).
Libytheine eggs are reticulate, barrel-shaped,
and rounded or truncate apically. The smooth
larvae, which feed on Ulmaceae, superficially re-
semble those of Pieridae in lacking head orna-
mentation or a bifid tail.
Heticoniinae. This subfamily now includes
Cethosia and Pardopsis, together with the
Acraeinae (Pierre, 1987) and Argynninae (Dos
Passos & Grey, 1945; Warren, 1944) of authors,
as well as the traditionally recognized Heliconii-
nae (Michener, 1942; Emsley, 1963). According
to Harvey (1991), this extension is supported by
three larval apomorphies: Larva with lateral scoli
arising between segments T1 and T2 and between
segments T2 and T3. When present in other
nymphalids, these scoli are located at the mid-
points of segments T2 and T3. Filiform setae on
segments A28 of larva arise from the sclerotized
base of the lateral scoli. Elsewhere in the nymph-
alids these filiform setae arise directly from the
body, even in those other nymphalids that have
lateral scoli. Scolus on segment A9 of larva dorsal
to the filiform seta. Such a scolus is also present in
Nymphalinae, where it lies posteroventrad of the
filiform seta. Harvey (1991) divides Heliconiinae
into three tribes: Acraeini (including Cethosia,
initially thought by Harvey to constitute a fourth
tribe, Cethosiini), Heliconiini and the monobasic
Pardopsini. Of these, the females of Acraeini
possess subpapillary glands (Pierre, 1987) at the
tip of the abdomen, while Heliconiini possess one
or more organs comparable to the clavatia de-
scribed by Parsons (1989). The homologies of
these various organs are uncertain, but poten-
tially they may provide further synapomorphies.
Now expanded, Heliconiinae embraces about
400 species in some 40 genera (including Helicon-
ius, Eueides, Dione, Acraea, Vindula, Cupha, Pha-
lanta, Argyreus, Argynnis, Speyeria, Issoria, Bolo-
ria and Yramea). Primarily tropical and north
temperate, the subfamily is unrepresented in
New Zealand. Eggs are reticulate and the spinose
larvae, in which the head may be ornamented or
bare, feed on many plants, notably Urticaceae,
Asteraceae and the related Violaceae, Flacourtia-
ceae and Passifloraceae. This group is of great
biological interest, notably for mimicry (Turner,
1975, 1984; Papageorgis, 1975; Sheppard et al.,
1985), chemical defence (Nahrstedt & Davis,
1981, 1983; Raubenheimer, 1989; Chai, 1986,
1990), coevolution (Benson et al., 1976; Gilbert,
1983), biogeography (Brown et al., 1974), and
mate-location behaviour (Bellinger, 1954; Crane,
1957; Magnus, 1958).
Nymphalinae. In the restricted sense employed
here, the subfamily now excludes Apaturinae, Li-
menitinae and Argynninae (the last treated as a
subtribe of Heliconiinae by Harvey, 1991). How-
ever, Nymphalinae may still be an unnatural as-
semblage. The position of the dorsolateral scoli
on A9 of the larva, being more ventral than the
condition in Heliconiinae, is unique; however,
the position of these scoli has already been used
to help characterize Heliconiinae, and it is un-
likely that both conditions can be apomorphous.
In practice, the group Nymphalinae can hardly
be diagnosed. Absence of a distinct mesothoracic
anepisternum distinguishes them from Apaturi-
nae and Charaxinae, absence of a hypandrium
distinguishes them from Biblidini (a major tribe
of the Limenitinae), absence of a basal spur on
the forewing cubitus distinguishes them from all
Limenitini except Neptis, and they also lack the
spiny eggs characteristic of many limenitines (in-
cluding Neptis). Their spiny larvae are reminis-
cent to those of Heliconiinae, but lack charac-
ters, including larval features, specified for that
group. The subfamily is subdivided into three
tribes, Nymphalini (= Vanessini), Kallimini and
Melitaeini. Nymphalini and Melitaeini uniquely
share presence of filiform setae on larval seg-
ments Ai and A2, while Kallimini and Melitaeini
share presence of a filiform seta on the sclero-
tized base of the scolus on larval segment A9
(Harvey, 1991). We are unable to resolve this
conflict.
This cosmopolitan group (some 350 species in
about 60 genera) includes species endemic to
New Zealand and Hawaii. Representatives of the
more dispersive genera (Hypolimnas, Cynthia,
Junonia) reach even remote oceanic islands.
Other well-known genera include Agais, Vanessa,
Nymphalis, Polygonia, Araschnia, Anartia, Kal-
lima, Doleschallia, Melitaea, Chlosyne, Phyciodes
and Eresia. Adults are mostly colourful, small to
medium-sized butterflies, much attracted to
flowers and, in some cases, fallen fruit and car-
rion. The group is well represented in temperate
regions, including a number of familiar species
that overwinter as adults. Certain groups have
been studied extensively with regard to behavi-
our and population biology (notably the Ameri-
can Euphydryas, Ehrlich, 1984), mimicry and
polymorphism (Hypolimnas; Vane-Wright et al.,
1977; Smith, 1976; Smith & Gordon, 1987), and
The Butterflies: Hedyloidea, Hesperioidea and Papilionoidea
seasonal variation, together with the develop-
ment of colour patterns (Junonia; Nijhout, 1991;
Junonia and Bicyclus; French, 1997). The reticu-
late eggs are roughly barrel-shaped and the ca-
terpillars spiny, covered with rows of scoli. The
diverse larval hosts notably include many Astera-
ceae, Urticaceae, Acanthaceae, Lamiaceae and
Scrophulariaceae. Gregarious behaviour is quite
common, and may involve the formation of
communal feeding webs.
Limenitinae. This subfamily has no recognized
autapomorphies and "is used here for conven-
ience only" (Harvey, 1991; see discussion above
under Nymphalinae). While eggs of many Li-
menitini are characteristically spined, appearing
like "minute sea urchins" (Corbet & Pendlebury,
1992), those of other subgroups (e. g. Cyrestini,
Biblini, Coloburini) are variable in form, with-
out spines.
With well over 1000 species, in about 100 gen-
era (including Historis, Biblis, Ariadne, Cato-
nephele, Asterope, Dynamine, Callicore, Limen-
itis, Cymothoe, Neptis, Parthenos, Euthalia, Eu-
phaedra, Cyrestis, Pseudergolis, Stibochiona), Li-
menitinae are virtually cosmopolitan, but the
group includes only four Madagascan species,
and lacks representation in New Zealand. Adults
are mostly of medium build, often with a sailing
flight, and include convincing examples of mim-
icry (e. g. species of Limenitis (Basilarchia), Adel-
pha, Pseudacraea). South American "cracker
butterflies" (Hamadryas) are remarkable for
their noisy aerial displays (Otero, 1990). Limeni-
tine larvae feed on a wide variety of flowering
plants, with members of one genus, Bebearia,
surprisingly on monocots (Marantaceae and
Arecaceae; Ackery, 1988). Major foodplant fami-
lies include Euphorbiaceae, Sapindaceae and Ca-
prifoliaceae. Larval form is varied: large median
dorsal spines only (Cyrestini - see Fig. 16.6 E),
some have pectinate lateral spines (Euthaliina
Fig. 16.7 A), while many Limenitini have incom-
plete but prominent dorso-lateral rows of spines
(Fig. 16.6 D).
Charaxinae. This subfamily is characterized by
two synapomorphies: Parapatagia partly or en-
tirely sclerotized (Ehrlich, 1958 b). They are
membranous in other butterflies. Forewing veins
Rs3 and Rs4 shortly stalked (Rydon, 1971). In
other nymphalids, according to Rydon, Rs3+4
is significantly longer. Rydon (1971) divided the
group into seven tribes (Pallini, Euxanthini,
Charaxini, Anaeini, Zaretini, Preponini and Pro-
thoini), of which the first three can be grouped
by a striking synapomorphy, the serrated fore-
wing costa (the so-called prionopterous genera).
Charaxinae includes about 400 species, in 28
genera, distributed throughout the tropics, with
just a few extending into more temperate regions
in North America, Europe, China and southern
Australia; absent from New Zealand. Well
known genera include Charaxes, Polyura, Eu-
289
xanthe, Palla, Prothoe, Agrias, Prepona, Zaretis,
Anaea and Memphis. Adults, which are often ro-
bust and fly with extraordinary rapidity (Mik-
kola, 1977), are strongly attracted to rotting
fruit, carrion and dung, but rarely (if ever) 'nec-
tar' at flowers. Many species have sumptuous
(e. g. Agrias) and often very complex (e. g. Char-
axes jasius) colour patterns (Nijhout, 1991).
Some mimic other butterflies (e. g. Consul) or
form co-mimetic complexes (e. g. Charaxes,
Swynnerton, 1926). Eggs are generally smooth
and globular. Larvae, which exhibit a range of
form (Rydon, 1971), usually have head ornamen-
tation and a bifid tail. A very wide range of plant
families (including many trees in Fabaceae, Lau-
raceae, Piperaceae and Sapindaceae) are used as
hosts (see Ackery, 1988).
Apaturinae. This subfamily can be recognized
by its distinctive male genitalia: Aedeagus long,
and saccus extremely long and thin, occupying
most of the length of the abdomen. A long aedea-
gus occurs in many Charaxinae, but the saccus
is relatively short. Apaturinae share a number of
other characteristics with Charaxinae, such as
robust build, feeding from dung, carrion and rot-
ting fruits, and a bifid larval tail. In the analysis
of DeVries et al. (1985) both Apatura and Chara-
xinae clustered with Satyrinae. The entirely
membranous parapatagia of Apaturinae distin-
guish them from Charaxinae.
Although absent from New Zealand and Aus-
tralia, this group (about 430 species, in 20 gen-
era; Le Moult, 1950) otherwise has a virtually
cosmopolitan distribution. Some species possess
beautiful iridescent wing patterns, including the
large and spectacular national butterfly of Ja-
pan, Sasakia charonda, and males of the familiar
European purple emperor, Apatura iris. Other
genera include remarkable mimics, such as spe-
cies of Hestina (Morishita, 1980), while Idrusia
nyctelius exhibits complex female-limited poly-
morphism (Nguyen, 1985). Other better-known
genera include Doxocopa, Euripus and Astero-
campa. Eggs are polygonal and dome-shaped.
The slug-like larvae have bifid tails, but are
sometimes spinose, and usually have an armed
head capsule. Hostplant records are dominated
by Ulmaceae, with occasional records for Betula-
ceae, Salicaceae and Fagaceae.
Morphinae. Ehrlich (1958 ) included three
well-known groups within this subfamily: Indo-
Australian amathusiids, and Neotropical bras-
solids and morphos. Miller (1968), however,
placed brassolids within Satyrinae. DeVries et al.
(1985) added Antirrhea and Caerois (from Satyri-
nae), and presented new evidence in favour of
Ehrlich's grouping. However, misled by an un-
fortunate misunderstanding in Ackery's (1988)
account regarding the characters involved, Har-
vey (1991) continued to separate Brassolinae and
Morphinae (including amathusiids) as distinct
subfamilies. Thus Harvey recognized two un-
290 Phillip R. Ackery, Rienk d e J o n g & R i c h a r d I. Vane-Wright
characterized groups, whereas Morphinae sensu
Ehrlich are currently supported by three larval
characters (DeVries et ai, 1985). However, these
are all in need of confirmation across a much
wider range of taxa: Post-occiput of larva with
well-sclerotized margins. In other larvae exam-
ined by DeVries et al. (1985) the edges of the
post-occiput were not strongly sclerotized. Ovoid
muscle scar mesad of lateral spine plate. Other
larvae examined lacked a muscle scar in this po-
sition. Wart-like pinacula restricted to the epi-
proct. In other nymphalid larvae the pinacula are
uniformly distributed on the body. In our parsi-
mony analyses (de Jong et al., 1996), Morpho,
Amathusia and Brassolis frequently cluster to-
gether, but only as a paraphyletic grouping.
Morphinae are conventionally separated from
the rest of Nymphalidae by a combination of
mostly homoplasious or plesiomorphic features,
including: antennae only weakly d a v a t e and gen-
erally unsealed, parapatagia membranous, an-
episternum well-developed, forewing discal cell
closed and without swollen veins, larvae (which
are often gregarious) finely pubescent, and usu-
ally with hair-tufts and bifid tails. Within the
subfamily, Morphini form a well-characterized
group on the basis of a number of features, in-
cluding hemispherical eggs, male abdominal
hairbrushes, and anal plates of the female genita-
lia (DeVries et al., 1985), while Brassolini (with
the possible exception of Brassolis and Penetes)
and Amathusiini, although probably natural
groups, currently lack convincing autapomor-
phies.
As recognized here, Morphinae includes ap-
proximately 230 species, in about 40 genera con-
fined to the Indo-Australian tropics and Neo-
tropics (being completely absent from the Afro-
tropical region). Most species are large, 'satyrid'-
like butterflies (at least on the underside), char-
acteristically attracted to fallen fruit. The group
includes many spectacular insects, such as bril-
liant blue Morpho species, huge owl-butterflies
{Caligo), heavy-bodied Brassolis, and ocellated
Taenaris. Many species are crepuscular (but Mor-
pho fly in broad daylight, while Morphopsis ap-
pears to be nocturnal), often possessing well-de-
veloped androconial organs. These attain great
complexity in genera such as Zeuxidia and Antir-
rhea (e. g. Vane-Wright, 1972). Eggs are mostly
globular, but include the unusual hemispherical
forms of Morphini. Larvae usually have bifid
tails, and can appear smooth (Fig. 16.6F), dis-
tinctly hairy or tufted (e. g. Morpho - Fig.
16.7 B). Most species feed exclusively on mono-
cots (including Arecaceae, Bromeliaceae, Heli-
coniaceae, Musaceae, Poaceae and Smilacaceae),
some being significant pests of bananas (e. g. Op-
siphanes, Caligo). However, the majority of Mor-
pho feed on legumes, with just one subgroup re-
stricted to bamboos.
Satyrinae. Traditionally, this group has been
defined by presence of swollen veins at the fore-
wing base (e. g. Ford, 1945). Although most sa-
tyrines do have one or more such veins, some
included genera do not (e. g. Melanitis, Lethe),
while the character recurs in other Nymphalidae
(e. g. Mestra, Ariadne, etc.). Other traditional
features include a closed hindwing discal cell
(open in Ragadiini), feeding on monocots, and
the fleshy, bifid larval tail. However, many
nymphalids have a closed hindwing cell, most
Morphinae and some Charaxinae feed on mono-
cots (while some Satyrinae never do!), and the
bifid larval tail recurs in Morphinae and Chara-
xinae together with Apaturinae and Calinaginae,
and could be plesiomorphic for the whole
Nymphalidae. Scott & Wright (1990) record sev-
eral other larval characters that could be apo-
morphic, but give no indication of how wide-
spread they are. According to Harvey (1991),
there is no single derived character that defines
Satyrinae as used here (equivalent to Satyridae
of Miller, 1968, less Antirrheini and Brassolini,
now both included within Morphinae). This is
reflected in our parsimony analyses (de Jong
et al., 1996), in which the two exemplar genera,
Satyrus and Haetera, never clustered. In practice
most Satyrinae can be recognized by a combina-
tion of inflated veins, a small anepisternum, and
closed hindwing discal cell.
With approximately 2400 species in about 280
genera, this subfamily is cosmopolitan in range,
with high diversity in both temperate and tropi-
cal regions, including endemic genera of New
Zealand, New Caledonia and Fiji. Extensive use
of grasses as larval hosts may explain the con-
siderable extratropical diversity of the group.
Larger or more familiar genera include Pierella,
Melanitis, Lethe, Neope, Pararge, Mycalesis, Bi-
cyclus, Henotesia, Erites, Ragada, Hypocysta,
Oreixenica, Ypthima, Neocoenyra, Callerebia,
Euptychia, Cissia, Coenonympha, Manila, Ere-
bia, Cassus, Calisto, Corades, Eretris, Pedaliodes,
Pronophila, Argyrophorus, Hipparchia, Oeneis,
Satyrus and Melanargia. The great majority of
Satyrinae are brown, of small to medium size,
and have prominent ocelli along the wing mar-
gins. However, striking exceptions include mi-
metic species (e. g. Elymnias and Zethera), occa-
sional brightly coloured forms (such as white or
blue Lymanopoda and Ptychandra species), or
clearwing groups (e. g. Haetera). Adults of some
groups (e. g. Mycalesini) find rotting fruit
strongly attractive, while others (e. g. Satyrini)
visit flowers. Satyrinae have played an important
role in comparative anatomy studies of butterfly
wing patterns (e. g. Schwanwitsch, 1924; Nij-
hout, 1991), and in ecological and developmental
genetics (e.g. Ford, 1975; Brakefield, 1984;
Brakefield et al., 1996). Eggs are generally globu-
lar, sometimes flattened apically. The smooth,
often green larvae normally have longitudinal
The Butterflies: Hedyloidea, Hesperioidea and Papilionoidea 291

Fig. 16.9. A D: wing venation. A, Hesperiidae (Trapezitinae), Trapezites sp. B, Papilionidae (Papilioninae),
Graphium sp. (A, after Nielsen & Common 1991). C, Nymphalidae (Satyrinae), Physcaeneura panda, forewing
(after Van Son 1955, Transv. Mus. Mem. 8). D, Nymphalidae (Nymphalinae), Catacroptera cloanthe, hindwing
(after Van Son 1979). E, Pieridae (Coliadinae), Eurema hecabe, pretarsal structure of foreleg (after Yata 1989, Bull.
Kitakyushu Mus. nat. Hist. 9). F, Nymphalidae (Nymphalinae), Melitaea athalia, female genitalia (BC = bursa
copulatrix, G R = gianduia receptaculi, G S = gianduia sebaceae, LG = lagena receptaculi, OV = ovary bases,
SD = von Siebold's organ) (after Huser 1993 c). G, Lycaenidae (Riodininae), Dicallaneura pulchra, male foreleg
(after Harvey 1987). H, Nymphalidae (Limenitinae), Crenidomimas concordia, female foreleg (after Van Son 1979).
I, Nymphalidae (Satyrinae), Lopinga achine, antenna! segment, ventral surface showing three carinae (redrawn
after Jordan 1898).

stripes and bifid tails. However, those feeding on
Selaginellaceae (e. g. the unrelated Ragada and
Euptychia species) bear foliose lateral processes.
Apart from these exceptions, other satyrines feed
on monocotyledons, mainly Poaceae and Cyper-
aceae but also members of Arecaceae (Elymnias,
and Haetera and its relatives). Pupae may either
be suspended or terrestrial.
Calinaginae. This small subfamily is supported
by two characters of the male genitalia: presence
of a superuncus, and absence of a gnathos (Ehr-
lich, 19586). Loss of the gnathos has also oc-
curred in some Lycaenidae (Eliot, 1973, as
brachia) and Hesperiidae. The superuncus is pre-
sumably not homologous with comparable struc-
tures found in Colias, Libytheinae or Papilioni-
nae. Two further unusual features seem diagnos-
tic: Thorax clothed in golden, orange or red hairs.
This peculiarity makes it possible to separate
Calinaga immediately from similar-looking Pier-
idae and other butterflies, but a similar condition
does occur in other Pieridae (e. g. certain Colias).
Antennae naked, with two circular shallow depres-
sions between the carinae on every flagellar seg-
ment. The antennae are very distinctive under the
light microscope, and even more so under scan-
ning electron microscopy.
Represented only by the Sino-Himalayan ge-
nus Calinaga (comprising about eight species;
Vane-Wright, unpublished; H. Gaonkar, pers.
comm.), these medium to large butterflies resem-
ble certain zygaenid moths (e. g. Agalope), and
to some extent Parantica species (Danainae) to-
gether with a variety of Pieridae. Eggs are globu-
lar and reticulate; the finely pubescent larva
bears prominent head horns and a weakly bifid
tail. The only known larval hosts are mulberry
(Moraceae).
292 Phillip R. Ackery, Rienk de Jong & Richard I. Vane-Wright
Danainae. As characterized here, Danainae
comprise over 450 species in more than 60 gen-
era, divided amongst 3 tribes (each is dealt with
separately). A close relationship between these
three groups (Danaini, Tellervini and Ithomiini),
widely treated as separate subfamilies (e. g. Ack-
ery & Vane-Wright, 1984; Harvey, 1991), has
long been recognized (e.g. Bates, 1862; Kirby,
1871) and can be supported by at least one syna-
pomorphy: Adults imbibe pyrrolizidine alkaloids
from damaged or withered plants, or from nectar,
and use them to make sex pheromones and/or for
defence (Ackery & Vane-Wright, 1984). Origi-
nally based on records for many genera of Da-
naini and Ithomiini, but available evidence for
Tellervini remains equivocal (Ackery, 1987). This
pharmacophagous relationship (Boppr, 1984) is
not known elsewhere in butterflies, but certain
moths (notably various Arctiidae) imbibe pyrrol-
izidine alkaloids for similar purposes (although
there are many differences in detail, strongly sug-
gestive that these are independent develop-
ments). Further possible synapomorphies for
Danainae sensu lato include clubbed female fore-
tarsi (but fairly similar structures do occur else-
where in nymphalids), and loss of the tergal mus-
cle in abdominal segment 2 (Ehrlich & Ehrlich,
1963) but this needs further examination. Pos-
sible larval synapomorphies include presence of
an extra primary seta, 1 a, located directly below
seta D l , in the first instar larva (Harvey, 1991).
This currently appears universal, but has only
been confirmed for a few species and has not
been examined in Tellervo. Another larval pecu-
liarity is possession of curious fleshy filaments
(see Fig. 16.6C), seen in all known danaine and
tellervine caterpillars, and in those of a few
"primitive" ithomiines. It is on the basis of these
findings that we propose to downgrade the three
groups to co-ordinate tribes within the single
subfamily Danainae.
Danaini. This tribe is characterized by two
autapomorphies: Males with eversible hairpencils
enclosed in elongate membranous sheaths opening
between abdominal segments 8 and 9. Hairpencils
of the form found in Danaini are not found in
any other butterflies (Ackery & Vane-Wright,
1984). Female with clubbed, ankylosed foretarsi.
Although not found in this precise form in other
butterflies, a broadly comparable condition is
approached in a few other groups, including the
Ithomiini and Tellervini (Ackery & Vane-Wright,
1984). In addition, the naked, unsealed antennae
(Jordan, 1898) are often considered to be unique
within Nymphalidae. However, other nymph-
alids (e. g. Calinaginae, Morphinae, some Heli-
coniinae, some Satyrinae) share this trait, and it
recurs in Papilionidae. Huser (1993 c) has re-
cently discovered an apparently autapomorphic
feature of the spermatheca.
As widely recognized, this group (162 species,
in 11 genera, of which Parantica, Amauris, Tiru-
mala, Danaus, Euploea and Idea are the most
species-rich) is pantropical, with greatest diver-
sity in the Indo-Australian region, while a few
migrant species extend into temperate zones. The
medium to large adults have generally apose-
matic patterns, although the fritillary-like color-
ation of some Anetia species led to their former
misclassification. Flight is generally slow. Adults
are strongly attracted to flowers (notably those
of plants which produce pyrrolizidine alkaloids,
such as Ageratum), other pyrrolizidine alkaloid
sources (e. g., withered heliotropes, etc.), salt,
and sometimes damp wood, rocks and a variety
of other substances. Danaini include outstanding
examples of chemical protection and communi-
cation, mimicry, migration, and sexual and social
behaviour (Ackery & Vane-Wright, 1984; papers
in Vane-Wright & Ackery, 1984; Vane-Wright &
Boppr, 1993; Schulz tal., 1993), including per-
haps the most widely studied of all butterflies,
the Monarch, Danaus plexippus (Ehrlich, 1958 ;
Brower, 1984; see Malcolm & Zalucki, 1993, and
references therein). Eggs are barrel-shaped,
pointed apically and the brightly coloured larvae
bear one or more pairs of flexible filaments. Pu-
pae of many species are remarkable for their bril-
liant silver or gold multiple thin-layer reflections
(Steinbrecht & Pulker, 1980). Hosts largely com-
prise plants with milky sap belonging to Asclepi-
adaceae, Apocynaceae and Moraceae.
Tellervini. This small group can safely be con-
sidered monophyletic, but it is not supported by
any convincing autapomorphies. Larva with
single pair of fleshy tubercles on metathorax (Ack-
ery, 1987). Although this is a unique feature of
Tellervini (none of the filamented larvae of other
Danainae has this particular "formula"), it needs
confirmation, larvae of only two species being
known.
Tellervini includes six species in a single genus
{Tellervo), restricted to the New Guinea subre-
gion (Ackery, 1987). The rather small, weak fly-
ing, largely black and white adults characteristi-
cally have bright yellow eyes (a feature surpris-
ingly shared by some of their mimics; Ackery,
1987). Early stages are very 'danaine', as are the
larval hosts (Parsonsia, Apocynaceae).
Ithomiini. Just one reliable autapomorphy
characterizes this group: Male hindwing with cos-
tal hairpencil. Although costal androconia occur
widely in Nymphalidae, the form in ithomiines is
diagnostic. In particular, such a structure never
occurs in other Danainae (while the abdominal
hairpencil of Danaini does not occur in Ithomi-
ini). The two major groups can also be separated
by the antennae (scaled in Ithomiini, unsealed
in Danaini).
Ithomiini, or glasswing butterflies (so-called
because many have diaphanous or even fully
transparent wings), include more than 300 spe-
cies, in about 50 genera (Lamas, in prep.), all
entirely restricted to the Neotropics. Major or
The Butterflies: Hedyloidea, Hesperioidea and Papilionoidea
better-known genera include Napeogenes, Hypo-
thyris, Oleria, Ithomia, Pteronymia and Hypole-
ria. The group is remarkable for the formation
of dense aggregations, so-called ithomiine
"pockets", which may be aerial leks, but is best
known for its involvement in large and complex
Mllerian mimicry rings (e. g. Papageorgis,
1975), first described in the classical work of
Bates (1862). Like other danaines, the chemical
communication, defence and behaviour of these
butterflies are of outstanding interest (e. g.
Brown et al., 1991). Eggs are barrel-shaped and
ribbed. Larvae occur in three principal forms:
the majority are smooth, but a few possess fila-
ments (e. g. Melinaea, Tithorea; Ackery & Vane-
Wright, 1984), while those of one group (Mecha-
nics) have been described as "cog-wheeled"
(Fox, 1967). Solanaceae are primarily exploited
as larval hosts (Drummond, 1986), but a few
groups (including most species with filamented
larvae) feed on Apocynaceae, with Hyposcada on
Gesneriaceae (Ackery, 1988; G. Beccaloni, pers.
comm.). The glasswings have been used exten-
sively in work on South American biogeography
(e. g. Brown, 1982), and are now considered to
be outstanding indicators for evaluating diversity
in Neotropical forests (Brown, 1991; Beccaloni &
Gaston, 1994).
HEDYLOIDEA
The superfamily Hedyloidea contains a single
family, Hedylidae.
Hedylidae. All hedylid species (about 40 in
number - see Scoble, 1990 a, b), are now placed
in a single genus 'Macrosoma', and confined to
Central and South America. These small, rather
dull lepidopterans, are predominantly nocturnal
although occasionally noted as day-flying. Their
resting posture is said to be characteristic, with
the thorax tilted so that the posterior edges of
the hindwings almost touch the substrate on
which the insect rests (Scoble, 1986). Curiously,
like nymphalids, the male forelegs are reduced,
and in both sexes only the pterothoracic legs are
used for perching (Scoble, 1992). Hedylid eggs
are remarkably pierid-like, being upright, spin-
dle-shaped and distinctly ribbed. The smooth,
slender caterpillar has both head ornamentation
and a bifid tail; the pupa is distinctly girdled
(Scoble & Aiello, 1990).
Although previously treated as geometrid
moths, Hedylidae are included here following
Scoble's suggestion that the Hedylidae either
form a trichotomy with the Hesperioidea and
Papilionoidea, or form the sister group of the
butterflies contentions which have aroused
dispute (see Weintraub & Miller, 1987; Scoble,
1988). Hedylidae indisputably possess several
butterfly-like 'traits'. In our parsimony analyses
(de Jong et al, 1996), when taking one moth ge-
nus at a time, Macrosoma most often groups
293
with Hesperioidea or, when taking all butterflies
and moths together, forms the sister group of
Papilionoidea. Debate will continue as to the
true relationships of Macrosoma. While mono-
phyly of Hedyloidea has never been formally de-
monstrated, probable synapomorphies include
resting posture, forewing veins Rsl and Rs2 sin-
uate, forewing shape, and fusion of tarsomeres
of male foreleg to form two elements (M. Scoble,
pers. comm.).
Acknowledgments
Bob Robbins (Washington) and Don Harvey
(Washington) made many helpful comments on
an early draft, notably on the lycaenid section.
Bob Robbins also contributed a detailed critique
of a later version, leading to many further im-
provements. Christoph Huser (Bonn) kindly
made various draft manuscripts available to us,
particularly concerning his original work on
adult anatomy, and discussed ideas on relation-
ships. Carla Penz commented on the draft sec-
tion on Heliconiinae. Andrew Atkins generously
shared information on hesperiid morphology.
R D J also wishes to acknowledge discussions
with David Hancock (Brisbane), Pim Schoorl
(Amsterdam) and Osamu Yata (Fukuoka) on
matters of anatomy and relationships, and col-
leagues at the Nationaal Natuurhistorisch Mu-
seum, particularly Kees van Achterberg, for vari-
ous advice, and discussion on phylogenetics.
P R A and RIVW are grateful to many colleagues
at The Natural History Museum, including
George Beccaloni, David Carter, David Good-
ger, Jeremy Holloway, Ian Kitching, Martin
Honey, Djunijanti Peggie (Bogor), Gaden Rob-
inson, Klaus Sattler, Malcolm Scoble, Michael
Shaffer, Campbell Smith and Kevin Tuck, for
comments on the manuscript and help with vari-
ous matters lepidopterological. Sarah Turner
made the early stage illustrations, and the Muse-
um's Photographic Studio made the original
photographs of adults. RIVW would also like to
thank the Wissenschaftskolleg zu Berlin for facil-
ities during preparation of the final manuscript.
All three of us are deeply indebted to Niels
Peder Kristensen for his patience, faith and guid-
ance.
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Addenda
Since the c o m p l e t i o n a n d s u b m i s s i o n of this
m a n u s c r i p t in 1994, a n u m b e r of f u r t h e r i m p o r -
tant works have come to our attention. Recent
monographs treat N e w - W o r l d Papilionidae
(Tyler et al., 1994) a n d A f r o t r o p i c a l C h a r a x i n a e
( H e n n i n g , 1989). F o r s t u d e n t s of N e o t r o p i c a l
butterflies, t h e b i b l i o g r a p h y b y L a m a s et al.
(1995) p r o v i d e s a n i n c o m p a r a b l e overview of the
available literature. Elsewhere, recent f a u n i s t i c
studies c o v e r C h i n a (Io, 1994), T h a i l a n d (Pinra-
t a n a & Eliot, 1996), t h e P a m i r s (Tshikolovets,
1997), T u r k e y ( H e s s e l b a r t h et al., 1995), N o r t h
A f r i c a (Tennent, 1996), as well as C o s t a R i c a
(DeVries, 1997) a n d Venezuela (Neild, 1996).
M a n y of these w o r k s also c o n t a i n a w e a l t h o f
life h i s t o r y d a t a (see also I g a r a s h i & F u k u d a ,
1997, a n d K o i w a y a , 1 9 8 9 - 9 6 ) .
Weller & Pashley (1995) h a v e a g a i n p r o v i d e d
s u p p o r t f o r the suggested close r e l a t i o n s h i p be-
tween butterflies a n d hedylid m o t h s . W i t h i n the
butterflies themselves, p a r t i c u l a r a t t e n t i o n h a s
f o c u s s e d o n t h e classification of t w o g r o u p s , the
t r o i d i n e P a p i l i o n i d a e ( P a r s o n s , 1996 a; b) a n d t h e
Heliconiini (Brower, 1997; B r o w e r & E g e n , 1997;
A u b e r t et al., 1996).
300
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Jol Minet & Malcolm J. Scoble
(J. Minet: Epicopeiidae, Drepanidae [part], Sematuridae, Uraniidae; M. J. Scoble: Drepanidae [part], Geometridae)
The composition adopted here for Drepanoidea
and Geometroidea follows that of Minet (1991):
Drepanoidea includes Epicopeiidae and Drepa-
nidae, and Geometroidea includes Sematuridae,
Uraniidae, and Geometridae. The Epicopeiidae
and Drepanidae share a few synapomorphies
(Minet 1991 and in press), such as the complete/
nearly complete, prespiracular, tergosternal scle-
rites at the base of the adult abdomen (also oc-
curring independently in several other Ditrysia),
or the modified mandibles of the larva, each of
them with a large, flat, lateral area, which is de-
limited ventrally by a well defined carina. For
Geometroidea (sensu Minet 1991), the most
reliable a u t a p o m o r p h y lies in the shape of the
larval labium where the spinneret is shorter than
the prementum along its midline (the spinneret
is secondarily fairly long in certain advanced
Geometridae). Drepanoidea and Geometroidea
are likely to represent sister groups based on a
few shared derived traits belonging to the ground
plan of their pupae: strong anterior extension of
the forelegs (as already noted for Geometridae
by Mosher, 1916); and presence of a transverse,
dorsal groove on AIO. Secondary disappearance
of one or b o t h of these apomorphies has oc-
curred on several occasions.
The clade Drepanoidea + Geometroidea has
few distinctive characters. The eggs may be flat
or upright, smooth or variously sculptured. The
larva usually bears a few secondary setae; in all
families, excepting Sematuridae, most abdominal
segments possess a secondary seta cephalad of
L3 (see also 1-15); certain prolegs may be vestig-
ial or entirely absent in Drepaninae (Drepanidae)
and Geometridae. The p u p a has long or rather
long antennae, and its abdomen is often punc-
tate. In the adult, the metathorax lacks tympanal
organs; the fenestrae laterales are at least
strongly reduced (see definition in 115), but the
fenestra media is often large; abdominal tympa-
nal organs occur in three families (Drepanidae,
Uraniidae, Geometridae). According to E h n b o m
(1948), the shape of the adult brain varies f r o m
primitive (Drepanidae) to advanced i. e. of the
"macrofrenate type" (Geometridae).
Key to the families: adults
1 Apterous or micropterous (females)
Geometridae (in part)
Fully winged 2 the forewing venation; (c) Rs4 separated f r o m
2 Abdominal sternum 2 connected to lateral
bars of tergum 1 by complete/nearly com-
plete tergosternal sclerites (which may be
part of tympanal organs); chaetosemata
often absent, never forming large areas on
vertex (Drepanoidea) 3
- Abdominal sternum 2 free, at most provided
with short anterolateral processes; chaeto-
semata always present, sometimes delimiting
fairly large areas (which may meet on the
meson) (Geometroidea) 4
3 Forewing (Fig. 17.1 A) lacking areole; Rs4
never stalked with R s l + Rs2 + Rs3 (al-
though it may be stalked with M l ) ; hind-
wing with Se + R and Rs always remote
f r o m each other beyond discal cell; abdomen
without tympanal organs . . . . Epicopeiidae
- Forewing (Figs. 17.1 B, C) with Rs4 stalked
at least with Rs3, and Rs3 + Rs4 often in-
volved in formation of an areole; hindwing
with Se + R and Rs usually approximated
or fused for a varying distance beyond discal
cell; base of abdomen with tympanal organs
Drepanidae
4 Rs4 (forewing) usually stalked with M l , but
never with the other branches of Rs (Figs.
17.1 E, F); abdomen with sexually dimor-
phic tympanal organs, those of female lying
on sternum 2, those of the male (laterally)
at junction of terga 2 and 3 (tympana on
tergum 2) Uraniidae
- Rs4 either involved in formation of an
areole (Fig. 17.6 A) or stalked at least with
Rs3 (Fig. 17.1 D); abdominal tympanal or-
gans either absent or situated on sternum 2
in b o t h sexes 5
5 Antennal "shaft" usually thickened near the
tapering apex; abdomen without tympanal
organs Sematuridae
- Antennal shaft without distal thickening; ab-
domen with a pair of tympanal organs on
sternum 2 Geometridae
DREPANOIDEA
Epicopeiidae. This small family is regarded as the
sister-group of the Drepanidae by Minet (1991).
It is characterized by several autapomorphies
(Minet in press): e. g. (a) c o m p o u n d eyes reduced
in size, their posterior margins never extending
distinctly - backwards - beyond the posterior
parts of the antennal foramina; (b) no areole in
R s l + Rs2 + Rs3, but primarily stalked with
M l (in the forewing). Study of the early stages
302
reveals further, potential autapomorphies, such
as: (d) larva with waxy secretions (Minet 1986;
Scoble 1992); (e) galeae of the pupa very broad
anteriorly, being partly adjacent to the eyes.
Medium-sized to large moths, with broad,
often brightly coloured wings (Figs. 17.2 A,
and R). Antennae either filiform (sometimes al-
most claviform), lamellate, serrate or bipectinate;
flagellum provided with sensilla trichodea
ventrally, laterally and, occasionally, also dor-
sally (these may be long or short in the male).
Ocelli absent. Compound eyes with long or short
interommatidial setae; the ocular margin does
not touch the antennal foramen, and tends to be
angulate dorsally. Chaetosema: usually a small
bundle of long setae. Pilifers present. Proboscis
well developed, with a broad base. Maxillary
palpi minute (2-segmented in Epicopeia). Labial
palpi curved, short to medium-sized; their last
two segments porrect or slightly upturned.
Epiphysis well developed; tibial spurs: 0-2-4. Ar-
olium present; claws simple (i. e. without a me-
dial tooth). Frenulum and subcostal retinaculum
usually present in males (though often absent in
Epicopeia), female frenulum absent or strongly
reduced. Spinarea absent. Forewing venation
(Fig. 17.1 A) lacking an areole; Rsl, Rs2 and Rs3
stalked together; Rs4 either free or stalked with
M l ; stem of M non-tubular, at least for most
of its length; M2 usually arising about midway
between Ml and M3 (rarely nearer M3 than
Ml); CuAl separated from M l ; tubular CuP ab-
sent; basal fork of 1 A + 2 A with very thin lower
branch. Hindwing: Sc diverging from upper edge
of discal cell (except near base where it briefly
touches cell, or is connected to it by an oblique
crossvein: R); veins Rs and Ml either free or
short-stalked; M2 rarely arising nearer M3 than
M l ; tubular CuP absent; 1A + 2 A without
basal fork; 3 A moderately long to very short.
Abdomen bicoloured; tympanal organs absent;
lateral bars of dorsum 1 connected to sternum
2 (anteriorly) by complete, or almost complete,
tergosternal sclerites; venulae absent; sternum 2
with short to minute apodemes and sometimes,
in males, posterolateral arms that bear - distally
long pencils of hair-scales (which lie in pleural
folds; e. g. Inoue 1992). Male genitalia: tegumen
usually delimited by distinct sutures, ventrally
and posteriorly (e. g. Kuznetzov & Stekolnikov
1988, 1993); socii sometimes present (small, pad-
like); gnathos when present strongly bent,
often with proximal half fused to tegumen. Fe-
male genitalia: ostium bursae lying just caudad
of sternum A8; this sternum developed into
strong projection beneath ostium (Okano 1973);
apophyses present (2 pairs).
Eggs of the upright type (Alin 1942, Issiki
et al. 1969, Yen et al. 1995), hemispherical in
Psychostrophia, practically spherical with radiat-
ing vertical ribs in Epicopeia. Mature larva with
white waxy secretions, often entirely covering its
Jol Minet & Malcolm J. Scoble
skin (Janet & Wytsman 1903, Issiki et al. 1969,
Sugi etal. 1987, Yen et al. 1995); mandible with
a large, flat, lateral area, the ventral edge of
which is more or less projecting; TI, T2 and T3
with more than two setae in each group SV; ab-
domen with several secondary setae, and with LI
and L2 far apart; five pairs of prolegs, all well
developed: crochets in curved, bi- to multiordi-
nal mesoseries). Pupa (often waxy): forefemora
and major part of the labial palpi concealed;
maxillary palpi occasionally exposed (as small
triangles in Psychostrophia); proboscis cases ex-
tending, laterad, beyond the ocular sutures; ab-
domen either punctate (Psychostrophia) or dor-
sally granulate (Epicopeia); A5 and A6 movea-
ble; cremaster typically with eight hooklets (api-
cal pair stronger). Cocoon waxy, either spun on/
between leaves (Psychostrophia, Schistomitra), or
subterranean or hidden in leaf litter (Epicopeia:
Strelkov 1932, Yen etal. 1995).
The colourful adults are day-flying and often
mimic various moths or butterflies, such as diur-
nal Geometridae and Arctiidae, the pierid genus
Aporia, or the papilionid genera Pachliopta and
Parides. At rest, wings are outspread and closely
appressed to the substrate. The known larvae
feed externally on plants of the following fami-
lies: Clethraceae, Cornaceae, Ericaceae, Thea-
ceae and Ulmaceae (Sugi et al. 1987).
The family is not divided into subfamilies (Mi-
net in press). Until recently, it was regarded as
monotypic (e. g. Laithwaite etal. 1975). Fletcher
(1979) placed two genera in the Epicopeiidae
(Nossa; Epicopeia), maintaining in the "Epiple-
midae" Psychostrophia, Schistomitra, Amana,
etc. However, Nakamura (1972) had already sug-
gested epicopeiid affinities for Schistomitra fu-
neralis, mainly on the basis of its pupal morpho-
logy. Three papers give a more complete outline
of the Epicopeiidae (Minet 1983, 1986 and in
press). The family is known only from palaearc-
tic and tropical Asia. It contains about 25 spe-
cies, placed in nine genera, for example Psycho-
strophia (Fig. 17.2 A), Parabraxas (Fig. 17.2 B),
Nossa, and Epicopeia (Fig. 17.2R). In the last
genus, the species have a large wingspan and
strongly caudate hindwings, and seem to be
Mllerian mimics of certain Papilionidae; their
distribution ranges from northern India and Su-
matra to northeastern China, Korea and Japan
(see e. g. Inoue 1978).
Drepanidae. Long regarded as separate families
("Thyatiridae", "Cyclidiidae" and "Drepani-
dae"), the three main subdivisions of the Drepa-
nidae share a distinctive synapomorphy in the
adult, viz. the possession of abdominal tympanal
organs associated with the tergosternal sclerites,
which connect tergum 1 with sternum 2. The
characteristic structure of these organs (they
have internal tympana) is invariable throughout
the family, but is found nowhere else in the Lepi-
The Drepanoid/Geometroid Assemblage
Fig. 17.1 Wing venation of Epicopeiidae (A), Drepanidae (B, C), Sematuridae (D) and Uraniidae (E, F): A,
Parabraxas davidi; B, Ochropacha duplaris; C, Watsonalla binaria, D, Homidiana japet; E, Erosia incendiata 9; F,
Urapterilra piperita. Scales = 5 mm. sp: spinarea. (G. Hodebert).
doptera; see also 211. In the larva, the presence
of a secondary seta on A l - A 8 , which is situated
dorso-caudad of the spiracle, is likely to repre-
sent a further autapomorphy of the Drepanidae
(Minet 1991: 87).
Small to fairly large moths, usually medium-
sized, with broad or moderately broad wings
(Figs. 17.2 C - J ) ; forewings often angulose or
falcate (hence the name "Hook-tip moths" given
to many Drepaninae); body either stout or slen-
der. Antennae often lamellate or bipectinate (to
the tip or for most of their length), sometimes
filiform, serrate or unipectinate; bipectinate con-
dition not rare in females; flagellum usually
scaled dorsally; rami often scaled externally.
Compound eyes naked or with long interomma-
tidial setae. Ocelli distinctly present only in a few
genera (Chaeopsestis, Spica, some Hypsidia).
Chaetosemata absent or inconspicuous. Pilifers
and proboscis variously developed (sometimes
absent); the latter always unsealed. Maxillary
palpi vestigial or minute, at most 3-segmented.
Labial palpi usually 3-segmented, weak to very
small in Drepaninae, robust or rather well devel-
303
oped in other members of the family; these palps
upturned, porrect, or even drooping (sometimes
only segment 3 is porrect); segment 3 smooth-
scaled, either tubular or button-like. Mesotho-
rax: ventral process of tegula never ending in a
sharp point; anepisternum often large; meron
frequently with a posterodorsal extension (espe-
cially well developed in Thyatirinae). Metatho-
rax: scutum divided or almost divided mesally;
tergal rim (sensu Brock 1971) distinctly enlarged
in most Thyatirinae. Epiphysis rarely absent;
tibial spurs: 0-2-4, 0-2-2, 0-0-4 or 0-0-0. Pretar-
sus: arolium and pulvilli normally well devel-
oped; claws frequently with a medial tooth. Male
wing-coupling: usually a subcostal retinaculum
and a frenulum, which may have an apical club
(also observable on the frenular bristles of cer-
tain females: Scoble & Edwards 1988); frenulum
and retinaculum absent from both sexes in Ore-
tini. Spinarea often absent, but preserved dis-
tinctly in Thyatirinae (Fig. 17.1 B, sp.). Forewing
frequently with long narrow areole (running par-
allel to costa: Figs. 17.1 B, C); Rs with 3 or 4
branches (when areole is lacking, Rsl is stalked
304
with Rs2 or occasionally with R and Rs3
with Rs4, or all Rs veins may share a common
stem); M2 arising either about midway between
Ml and M3 (most Cyclidiinae and Thyatirinae)
or nearer M3 than Ml (e. g. Drepaninae); tubu-
lar CuP absent; 1A + 2 A usually forked.
Hindwing: Se + R approximated to Rs (or partly
fused with it) before or beyond point at which
Rs separates from discal cell (this point usually
remote from origin of M l , although approxi-
mated to it in Hypsidia and Spica); M2 arising
nearer M3 than Ml or, less frequently, about
midway between Ml and M3; CuP absent or re-
placed by a fold; 3 A varying from well devel-
oped to reduced or entirely absent. Abdomen:
pleura of A2 sometimes with long androconial
pencils, transversely held under sternum 2 when
moth is at rest (Cyclidiinae; a few Oretini: see
e. g. Fig. 4 in Watson 1965 b); tergosternal scle-
rites (connecting tergum 1 with sternum 2) modi-
fied into tympanal organs with a unique mor-
phology: rounded ventral part of each sclerite
enclosing two tympanal chambers, partly sepa-
rated by an internal transverse tympanum;
arched dorsal part enclosing pleural chamber
posteriorly delimited by a large counter-tympa-
num (e. g. Gohrbandt 1937; Minet 1983; Scoble
1992; 2 - 1 1 ) . In male, A8 (or even, occasionally,
sternum 7), modified to varying degree; saccus
sometimes present; vinculum and tegumen not
always fused; gnathos complete in a few taxa
(e. g. Cyclidiinae); uncus either simple, bifid, or
absent; socii often sclerotized and elongated
("subunci" of, e. g., Werny 1966); phallus some-
times with a coecum (e. g. in Thyatirinae). Fe-
male genitalia: signum/signa, when present, usu-
ally small and rounded or roughly linear; ostium
bursae on venter 8 or, occasionally (Paralbara),
just caudad of venter 7; apophyses anteriores
sometimes rather long, apophyses posteriores
never very long; segment 9 + 1 0 often with a
complex structure: e. g. in Drepaninae (papillae
anales tending to develop a ventral pair of dif-
ferentiated lobes).
Egg of the "flat" type, roughly oval or ellipti-
cal; chorion either smooth or variously sculp-
tured, often with parallel, longitudinal ribs (eggs
usually laid in small groups, occasionally singly
or in short rows). Last instar larva: head globu-
lar or dorsally bifid; segments T2 to A8 with at
least two pairs of secondary setae; numerous
secondary setae occasionally present (e. g. in Cy-
clidiinae where they are very short); Verrucae
normally absent (present in Mimopsestis); Dre-
panini with a prothoracic, eversible vesicle be-
tween setae SV and the coxa (Minet 1985); tho-
rax and abdomen sometimes with dorsal protu-
berances or elongated processes (e. g. in Tridre-
pana); Drepanini often with a few modified setae
(see e. g. Nakajima 1970 and Minet 1985); anal
prolegs vestigial without crochets in Drepa-
ninae; ventral prolegs well developed on A 3 - A 6 ;
Jol Minet & Malcolm J. Scoble
proleg crochets in uniordinal or, more often,
biordinal mesoseries (in addition, there are often
uniordinal lateroseries of small crochets); most
Drepaninae with anal shield conspicuously elon-
gate caudad. Pupa: sometimes covered with
waxy secretion; labial palpi and forefemora con-
cealed or very slightly exposed; maxillary palpi
concealed; base of proboscis usually broad
(though it does not reach the eyes); forelegs tend-
ing to project cephalad as in Geometroidea; an-
tennae often extending, posteriorly, as far as
midlegs, or even further; Al dorsum with blunt,
obtuse, anterior angles; spiracles reduced on
both A7 and A8 in many Thyatirinae (Naka-
mura 1980); an irregular furrow sometimes pre-
sent, dorsally, between A9 and AIO. Cocoon
usually spun between leaves, occasionally almost
absent or subterranean.
Adult Drepanidae are usually nocturnal, but a
few species fly readily during the day (e. g. Ni-
dara). Their resting posture varies from tecti-
form, where the wings are positioned roof-like
over the body (e. g., as in Thyatirinae), to plani-
form, where the wings are outspread (in Cyclidii-
nae and most Drepaninae). However, peculiar
postures have been noted for some species (see
Seitz 1922 and Kaishoven et al. 1951). Posture is
generally related to the presence or absence of
the spinarea, which varies within the family.
Most larvae are arboreal feeders. They may be
gregarious when young. By day, they live either
exposed or concealed in rolled leaves or between
leaves tied together.
As conceived by Minet (1983) and Scoble &
Edwards (1988), the family contains approxi-
mately 120 genera and 650 described species. It
consists of three subfamilies Thyatirinae, Cy-
clidiinae and Drepaninae and at least one ge-
nus (Hypsidia) with undetermined subfamily as-
signment. In the adult, these taxa can be distin-
guished as follows:
Key to the subfamilies
1 Forewing with M2 arising about midway be-
tween Ml and M3 (Fig. 17.1 B) 2
Forewing with M2 arising distinctly closer
to M3 than to Ml (Fig. 17.1 C) 3
2. Tarsi conspicuously spinose ventrally . . . .
Thyatirinae (in part)
Tarsi without obvious spines
Cyclidiinae (in part)
3 Forewing with a well developed spur arising
distally from the "anal loop"
Cyclidiinae (in part)
Forewing without such a spur 4
4 Head without ocelli; labial palpi short or, if
moderately long (i. e. length slightly exceed-
ing diameter of the eye), then with segment
2 rather thin (i. e. not distinctly thicker than
segment 3) Drepaninae
The Drepanoid/Geometroid Assemblage
Ocelli present or, if absent, labial palpi long,
with segment 2 distinctly thicker than the
smooth-scaled third segment
. . . genus Hypsidia and a few Thyatirinae
Recently assigned to the Drepanidae (Scoble &
Edwards 1988), Hypsidia, with 6 species, is con-
fined to Australia. Its distribution is however dis-
junct, with some species occurring in sclerophyll
forests of southern Western Australia and others
in the rainforests of northeastern Queensland
(Fig. 17.2 E). The early stages are unknown.
Thyatirinae are nocturnal, noctuid-like moths.
The larvae have crochet-bearing anal prolegs and
usually feed at night. K n o w n hostplants belong
to the following families: Betulaceae, Caprifolia-
ceae, Cornaceae, Ebenaceae, Fagaceae, Juglan-
daceae, Rosaceae and Salicaceae (e. g. Forbes
1923, Watson 1965 a, Skinner 1984 and Sugi
et al. 1987). Thyatirinae occur mostly in the ho-
larctic and oriental regions, but they are also
represented in Africa, New Guinea and in the
neotropical region. They contain about 50 genera
and 200 species (Munroe 1982). Thyatirine tribes
are still unsatisfactorily defined, but the
following names are available: Aethiopsestini,
Ceranemotini, Demopsestini, Euparyphasmini
( = Lithocharini auct.), Gaurenini, Habrosynini,
Macrothyatirini, Plusiniini, Polydactylini, Poly-
plocini, Psidopalini, Tetheini ( = Bombyciini,
= Palimpsestini) and Thyatirini (see notably
Houlbert 1921, Forbes 1936 and 1940, Werny
1966, and Yoshimoto 1984). True Thyatirinae in-
clude, e. g., Thy atira, Gaurena, Habrosyne (Fig.
17.2 C), Tethea (Fig. 17.2 D), Ochropacha, Para-
psestis, Epipsestis and Polyploca. However, for a
few taxa (e. g. Spica), placement in Thyatirinae
remains questionable.
Cyclidiinae include fewer than 15 species, as-
signed to two or three genera (Nelcynda being
either a valid genus, or a synonym of Cyclidia).
The subfamily is represented in palaearctic Asia
and the oriental region. Medium-sized (Mimo-
zethes: Fig. 17.2 F) to rather large (Cyclidia: Fig.
17.2 G), m o t h s are geometrid-like, with broad
wings and slender bodies. The larvae have nu-
merous, minute, secondary setae and normally
developed anal prolegs. In Mimozethes argentili-
nearia and Cyclidia substigmaria the larvae feed
on Alangium platanifolium (Alangiaceae), which
is the only recorded cyclidiine hostplant (Sugi
et al. 1987).
Drepaninae are geometrid-like, usually noctur-
nal moths; their forewings are often falcate (see
e. g. Yazaki 1992: PI. 1). The subfamily is absent
f r o m the neotropical region, and f r o m New
Zealand, and is particularly well represented in
palaearctic Asia and in the oriental region. Dre-
paninae include nearly 450 described species re-
ferred to about 70 genera (see notably Watson
1957 a, b, 1959, 1961, 1965 b, 1967, 1968; and
Wilkinson 1967 a, b, 1968, 1972). Drepanine lar-
305
vae have reduced anal prolegs, which lack
crochets, and usually an anal shield ending in a
pointed, posterior process (e. g. N a k a j i m a 1970,
1972; Stehr et al. 1987; and Sugi et al. 1987).
Hostplants have been recorded f r o m Anacardia-
ceae, Arecaceae, Betulaceae, Caprifoliaceae,
Cornaceae, Daphniphyllaceae, Fagaceae, Ju-
glandaceae, Musaceae, Myrtaceae, Rosaceae,
Rubiaceae, Salicaceae, Sapindaceae, Theaceae,
Tiliaceae and Zingiberaceae (see, in particular,
Forbes 1923, L h o m m e 1935, Sevastopulo 1940,
Kaishoven et al. 1951, Laithwaite et al. 1975,
Skou 1986 and Sugi et al. 1987). Three tribes are
recognized. The monotypic Nidarini, f r o m Ma-
dagascar, include five described species and
might be no more than specialized Drepanini.
They have non-falcate forewings (mainly yellow
or orange), and their tibiae lack spurs. Mostly
tropical, the Oretini (Fig. 17.2 H) include 15 gen-
era. Amplexiform wing-coupling occurs in both
sexes. In Ore ta and especially Epicampoptera,
certain species are well known pests of coffee
(Coffea). Drepanini, with about 50 genera and
more than 300 described species, are the largest
tribe. This g r o u p is defined by a pair of lateral,
eversible vesicles, which occur on the larval pro-
thorax near setae SV (Minet 1985). A m o n g the
main genera are Drepana (Fig. 17.21), Tridre-
pana, Nordstromia, Microblepsis, Cilix, Agnidra,
Callidrepana, Macrocilix (Fig. 17.2 J), Teldenia,
Ditrigona, Drapetodes, Phalacra and Ametro-
ptila.
GEOMETROIDEA
Sematuridae. Unlike other members of the Geo-
metroidea (Uraniidae and Geometridae), the
Sematuridae invariably lack abdominal tympa-
nal organs. In the adult, sematurid a u t a p o m o r -
phies include: (a) antenna distally thickened (not
distinctly so in a few advanced species), with
swollen scape and elongate first flagellomere; (b)
median, dorsal sclerite of A l distinctly narrowed
on the meson; (c) lateral bars of A l d o r s u m very
broad and adjacent to strong anterolateral exten-
sions of tergum 2; (d) vinculum with a spinulose
or finely speckled area ventrad of the juxta.
Medium-sized to large broad-winged moths,
often with tails to the hindwings (Figs. 17.2 K,
L). Antennae usually more or less clubbed, with
a tapering apex; scape bulky; pedicel short; first
flagellomere long and narrow; flagellum scaled
dorsally; antennae sometimes bipectinate in fe-
males (Coronidia). Ocelli often present. Chaeto-
semata well developed, their long setae over-
hanging the eyes (that usually bear conspicuous
interommatidial setae). Pilifers fairly strong. Pro-
boscis long, unsealed. Maxillary palpi minute.
Labial palpi well developed, upturned or nearly
porrect; segment 1 with an internal patch of
microtrichia (at least in Ilomidiana): segment 3
tubular, smooth-scaled (tending to be shorter in
306 Jol Minet & Malcolm J. Scoble

Fig. 17.2. Adults of Epicopeiidae (A, B, R), Drepanidae ( C - J ) , Sematuridae (K, L) and Uraniidae ( M - Q ) :
A, Psychostrophia nymphidiaria; B, Parabraxas davidi; C, Habrosyne pyritoides; D, Tethea ocularis; E, Hypsidia
erythropsalis; F, Mimozethes argentilinearia; G, Cyclidia substigmaria; H, Gonoretodes timea; I, Drepana curvatala;
J, Macrocilix mysticata; K, Homidiana japet <J; L, Sematura lunus; M, Auzea rufifrontata; N, Erosia incendiata <?;
O, Dissoprumna erycinaria: P, Urapteritra piperita; Q, Urania boisduvalii; R, Epicopeia mencia. Scales A J,
M - P = 5 mm; K, L, Q, R = 10 mm. (J. Minet).
The Drepanoid/Geometroid Assemblage
the male). Tibiae smooth-scaled, with or without
spines; epiphysis present; spurs: 0-2-4. Tarsi
spined ventrally. Arolium and pulvilli well devel-
oped; claws simple. Frenulum long to very short
(sometimes distally clubbed in male); male sub-
costal retinaculum sometimes absent (Sematura).
Spinarea absent. Forewing venation (Fig.
17.1 D): R free; R s l - R s 4 on a common stem;
M2 arising closer to Ml than to M3, or about
midway between Ml and M3; a transverse spur
or fold usually visible, near the base, between
CuA and the fold replacing CuP. Hindwing:
basal cell small, delimited by upper edge of discal
cell, swollen base of Sc, and a short crossvein
(R), which is often reduced to a point; M2 never
arising closer to M3 than to Ml; M3 and CuAl
sometimes stalked (CuA2 may be stalked with
M3 + CuAl); no tubular CuR Abdomen: tym-
panal organs absent; lateral bars of dorsum 1 en-
larged, adjacent to broad anterolateral exten-
sions of tergum 2 (which are very long and reach
a pair of androconial brushes in the male of Apo-
progones hesperistis); sternum 2 with the apo-
demes and anterolateral processes rather short;
venulae present in Apoprogones. Male genitalia
normally with a spinulate area on the vinculum
near juxta; valvae often with several sensilla
chaetica arising from tubular projections;
gnathos present or absent; phallus often with a
coecum. Female genitalia sometimes with a well
defined appendix bursae; ostium bursae on ven-
ter 8 (anteriorly): apophyses (both pairs) of
moderate length.
Egg of the upright type, fairly high, with many
radiating vertical ribs and finer transverse rib-
bing. Larva and pupa known only in Sematuri-
nae (e. g. Westwood 1879, Fassl 1910). Last in-
star larva: head hypognathous, apparently with-
out secondary setae; most "trunk" segments with
the base of D1 (sometimes also of D2) trans-
formed into a conical projection; T1 with a
secondary seta near XD2, but no other second-
ary setae in thoracic/abdominal segments (at
least in Homidiana); LI and L2 remote from
each other on A1-A8; five pairs of prolegs (all
well developed); proleg crochets roughly triordi-
nal, in mesoseries. Pupa: proboscis long, ex-
tended anteriorly, prolonged caudad beyond
forewing apices, and ending as a free, straight
process; maxillary palpi, major part of the labial
palpi, and forefemora, concealed; antennae long;
epicranial suture present; hindwing margin ex-
posed along forewing termen; A5 and A6 movea-
ble; AIO sometimes deflexed; cremaster com-
posed of eight hooklets in Homidiana. Pupation
occurs in a loose cocoon, probably in leaf litter.
The adults are mostly nocturnal, but may also
be diurnal in some genera. They are robust fly-
ers. At rest, the wings are widely outspread and
the antennae are directed antero-laterad. Larval
hostplants remain almost completely unknown.
307
Sematuridae contain approximately six genera
and 40 species. Often regarded as separate fami-
lies, Apoprogoninae and Sematurinae are here
placed in a single family (see Minet 1986: 307).
These subfamilies can be easily recognized:
Key to the subfamilies
1 Ocelli absent; compound eyes without long
interommatidial setae; forewing with vein
Ml free Apoprogoninae
Ocelli present; eyes distinctly hairy; forewing
with Ml and Rs sharing short stalk (Fig.
17.1 D) Sematurinae
The only member of the Apoprogoninae is Apo-
progones, a monotypic genus from South Africa
(Janse 1932).
Sematurinae are neotropical, except for one
species living in Arizona (Gaede 1930). Sematura
includes the largest and best known species (Fig.
17.2 L). In Coronidia, the hindwing has an irides-
cent blue band, whereas this wing is usually
marked with pink, orange or yellow in Homidi-
ana (although not in some males: Fig. 17.2 K).
Uraniidae. The present concept of the family is
based mainly on several autapomorphies of the
tympanal organs (Minet 1983; 2-11). These
structures show a pronounced sexual dimor-
phism: in males, they are invariably found on the
posterolateral parts of abdominal tergum 2 and
include anterolateral elements of tergum 3; in fe-
males, tympanal organs occur on the antero-
lateral parts of sternum 2, and their skeletal com-
ponents are not homologous with those of males.
Another derived trait of Uraniidae is that, in the
forewing, the base of Rs4 is connate or stalked
with M l , but separate from the other branches
of Rs (this apomorphy also occurs, through
parallel evolution, in many Epicopeiidae).
Small to very large broad-winged moths, usu-
ally having rather slender bodies (Figs. 17.2
M - Q ) . Hindwing termen often angulate or
tailed at M3, sometimes variously notched or
with several tails. Wings of day-flying species fre-
quently with brilliant, iridescent colours. Anten-
nae often lamellate or serrate, sometimes filiform
or unipectinate, rarely bipectinate (e. g. Aorista);
shaft scaled dorsally, with its subapical section
occasionally thickened (usually thickened in Al-
cides and Urania where scape is short and in-
flated). Compound eyes rarely with long inter-
ommatidial setae (Chrysiridia rhipheus). Ocelli
absent or rather small. Chaetosemata well devel-
oped, usually mixed with minute scales (chaeto-
semata may meet mesally). Frontoclypeus some-
times very narrow. Pilifer bristles well developed.
Proboscis present, unsealed; base often with long
sensilla trichodea. Maxillary palpi small, having
usually one segment (there may be a second, ill-
defined segment). Labial palpi 3-segmented, long
308
to short, porrect or ascending (sometimes: seg-
ment 2 ascending and segment 3 porrect). Meso-
thorax: ventral process of tegula elongate, often
with apex spatulate; anepisternum sometimes re-
duced or absent (Uraniinae). Metascutum di-
vided or strongly constricted mesally. Fenestra
media (sensu Minet 1982) narrow to fairly wide.
In female Uraniinae, posterior part of metepime-
ron modified into a concave, oval sclerite (which
is usually "replaced" by erect scales in other Ura-
niidae). Forelegs with stridulatory scales in male
Urania (Lees 1992). Tibiae and tarsi smooth-
scaled, except when tibiae possess androconial
hair-scales (some Microniinae); epiphysis pre-
sent; tibial spurs: 0-2-4, occasionally 0-2-3 or 0-
2-2 (males of some Microniinae). Pretarsus:
claws simple (no medial tooth); arolium present;
pulvilli bifid or with long dorsal outgrowths.
Wing-coupling either retinaculo-frenate or am-
plexiform (occasionally, male with frenulum, but
lacking subcostal retinaculum: e. g. in Dicro-
plema). Spinarea absent. Wing scales never
deeply cleft; several Epipleminae with andro-
conia in modified areas of hindwings or among
fringe scales. Forewing venation (Figs. 17.1 E,
F): Rsl often free; Rs2 and Rs3 stalked; Rsl and
Rs2 + Rs3 occasionally arising from a small,
narrow areole (other "accessory cells" may oc-
cur, notably because R may anastomose with Sc
and/or Rsl: e. g. Fig. 67 in Hampson 1895); Rs4
separated from other branches of Rs, but con-
nate or stalked with M l ; M2 usually arising mid-
way between Ml and M3 or closer to Ml than
to M3 (Auzea exceptional in having M2 slightly
closer to M3 than to Ml); CuAl and CuA2
stalked in males of certain Microniinae; no tubu-
lar CuP; a depression or transverse fold usually
present between bases of CuA and CuP; 1A +
2 A often forked (a free spur occasionally arises
from the apex of the "fork": Auzea). Hindwing:
humeral angle, when expanded, usually contain-
ing a humeral vein; subcosta diverging from up-
per edge of discal cell, usually after a short basal
connection (junction replaced by a crossvein in
Auzea)', position of M2 varying as in forewing;
Epipleminae sometimes lacking one or two M
veins (hindwing venation occasionally highly
modified: e. g. Forbes 1942, Fig. 100); no tubular
CuP; 3 A either well developed or reduced to ab-
sent. Abdomen: sternum 2 of male provided with
subtriangular or linear apodemes, which are
sometimes connected to fairly long venulae; in
female, anterior area of sternum 2 with a pair of
tympanal organs (see e. g. Sick 1937; 2 - 1 1 ) that
are adjacent to well defined apodemes (e. g. Mi-
net 1983: Fig. 96) and have clearly "exposed",
rounded tympana; male tympanal organs lying
laterally, at the junction between terga 2 and 3
(tympana on tergum 2, only protected by dense
fringes of scales: see e. g. Scoble 1992, Fig. 156);
in male, sternum 8 varying from well developed
and rectangular to reduced and/or modified (ter-
Jol Minet & Malcolm J. Scoble
gum 8 may also be modified and androconial
brushes may occur in diverse areas of genital/
pregenital segments). Male genitalia: tegumen
and vinculum frequently synscleritous; juxta
often fused, ventrally, with the valvae; gnathos
rarely well developed (Holloway 1976: Fig. 684),
usually reduced to absent (Figs, in Kuznetzov &
Stekolnikov 1981 and Boudinot 1982); phallus
sometimes with an anterior coecum, occasionally
with a ventral projection (Kuznetzov & Stekolni-
kov 1988: Fig. 1). Female genitalia: corpus bur-
sae variously ornamented, occasionally provided
with appendix bursae (Viette 1972); ostium bur-
sae on venter 8; apophyses posteriores moder-
ately developed, usually longer than apophyses
anteriores.
Eggs domed to subspherical (depressed in
Epipleminae: Dyar 1901, 1903), always of the
upright type (Guppy 1907; Eltringham 1924; Ca-
tala 1940; Coleman & Monteith 1981; Common
1990; Smith 1992). Chorion with many radiating
vertical ribs, sometimes obliterated in the upper
area. Last instar larva with only a few secondary
setae (e.g. Dyar 1898; Fracker 1915; Kodama
1959; Stehr 1987; Nakamura & Yoshiyasu 1992);
a number of spatulate setae in Chrysiridia and
most Urania species (e. g. Guppy 1907; Eltrin-
gham 1924; Catala 1940; Smith 1992); T1 with
SD and SD2 on dorsal shield; pro thoracic L
group bisetose; LI and L2 sometimes approxi-
mated on A l - A 3 , but always remote from each
other on A 4 - A 8 ; most abdominal segments with
a secondary seta cephalad of L3; ventral prolegs
usually also with a secondary seta (in SV group);
prolegs all well developed (5 pairs); in Epiplemi-
nae crochets bi- or uniordinal, arranged in a
semicircular mesoseries, which may be narrowed
mesally or even, occasionally, divided into two
bands; in Microniinae and Uraniinae, mesoseries
biordinal and relatively straight. Pupa: maxillary
palpi and forefemora concealed (Common 1990;
Nakamura & Yoshiyasu 1992); labial palpi con-
cealed or very slightly exposed; galeae long to
short; antennae long; forelegs tending to be
strongly extended cephalad; exposed part of
hindwing often broader near A3A4 suture than
near A3 spiracle; abdomen with two moveable
segments (A5; A6); anterior edge of AIO some-
times denticulate dorsally; cremaster present,
usually with hooklets of different sizes. Known
cocoons netlike, with large meshes.
Adults usually come to light at night, but three
colourful, uraniine genera are exclusively diurnal
(Alcides, Chrysiridia and Urania). At rest, the
wings are often outspread or rolled to a varying
degree, the forewings being then markedly re-
mote from the hindwings. The wings are never
held in a roof-like arrangement. The larvae are
external feeders and pupation often occurs in
leaf litter.
Worldwide, there are around 700 described
species, assigned to about 90 genera. As now
The Drepanoid/Geometroid Assemblage
conceived (Minet 1995), the family is divided
into four subfamilies that are distinguishable
with the help of the following key (imaginai
characters):
Key to the subfamilies
1 Male hindwing with a frenulum (usually also
present in female: Fig. 17.1 E) or forewing
with apex produced and termen distinctly
concave below it (Fig. 17.2 M) 2 of them being held remote from the substrate
Both sexes without frenulum (Fig. 17.1 F);
forewing termen straight or convex near
apex (Figs. 17.2 O - Q ) 3 by the forewings; in another case, wings are sym-
2 Male tympanal organ (on A2) with elongate,
internal projection arising from dorsal part
of tympanal frame (this projection visible
through the tympanum, after removal of
scales which protect the latter); forewing
with base of M2 always separate from Rs4
+ Ml (or from Ml); hindwing termen
rather regular (Fig. 17.2 M), at most slightly
convex at M3 Auzeinae
- Male tympanal organ (on A2) with internal
projection which is basally broad and arises
from anterior part of tympanal frame; fore-
wing with M2 and Rs4 + M1 often connate
(however, Ml is not necessarily stalked with
Rs4); hindwing frequently with an irregular
termen (i. e. variously notched and/or tailed:
Fig. 17.1 E) Epipleminae
3 Tarsi without obvious spines on tarsomeres
1 - 4 (spines, when present, minute and hid-
den by lanceolate scales); hindwing often an-
gulate (Fig. 17.20), at most provided with a
very short tail at M3 Microniinae
Ventral surface of tarsomeres 1 4 distinctly
spined; hindwing often with a conspicuous
tail at M3 (Figs. 17.2P, Q) (sometimes also
with other tails) Uraniinae
Auzeinae were recently proposed as a new sub-
family, and probably represent the most primi-
tive lineage within Uraniidae (Minet 1995). Pre-
viously, these insects were usually placed in
"Epipleminae" (e. g. Dalla Torre 1924) or, occa-
sionally, in Microniinae (Sick 1937; Minet 1983).
Auzeinae contain about 25 species, most of them
described in Deceda. They are mainly Indo-Aus-
tralian, but possess at least one species in tropi-
cal Africa. The wings often have oblique stripes
(Fig. 17.2M) perpendicular to the body axis
when the moth is at rest. The known larval host-
plants belong to Olacaceae (Lees & Smith 1992;
Minet 1995).
Epipleminae, with more than 550 described
species, are the largest subfamily of Uraniidae.
They occur on all continents except Europe and
Antarctica, but the group is mainly tropical.
Epiplemines are absent from New Zealand and
North Africa. The subfamily has been divided
into about 70 genera, but many of these are un-
309
satisfactorily defined (the main genera include
Dirades, Epiplema and Gathynia). The moths are
usually small and nocturnal, although a few me-
dium-sized insects are known, especially in the
neotropical region (e. g. Syngria, Symphytophleps
and Erosia: Fig. 17.2 N). Hindwings may be ex-
cised in different places (see e. g. Forbes 1942:
PI. 13) and their termen is often provided with
acute teeth or short tails. Resting postures vary:
for instance, the wings may be rather flat, three
(and one forewing appressed to the substrate),
with the hindwing costal areas being concealed
metrically outspread, with the flat forewings re-
mote from the hind pair; sometimes, fore- and
hindwings are conspicuously rolled, the two
pairs being held far apart from each other (see
e.g. Figs. 53.10, 11, in Common 1990) etc.
Young larvae tend to have gregarious habits and
usually live in a web. Recorded hostplants be-
long to various families: e. g. Bignoniaceae, Ca-
prifoliaceae, Daphniphyllaceae, Loganiaceae,
Oleaceae, Rosaceae, Rubiaceae and Verbenaceae
(Sevastopulo 1940; Pinhey 1975; Holloway et al.
1987; Stehr et al. 1987; Sugi et al. 1987; Lees &
Smith 1992).
Microniinae contain approximately 50 species,
referred to less than 10 genera. They are known
only from tropical Africa, Madagascar and the
Indo-Australian region (with a weak penetration
into palaearctic Asia). The adults are nocturnal
geometrid-like moths, with slender bodies and
white or whitish wings that usually exhibit
several darker stripes (Fig. 17.20). The forewing
venation is often sexually dimorphic, with CuA 1
and CuA2 separate in females, connate or
stalked in males. Acropteris and Micronia are the
main genera. At rest, wings are outspread and
appressed to the substrate; as in the preceding
subfamilies, the antennae are hidden under the
wings. Known larvae live on Asclepiadaceae
(Holloway et al. 1987; Yoshiyasu 1991; Lees &
Smith 1992; Nakamura & Yoshiyasu 1992).
Uraniinae adults, unlike Microniinae, lack a
well developed mesanepisternum, this sclerite
varying from reduced to absent (Brock 1971: Fig.
43 c). As mentioned above, female Uraniinae also
show a characteristic modification of their mete-
pimeron (Sick 1937: Figs. 34-36). With such a
definition, the subfamily contains about 50 de-
scribed species (Lees & Smith 1992) and in-
cludes, in addition to the Indo-Australian Lyssa
and the well known day-flying taxa, three micro-
niine-like genera from the Old World tropics, viz.
Urapteritra (Fig. 17.2P), Urapteroides and Cy-
phura (Sick 1937; Minet 1983; Kite et al. 1991).
At rest, the moths usually have their wings
broadly outspread and more or less appressed to
the substrate (e. g. Altena 1953: Figs. 2 and 3;
Laithwaite et al. 1975: Figs. 315 and 316); how-
ever, in at least one genus, Chrysiridia, the imago
310
rests at night with wings held vertically over the
back (Lees & Smith 1992). The diurnal, bril-
liantly coloured genera are closely related: Al-
cides inhabits the Australian region, Chrysiridia
occurs in Madagascar and East Africa, and Ura-
nia (Fig. 17.2Q) is neotropical. These three
genera are famous for their migratory flights (see
e. g. Smith 1992). All uraniine larvae feed on Eu-
phorbiaceae: Endospermum (Alcides and three
nocturnal genera), Suregada (Urapteritra) and
Omphalea (Alcides, Chrysiridia and Urania) (e. g.
Kite et al. 1991; Lees & Smith 1992; Forster &
Sankowsky 1995).
Geometridae. Geometridae are macrolepidopter-
ans characterized by the presence of structurally
unique tympanal organs at the base of the abdo-
men. Only rarely are these structures reduced or
lost. Their larvae are frequently easy to distin-
guish from those of other families since the
number of prolegs on the abdomen is typically,
but not exclusively, reduced to two pairs causing
the organisms to move by 'looping' (e. g., as in
Fig. 17.5 A). The name Geometridae is derived
from this 'earth-measuring', looping habit of the
larvae. The family, which is represented globally,
includes about 21000 described species (Scoble
et al. 1995), making it one of the three largest
families of Lepidoptera. Many geometrid moths
are brightly coloured, but most are drab. Fre-
quently, the wings bear transverse, wavy-lines,
but bold, distinct patterns occur in many species.
Sometimes the moths may be confused with
other families, particularly Noctuidae; however,
structurally they are distinct.
Adults (Figs. 17.3 A - L ; 17.4A, B): small
(forewing length around 5 mm) to medium-large
(forewing length reaching 50 mm or more)
macrolepidopterans; generally rather slender-
bodied, broad-winged, and somewhat delicate,
but several more 'robust'-bodied species exist;
wing reduction occurring often and indepen-
dently. Head densely scaled. Antenna filiform,
serrate or bipectinate; pectinations often more
prominent in male than in female. Chaetosemata
present, typically small. External ocelli generally
absent. Head thickly scaled. Proboscis usually
well developed, not scaled at base; sometimes re-
duced or even absent; maxillary palpi minute, 1 -
or (occasionally) 2-segmented. Labial palpi up-
turned, but not strongly ascending. Foreleg with
epiphysis. Tibial spurs lacking on foreleg, typi-
cally one pair occurring on midleg and two pairs
on hindleg (0-2-4); occasionally tibial spur for-
mula is 0-2-2, 0-2-3, or 0-2-0. Hindtibia of male
sometimes swollen and bearing scent-brush lying
in groove; presence of brush may be correlated
with the occurrence of a comb, or two patches
of setae, on sternum A3 in male sex (B. Hua,
pers. comm.). Wings usually broad, but brachyp-
tery, microptery, and aptery also occurring with
such reduction having evolved often and inde-
Jol Minet & Malcolm J. Scoble
pendently (Sattler, 1991). Forewing (Fig. 17.6 A):
one or two areoles usually formed by veins R1
and Rs; vein Rs4 stalked with Rs2 and Rs3; vein
M2 usually arising nearer M l than M3, some-
times arising midway between M l and M 3 or,
rarely, close to M3; vein CuP absent. Hindwing
(Figs. 1 7 . 6 A - E ) : vein Sc may be bent strongly
at its base; it may run close to Rs or may fuse
with Rs for part of its length, or may be con-
nected to Rs via R; vein Sc diverging before or at
end of the cell; vein M2 usually arising midway
between M l and M3 or nearer M l , or absent or
very weak; occasionally, vein M2 arising nearer
M3 than M l . Spinarea (i. e., forewing-metathor-
acic, aculeate locking device) absent, yet, re-
markably, wings sometimes tightly folded over
abdomen when moth is at rest (e. g., as in Chlen-
ias, see McFarland, 1988). Wing coupling usually
effected by frenulum and retinaculum but these
structures often reduced. Metapostnotum with
enlarged, membranous fenestra media, probably
functioning as accessory tympanum, in all Geo-
metridae (e. g., Fig. 17.5 B) except Archiearinae
where fenestra media are narrow (Minet 1983).
Paired tympanal organs, structurally unique
(Kennel & Eggers, 1933; Minet 1983; Cook &
Scoble 1992; and see 211), present at base of
abdomen, reduced or lost in brachypterous fe-
males; opening ventrolaterally; lying in deep,
ventrolateral walled cavities (cavi tympani of
Cook & Scoble 1992), which are not homologous
with bullae tympani of Pyralidae; circumference
of most of tympanum supported by peritympa-
nal frame of cavus; where not supported by this
frame (i. e., along its anterior margin) then bor-
dering the thoraco-abdominal membrane; scolo-
parium with four sensory cells, strung between
tympanum and ansa (the sclerotized arm curved
over the tympanum, which represents a modifi-
cation unique to Geometridae). Sternum A3 of
abdomen frequently bearing two fields of bris-
tles, often united to form a row. Male genitalia:
most basic lepidopteran components present, but
characters reliably distinguishing Geometridae
from other families not apparent; valvae often
fairly simple, sometimes with complex modifica-
tions. Female genitalia: ovipositor not typically
extended, but exceptions exist, as in many mem-
bers of the Hypomecis-group (see figures of Sato,
1984) and a number of other Boarmiini (Hollo-
way 1994).
General treatments in e. g., Zerny & Beier,
1936; Bourgogne, 1951; Inoue 1961; Munroe
1982; McGuffin 1987; Common 1990; Nielsen &
Common 1991; Scoble 1992.
Eggs (Salkeld 1983) usually of 'flat' variety
with longitudinal axis lying parallel, or almost
parallel, to substrate; in some species 'upright',
with longitudinal axis running perpendicular to
substrate; both kinds of egg sometimes occurring
within same genus (e. g., Sterrha, Biston) (Hin-
ten, 1981); eggs of some species lying at angle to
The Drepanoid/Geometroid Assemblage 311

Fig. 17.3. Adults of Geometridae: A, Archiearis parthenias (Archiearinae); B, Thalaina clara (Ennominae); C,
Semiothisa gambaria (Ennominae); D, Bistort betularia (typical form) (Ennominae); E, Bistort betularia (melanic
form) (Ennominae); F, Epimecis hortaria (Ennominae); G, Oertochroma vinaria (Oenochrominae s. 1.); H, Crypsi-
phorta ocultara (Geometrinae); I, Omphaxplantara (Geometrinae); J, Anisodes rufulata (Sterrhinae); K, Cyllopoda
claudicla (Sterrhinae); L, Scopula aniara (Sterrhinae). Scale lines: 5.0 mm.
312 Jol Minet & Malcolm J. Scoble

Fig. 17.4. Adults of Geometridae: A, Eupithecia mille/oliata (Larentiinae); B, Operophtera brumaio, male (Larentii-
nae); C, Operophtera brumata, female (Larentiinae). Scale lines: 5.0 mm.

Fig. 17.5. Geometridae: A, larva of Erannis defoliaria (Ennominae); B, posterior aspect of thorax (abdomen re-
moved) of Scingala (Ennominae) showing accessory tympanum (i. e., enlarged, membranous fenestra media).

substrate, in some others not attached to sub-
strate; shape in lateral view usually ovoid, ellipti-
cal or bluntly elliptical, when viewed from either
end then circular or oval (Salkeld 1983); sculp-
turing present or absent. Larva (e. g., as in Fig.
17.5 B) (McGuffin 1987; McFarland 1988; Stehr
1987) generally cylindrical and slender-bodied
(length about 10 times the width), less frequently
stout, but rarely flattened; cuticle smooth or
granular, often with longitudinal stripes running
length of body; various humps, filaments, or
other processes may occur. Head rounded in first
instars, in later instars more rectangular or bifid
(particularly in Geometrinae); spinneret typically
shorter than prementum (Minet 1991); full com-
plement of primary setae present, fourth SV seta
occurring on abdominal segments (Stehr 1987)
(interpreted as L3 by Minet, unpublished); sec-
ondary setae not usually prominent or dense.
Prolegs typically only on abdominal segments
A6 and AIO; certain species with full comple-
ment of prolegs with some reduced in size, or less
than a full complement but with more than the
two pairs typical of the family; crochets may be
lacking from segments other than on A6 and
AIO. Pupa (McGuffin 1987): maxillary palpi not
exposed; cremaster usually well developed.
In several species of 'winter moths' the wings
are virtually lost in the female. These moths are
mated almost immediately after emergence from
the pupa and lay their eggs soon after. Wing re-
duction occurs in far more species of Geome-
tridae than in any other family, and has evolved
independently many times in Oenochrominae
(sensu lato), Larentiinae (tribes Xanthorhoini
and Operophterini), and Ennominae (tribes Ma-
cariini (= Semiothisini), Boarmiini (in the broad
sense of Holloway 1994, and Gnophini). (See
Sattler, 1991 for a detailed review of wing reduc-
tion in Lepidoptera.)
The major ecological impact of Geometridae
is observed in the tropics or subtropics where
most species (other than those belonging to Lar-
entiinae) exist. Under appropriate conditions,
certain species may reach pest status through the
defoliating activities of the larvae; but the Geo-
metridae are not a major pest group. Neverthe-
less, their environmental significance is profound
in terms of their activities as primary consumers
of green plants and as potential items of prey for
The Drepanoid/Geometroid Assemblage
parasitoids and vertebrate predators. Also, their
tendency towards habitat specificity, gives Geo-
metridae a special value as indicators of environ-
mental change (e. g., Holloway 1984).
Eggs are laid singly, or in batches wrapped
around twigs. They may be glued to plant sub-
strate, inserted in crevices of bark, or simply
dropped. As a very broad generalization, the lar-
vae tend to feed on the leaves of trees or shrubs;
their association with herbaceous plants is less
pronounced. They are often camouflaged, and
frequently assume an outstretched twig- or stick-
like position on the plant while grasping the sub-
strate with their prolegs on A6 and AIO. Loco-
motion occurs by 'looping' (Fig. 17.5 A) even in
species without reduced prolegs. Pupation occurs
typically within a flimsy cocoon in debris, but
some pupae are merely girdled.
The taxonomy of the family has developed
over a long period, but the infrafamilial classifi-
cation has not yet received much critical atten-
tion. The history of classification of Geometridae
to early in this century was chronicled by Prout
(1910). Six subfamilies are generally recognized,
but the system is unsatisfactory with the mono-
phyly of all being, at the least, in some doubt.
Oenochrominae sensu lato is a polyphyletic
grouping (Scoble & Edwards 1990). We are far
from achieving a resolved system, so the subfam-
ily classification in general current use is best re-
tained at present. Many monophyletic generic
groupings are evident within the family (Hollo-
way et al. 1987, and Holloway 1994) and tribal
names have frequently been applied (e. g., by
Forbes, 1948; McGuffin 1967, 1972, 1977, 1981,
1987; Holloway [1994]).
The status of the subfamilies is uncertain so
the key presented below should be treated as a
preliminary guide of the most general kind only,
for numerous exceptions to it exist; its value as
an identification tool is limited.
Important tribal names currently in use are
listed under the appropriate subfamilies, and ref-
erences are given to works that provide a concept
of a given tribe. Many tribal names have been
used in Geometridae classification (J. D. Hollo-
way in prep. ) and considerable change in nomen-
clature at this level is expected. The tribes men-
tioned here were selected from studies that have
taken at least some account of the Geometridae
as a whole, even if they are themselves essentially
regional works.
Key to subfamilies
(Based on Holloway et al., 1987, but with modi-
fication. Because they are certainly not mono-
phyletic, Oenochrominae sensu lato have been
omitted.)
1 Vein M2 of hindwing weak (not existing as
a tubular vein) (as in Figs. 17.6 A, B) or not
visible 2 (Carter & Hargreaves 1986) although a full com-
313
- Vein M2 of hindwing strong, present as a
tubular vein (as in Fig. 7.6 CE) 3
2 Fenestra media of metapostnotum small,
not expanded into accessory tympanum.
Face 'hairy'. Restricted to temperate or sub-
alpine areas. Day flying . . . . Archiearinae
- Fenestra media of metapostnotum large (as
in Fig. 17.5 B), forming accessory tympa-
num. Face typically not 'hairy'. Not re-
stricted to temperate or subalpine areas;
mostly tropical or subtropical. Typically
nocturnal Ennominae
3 Vein Sc of hindwing strongly anastomosing
with Rs for most of length of cell (as in Fig.
17.6E) or, rarely, connected via vein R l .
Fasciae of forewing usually composite,
formed from numerous lines (e. g., as in Fig.
17.4 A) Larentiinae
- Vein Sc of hindwing typically free from or
approximating Rs at or near its base. Fas-
ciae of forewing, when present, not compos-
ite (e. g., as in Fig. 17.3 L) 4
4 Most species green. Socii usually present,
sometimes more prominent than uncus; un-
cus often bifid. Vein M2 of hindwing arising
nearer M l than M3 (as in Fig. 17.6C) . . .
Geometrinae
- Species not green. Socii not usually present.
Vein M2 of hindwing tends to arise approxi-
mately midway between Ml and M3 (e. g.,
as in Fig. 17.6E) 5
5 Moths mostly slender-bodied, pink or pale
fawn, occasionally yellowish or white.
Wings often with many fasciae (e. g., as in
Fig. 17.3 L) (fasciae not typically composite,
unlike Larentiinae). Uncus or gnathos often
reduced Sterrhinae
- Moths robust-bodied, mostly grey, yellow,
or brownish. Wings without numerous fas-
ciae. Uncus and gnathos typically promi-
nent, not usually reduced
Oenochrominae sensu stricto
Archiearinae (= Brephinae) (Fig. 17.3 A) (see
Prout 1910; Inoue 1961), with 12 species of me-
dium-sized, temperate Geometridae, occur in the
northern Palaearctic and Nearctic (Archiearis
and Leucobrephos), the southern Andes in Chile
(Archiearides and Lachnocephala, see Fletcher
1953) and subalpine regions of Tasmania (Dirce;
Acalyphes, see Common 1990). The subfamily
has, therefore, a strikingly disjunct distribution.
These rather colourful diurnal moths bear nu-
merous hair-like scales on the front of the head,
and have small, oval compound eyes and densely
hairy labial palpi. Vein M2 of the hindwing is
usually present but weak. The accessory tympa-
num is lacking from the metapostnotum the fe-
nestra media being narrow (Minet 1983). Eggs of
Archiearis are of the upright variety. The larvae
(at least in Archiearis) progress by 'looping'
314
Fig. 17.6. Wing venation of Geometridae. A, forewing and hindwing of Ennomos autumnaria (Ennominae); B - E ,
hindwings: B, Archiearis notha (Archiearinae); C, Oospila albicoma (Geometrinae); D, Idaea sericeata (Sterrhinae);
E, Larentia clavaria (Larentiinae). Scale lines: A, 0.5 mm; D, 0.25 mm. Scales of B, C, E as for A.
plement of prolegs, the anterior pairs of which
are nevertheless reduced in size, is usually pre-
sent. Crochet-bearing prolegs are absent, how-
ever, from abdominal segments A 3 - 6 in larvae
of Dirce (McQuillan in Common 1990).
In the Palaearctic, the males fly mainly in sun-
shine early in spring. Females of Archiearis tend
to remain near the tops of their hostplant trees,
aspen or birch. In Tasmania, males of Acalyphes
also fly mainly in sunshine around pencil pines
(Arthrotaxis cupressoides: Pinaceae) (McQuil-
lan & Scoble, unpublished), although this tree
has not been established as the foodplant.
Ennominae (Fig. 17.3 BF) (see Janse 1932;
Forbes 1948; McGuffin 1987; Holloway [1994]),
with over 9700 name species, include about 45%
of all Geometridae. The monophyly of the sub-
family is not well established: currently, the defi-
nition of Ennominae is based largely on the ab-
sence of M2 as a tubular vein in the hindwing
(e. g., as in Fig. 17.6 A), but this character is not
entirely reliable. The presence of a transverse
band of posteriorly directed setae on segment A3
of the male occurs widely in the subfamily as
currently constituted, but is absent from a high
proportion of genera (Holloway et al. 1987). En-
nominae are mainly slender-bodied, medium to
large Geometridae, but a group of robust-bodied
species, often with adults that fly in autumn, is
represented mainly in southern Australia (e. g.,
McQuillan 1981, 1984; McFarland 1988).
Both sexes are generally fully winged, but fe-
males of some species are almost wingless. In
males of several species, and in females of some,
a blister-like fovea occurs near the base of the
Jol Minet & Malcolm J. Scoble
forewing between veins Cu and 1 A + 2 A. Fo-
veae differ in structure and position within En-
nominae so these structures may not be homolo-
gous throughout the group (J. D. Holloway,
pers. comm.). In the forewing (e. g., as in Fig.
17.6 A), vein R commonly anastomoses with Sc.
A hairpencil, usually concealed in a deep groove,
is often present on the tibia of the hindleg. Its
occurrence seems to be correlated with the pres-
ence of a setal comb or setal patches on sternum
A3 in the male sex.
In some Ennominae (particularly from the
southern hemisphere) the number of larval pro-
legs is not reduced to the extent typical of Geo-
metridae. Abraxas grossulariata, which exhibits
some colour variation in the larval and pupal
stages, and considerable variation in adult wing
colour, has been the subject of numerous genetic
experiments (summary in Robinson 1971). The
classic studies on industrial melanism were un-
dertaken on Biston betularia (Fig. 17.3 D, E) (see
Kettlewell 1973). Despite the size of the subfam-
ily, there are relatively few serious ennomine
pests. In Europe (see Carter 1984) larvae of Bu-
palus piniaria defoliate conifers, and larvae of Er-
annis defoliaria (Fig. 17.5 A) cause damage, also
by defoliation, to orchard trees and trees in for-
est nurseries. Ornamentals and fruit trees in gar-
dens may be damaged by larvae of Abraxes
grossulariata, and larvae of Biston betularia
sometimes attack nursery plantations. Pests of
hardwood trees in North America include Enno-
mos subsignaria, Erannis tiliaria, and Phigalia ti-
tea', Lambdina scellaria causes serious damage
particularly to conifers (Stehr 1987).
The Drepanoid/Geometroid Assemblage
The great number of species and the variation
exhibited across the subfamily have defied effec-
tive systematization. Tribal names with some
substantiation (see Forbes 1948; and particularly
Holloway 1994, for a recent review) are: Maca-
riini (= Semiothisini) (e.g., as in Fig. 17.3C)
(McGuffin 1987; Holloway [1994]); Abraxini
(McGuffin 1987; Holloway [1994]); Bistonini
(e.g., as in Fig. 17.3D, E) (McGuffin 1987;
Rindge 1985); Boarmiini (e. g., as in Fig. 17.3 F)
(McDunnough 1920, as Cleorini; Hypomecis-
complex of Sato 1984; McGuffin 1987; Holloway
[1994]); Melanolophini (McGuffin 1987; Rindge
1990); Ennomini (Forbes 1948; McGuffin 1987);
Caberini (McGuffin 1987; Holloway [1994]);
Baptini (McGuffin 1987; Holloway [1994]); Azel-
inini (Poole 1987, revision of Pero; McGuffin
1987); Campaeini (McGuffin 1987); Nacophorini
(Forbes 1948; Rindge 1983; McGuffin 1987);
Epirranthini (McGuffin 1987); Angeronini
(McGuffin 1987); Lithinini (McGuffin 1987;
Rindge 1986; Holloway 1987, 1994); Hypochros-
ini (sensu Holloway [1994]) (Anagogini of
McGuffin 1987, Hypochrosis-group of Holloway
et al. 1987, Hypochrosidae of Guene 1857);
Cingiliini (McGuffin 1987); Ourapterygini
(McGuffin 1987; Holloway [1994]); Gonodontini
(in the redefined sense of Holloway [1994]);
Sphacelodini (Forbes 1948).
Oenochrominae, as generally constituted (e. g.,
Prout 1910; Turner 1929-1930), are an unnatu-
ral group of Geometridae largely encompassing
those genera that cannot be placed in any other
subfamily. They include both slender- and stout-
bodied forms. Given the unnatural status of the
'group', useful generalizations cannot be made
about it.
Tribes listed by Prout (1910), in his world
treatment, include Oenochromini (as 'Oenochro-
micae') and Ametridini (as 'Ametridicae'). (The
tribe 'Hedylicae' was removed as superfamily
Hedyloidea by Scoble 1986.)
Oenochrominae sensu stricto (Fig. 17.3 G)
(Prout 1910, in part; Turner 1930, in part) are
composed of the robust-bodied and almost
exclusively Australasian moths belonging to
Oenochroma or related genera (Scoble & Ed-
wards 1990). Although the exact composition
of this group remains uncertain, the genera in-
cluded are similar in their general appearance, in
the structure of their male genitalia, and in their
pattern of wing venation. The main hostplants
are Proteaceae and Myrtaceae (McFarland 1979;
1988). The larvae often bear two pairs of ventral
prolegs in addition to the anal pair. The extra
pair, which occurs on segment A5, are much re-
duced.
Alsophilinae. The genus Alsophila, previously
included in Oenochrominae sensu lato, was as-
signed to a subfamily of its own by Holloway
(1996). Its members may be serious defoliators
of trees. Larvae of A. aescularia attack fruit trees
315
in Europe (Carter 1984), and those of A. pomet-
aria damage hardwoods, particularly in urban
areas, in North America (Stehr 1987).
Desmobathrinae. A pantropical grouping of
slender-bodied taxa formerly included in the Oe-
nochrominae sensu lato. Two tribes are recog-
nized, Desmobathrini and Eumeleini (Holloway
1996).
Geometrinae (= Hemitheinae) (Fig. 17.3 H, I)
(see Prout 1912; Janse 1934; Forbes 1948; Fergu-
son 1969, 1985) include around 2300 named spe-
cies in about 200 genera worldwide, and the
group is probably monophyletic. Most species
are tropical. In the tropics of the Old World oc-
curs a group of species with lichen-like or mot-
tled wings and stout bodies (Fig. 17.3 H). How-
ever, most species are green and have slender-
bodies (e.g., Fig. 17.31). The green pigment is
not a bile pigment but may be a derivative of
chlorophyll, although it is not chlorophyll a or
b (Cook et al., 1994). Although predominant in
Geometrinae, the pigment also exist in other
moths but usually only as a trace, and it is likely
that the presence of a large quantity of the pig-
ment is apomorphic for the subfamily. In the
forewing, vein R is usually separate from Rsl,
and anastomoses with Sc for a short distance. In
the hindwing (Fig. 17.6 C), vein M2 arises closer
to Ml than to M3. Two fields of deciduous
spines are found on abdominal sternum A3. The
frenulum is typically much reduced. In the male
genitalia, large socii very often occur (Holloway
et al., 1987).
The eggs (Salkeld 1983; Ferguson 1985;
McFarland 1988) are usually lozenge-like, and
seem to be dropped freely - rather than inserted
into crevices. The larval head is often bifid, and
prolegs are represented only on abdominal seg-
ments A6 and AIO. Synchlorini larvae attach,
with silk, pieces of plant material to well devel-
oped chalazae on their dorsal surfaces. Larvae of
Nemoriini bear prominent dorsolateral protu-
berances. Geometrinae larvae pupate in a flimsy
cocoon, usually on the hostplant but sometimes
on the ground among loose debris - not in the
soil (Ferguson 1985; McFarland 1988).
Concentration on regional faunas means that
an integrated tribal classification of the subfamily
on a global basis has not been achieved. Tribal
names in recent literature include: Nemoriini,
Dichordophorini, Synchlorini, Lophochoristini,
Hemitheini (all of which were used in a Nearctic
treatment by Ferguson 1985); and, in addition to
those of Ferguson, Terpnini (= Pachyodes-
group, based on 'Terpna\ a misspelling of
Terpne, itself an unavailable name, see Fletcher
1979), Aracimini, Timandromorphini, Geomet-
rini, Neohipparchini, Ochrognesiini, Thalassod-
ini, Jodiini, Rhomboristini, Comibaenini, Hemi-
ostolini, Comostolini, which were used by Inoue
(1961) in his work on the Japanese species. Re-
cent studies on species-rich groups of neotropical
316
Geometrinae have been carried out for Nemoria
and its relatives (Pitkin 1993) and Oospila
(Cook & Scoble 1995). A review of neotropical
geometrines at the generic level is given by Pit-
kin (1996).
Sterrhinae (= Acidaliinae), probably more ap-
propriately named Scopulinae (see Holloway
1994), (Fig. 17.3 JL) (see Janse, 1934; Forbes
1948; McGuffin 1967; Covell 1983; Holloway
et al., 1987), with around 2800 named species,
are generally small, rather inconspicuous Geo-
metridae with wings bearing numerous wavy fas-
ciae running from forewing to hindwing. The dis-
cal spot is usually prominent in both pairs of
wings and, in Anisodes (e. g., Fig. 17.3 J) and its
relatives, often larger in the hindwing and ap-
pearing as an open circle rather than a dark dot
or dash. Most species are nocturnal, but some
are colourful and diurnal (e. g., Cyllopoda, Fig.
17.3 K, and its relatives, with striking black and
yellow markings Covell, 1983). Although
mostly tropical, many Sterrhinae occur in tem-
perate areas. While members of the Anisodes-
group are typically arboreal, living in forest habi-
tats, those in the Scopula-group (e. g., Fig.
17.3 L) and in Idaea tend to live in open habitats
and feed on herbs (Holloway et al. 1987). Vein
Sc + RI of the hindwing typically fuses with Rs
near the wing base for only a short distance be-
fore it diverges sharply (Fig. 17.6D), but it is
by no means certain that this feature effectively
defines the group, and in Rhodometra the anas-
tomosis is long.
The eggs (Salkeld 1983) are flattened, vari-
ously ribbed and sculptured, and deposited
loosely. The larvae (McGuffin 1967; McFarland
1988) are usually cryptic with some having a bi-
lobed head and a long, slender body. Only two
pairs of prolegs are present. The larvae of most
species feed on low herbs, but some (e. g., Ani-
sodes and its relatives) are arboreal. Pupation
generally occurs in a flimsy cocoon, but in Ani-
sodes the pupae are girdled (Common 1986) as
they are in the related genus Cyclophora (and at
least one of its relatives), the girdle traversing
abdominal segment A3. Adults of some S. E.
Asian Species of Scopula feed on blood, from
mammalian wounds, on sweat, and on tears
(Bnziger & Fletcher 1985).
Tribal names (see Holloway 1994) include
Scopulini (= Acidaliini); Cyclophorini (= Zono-
somini; Cosymbiini); Sterrhini (= Idaeini); Ti-
mandrini (= Calothysanini); Rhodometrini; Cyl-
lopodini; Rhodostrophini.
Larentiinae (= Hydriomeninae) (Fig. 17.4 A,
B) (see Janse 1933; Forbes 1948; Holloway et al.,
1987), with over 5700 species, are widely distrib-
uted, but most occur in temperate regions and at
high altitudes in the tropics (e. g., Holloway
1986; McQuillan 1986). A discal spot is fre-
quently present on both forewing and hindwing.
Fasciae on the forewings are often composed of
Jol Minet & Malcolm J. Scoble
bands of closely juxtaposed lines. In females of
some species the wings are much reduced. In the
hindwing (e. g., Fig. 17.6 E), vein Se + R usually
fuses with Rs for at least half the length of the
cell, and vein M2 is fully developed. Wing reduc-
tion has been recorded in many species belonging
to tribes Xanthorhoini and Operophterini (in-
cluding the 'winter moths', where males fly in
cold weather to flightless females) (Sattler 1991).
The eggs (Salkeld 1983) are smooth, weakly or
heavily sculptured, and usually oblong and vari-
ously flattened. The larvae (McFarland 1988) are
generally short and relatively stout. Some con-
struct nests of leaves and silk, some resemble
twigs or stems, and others hide in crevices. In
nearly all Hawaiian species of the genus Eupi-
thecia, the larva is carnivorous and ambushes
small insects and spiders (Montgomery 1982),
whereas in other members of this widespread
genus the larvae typically feed concealed in buds
and flowers or even bore in cones. Concealed
feeding is extreme in the larva of Coenotephria
incultaria, which mines the leaves of Primula (see
Hering 1951). Larvae of Operophtera brumata
are often serious pests of fruit trees, and some-
times forest trees, attacking leaves, buds, blos-
som and developing fruit in Europe (Carter
1984). This species has been introduced on both
east and west coasts of North America where it
has become a pest of hardwoods (Stehr 1987).
The larvae of Chloroclystis rectangulata, which
feed on the blossom of fruit trees, also frequently
reach pest status in Europe (Carter 1984). This
species has been introduced also into North
America where it has become well established
(Ferguson 1972).
Tribes included by Forbes (1948) include Hy-
driomenini, Stamnodini, Xanthorhoini, Asthen-
ini, Operophterini, Eudulini, Eupitheciini, and
Lobophorini (this last tribe falling into Trichopt-
erygini, see Dugdale 1980).
Acknowledgements
For their valued comments on sections of the
manuscript, we thank Dr J. D. Holloway, In-
ternational Institute of Entomology, London, Dr
D. C. Lees, The Natural History Museum, Lon-
don, and an anonymous referee. We are grateful
to Monsieur G. Hodebert, Musum National
d'Histoire Naturelle, Paris, for figuring the wing
venation of Epicopeiidae, Drepanidae, Sematuri-
dae and Uraniidae, and to members of the Pho-
tographic Unit at The Natural History Museum,
London, for their assistance in preparing the
halftone figures for the Geometridae. For pro-
viding an English translation of one of his papers
from the Japanese, we thank Mr M. Nakamura,
Shizuoka Pref., and we are grateful to Mr S.
Sugi, Tokyo, for gifting larvae and pupae of Epi-
copeiidae.
The Drepanoid/Geometroid Assemblage
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 18. The Bombycoidea and their Relatives
Claude Lemaire & Jol Minet
(C. Lemaire: Saturniidae; J. Minet: Mimallonoidea, Lasiocampoidea and non-saturniid Bombycoidea)
Taken together, these taxa probably form a
monophyletic group (Minet 1991, 1994) and they
have been occasionally regarded as parts of a
single superfamily (Brock 1971). Yet there are
few known apomorphies to support the mono-
phyly of this group. Among the most convincing
is the relative closeness on the mesothorax of the
dorsal extremeties of the prescutal clefts (Minet
1991: Figs. 3138). In several advanced taxa,
these clefts may even merge, with the prescutum
becoming entirely separated from the mesoscu-
tum. However, in a few genera (e. g. Mirino),
they are only slightly closer to each other dor-
sally than in the hypothetical ground plan of the
Macrolepidoptera.
Mimallonidae retain several larval plesiomor-
phies of rare occurrence within Macrolepido-
ptera. For example, the prolegs are coronate
(also found in primitive butterflies), and setae LI
and L2 are approximated on A 1 - A 8 (as in some
Bombycoidea; see Franclemont 1973: 16). In
contrast with those of Lasiocampoidea and
Bombycoidea, their mature larvae lack second-
ary setae on most parts of their integument. Ac-
cordingly, modern authors usually place Mimal-
lonidae in a superfamily of their own (Francle-
mont 1973, Fletcher & Nye 1982, Minet 1986,
Holloway et al. 1987, etc.). Earlier authors pro-
posed inclusion in such superfamilies as Urani-
oidea (Forbes 1923), Tineoidea s. I. (Handlirsch
1925, Borner 1939), Mimallonoidea (Lima 1950)
or Bombycoidea (Bourgogne 1951). The family
was even recently viewed as a member of Pyra-
loidea (Scott 1986).
Often placed in Bombycoidea, Lasiocampidae
deserve a separate superfamily according to cer-
tain authors (in particular Tillyard 1926, Lima
1950, and Kuznetzov & Stekolnikov 1985). The
present concept of Lasiocampoidea includes An-
thelidae (Minet 1991 and 1994), based on five
possible synapomorphies. None of these traits is
restricted to Lasiocampoidea, so the systematic
position of Anthelidae clearly needs further in-
vestigation. In Lasiocampoidea, larvae never
possess fused forecoxae and the adults have fore-
wings with the base of vein M2 invariably closer
to M3 than to M l .
Sphingidae were long assigned to a monotypic
superfamily (Sphingoidea) but they are genuine
Bombycoidea, as indicated by characters of their
larvae and adults (e. g. Brock 1971, Minet 1986
and Holloway 1987). Redefined by Minet (1991),
Bombycoidea s. str. include the following nine
families: Eupterotidae (syn. : Hibrildidae), Bom-
bycidae (syn. : Apatelodidae), Endromidae, Miri-
nidae, Saturniidae (syn. : Attacidae, Cercophani-
dae and Oxytenidae), Carthaeidae, Lemoniidae,
Brahmaeidae and Sphingidae (see Minet 1994 for
discussion of familial limits). Bombycoidea are
considered monophyletic, based on such autapo-
morphies as: (a) forecoxae anteriorly fused in the
last instar larva; (b) A8 of the larva with D1 se-
tae arising from a middorsal protuberance (usu-
ally a scolus); (c) forewing with Rsl + Rs2
closely parallel to stem Rs3 + Rs4, or even fused
to it; (d) male genitalia with a modified position
of "muscles 4" (Kuznetzov & Stekolnikov 1985).
From what is known about the phylogeny of
Bombycoidea, the absence of (a) in Apatelodinae
and certain Eupterotidae (Oberprieler & Duke
1994) is likely to represent a reversal, as is the
absence of (b) in Eupterotidae and a few other
taxa.
Mimallonoidea, Lasiocampoidea and Bomby-
coidea do not share many distinctive traits. The
eggs are usually of the flat type, with a rather
smooth chorion, but upright eggs occur in sev-
eral Bombycoidea (e. g. Eupterotidae, Lemonii-
dae and Brahmaeidae), and the chorion is sculp-
tured in certain Mimallonidae. The mature larva
has five pairs of prolegs, the anal pair often be-
ing distinctly enlarged. There may be only a few
secondary setae (Mimallonoidea) but these are
usually numerous (Lasiocampoidea, Bomby-
coidea), varying from long to minute. Verrucae
and scoli are frequent. The stout pupa often has
short antennae; its abdomen has two moveable
segments (A5, A6) and occasionally is provided
with dorsal, acute teeth. There is often an elabo-
rate, silken cocoon (used for commercial silk
production in certain species). The imago lacks
functional ocelli and genuine chaetosemata. The
male antennae are bi-, tri- or quadripectinate, ex-
cept in most Sphingidae where they are second-
arily modified. In the forewing, Rsl and Rs2 are
either stalked or entirely fused, Rsl + Rs2 occa-
sionally forming the "anterior" edge of an
areole. Although present in most Lasiocampidae
(Figs. 18.1 E and F, sp), the spinarea is entirely
absent elsewhere (term defined by Minet: 1990:
355). On the mesothorax, the meron and epi-
meron have a long, or fairly long, line of junc-
tion. The metascutum is divided or almost di-
vided mesally. The thorax and the abdomen al-
ways lack tympanal organs.
322
Key to the families: adults
1 Micropterous (a few female moths) . . . . 2
Fully winged 4 Hindwing with vein Se + R remote from
2 Australian Anthelidae (in part)
Non-Australian 3 18.3A-C, 1 8 . 5 A - G )
3 Antenna dorsally scaled to the apex
Lasiocampidae (in part)
Antenna apically devoid of scales
Saturniidae (in part)
4 Forewing with stem Rsl + Rs2 usually free
from other branches of Rs, and either Rs3
free (Fig. 18.1 E) or the apex of stem Rs3 +
Rs4 more or less remote from Rsl + Rs2
(e.g. Fig. 18.1 A); if Rs branches are ar-
ranged differently, an areole is associated
with them or a chaetosema-like organ is pre-
sent on labial palpus segment 1 5 (Figs. 18.51, J)
Forewing stem Rs3 + Rs4 closely parallel to
Rsl + Rs2 (Figs. 18.5G, I, J), or all
Rs branches stalked together (Figs. 18.1
GI, 18.3 A - C , 18.5 A - F , and 18.5H); ar-
eole absent; labial palpus segment 1 never
with a chaetosema-like organ (Bomby-
co idea) 7 ways absent, both sexes having amplexiform
5 Forewing without areole; M2 (forewing)
often arising approximately midway be-
tween Ml and M3 (Figs. 18.1 A, B); hind-
wing subcosta diverging strongly from upper
edge of discal cell (after a short fusion with
the base of this edge) and remote from the
postdiscal section of Rs (i. e. that section of
Rs which leaves the discal cell); no chaeto-
sema-like organ at base of labial palpus (Mi-
mallonoidea) Mimallonidae
Forewing with areole sometimes present;
M2 (forewing) always arising nearer M3
than M l (Figs. 18.1 C - F ) ; when hindwing
subcosta is fused only with the base of the
upper edge of the discal cell, it tends to di-
verge moderately from this edge (Fig.
18.1 D); crossvein sometimes occurring be-
tween the latter and the subcosta, or be-
tween the subcosta and the postdiscal sec-
tion of Rs (which is frequently approxi-
mated to, or fused with, a section of the sub-
costa); basally, labial palpus often provided
with a chaetosema-like organ (Lasiocam-
poidea) 6 Forewing with stem common to Rs3 and
6 Forewing usually with long areole between
branches of Rs (Fig. 18.1 C); male with sub-
costal retinaculum and well developed fre-
nulum; humeral vein absent from hindwing;
chaetosema-like organ absent from base of
labial palpus Anthelidae (in part)
Forewing without areole (Figs. 18.1 DF);
frenulum and retinaculum absent in both
sexes; one to several humeral veins usually
present in hindwing; segment 1 of labial pal-
pus often provided with ventral, or latero-
ventral, chaetosema-like organ
Lasiocampidae (in part)
Claude Lemaire & Jol Minet
7 Hindwing with Se + R approximated to
postdiscal section of Rs (Figs. 18.1 G and
18.5H-J) 8
postdiscal section of Rs (Figs. 18.1 H I,
10
8 Antennae always bi- or tripectinate in both
sexes; forewing with all branches of Rs
stalked together and with base of M2 equi-
distant from M l and M3 (Fig. 18.1 G), or
closer to Ml (Fig. 18.5 H) 9
Antennae rarely pectinate, often thick and
roughly filiform or davate (usually with a
lamellate ventral edge and a tapering, up-
turned apex); forewing with Rsl + Rs2
never stalked with Rs3 + Rs4, and with base
of M2 slightly closer to M3 than to M l
Sphingidae
9 Foreleg more or less shortened, its basitar-
sus (externally and/or ventrally) with two
stronger spines, which may be tooth-like
and remote from each other (enlarged spines
are sometimes also observable beyond the
basitarsus); frenulum and retinaculum al-
wing-coupling (Fig. 18.1 G) . . Lemoniidae
Foreleg never distinctly shortened, its tarsus
with normally developed spines; male wing-
coupling sometimes of retinaculo-frenate
type (Fig. 18.5 H) Brahmaeidae
10 Forewing stems Rsl + Rs2 and Rs3 + Rs4
separate, although closely parallel (Fig.
18.5 G) Carthaeidae
All branches of forewing Rs stalked to-
gether 11
11 Hindwing with frenulum (Figs. 18.11,
18.3 A - C ) or at least with the tubular base
of this organ (Fig. 18.1 H) 12
Frenulum completely absent (Figs. 18.5
A-F) 13
12 In the forewing, stem of Rs3 and Rs4 ap-
proximately as long as free section of Rs4
(Fig. 18.3 B), or distinctly longer than it
(Figs. 18.3 A, C); if Rs4 stalked with Rs3 for
a shorter distance, then compound eyes are
hairy, or the male antennae distally lack well
developed rami; discal cells never very small
Bombycidae
Rs4 invariably shorter than free section of
Rs4 (Figs. 18.1 H, I); eyes always "naked";
antennae with the flagellum bi- or tripecti-
nate to the apex; discal cells often very small
Eupterotidae
13 In both pairs of wings, M2 arises closer to
M3 than to Ml (Fig. 18.5E) . Endromidae
In fore- and hindwings, M2 arises midway
between M l and M3, or closer to Ml than
to M3 (see e. g. Fig. 18.5 A) 14
14 Forewing with four Rs branches and a very
long "anal loop" (Fig. 18.5 F); male and fe-
male antennae bipectinate to apex, with
The Bombycoidea and their Relatives
rami directed ventrad; foreleg tarsomere 4
with similar, "normal" spines in both sexes
Mirinidae
In forewing, Rs with at most three branches,
and anal loop moderately developed to
practically absent (Figs. 18.5 - D ) ; anten-
nae various, often quadripectinate, and fre-
quently with rami directed laterad or latero-
dorsad; tarsomere 4 of female foreleg usu-
ally with distal pair of tooth-like structures
(absent in the male) . . Saturniidae (in part)
MIMALLONOIDEA
Mimallonidae. Being monotypic, Mimallonoidea
are based on the same autapomorphies as Mi-
mallonidae (Minet 1994). Among these apomor-
phies are: (a) distal section of male antenna pro-
vided with short rami (distinctly shorter than
those of the proximal section) or even wholly de-
void of rami; (b) proboscis short to absent; (c)
segment 2 of labial palpus short, its length never
exceeding that of segment 1 ; (d) hindtibiae with-
out medial spurs; (e) hindwing with Se + R
strongly arched beyond wing base; (f) presence
of dorsal grooves with dentate edges on pupal
abdomen, along the anterior margins of seg-
ments A 2 - A 7 .
Adults stout-bodied, broad-winged, medium-
sized to fairly large (Figs. 18.2A-D). Male an-
tennae bipectinate, at least proximally; distal sec-
tion of flagcllum either serrate or provided with
shorter rami; transition between proximal and
distal flagellum sections usually abrupt (often
with rami shorter than those of distal section
itself); shaft dorsally scaled, distinctly thicker
near base; rami normally scaled outwards. Fe-
male antennae usually bipectinate, occasionally
serrate. Compound eyes naked. Ocelli and chae-
tosemata absent (or vestigial?). No distinct pilifer
bristles (they may be replaced by scales). Probos-
cis short (e. g. Alheita) to entirely absent. Maxil-
lary palpi with at most one segment. Labial palpi
usually small and 3-segmented, occasionally me-
dium-sized (e. g. Ulmara) or 2-segmented; seg-
ment 1 as long as, or more often longer than,
segment 2; segment 3 rarely long. Ventral process
of tegula ending in a rather sharp point. Tibiae
unspined (unlike the tarsi); mid- and hindtibiae
with a dorsal fringe of scales; epiphysis present,
often as long in female as in male; tibial spurs:
0-2-2. Arolium and pulvilli present; claws usually
simple, but with a ventral, subapical tooth in 7o-
lypida. Male occasionally with a subcostal reti-
naculum; frenulum present or absent. Spinarea
absent. Wings often dotted with large, petiolate,
subtriangular scales. Forewing venation (Figs.
18.1 A, B): Rsl and Rs2 long-stalked, Rs3 and
Rs4 stalked for a shorter distance; stem Rs3 +
Rs4 remote from Rsl + Rs2; Ml sometimes
stalked with Rs3 + Rs4 (e. g. Roelmana); M2
arising either midway between Ml and M3, or
323
closer to M3 than to M l ; stem of M vestigial
(not a tubular vein); CuP usually absent (present
distally, but thin, in Tolypida: see Fig. 1 in Pear-
son 1984); 1 A + 2 A with a basal fork. Hind-
wing: humeral area somewhat expanded (even
when frenulum is well developed); subcosta
strongly diverging from discal cell upper edge
(after a short connection near wing base); M2
usually arising midway between Ml and M3 or
closer to the latter (occasionally slightly closer to
Ml than to M3); no distinct M stem; M3 occa-
sionally stalked with CuAl, and sometimes also
with CuA2; CuP either present or absent; 3 A
well developed. Abdomen: median sclerite of
dorsum 1 never very broad, sometimes divided
in two; sternum 2 free, with very short apodemes
and without distinct venulae; sternum 7 some-
times distinctly modified in male, often divided
into three sclerites in female (then median, mem-
branous part bearing erect setae among a dense
group of scales); A8 variously modified in male,
sternum being e. g. divided or very narrow, or
even fused with vinculum (terminal tufts of long
scales may arise from dorsum 8 and/or from the
valvae). Male genitalia: no saccus; vinculum usu-
ally notched mesally (anteriorly); vinculum and
tegumen synscleritous; gnathos either entire (dis-
tally undivided or bifid), divided into a pair of
lateral arms, or vestigial to absent (e. g. in Al-
heita), phallus (often with a coecum) fused to the
juxta, that may be highly modified and/or associ-
ated with complex structures of various origins.
Female genitalia: corpus bursae variously devel-
oped, sometimes with an appendix bursae; os-
tium bursae on venter 8 or at its anterior border;
dorsum 8 often with a strong sclerotized pro-
jection (directed caudad); apophyses anteriores
of moderate length, apophyses posteriores
moderately long to very short.
Eggs of the flat type, elongate, subcylindrical
and often asymmetrical; chorion tough, trans-
parent or translucent, sometimes with well de-
fined longitudinal ribs and numerous, thin, trans-
verse ribs (e. g. Roelofa). Most setae clubbed in
first instar larvae. Mature larva: no secondary
setae, except in abdominal SV groups (e. g. Stehr
et al. 1987); head almost prognathous, with PI
and A3 sometimes long and spatulate; on T l ,
dorsal shield wide, well sclerotized, and coxae
free, though approximate; T2 and T3 with one
or two pairs of ventral, toothed protuberances
caudad of the coxae; LI and L2 approximate on
AlA8; last abdominal segments tending to de-
velop a sclerotized, oblique, dorsal plate (Forbes
1923: Fig. 412); ventral proleg crochets biordi-
nal, arranged in transverse, oval rings (anal pro-
legs similar, but with uniordinal posterior bor-
ders). Pupa: antennae and proboscis short or
rather short; maxillary palpi concealed; fore-
coxae and femora often exposed; A2A7 show-
ing characteristic grooves (with dentate edges)
along their anterior margins (dorsally); no genu-
324
ine cremaster (small teeth may occur on the last
two or three segments).
Adults nocturnal. At rest, the forewings are
held remote from the abdomen and thus reveal
part of the hindwings; the antennae are hidden
under the wings and the end of the abdomen
tends to be flexed upwards. In Mimallo, when an
imago is at rest on a branch, its wings are
strongly deflexed, the forewing inner margin be-
ing held perpendicular to the abdomen, and the
hindwing dorsal surface being almost wholly ex-
posed.
The larvae are often brightly coloured. When
young, they usually live in a folded, perforated
leaf, or in a silken nest between two leaves. Older
larvae construct portable cases of silk, plant frag-
ments and often frass pellets (e. g. Newman 1854,
Schreiter 1943, Lima 1950, Dinther 1960, and
Stehr et al. 1987). These cases vary from irregu-
larly shaped (e. g. Mimallo) to spindle-shaped
and extremely tough (e. g. Menevia). Each case
always has two openings, which can be closed
simultaneously by the head and the sclerotized,
postero-abdominal plate of the larva. Pupation
occurs in the last larval case, after closure of the
openings. Hostplants belong to various families:
e. g. Anacardiaceae, Clusiaceae, Combretaceae,
Fagaceae, Melastomataceae, Myrtaceae and
Rubiaceae (see in particular Silva et al. 1968).
The family was divided into two subfamilies
by Schaus (1928), viz. Mimalloninae and Laco-
sominae (or Lacosomatinae). However, as
pointed out by Pearson (1951) and Franclemont
(1973), neither of these represents a monophy-
letic lineage. Mimallonidae contain approxi-
mately 30 genera and 200 described species
(Gaede 1931, Becker 1996). They are mostly neo-
tropical but four species occur in the nearctic re-
gion. Mimallo amilia is recorded as a pest of
guava but may also feed on other Myrtaceae.
Main genera include Trogoptera (Fig. 18.2 A),
Lacosoma, Druentica, Bedosia, Alheita, Psycho-
campa and Cicinnus.
LASIOCAMPOIDEA
Anthelidae. If one accepts the sister group rela-
tionship Anthelidae/Lasiocampidae (Minet 1991),
four imaginai autapomorphies can be assigned
to the anthelid ground plan: (a) forewing with an
oblique fold or cross-bar between R and Rsl,
near the apex of the areole; (b) forewing without
spinarea; (c) hindwing with base of M2 distinctly
closer to M3 than to M l ; (d) Al of male with
postspiracular conical projections of tergal origin.
Fairly small to very large moths, with broad
wings and a stout, or rather stout, body (Figs.
18.2E, F and Q); females occasionally micro-
pterous (most species of Pterolocera). Male an-
tennae bipectinate to the tip, or sometimes even
tripectinate (when the shaft bears a ventral row
of fairly long teeth); rami forming lateroventral
Claude Lemaire & Jol Minet
rows that are never clearly enlarged proximally;
shaft densely scaled. Antennae of females either
similar to those of males (though with shorter
rami) or quite different (unipectinate, dentate or
nearly filiform). Compound eyes rarely with con-
spicuous interommatidial setae (but present in,
e. g., Chelepteryx chalepteryx). Distinct ocelli
and chaetosemata absent. Pilifer bristles well de-
veloped in Munychryia, reduced to absent in
other genera. Proboscis absent or very short, ex-
cept in Munychryia, where it is long, unsealed,
and has carnate sensilla. Maxillary palpi vestig-
ial. Labial palpi fairly long to very short, either
ascending, porrect or drooping (usually longer in
male than in female); segment 1 without chaeto-
sema-like organ. Ventral process of tegula some-
times ending in a rather sharp point. Tibiae usu-
ally with several spines (absent in Munychryia);
epiphysis present in males, reduced or absent in
females; tibial spurs: 0-2-4, 0-2-3 or 0-2-2. Tarsi
distinctly spined. Arolium present; pulvilli with a
dorsal lobe (or instead fairly long setiform
outgrowths); claws simple (i. e. without a
ventral, medial tooth). Wing-coupling reti-
naculo-frenate in the male, amplexiform in the
female (preserving only the base of the frenu-
lum). Spinarea absent. Forewing venation (Fig.
18.1 C): usually with a long areole from which
arise the distal sections of Rsl, Rs2, Rs3 and Rs4
(in Gephyroneura, this areole is small because
Rsl + Rs2 and Rs3 + Rs4 are coalescent proxi-
mally); near apex of areole, extra cross-bars
often present (one between Rsl and Rs2, a se-
cond usually incomplete or replaced by a fold
- between R and Rsl); M2 closer to M3 than to
M l ; M3, CuAl and CuA2 distinctly separate; no
tubular CuP, 1 A + 2 A with basal fork. Hind-
wing: subcosta connected to discal cell upper
edge by a crossvein (R) that is sometimes very
short or extremely thin; Rs and M l often short-
stalked; M2 arising closer to M3 than to M l ; no
tubular CuP; 3 A present. Abdomen: lateral bars
of dorsum 1 with posterolateral extensions
which, in the male, end in conical projections
caudad of Al spiracles; sternum 2 free from
terga, with short anterior apodemes; no well de-
veloped venulae; male sternum 8 plate-like, al-
though sometimes reduced and often only lightly
sclerotized. Male genitalia: tegumen and vincu-
lum synscleritous; valvae rather well developed;
gnathos either present (Chelepteryx), reduced or
entirely absent; uncus either simple, truncate or
bilobed; phallus usually with anterior coecum.
Female genitalia: bursa copulatrix never very
large; signum sometimes present; ostium bursae
lying in the anterior or median area of venter
8; apophyses short to moderately long; papillae
anales either small (Munychryia) or well devel-
oped.
Egg of the flat type, roughly ovoid but slightly
flattened, with a broader end which bears the
micropyle. Chorion smooth, occasionally trans-
The Bombycoidea and their Relatives 325

Fig. 18.1. Wing venation of Mimallonidae (A, B), Anthelidae (C), Lasiocampidae ( D - F ) , Lemoniidae (G) and
Eupterotidae (H, I): A, Trogoptera notata; , Cicinnus anysia\ C, Anthela ocellata-, D, Chondrostega tingitana <J;
E, Eriogaster lanestris; F, Napta Serratilinea V; G, Lemonia sardanapalus; H, Hibrildes norax; I, Phiala fuscodorsata.
Scales = 5 mm. sp: spinarea (G. Hodebert).

lucent, sometimes with dark markings. First in-
star larva with few (Munychryiinae) or many
(Anthelinae) secondary setae. Mature larva with
many secondary setae, which are minute and
spatulate (scattered over the integument) in Mu-
nychryia, mostly barbed, plumose or spine-like
in Anthelinae (the larva having then well defined
Verrucae); in most (all?) Anthelinae, one pair of
dorsal Verrucae on A l , two pairs (Dl, D2) on
A2A7, and a single middorsal verruca on A8;
326
proleg crochets in mesoseries, usually bi- or
multiordinal, but uniordinal in Munychryia
(Common & McFarland 1970); anal prolegs pro-
duced posteriorly (or even acutely pointed: Mu-
nychryia). Pupa (see e. g. Common 1990): galeae
usually short, but long (reaching forewing api-
ces) in Munychryia\ maxillary palpi and forefem-
ora concealed; labial palpi only represented by a
small area caudad of the labrum; abdomen
smooth; AIO usually with a number of crem-
astral setae (either hooked or straight and spine-
like). Cocoon opaque, often incorporating larval
setae; occasionally subterranean.
The adults are nocturnal, except for the day-
flying males of Anthela connexa. In general, fe-
males are more sedentary than males. As far as is
known, the larvae never have gregarious habits.
Many of them are urticating.
The family was erected by Turner (1920), and
revised by him in 1921. It was long regarded as
a subfamily within the Lymantriidae, and even
when considered an independent family (e. g. by
Hulstaert 1928), Anthelidae were often thought
to be closely related to Lymantriidae. Common
(1970) first clearly recognized the bombycoid
(s. I. ) affinities of Anthelidae. The family in-
cludes about 75 described species in 8 genera and
2 subfamilies (Common & McFarland 1970).
The subfamilies can be distinguished as follows
(imagines):
Key to the subfamilies
1 Proboscis often long; forewing with a com-
plete crossvein between R and Rsl
Munychryiinae
Proboscis always reduced or absent; fore-
wing at most with an incomplete crossvein
between R and Rsl, usually with a mere
transverse fold (Fig. 18.1 C) . . . Anthelinae
Munychryiinae are a small Australian group,
with 3 species placed in two genera: Munychryia
(Fig. 18.2 E) and Gephyroneura. . The adults do
not exceed 37 mm in wingspan. Munychryia lar-
vae feed on Casuarina (Casuarinaceae) and are
green with white longitudinal stripes, thus hav-
ing a cryptic appearance among hostplant
branchlets.
Anthelinae are restricted to Australia and New
Guinea. Adult wingspan varies from moderately
small to very large. The hostplants belong to var-
ious families, but are mostly Poaceae, Fabaceae
(especially Acacia) and Myrtaceae (especially Eu-
calyptus). Chelepteryx (Fig. 18.2 Q) contains two
very large species, the tibial spurs of which
number 0-2-4, contrary to the condition found
in the other Anthelinae (0-2-2). Omphaliodes, Na-
taxa and Chenuala are monotypic or small gen-
era, the last two including sexually dimorphic
species. Anthela (Fig. 18.2 F) is the largest genus,
with about 60 described species. In Pterolocera,
Claude Lemaire & Jol Minet
the females are usually micropterous (see, e. g.,
Common 1990 and Heppner 1991).
Lasiocampidae. This family can be recognized by
the following autapomorphies: (a) pilifer bristles
absent; (b) forewing with the distal section of R
distinctly approximated to Sc; (c) no areole; (d)
CuA2 of forewing arising closer to the base of
the discal cell than to the origin of M3; (e) am-
plexiform wing-coupling; (f) pupal proboscis
short to strongly reduced. A secondary loss of
(b) only occurs in a few specialized taxa, such as
Lasiocampa quercus.
Rather small to very large moths, with broad
wings and stout bodies (Figs. 18.2G-N); fe-
males occasionally micropterous (Heppner 1991;
Sattler 1991). Male antennae bipectinate to the
tip; shaft densely scaled, tending to be distinctly
stronger proximally; rami directed ventrad or
latero-ventrad, and sometimes scaled outwards;
external row of rami basally enlarged in many
genera; no midventral row of teeth. Female an-
tennae nearly always bipectinate (though with
shorter rami than in male). Distinct ocelli and
chaetosemata absent. Compound eyes occasion-
ally fairly small (in day-flying males), often with
long interommatidial setae. Frontoclypeus some-
times with conspicuous central protrusion (e. g.
Chondrostega). No pilifer bristles. Maxillary
palpi reduced to absent, proboscis short (e. g.
Odonestis) to entirely absent. Labial palpi short
to rather long, usually 3-segmented (occasionally
2-segmented) and either drooping, porrect or
slightly ascending; in all genera except Chiono-
psyche, segment 1 has a ventral or lateroventral,
chaetosema-like, group of setae. Ventral arms of
laterocervicalia either free and very close to pro-
basisternum (Chondrosteginae, Poecilocampi-
nae), or more or less fused mesally and remote
from basisternum (Macromphaliinae, Lasiocam-
pinae). Ventral process of tegula rarely ending
in a sharp point. Mesanepisterna large. Narrow
median strip of metascutum nearly always deeply
concave. Foretibia sometimes with one or two
strong apical teeth (tibia then often shortened;
teeth sometimes also present on other legs: e. g.
in Chondrostega), epiphysis often well developed
in male, always reduced or absent in female; tib-
ial spurs: 0-2-2 or, occasionally, 0-0-0 (e. g. Poe-
cilocampa) or 0-2-4 (Chionopsyche only). Tarsi
ventrally spinose (spines may also occur on tib-
iae). Pretarsus: claws simple; arolium present;
pulvilli entire or slightly bifid. Frenulum and
retinaculum absent in both sexes. Spinarea nor-
mally well developed, rarely absent (Chondros-
tega). Forewing venation (Figs. 18.1 D - F ) : dis-
tal section of R usually closely approximated to
Sc; Rsl and Rs2 stalked or, rarely, entirely fused;
Rs4 often stalked with Ml (not stalked in lasio-
campid ground plan: Ml free in the primitive
subfamily Chionopsychinae); Rs3 sometimes
free; discal cell closed or open; M2 closer to M3
The Bombycoidea and their Relatives
than to Ml (M2 and M3 may even be stalked);
CuAl usually remote from M3, but stalked with
it in Chionopsyche\ CuA2 arising from basal half
of discal cell lower edge; CuP sometimes repre-
sented (distally) by a strong, sclerotized fold
(e.g. Euthrix); 1A + 2A forked or simple.
Hindwing: humeral area expanded, usually rein-
forced by one to several humeral veins (especially
numerous in Gastropacha), if single, humeral
vein then rather long and arising from base of
Se; subcosta sometimes fused with a basal or
subbasal section of discal cell upper edge, often
approximated to, or partly fused with, postdiscal
section of Rs, or even connected to Rs by a
crossvein (R); "humeral cell" variously devel-
oped (i. e. the cell distally closed by crossvein R:
"cellule accessoire" of Viette 1965); discal cell oc-
casionally open; M2 and M3 usually approxi-
mate, connate or stalked (only remote and paral-
lel in Chionopsychinae); CuAl free or stalked;
CuP absent or represented by sclerotized fold;
3 A relatively long. Abdomen: dorsum 1 with
rather narrow lateral bars; sternum 2 fairly
broad, with moderately developed apodemes and
short anterolateral processes (which do not reach
dorsum 1); no venulae; sternum 8 of male usually
plate-like and variously modified; in males of
Maiacosoma, sternum 8 U-shaped, ending in
pointed arms (see Kuznetzov & Stekolnikov
1985), sternum 7 more or less modified and ter-
gum 8 divided into a pair of dentate plates that
may represent accessory claspers (Franclemont
1973). Male genitalia: vinculum and tegumen not
always synscleritous; in many Lasiocampinae,
median part of vinculum with a cubile (La-
jonquire 1968), i. e. a ventral - often bifid -
sclerotized extension that pertains to interseg-
mental membrane 89; valvae well developed to
more or less reduced (often mainly consisting of
a tubular or sickle-shaped process); gnathos en-
tire only in primitive taxa (e. g. Poecilocampinae:
see Chistyakov & Zolotuhin 1994); uncus fre-
quently absent; socii usually well developed;
phallus often sharply pointed caudad and closely
associated or joined with juxta. Female genitalia:
corpus bursae rather small, occasionally with a
signum (e. g. Euthrix), ductus bursae short, often
forming a wide antrum; apophyses moderately
long to very short (anterior pair sometimes en-
tirely absent); corethrogyne sometimes present.
Eggs of "flat" type (easily mistaken for up-
right eggs when set side by side in a cylindrical
layer encircling a twig: Maiacosoma), ovoid or
roughly subcylindrical with rounded extremities
and, often, a flattened upper surface; chorion
tough, smooth, often distinctively patterned.
Eggs laid singly or in groups (sometimes pro-
tected by hair-scales), or even dropped at ran-
dom, in flight (Lasiocampa). First instar larva
with secondary setae, mostly located on Verru-
cae. Mature larva densely covered with second-
ary setae of varying lengths, usually hair-like and
327
never barbed or plumose (there may be a number
of modified setae: e. g. spine-like, spatulate,
scale-like or fan-shaped); setae often urticating;
middorsal tufts sometimes present; Verrucae usu-
ally indistinct (but conspicuous in, e. g., Chon-
dros tega), scoli and flexible filaments quite un-
common; SV and L often forming prominent
lobes on Tl; SV groups of T 2 - T 3 and abdomi-
nal L3 groups frequently also forming projecting
lappets with long setae (hence the common name
of many Lasiocampidae: "lappet moths"); on T2
and T3, D1 and D2 groups separate, D2 lying
caudad of D l ; T2 and/or T3 often with a col-
oured transverse band (D2) which is usually
groove-like, erectile and provided with modified
setae; on AlA8, LI and L2 usually separate
(when visible), except on Al of Chondrostega; on
A8, D1 groups sometimes slightly prominent
(they may form a single middorsal protuber-
ance); prolegs typically rather long, with SV area
partly membranous (posteriorly or laterally);
proleg crochets usually in biordinal mesoseries;
venter 10 posteriorly ending in a conical pro-
jection (Stehr et al. 1987). Pupa: covered with
many, very short, secondary setae (except on ap-
pendages); antennae sometimes long (e. g. Poeci-
locampa and Eriogaster), but usually fairly short
and often strongly bent near their middle; maxil-
lary palpi concealed; proboscis short, or even re-
duced to pair of small, separate lobes; labial
palpi often largely exposed (e. g. Nakamura &
Yamamoto 1990: Fig. 17); forefemora concealed
when galeae broadly meet on the meson; abdo-
men without dorsal teeth, occasionally with
transverse ridges (Poecilocampa); anterior angles
of Al very obtuse to nearly absent; end of abdo-
men usually broadly rounded and provided with
hooklets or strong, straight setae (in Euglyphis,
there is a conical cremaster, with many hook-
lets). Cocoon well developed (occasionally sub-
terranean); its silk often mixed with larval setae.
Most adults come to light at night, but in cer-
tain species, the males are diurnal and fly rapidly
in sunshine. At rest, the forewings are held roof-
wise and the hindwing humeral/subcostal area is
frequently uncovered; the antennae are con-
cealed by the wings and the vestiture of the
thorax.
Lasiocampidae include about 1500 species, re-
ferred to about 150 genera (Munroe 1982). The
family occurs almost worldwide, but is best rep-
resented in tropical countries. It was divided into
seven subfamilies by Aurivillius (1927): Chon-
drosteginae, Malacosomatinae, Archaeopachi-
nae, Gastropachinae, Chionopsychinae, Lasio-
campinae and Gonometinae. This classification
was accepted by Lajonquire (1972), but partly
modified by Franclemont (1973), who erected a
new subfamily, Macromphaliinae, and regarded
the Malacosomatini s. str. as a tribe of the Lasio-
campinae. Taking account of observations made
by Franclemont (1973) and Holloway (1987), I
328 Claude Lemaire & Jol Minet

Fig. 18.2. Adults of Mimallonidae ( A D ) , Anthelidae (E, F, Q), Lasiocampidae ( G - N ) , Lemoniidae (O, R) and
Eupterotidae (P, S): A, Trogoptera erosa; B, Ulmara rotunda; C, Tolypida amaryllis, D, Roelmana maloba; E,
Munychryia senicula; F, Alititela connexa; G, Chionopsyche montana; H, Chondrostega tingitana <?; I, Poecilocampa
alpina-, J, Euglyphis marna; , Malacosoma castrensis; L, Eriogaster rimicela. M, Borocera marginepunctata ; N,
ditto 9; O, Lemonia sardanapalus; P, Hibrildes sp. ; Q, Chelepleryx collesi; R, Sabalia tippelskirchi; S, Janomima
karschi. Scales A - P = 5 mm; Q S = 10 mm (J. Minet).

propose to treat Gastropachinae and Gonometi- cation being based on the present work). More-
nae as synonyms of Lasiocampinae (syn. rev.) over, I examined the holotype of Archaeopacha
and to reinstate Poecilocampinae (Tutt 1900), obsoleta, the only member of so-called "Archae-
but in a new sense (Schintlmeister's 1996 classifi- opachinae". From a study of its morphology
The Bombycoidea and their Relatives
(wing venation in particular), it became obvious
that this taxon did not belong to Lasiocampidae,
but should be placed in Cossidae: Metarbelinae.
I therefore here transfer Archaeopacha to this
group and retain only five subfamilies in Lasio-
campidae, viz. Chionopsychinae, Chondrostegi-
nae, Poecilocampinae, Macromphaliinae and
Lasiocampinae. The main autapomorphy of
Poecilocampinae is the crown-shaped extremity
of their frontal protuberance; Lasiocampinae
can be delimited by the absence or strong reduc-
tion of both uncus and gnathos. The five sub-
families are distinguishable using the following
key:
Key to the subfamilies
1 Labial palpi without chaetosema-like or-
gans; tibial spurs fairly long, numbering 0-
2-4; in the hindwing, vein M2 arising about
midway between M l and M3
Chionopsychinae
Basal segment of labial palpi with a chaeto-
sema-like group of setae; tibial spurs num-
bering 0-2-2 or 0-0-0; in the hindwing, base
of M2 distinctly closer to M3 than to M l
(Figs. 18.1 D - F ) 2 the presence of a "cubile" in the male genitalia
2 Hindwing Se + R neither partly fused with
the postdiscal section of Rs, nor connected
to it by a crossvein; male sternum 8 with
lateral edges angulate, strongly convex; un-
cus well developed or replaced by a pair of
long, sclerotized arms . . . Macromphaliinae
Different from above 3 ara). In Malacosoma, the larvae often live gre-
3 Frontoclypeus always with a median protu-
berance; uncus and gnathos well developed
4 worldwide (key to most oriental genera in Hollo-
Frontal protuberance often absent; both un-
cus and gnathos absent or strongly reduced
Lasiocampinae
4 Hindwing with Se + R never partly fused
with postdiscal section of Rs (Fig. 18.1 D);
female moth often micropterous
Chondrosteginae
Hindwing with Se + R partly fused with
both upper edge of closed discal cell and
postdiscal section of vein Rs; female moth
never micropterous . . . . Poecilocampinae
Chionopsychinae contain only one African ge-
nus, Chionopsyche (Fig. 18.2G), with two de-
scribed species that resemble certain Eupteroti-
dae. Nothing is known of their early stages and
their genitalia are undescribed.
Chondrosteginae inhabit North Africa, South
Africa, and the Iberian and Arabian peninsulas.
Chondrostegoides differs from Chondrostega (Fig.
18.2H) in its fully winged females. The known
larvae feed on low-growing plants of various
families: e. g. Asteraceae, Brassicaceae, Fabaceae
and Poaceae (Rougeot & Viette 1978).
329
Poecilocampinae comprise the palaearctic Poe-
cilocampa (Fig. 18.21) and Trichiura. Adults are
medium-sized to rather small, lack the epiphysis
and may also be devoid of tibial spurs. Larvae
feed chiefly on trees and shrubs, mainly Betula-
ceae, Fagaceae, Rosaceae, Salicaceae and Tilia-
ceae (Freina & Witt 1987).
Macromphaliinae occur only in the New World
and contain about 15 genera (Franclemont 1973,
Becker & Heppner 1996). In North America, 7o-
lype is the most speciose genus: the adults are
usually greyish, and the flattened larvae -
known as "lappet caterpillars" live either on
Pinaceae or various deciduous trees: Betulaceae,
Fagaceae, Rosaceae, Salicaceae, etc. (hostplants
are also very diverse for the neotropical species
of Tolype: Silva et al. 1968). The main neotropi-
cal genus, Euglyphis (Fig. 18.2 J), is extremely
speciose; its larvae often feed on Lauraceae or
Myrtaceae (Silva et al. 1968). Other important
genera are Artace, Macromphalia, Nesara and
Titya.
Lasiocampinae are by far the most diverse sub-
family, and should be divided into several tribes,
such as Lachneini (Fig. 18.2L), Malacosomatini
(Fig. 18.2 K), Gastropachini and Lasiocampini
(Figs. 18.2 M, N). The latter can be diagnosed by
and thus must include those genera usually re-
ferred to "Gonometinae". The tribal classifica-
tion of the Lasiocampinae needs further re-
search. In this subfamily, male moths often have
shortened hindwings; occasionally their wings
are nearly transparent (most species of Gendu-
gariously in conspicuous silken tents ("tent ca-
terpillars"). Lasiocampinae occur practically
way & Bender 1990). Hostplants are very di-
verse. The cocoons are sometimes tough and
ovoid (in "eggars"), and their silk is occasionally
used for textiles (e. g. certain species of Borocera:
see Lajonquire 1972 and Peigler 1993). Main
genera: Anadiasa, Arguda, Beralade, Bomby-
copsis, Borocera (Figs. 18.2 M, N), Chrysopsyche,
Closterothrix, Cyclophragma, Dendrolimus, Dig-
glesia, Eriogaster (Fig. 18.2 L), Euthrix, Gastro-
pacha, Gastroplakaeis, Gloveria, Gonometa, Ku-
nugia, Lasiocampa, Lebeda, Lechriolepis, Leipox-
ais, Malacosoma (Fig. 18.2K), Pachymeta,
Pachymetana, Pachypasa, Philotherma, Phoe-
nicladocera, Phyllodesma, Porela, Prorifrons,
Pseudometa, Sena, Streblote and Trabala.
BOMBYCOIDEA
Eupterotidae. As now conceived (Minet 1994),
Eupterotidae include Hibrildes, a genus pre-
viously placed either in "Pterothysanidae" (e. g.
Zerny & Beier 1936) or in a family of its own
(Berger 1958; Minet 1983: 204). The following
three autapomorphies can be ascribed to the
330
ground plan of the Eupterotidae: (a) metathora-
cic distitarsus of female with a conspicuous, mid-
ventral row of spines; (b) "notai incisions" (sensu
Sharplin 1963) elongate and dorsally pointed on
T2; (c) mesal part of uncus distinctly reduced.
Fairly small to very large moths, with broad
wings often displaying several transverse wavy
lines (Figs. 18.2P, S). Male antennae bipectinate
to the apex or, occasionally (in Melanothrix),
more or less tripectinate, i. e. with a ventral row
of elongate teeth; rami directed latero-ventrad;
"shaft" scaled dorsally. Female antennae either
bipectinate, tripectinate or roughly serrate; rami
rarely as long as those of male. Compound eyes
"naked". Distinct ocelli and chaetosemata ab-
sent. Ventral part of frontoclypeus often pro-
duced. Pilifer bristles absent or replaced by
scales. Proboscis very short to absent; no well
developed maxillary palpi. Labial palpi usually
3-segmented (occasionally with only one or two
segments), varying from moderately long and as-
cending to very short and drooping; segment 3
always short. Hindcoxa: meron delimited ante-
riorly by a long suture. Fore- and/or midtibiae
sometimes shortened (if so usually ending in
pointed teeth). Epiphysis often well developed in
male, varying in female from fairly long (though
shorter than in male) to entirely absent; epiphysis
absent from both sexes in a few taxa. Tibial spurs
(long to short): 0-2-4 or 0-2-2. Spines absent
from tibiae, frequently absent from tarsi; meta-
thoracic distitarsus of female usually with a me-
diolongitudinal, ventral row of spines (lost or
vestigial in, e. g., Dreata and Palirisa); male lack-
ing this row of spines. Arolium well developed;
pulvilli usually with irregular dorsal edge; claws
with simple or serrulate ventral edge. Male
usually with subcostal retinaculum and fairly
long frenulum; female often with rather short,
multiple frenulum; frenular bristles occasionally
absent in both sexes (tubular base of frenulum
always preserved). Spinarea absent. Wing vesti-
ture: most lamellar scales with acute teeth that
may be very long (Meli 1930: Fig. 33). Forewing
venation (Figs. 18.1 H, I): Sc occasionally fused
with a section of R; branches of Rs stalked to-
gether (they may also be stalked with R); Rsl
and Rs2 either long-stalked (Fig. 18.11) or en-
tirely fused (Fig. 18.1 H); stem common to Rs3
and Rs4 always shorter than free section of Rs4;
M2 with a varying position; discal cell often very
short; tubular CuP lacking; fork of 1A + 2 A
occasionally vestigial. Hindwing: subcosta some-
times adjacent to, or fused with, a short section
of discal cell upper edge, but usually only con-
nected to it by a crossvein (R); Rs and M l either
separate, connate or stalked; discal cell often
short; tubular CuP lacking; 3 A variously devel-
oped. Abdomen: terga usually with numerous
spine-like scales (sometimes also visible on
sterna); lateral bars of dorsum 1 frequently en-
larged in the rear; sternum 2 with short apo-
Claude Lemaire & Jol Minet
demes, short anterior processes and often, in
male, posterolateral arms bearing eversible, an-
droconial pouches; venulae sometimes well de-
veloped; anterior angles of sternum 3 often with
narrow apodemes (directed cephalad); male ster-
num 8 plate-like. Male genitalia often reduced in
size (Forbes 1955), frequently with a long saccus;
vinculum and tegumen not always synscleritous;
valva often cleft, sometimes fused to vinculum;
gnathos entire to wholly absent; uncus primarily
reduced on the meson, usually flanked by a pair
of strong projections (socii?); phallus free or
fused to juxta. Sternum 7 of female varying from
large and well sclerotized to almost entirely
membranous; corpus bursae often unarmed,
tending to be small; ostium bursae on venter 8;
apophyses moderately long to very short; co-
rethrogyne sometimes present.
Egg of upright type, usually hemispherical or
dome-shaped (but nearly disc-like, with a flat cir-
cular margin, in Hibrildes); chorion fairly
smooth. First instar larva with verrucae/verri-
cules; setae LI and L2 remote from each other
on A 1 - A 8 ; each ventral proleg with two sepa-
rate, vertical, setose sclerites (SV). Last instar
larva: head with numerous secondary setae; tho-
rax and abdomen densely covered with long to
very long secondary setae, many of which arise
from Verrucae or verricules (e. g. Horsfield &
Moore 1859; Moore 1883; Froggatt 1923: 62;
Meli 1930; Holloway 1987; Sugi et al. 1987;
Common 1990); barbed and/or spine-like setae
sometimes present; T1 coxae firmly fused; proleg
crochets biordinal, in mesoseries or, occasionally,
in mesal penellipses. Pupa often with a stout,
rounded, anterior half (see e. g. Meli 1930; Hol-
loway 1987; Common 1990); maxillary palpi, la-
bial palpi and forefemora concealed; proboscis
sometimes very short (e. g. Acrojana), but usu-
ally reaching apices of antennae, or even extend-
ing slightly beyond them (e. g. Nakamura 1970);
forewings meeting broadly mesally; metanotum
sometimes with conspicuous sculpturing; end of
abdomen either rounded (with or without anal
spines) or provided with a cremaster (usually
short, with spines or hooklets). Pupation in the
earth or in a cocoon of silk mixed with larval
setae (often spun in leaf litter).
The adults are usually nocturnal, but diurnal
habits sometimes occur, especially in females of
Hibrildes and Melanothrix. At rest, the wings are
often broadly outspread, although sometimes
differently set: in Pseudojana, for instance, they
are directed backwards, forming a flattened roof
(Meli 1930: Pl. XI); the antennae are concealed
under the wings. The larvae are sometimes po-
lyphagous, and Nisaga simplex is regarded as a
pest of rice in India (Holloway et al. 1987). Re-
corded hostplants belong to a wide range of
families: e. g. Acanthaceae, Boraginaceae, Capri-
foliaceae, Fabaceae, Hamamelidaceae, Myrta-
ceae, Oleaceae, Pinaceae, Poaceae and Rubiaceae
The Bombycoidea and their Relatives
(see notably Meli 1930, Pinhey 1975, Barlow
1982, Beutelspacher B. 1983, Holloway 1987,
Sugi et al. 1987 and Common 1990).
Worldwide, there are almost 300 described
species referred to about 50 genera. The present
classification of the Eupterotidae is largely based
on Forbes (1955), with a few changes recently
proposed by Minet (1994), namely: inclusion of
Hibrildes in the family, due in particular to its
tarsal morphology; exclusion of Apatelodinae
(Bombycidae); raising of Forbes' tribes to sub-
family status; and assignment of Cotana and
relatives to Panacelinae, mainly on the basis of a
few synapomorphies of the wing venation. Five
subfamilies are accepted. The following key can
be used to determine the adults:
Key to the subfamilies
1 Hindwing discocellular M2M3 straight,
usually shorter than discocellular Ml M2
(Fig. 18.1 H), at most equal to it 2 Stenoglene and relatives (Minet 1990).
Hindwing discocellular M2M3 longer
than discocellular Ml M2 (which may even
disappear) and often distinctly bent or an-
gled (Fig. 18.11) 4 spurs number 0-2-2. The female has a distinct
2 Hindtibia with two pairs of spurs . . Janinae
Hindtibia with one pair of spurs 3 and Panacela. The latter is unique in having M3
3 Labial palpi short, at most 2-segmented; dis-
cal cells well developed, that of forewing ex-
ceeding one third length of costa (Fig.
18.1 H); frenular bristles absent from both
sexes; hindwing 3 A absent or vestigial . . .
Hibrildinae
Labial palpi 2- or 3-segmented, often fairly
long; discal cells short, that of forewing less
than one third of costa length; male with
subcostal retinaculum and well developed
frenulum; female frenulum with or without
bristles; hindwing with a rather long 3 A . .
Eupterotinae (in part)
4 Midtibia very short, without apical tooth,
but with long, strong, claw-like spurs . . . .
Tissanginae
Midtibia sometimes shortened and/or pro-
vided with long spurs, but these always
straight, scaled for most of their length,
never claw-like 5 saga (see in particular Forbes 1955). The largest
5 In forewing, stem Rsl + Rs2 has a sinuous
course, being distinctly bent towards R be-
yond the origin of the free section of Rs3
(short crossvein often occurring between R
and Rsl + Rs2; Rsl and Rs2 sometimes en-
tirely fused); in both pairs of wings, Ml
stalked with Rs, and M2 and Rs + M l
either connate or stalked (M2 perhaps occa-
sionally absent) Panacelinae
In forewing, stem Rsl 4- Rs2 not sinuous
(Fig. 18.11) or only slightly so (Rsl and Rs2
often entirely fused); in both pairs of wings,
origin of M2 separated from Ml, or from
Rs + M l Eupterotinae (in part)
331
Tissanginae and Hibrildinae are monotypic,
afrotropical taxa. Their larvae and hostplants are
unknown. In Tissanga, the moths have large
wings, mainly brownish, with contrasting, white
or yellow veins. In Hibrildes (Fig. 18.2 P), the
males resemble the saturniid Pseudaphelia, while
females mimic certain species of nymphalid gen-
era Acraea and Danaus (e. g. Janse 1937, 1948;
Laithwaite et al. 1975; Pinhey 1975).
Janinae are confined to tropical Africa (about
90 species) and Madagascar (1 species). In the
male genitalia, the valva has its costa and saccu-
lus separated by an elongate, rather unsclero-
tized area, and/or by a deep cleft; the phallus is
not fused to the juxta. The main genera, among
the 12 recognized, are Hoplojana, Jana and Ste-
noglene (genus revised by Dall'Asta & Poncin in
1980). Pterocerota Hampson, long placed in
Callidulidae-Pterothysaninae, is indisputably a
member of Janinae, its forewing venation show-
ing the same arrangement of Sc, R and Rs as in
Panacelinae are a small group (with about 25
species) from the Australian region. The adults
are usually sexually dimorphic, and their tibial
corethrogyne. The main two genera are Cotana
and CuAl stalked in both pairs of wings; the
fairly small adults are bombycid-like; the larvae
possess urticating setae, and live gregariously in
silken bags, spun in the branches of various
trees; they are sometimes of economic impor-
tance (see Froggatt 1923: 61-63; Common 1990;
Nielsen & Common 1991).
Eupterotinae (Fig. 18.2 S) are the largest sub-
family (about 160 species), but may be paraphy-
letic. They occur mainly in the oriental and afro-
tropical regions. However, they reach Australia
(1 species), palaearctic Asia (a few genera) and
there are two endemic genera in Central America
{Preptos, Neopreptos). The tibial spurs may
number 0-2-4 or 0-2-2, and the phallus may be
free or fused to the juxta. Indisputable Eupteroti-
nae include those taxa in which the valva has a
striate area on the sacculus: e. g. Phiala, Striph-
nopteryx, Dreata, Eupterote, Sarmalia and Ni-
genera are Phiala (afrotropical) and Eupterote
(mostly oriental; see e. g. Holloway 1987, Ki-
shida 1994). The larvae of the latter are often
polyphagous and are occasionally minor pests
(Kaishoven et al. 1951; Holloway 1987).
Bombycidae. The "Apatelodidae" of recent au-
thors proving to be diphyletic, the older broad
concept of Bombycidae, inclusive of Apatelodi-
nae, is tentatively reinstated, with an additional
subfamily, Phiditiinae (Minet 1994). Allowing for
a few assumptions about the phylogeny of Bom-
bycoidea, the following traits can be regarded as
possible autapomorphies of the Bombycidae: (a)
332
5 mm (G. Hodebert).
lower part of the mesepimeron strongly convex,
in cross-section, along the upper edge of the
meron; (b) metacoxa with the internal ridge of
the merocosta (or "meral suture") more or less
shortened (ridge length never exceeding three-
quarters length of coxa); (c) venulae vestigial to
absent; (d) pupa with short galeae that fail to
reach foreleg apices.
Adults broad-winged, small to fairly large,
usually medium-sized; forewing apex often fal-
cate or produced; hindwing anal margin usually
straight, or even concave, tending to be distinc-
tively patterned (Figs. 18.4A-D). Male anten-
nae bipectinate either proximally or for their
whole length (or almost to the apex) or "tri-
pectinate" (Phiditiinae, in which there is a mid-
ventral row of teeth); rami directed latero-
ventrad; "shaft" usually scaled to the apex (dor-
sally). In female, antennae often similar to those
of male (especially regarding length of rami), but
serrate or almost filiform in many Prismosticti-
nae. Compound eyes naked, except in Prismo-
sticta (eyes then with long interommatidial setae
and distinctly lashed anteriorly). Distinct ocelli
and chaetosemata absent. Pilifer bristles absent
or replaced by scales. Proboscis very short to ab-
sent (galeae sometimes bladder-like). Maxillary
palpi absent or vestigial. Labial palpi moderately
long to vestigial, usually 2- or 3-segmented, and
either upturned, porrect or drooping; segment 3
invariably short. Line of junction between mese-
pimeron and meron usually strongly ascending
caudad (Shepard 1930: Fig. 27). Longitudinal su-
ture of hindcoxa (merocosta) often markedly
shortened (i. e. with its ventral extremity lying
far from trochanter). Tibiae unspined, often with
erect scales (that may form conspicuous dorsal
fringe on hindtibia); epiphysis well developed in
male (very long in Bombycinae: e. g. Fig. 14 in
Common & Edwards 1991), usually as long, or
nearly as long, in female (reduced/vestigial in,
Claude Lemaire & Jol Minet
e. g., females of Andraca)\ spurs (moderately
long to very short): 0-2-4 or 0-2-2. Tarsi some-
times with ventral spines, often with erect scales.
Pretarsus: arolium present; claws with simple/
serrulate ventral edge; pulvilli produced dorsally.
Male often with a subcostal retinaculum and a
fairly long frenulum (if subcostal retinaculum
absent, frenulum distinctly shortened); female
frenulum multiple, fairly short to vestigial. Spi-
narea absent. Forewing venation (Figs. 18.3
- C ) : all Rs branches stalked together, one
branch sometimes absent (Figs. 18.3 A, B); free
section of Rs4 usually shorter than stem Rs3 +
Rs4, but longer than stem in several Prismosticti-
nae; distance separating Rs4 from Ml often
decreasing distad; M2 rarely closer to M3 than
to Ml (Phiditiinae: Fig. 18.3 A); in many Bom-
bycini, stem of M and CuP preserved as tubular
veins (e. g. Holloway 1987: Fig. 4, right). Hind-
wing: subcosta fused for a short distance with
discal cell upper edge or meeting it at a point
situated either near wing base, or approximately
in the middle of this edge (the point of contact
may be replaced by a crossvein: R); Rs and Ml
either free or stalked; M2 arising nearer M 3 than
M l only in some Phiditiinae; M3 and CuAl
sometimes stalked (e. g. Forbes 1942: Fig. 86;
Common & Edwards 1991: Fig. 13); CuP some-
times tubular; 3 A well developed. Abdomen:
dorsal surface occasionally with numerous spine-
like scales (Apatelodes); dorsum 1 with a median
sclerite varying from moderately broad to en-
tirely absent, and lateral bars that may be en-
larged in the rear; anterior processes of sternum
2 small to vestigial; sternum 2 apodemes usually
short (moderately long and spatulate in, e. g.,
Bombyx mori), venulae absent in most cases; in
male, A2 sometimes with pair of pleural, andro-
conial, tufts and sternum 8 often distinctly modi-
fied, sometimes fused with vinculum (modifica-
tion may also affect tergum 8 and, occasionally,
The Bombycoidea and their Relatives
tergum 7; A8 may develop a superuncus and/or
valva-like processes); intersegmental fold 8 - 9
sometimes with androconial tufts or a dorsal
gland (e. g. Dierl 1978: Fig. 3; 1979: Fig. 3). Male
genitalia: saccus varying from absent (e. g.
Prismostictinae) to long and often spatulate
(Holloway 1987: Figs. 95, 106, 107, etc.); vincu-
lum and tegumen often synscleritous; valvae var-
iously developed, entire or cleft; socii sometimes
present; uncus long to vestigial/absent; gnathos
complete, divided or absent; phallus sometimes
fused with juxta, but free and elongate in most
Bombycinae (it may have a long anterior coe-
cum: e. g. Kuznetzov & Stekolnikov 1985: Fig.
12). Female genitalia: length and ornamentation
of corpus bursae extremely variable; ostium bur-
sae either on venter 8 or on intersegmental mem-
brane A7A8; each pair of apophyses varying
from moderately long to entirely absent; in
Epiini, papillae anales tending to be closely
associated with A8; corethrogyne occasionally
present.
Egg of "upright" type in Apatelodinae
(though strongly flattened, disc-like, with a
sharp circular margin: see PI. 9 in Packard 1895),
of "flat" type and when seen from above
oval to nearly round in the other subfamilies
(e. g. Common & Edwards 1991); chorion with-
out conspicuous sculpturing. First instar larva
sometimes without secondary setae (e. g. Com-
mon & Edwards 1991), often with several Verru-
cae or verricules (e. g. Forbes 1923: Fig. 430);
head flattened anteriorly in Apatelodes; LI and
L2 distinctly separate on A l - A8; A8 often with
a middorsal scolus (short to long, sometimes
with an evaginable distal section); prolegs often
coronate. Mature larva with numerous second-
ary setae, either short/minute (Figs. 18.7 A, B)
or long (Apatelodinae); Apatelodinae often with
dorsal tufts of specialized setae (e. g. Packard
1895, Peterson 1948 and Stehr et al. 1987), some-
times with numerous spatulate setae (e. g. in
Tarchon: Forbes 1942); A8 showing a middorsal
scolus, except in Andraca (Fig. 18.7 B) and Apa-
telodinae; scoli or filaments occasionally present
elsewhere on the integument (swellings and small
gibbosities more frequent); forecoxae either free
(Apatelodinae) or fused (remaining subfamilies);
proleg crochets in biordinal mesoseries (that is
strongly curved in Prismosticla: Miyata 1970).
Pupa (Figs. 18.7 F, G): labial palpi concealed or
very slightly exposed (e. g. Miyata 1970: PI. 60);
proboscis short; maxillary palpi and forefemora
concealed (coxae exposed in Bombyx mori:
Mosher 1916); midlegs usually extending, cau-
dad, beyond apices of antennae; thoracic spira-
cles of Phiditiinae in deep, characteristic hollows;
abdominal integument sometimes soft and flexi-
ble (e. g. Bombyx mori); Apatelodinae with an
anterior row of crenulations or pits on A2A7;
cremaster usually absent (present in Phiditiinae).
Pupation either in the ground (many Apatelodi-
333
nae: see e. g. Peigler 1994) or in a silken cocoon,
usually tough and spun beneath a leaf; cocoon
of Tarchon incorporating larval setae.
The adults are nocturnal. At rest, they often
hold their fore- and hindwings remote from the
abdomen, most of the hindwing surface being
concealed, except for the anal margin, which is
usually folded upwards in a characteristic way
(e.g. Danesch & Dierl 1968: 147; Holloway
1987: 6; Chang 1989: 213-227). The hindwing
margin is often produced into a "tooth" near the
extremity of 3 A, caused by a fold between this
vein and 1 A + 2 A. The abdomen tends to be
curved upwards or sideways. In Phiditiinae, fore-
and hindwings are strongly folded, and they may
be rolled together (Sorocaba), more or less in a
pterophorid-like arrangement. A different rest-
ing posture is seen in many Apatelodinae: the
forewing inner margins are held along the abdo-
men, the hindwings being entirely concealed
(when the insect is viewed from above), and the
wings and the abdomen are held remote from
the substrate.
Worldwide, there are approximately 350 de-
scribed species, referred to about 40 genera.
Apart from the domesticated species, Bombyx
mori, the family is absent from Europe. It occurs
practically everywhere else, being better repre-
sented in the oriental and neotropical regions.
Four subfamilies can be recognized (Holloway
1987; Minet 1994). They are distinguishable as
follows (imaginai characters):
Key to the subfamilies
1 Hindtibia with two pairs of spurs 2
Hindtibia with one pair of spurs 4
2 Forewing with M2 arising closer to M3 than
to M l (Fig. 18.3 A) Phiditiinae
Forewing with M2 arising midway between
M l and M3, or closer to M l than to M3
3
3 Compound eyes "naked" and flagellum of
male antenna bipectinate to apex (New
World only) Apatelodinae
Compound eyes hairy or distal section of
male antenna serrate or unipectinate (Old
World only) . . . . Prismostictinae (in part)
4 Male antenna with a long or fairly long dis-
tal section, which is not bipectinate; female
antenna often serrate to almost filiform;
forewing with discocellular M2M3 dis-
tinctly angled inwards
Prismostictinae (in part)
Male antenna bipectinate to apex, or not bi-
pectinate only on a short apical section
(comprising less than 10 flagellomeres); fe-
male antenna always distinctly bipectinate;
forewing with discocellular M2M3
straight or weakly curved (Figs. 18.3 B, C)
Bombycinae
334
Apatelodinae are mainly neotropical, but five
species occur in America north of Mexico
(Franclemont 1973, Hodges et al. 1983). The
subfamily contains more than 10 genera, and
about 150 described species. Apatelodes (Fig.
18.4 A), Colabata and Olceclostera are the main
genera. The adults often possess four Rs
branches in the forewing (Figs, in, e. g., Forbes
1942, Lima 1950 and Franclemont 1973). Cer-
tain larvae are polyphagous (Packard 1895: 103).
The recorded hostplants thus belong to a wide
range of families: Aquifoliaceae, Betulaceae, Bi-
gnoniaceae, Lauraceae, Oleaceae, Rosaceae, and
others. However, Apatelodinae are of no eco-
nomic importance (Stehr et al. 1987). Pupation
usually occurs in the ground, but silken cocoons
are spun e. g. in Tarchon and Zanola (Forbes
1942).
Phiditiinae are a small neotropical group of
about 25 species, belonging to four genera: Phi-
ditia (Fig. 18.3 A), Rolepa, Sorocaba and Tepilia.
Long placed in Lymantriidae, these taxa were
transferred by Minet (1986) to the "Apatelodi-
dae" sensu auct. The first instar larvae have only
a short dorsal protuberance on A8, unlike the
sphingid-like mature larvae, which bear a long
middorsal scolus on this segment (Fig. 18.7 A)
(see Jrgensen 1928 and 1932). Known host-
plants belong to Bignoniaceae.
Prismostictinae are an Asian subfamily, rede-
fined by Holloway (1987: "Mustilia lineage") and
renamed "Oberthueriinae" by Kuznetzov &
Stekolnikov (1985). The group contains 5 genera
and about 15 species. It can be divided into two
tribes (Minet 1994), viz. Prismostictini and Ober-
thueriini. The former have adults provided with
hairy eyes and fully pectinate antennae (Prismos-
ticta), whereas the latter possess "naked" eyes
and, in males, antennae distally devoid of well
developed rami (the main genera are Andraca
and Mustilia: Fig. 18.4D). The larvae feed on
Symplocaceae in Prismostictini, mostly on Sym-
plocaceae and Theaceae in Oberthueriini (Hollo-
way 1987; Sugi et al. 1987; Chang 1989). The lar-
val morphology is variable (Miyata 1970) and
some species are reported to defoliate tea
(Roepke 1924; Kaishoven et al. 1951: 552-553).
Bombycinae (sensu Minet 1994) contain two
tribes, Bombycini (about 15 genera) and Epiini
(about 5 genera), and a genus that may deserve
a separate tribe (Sesquiluna: Figs, in Kishida
1992 and 1993). The subfamily includes about
150 described species. Bombycini occur mainly
in the Indo-Australian region, but also in pa-
laearctic Asia, tropical Africa and Madagascar.
Epiini are exclusively neotropical. In both tribes,
the imagines have soft, bladder-like galeae (at
least in the males), a long epiphysis on the fore-
tibia, many acute-toothed wing scales, and a
modified sternum 8 on the male abdomen (Figs,
in e. g. Forbes 1942; Dierl 1978, 1979; Holloway
1987); the larvae primarily feed on Moraceae
Claude Lemaire & Jol Minet
(e. g. Schade 1939; Holloway 1987; Common &
Edwards 1991). In Bombycini, the four Rs
branches are preserved in the forewing (Fig.
18.3 C), as also may be vein CuP in both wings,
and a section of the M stem in the forewing cell.
Bombyx, Gunda and Ocinara (Fig. 18.4C) are the
largest genera. Feeding mostly on mulberries
(Morus), Bombyx mori is the best known silk-
moth. It is closely related to the Asian Bombyx
mandarina, but is now only a cultivated species,
wholly unknown in the wild. Its variously col-
oured larva has the meso- and metathorax dis-
tinctly inflated. Several races exist, correspond-
ing to differences in the shape, size and colour of
the cocoons. Epiini lack one Rs branch in the
forewing (Fig. 18.3 B). Colla, Epia (Fig. 18.4 )
and Quentalia are their main genera. In Colla,
the larva has a middorsal scolus ( D l ) on A2, and
another on A8, which is long, flexible and move-
able (Schade 1939). This morphology is likely to
belong to the ground plan of the tribe.
Endromidae. Since it contains only a single spe-
cies, the family Endromidae is obviously mono-
phyletic, and several of its traits can be regarded
as autapomorphic: e. g. (a) proboscis vestigial,
practically absent; (b) labial palpus very short,
without distinct segmentation (at least on its in-
ner surface); (c) M2 distinctly closer to M3 than
to Ml in both pairs of wings; (d) cavity of the
"tergal rim" (sensu Brock 1971) broadly open
caudad; (e) in the male genitalia, valva with a
longitudinal sclerite ending in a free, acute point;
(f) female with the lamella postvaginalis repre-
sented by a strong, dorsally excavated, protuber-
ance.
Adult medium-sized to rather large (Fig.
18.6 A). Antennae bipectinate to apex in both
sexes; flagellum scaled dorsally, without teeth
ventrally, between the lateroventral rami. Com-
pound eyes naked. Distinct ocelli and chaeto-
semata absent. Pilifer elements numerous and
thin, more or less piliform. Proboscis vestigial;
maxillary palpi minute, unsegmented. Labial
palpi short, drooping, without distinct segmenta-
tion. Epiphysis either long and curved (male), or
strongly reduced (female); tibial spurs: 0-2-2.
Arolium well developed; pulvilli undivided, nar-
row distally; claws simple (i. e. without a ventral
tooth). Wing-coupling amplexiform in both
sexes. Spinarea absent. Forewing venation (Fig.
18.5 E): four Rs branches, all stalked together
(Rsl and Rs2 having an extremely long stem);
M2 arising close to M3; within the discal cell,
M stem with its lower branch distally sclerotized
(though thin); tubular CuP lacking; 1 A + 2 A
distinctly forked at base. Hindwing venation: hu-
meral vein absent; upper edge of discal cell con-
nected to Sc by an oblique crossvein (R); Rs free;
CuP replaced by a fold; 3 A present. Abdomen:
median sclerite of tergum 1 very narrow; no link
between tergum 1 and sternum 2; terga 3 to 7
The Bombycoidea and their Relatives

Fig. 18.4. Adults of Bombycidae ( - D ) and Saturniidae (EM): A, Apatelodes singularis; B, Epia muscosa; C,
Ocinara malagasy; D, Mustilia falcipennis; E, Asthenidia buckleyi; F, Neocercophana phiHppii <$; G, Tilaea timur \
H, Eacles masoni I, Dirphia lichyi 9; J, Aglia lau\ K, Salassa mesosa; L, Holocerina rhodesiensis 9; M, Antheraea
larissa. Scales AD = 5 mm; E - M = 10 mm (J. Minet).

anterolaterally produced; sternum 2 with short
apodemes; venulae absent; male sternum 8 plate-
like, m o r e or less crescentic. Male genitalia with
distinct sutures between vinculum a n d tegumen,
a n d between tegumen a n d uncus; saccus b r o a d
a n d short; valvae with a longitudinal sclerite
ending in an acute point; g n a t h o s reduced to a
pair of small lateral bars; phallus without a well
developed coecum. Female genitalia: signum ab-
sent; ostium bursae lying in middle of venter 8,
336
under a prominent lamella postvaginalis (cau-
dally rounded and dorsally concave); apophyses
anteriores short, apophyses posteriores long; pa-
pillae anales dorsally fused.
Eggs subcylindrical, of the flat type (Dring
1955), usually laid side by side in straight rows
along the twigs (Brooks 1991). Chorion smooth,
transparent. First instar larva with secondary se-
tae arising from prolegs, anal shield, and from
several Verrucae; A8 with a middorsal verruca
( D l ) set on a conical hump; LI and L2 rather
far apart on A 1 - A 8 . Last instar larva with sec-
ondary setae on head, as well as numerous short
secondary setae scattered over thorax and abdo-
men; proleg crochets in biordinal mesoseries; A8
with a well developed conical hump (e. g. Gmez
de Aizprua 1988). Pupa: head with ventral pro-
jections; galeae short; labial palpi, maxillary
palpi and forefemora concealed; forewings adja-
cent ventrally, on the meson; abdomen dorsally
covered with acute teeth, mainly on A5AIO;
AIO dorsum produced caudad and provided
with apical setae. Cocoon in the soil or in leaf
litter. Pupa protruding from cocoon just before
emergence (Brock 1990).
The palaearctic species, Endromis versicolora
(Fig. 18.6 A), is the only member of the family
(Nssig & Czipka 1994). Males are mainly day-
flying, but also come to light at night (Rougeot
1971). Females are nocturnal. At rest, the imago
holds its wings roofwise. The larvae are cryptic
and feed on Betula and Alnus (Betulaceae), occa-
sionally other Betulaceae or trees such as Salix
(Salicaceae), Tilia (Tiliaceae) and Ulmus (Ulma-
ceae).
Addendum According to Dr I. J. Kitching
(pers. comm.), a second taxon should be placed
in Endromidae, viz. the Tibetan genus Dalailama
(that "was long curated in the Endromidae in the
B M N H collection").
Mirinidae. In recent literature, this monotypic
taxon has been considered either a subfamily of
Endromidae (Kozlov 1985, Kuznetzov & Stekol-
nikov 1985), a separate family (Minet 1986, Hol-
loway 1987), or even a member of Bombycidae
(Fletcher & Nye 1982). Among the main autapo-
morphies of Mirinidae, three obvious ones are:
(a) uncus anteriorly provided with a pair of con-
spicuous, dorsal protuberances; (b) larva with
two pairs of long, thin, moveable scoli on the
dorsum of thoracic segments 2 and 3); (c) pres-
ence of eversible glands above the ventral prolegs
of the larva.
Imago medium-sized, broad-winged, with a
stout pilose body (Fig. 18.6M). Antennae bipec-
tinate to the tip in both sexes, with V-shaped
pairs of rami and a dorsally scaled shaft. Eyes
naked; ocelli absent or nearly so. No well devel-
oped pilifers. Proboscis short; maxillary palpi
small, unsegmented. Labial palpi 2-segmented
(distal segment moderately long, porrect or
Claude Lemaire & Jol Minet
slightly ascending). Epiphysis either well devel-
oped (male), or absent (female); tibial spurs: 0-2-
2. Tarsi with ventral spines. Arolium and pulvilli
present; claws strong, simple. Wing-coupling am-
plexiform in both sexes. Spinarea absent. Fore-
wing venation (Fig. 18.5 F): R free; Rs branches
all present and stalked (the free section of Rsl
being extremely short); M l and the Rs stem
either connate or short-stalked; M2 arising about
midway between M l and M3; CuP absent; 1A
+ 2 A with a long basal fork. Hindwing vena-
tion: Sc touches upper edge of discal cell just be-
yond wing base, then diverges strongly; M2
slightly closer to M l than to M3; tubular CuP
absent; 1 A + 2 A with a basal fork; 3 A present.
Abdomen: dorsum 1 with narrow posterior (me-
dian) sclerite, and with its lateral bars enlarged
posteriorly; sternum 2 free from the terga, with
short apodemes and without venulae; male ster-
num 8 large, plate-like, with concave posterior
edge. Male genitalia: vinculum and tegumen syn-
scleritous; a distinct suture between uncus and
tegumen; sacculus well defined; no sclerotized
gnathos; phallus with long anterior coecum. Fe-
male genitalia: corpus bursae small; signum ab-
sent; ductus bursae short and broad; ductus se-
minalis opening close to ostium bursae; apophy-
ses anteriores vestigial; apophyses posteriores of
moderate length.
Egg ovoid, without longitudinal ribs. There
are five larval instars, all with similar chaetotaxal
features (Kozlov 1985). Larva (Fig. 18.7 C): head
hypognathous; thorax and abdomen with many
lateral and dorsal scoli, the longest being the
middorsal scolus ( D l ) of A8 and the four pairs
of dorsal moveable scoli present on T2 +
T3; a characteristic, eversible, glandular organ
above each ventral proleg (these glands and the
moveable scoli are clearly defensive: Graeser
1888, Seitz 1911). Last instar larva with biordinal
proleg crochets. Pupa stout, without abdominal
teeth; no well defined cremaster. The strange,
tough cocoon (Kozlov 1985) is spun on the food
plant (Lonicera, Caprifoliaceae).
The family contains one genus, Mirina (Fig.
18.6 M), with two described species, both from
palaearctic Asia (Meli 1939; Laithwaite et al.
1975).
Saturniidae. As currently delimited (Minet 1994),
Saturniidae include "Oxytenidae" and "Cerco-
phanidae". However, such a concept of the fam-
ily is by no means a true innovation (see, e. g.,
Zerny & Beier 1936). Thus conceived, Saturnii-
dae are monophyletic, as indicated by several de-
rived traits of their ground plan, such as the sex-
ual dimorphism of tarsomere 4 of the adult fore-
leg (e. g. Lemaire 1978: Fig. 6).
Medium-sized to very large moths (wingspan
30280 mm); females sometimes micropterous
in Meroleuca (Dognin 1913, Bouvier 1935).
Ground colour extremely variable with predomi-
The Bombycoidea and their Relatives
nance of different shades of brown. Wing pattern
similarly variable (Figs. 18.4E-M): the eye spots
with concentric rings (from which the family de-
rives its name) and the equivalent crescentic dis-
cal spots (Attacini), although regarded as the
most characteristic markings, are actually want-
ing in more than 600 species; forewing ante- and
postmedian lines, hindwing postmedian line and
hindwing submarginai band are probably the
most stable pattern components. Male forewing
usually falcate; outer margin of hindwing either
rounded or angulate or, in several unrelated
groups, with a short to extremely long tail
(hindwing six times longer than forewing in some
Eudaemonia); in Ceratocampinae, wing shape
suggestive of that of Sphingidae or of some Lasi-
ocampidae. Body of male relatively slender (ex-
cept in Ceratocampinae), of female usually
much heavier.
Frontoclypeus usually flattened laterally, ex-
cept in Ceratocampinae. Antennae of male bi- or
quadripectinate, of female simple to quadripecti-
nate; 25 to 60 segments; quadripectinate anten-
nae with the apical rami of each flagellomere
arising near the basal rami of the following fla-
gellomere, except in Saturniinae and some Ludii-
nae; shaft in male normally scaled only on dor-
sum of basal segments, but densely scaled to the
apex in Oxyteninae and Periga. Compound eyes
usually large (reduced in day-flying males of var-
ious unrelated groups); external ocelli, if present,
extremely small (Dickens & Eaton 1973); chaeto-
semata absent. Pilifers without bristles except in
Arsenurinae (Michener 1952), some Oxyteninae
and Janiodes (Cercophaninae). Galeae, when
present, varying from rudimentary to longer
than head. Maxillary palpi rudimentary. Labial
palpi 1- to 3-segmented; when 3-segmented, seg-
ment 2 longer than either of the others. Mesa-
nepisternum large, but variable in shape and size,
depending on orientation of anepisternal suture.
Epiphysis either present or absent, more fre-
quently reduced/absent in female than in male,
when present varying from minute to nearly as
long as foretibia; tibial spurs: 0-2-2, 0-2-3 or 0-
2-4. Tarsi with or without ventral spines, in addi-
tion to the nearly universal apical pair on tar-
somere 4 of female foreleg. Tibiae unspined, but
fore- and even midtibiae sometimes with one or
more apical points. Arolium and pulvilli usually
present, but absent in Hemileuca and Coloradla.
Frenulum and retinaculum absent from both
sexes. Females of some Ludiinae with a large
patch of modified scales on forewing underside,
before the tornus (suggestive of a stridulating or-
gan: Jordan 1922a). Forewing (Figs. 18.5 A - D ) :
discal cell closed or open; Rs branches stalked
together, one or two of them absent; R free or
stalked with stem of Rs; M1 and Rs either con-
nate, stalked (e. g. distinctly stalked in almost all
Ceratocampinae: Fig. 18.5 C), or remote from
each other (Fig. 18.5 D); M2 arising midway be-
337
tween Ml and M3 or nearer Ml than M3; Ml
and M2 sometimes connate (e. g. Meroleuca), oc-
casionally stalked (some Hemileuca), CuP ab-
sent; 1 A + 2 A usually with a small basal loop
(a dorsal spur occasionally arises from apex of
loop, e.g. as in most Oxyteninae: Fig. 18.5A).
Hindwing: discal cell open in Attacini, otherwise
usually closed; there is sometimes an oblique
crossvein (R) between the subcosta and the up-
per edge of the discal cell (evident especially in
Cercophaninae: Fig. 18.5 B); Ml and M2 often
approximate, M3 arising from lower angle of
discal cell; CuAl and CuA2 usually remote from
each other; CuP absent; 1 A + 2 A without basal
loop; 3 A usually short to absent, but extending
to three-quarters length of 1A + 2 A in Co-
loradla and Hemileuca. Abdomen: in Agliinae,
Arsenurinae (Lemaire 1980) and most Oxyteni-
nae, male with an androconical tuft on each side
of A 2 (in a pleural fold); eight sternum and ter-
gum of male with strongly differentiated scleroti-
zations in several genera of Hemileucinae, Satur-
niinae and Ludiinae; sternum 8 sometimes with
long posterior points in Oxyteninae. Male and
female genitalia extremely variable. Phallus with
or without coecum; no sclerotized phallus in two
species of Saturnia (Calosaturnia), the vesica be-
ing then supported by long, caudal extension of
juxta (Lemaire 1978); length of bulbus ejacula-
torius exhibiting considerable interspecific varia-
tions in several genera of Hemileucinae (usually
in relation with length of female bursa Lem-
aire 197174); vesica with or without cornuti;
juxta either free or fused to valvae or vinculum,
sometimes replaced by projections of valvae or
by a pair of strongly sclerotized spines arising
from sternum 9 (e. g. Gamelia); anellus usually
membranous, but represented by strong tubular
process in Lonomia and Periga (Lemaire 1973);
tegumen fused to vinculum (it may bear dif-
ferently shaped processes protruding above the
uncus); saccus present or absent; valvae simple
to trilobed, either free or fused to vinculum/tegu-
men, and sometimes asymmetrical (e. g. Schau-
siella, Scolesa); uncus either simple or bilobed,
often with strong dorso-apical protuberances,
occasionally absent or membranous (e. g. Autom-
erina), typical gnathos and transtilla sometimes
present together (one or both of these structures
may be absent). Female genitalia: tergum 8 either
simple or bilobed, sometimes entirely membra-
nous, fused or not to sternum 8 laterally; the lat-
ter represented by more or less developed, usu-
ally circumvaginal, sclerite(s); ductus bursae
membranous or sclerotized; corpus bursae vari-
ously developed, with or without signum; two
pairs of apophyses (the postapophyses may be
very short and the anapophyses reduced to min-
ute, posteriorly produced, projections).
Egg usually of the flat type; micropyle espe-
cially conspicuous in most Hemileucinae;
chorion smooth, most frequently whitish and
338
more or less opaque, but green or yellow and
translucent in Ceratocampinae; in Arsenurinae,
eggs flattened, usually brown and encircled with
white. Eggs may be laid in large clusters encir-
cling twigs of the hostplant (e. g. Hemileuca); fe-
male Hylesia usually lay all their eggs simulta-
neously, with a dense covering of hairs around
the mass (Janzen 1984). Larva (Fig. 18.7 D):
head smooth, hypognathous; frontoclypeus ex-
tending usually less than half distance to epicra-
nial notch; thoracic and abdominal segments
usually with several scoli, normally better devel-
oped in first than in later instars (the arrange-
ment of the "scoli" and their structure, especially
in first instar larvae, are of great taxonomic sig-
nificance at supraspecific level: see e. g. Pease
1961); ventral prolegs present on A3A6, usu-
ally with mesoseries of 6 - 2 0 crochets; anal pro-
legs well developed, often very strong. First in-
star larvae of Arsenurinae, Ceratocampinae and
Hemileucinae typically with four pairs of "scoli"
s. I. on TI, T2, 3, Al, 2 and 7; 3 - 6 and
A9 without subventral scoli; AlAl in Hyp-
erchiria and Automerina, A3A6 in Polythysan-
ini with an extra dorsal pair (D2), caudad of
scoli D l ; D l of A8 (except in Anisota and Dryo-
campa) and D2 of A9 fused to form middorsal
horns; dorsal and subdorsal scoli of, at least, T2
and T3 and middorsal horns of A8 and A9 usu-
ally differently shaped than corresponding scoli
of other segments. In later instars of Hemileuci-
nae, scoli spine-like, with strongly urticating spi-
nules (these replaced, in Polythysanini, by very
long hair-like setae on the abnormally enlarged
subspiracular and paranal scoli). Last instar lar-
vae of Arsenurinae entirely devoid of scoli. In
Oxyteninae, last instar larvae sphingid-like, with-
out long protuberances, except for a middorsal
"horn" on A8; metathorax more or less enlarged,
often with a pair of eyespots (e. g. Nentwig 1985,
Aiello & Balczar L. 1997). In Cercophana (Cer-
cophaninae), first instar larvae with most D, SD
and L setae replaced by Verrucae (Wolfe & Bal-
czar L. 1994); last instar larvae with a single
"scolus" (a conical, middorsal projection on T3)
and anal shield posteriorly ending in an acute
angle. First instar larvae of Agliinae with pair of
long, apically forked, dorsal scoli on TI and T3,
and single middorsal scolus on A8 (other scoli
minute to absent; armature lost in last instar lar-
vae). Larvae of Ludiinae often clothed with
dense hair-like setae; scoli short, giving rise to
slender spine-like setae; subventral scoli absent;
Dl scoli of A8 fused, adjacent or separate (Ober-
prieler & Duke 1994: 236); A9 without middorsal
scolus. First instar larvae of Salassinae mainly
with primary setae and almost entirely devoid of
scoli; "armature" reduced to a few antero-dorsal,
prothoracic protuberances, and a caudal, conical
extension of the anal shield (Nssig 1994). In Sa-
turniinae, thorax and abdomen without sub-
ventral scoli, Dl scoli of A8 either separate (e. g.
Claude Lemaire & Jol Minet
Saturnia) or forming single, middorsal horn
(e. g. Attacus, Antheraea, Actias), and D2 always
separate on A9. Larvae of Australasian and Afri-
can Attacini covered with waxy powder in all but
first instars. Pupa: integument well sclerotized,
either smooth or, especially in Ceratocampinae,
roughened with spines on thorax (dorsally) and
abdomen (setae abundant on the same areas in
most Hemileucinae); maxillary palpi absent; ga-
leae varying from very short to roughly length
of forewings; antennae usually very broad, with
distinct pectinations and extending little more
than half length of forewing; labial palpi min-
utely exposed or absent; forefemora concealed;
midtarsi usually extending slightly beyond tips of
antennae; wings extending at least to posterior
half of A4, hindwings scarcely protruding from
forewings; metanotum with pair of prominent
tubercles in Ceratocampinae; cremaster present
or absent (when present, simple or bifid, giving
rise to a group of closely set hooklike spines in
several groups of Saturniinae and Hemileuci-
nae); in Caio (Arsenurinae), dorsum of AIO with
a pair of deep, well partitioned, twin depressions
(Wolfe & Pescador 1994).
Adults are primarily nocturnal, but males of
Agliinae (Fig. 18.4 J) and a few species of Satur-
niinae and Hemileucinae are diurnal. The resting
posture of the imagines is variable, but usually
(except in Saturniinae) stable within each of the
subfamilies. All larvae of Hemileucinae are gre-
garious and processionary, at least in first in-
stars; there are gregarious larvae in other sub-
families, e. g. Anisota and Dryocampa in Cerato-
campinae. The larvae of many Hylesia species
live in more or less loosely spun communal silk
pouches; L4 and L5 larvae of Lemaireia rest dur-
ing the day in groups within silken tubes between
leaves (Lampe & Nssig 1989). Pupation occurs
either in a silken cocoon or in the soil. The co-
coon may be single or double walled, solid or
more or less finely meshed, aerial (in a leaf or on
a twig of the hostplant) or spun in leaf litter (or
even underground). Subterranean pupae are
either in a cell or directly in contact with the
earth. Pupae of Pseudantheraea hang from the
foliage by the cremaster. Cocoons of several spe-
cies of Hylesia and Opodiphthera are spun in
mass. The overwintering stage of several species
in temperate climates is the ovum, but the pupa
is the more usual diapause stage. The duration
of the latter is extremely variable, from three
weeks to several years (up to 12 years in Epi-
phora atbarina: Rougeot et al. 1991). Most satur-
niids have highly polyphagous larvae; those of
Attacus were found to feed, in the wild and in
captivity, on at least 90 genera of plants in 48
families (Peigler 1989) - most of these records
applying to Attacus atlas. Forty-six plant species
in 18 families were recorded by Janzen (1984) as
hosts of Hylesia lineata in Parque Nacional
The Bombycoidea and their Relatives 339

Fig. 18.5. Wing venation of Saturniidae ( - D ) , Endromidae (E), Mirinidae (F), Carthaeidae (G), Brahmaeidae
(H) and Sphingidae (I, J): A, Asthenidia diffissa; B, Neocercophana philippii J; C, Adeloneivaia subangulata; D,
Copaxa decrescens; E, Endromis versicolora, F, Mirino christophi; G, Carthaea saturnioides; H, Dactyloceras lucina;
I, Leucophlebia afra; J, Hemaris fuciformis. Scales = 5 mm (G. Hodebert).

Santa Rosa in Costa Rica. Nevertheless, many
larvae are specialized feeders.
The long scoli of the larvae of Hemileucinae
bear poisonous spines; the sting is usually not
serious, but cases of a generalized, sometimes fa-
tal (Fraiha et al. 1986), haemorrhagic diathesis,
after contacts with larvae of Lonomia, have been
reported in Venezuela (Arocha-Piango & Lay-
risse 1969) and Brazil (Duarte et al. 1986). In
Hylesia, female moths protect egg batches with
extremely irritating hairs; these "flchettes"
(Lamy & Lemaire 1983) cause severe dermatitis,
340
known locally as "papillonite de Guyane" and
"Caripito itch".
Saturniidae occur almost worldwide, being ab-
sent only from northernmost and far southern
regions. Saturnia (Eudia) pavonia ranges as far
north as the vicinity of North Cape in Scandina-
via (70 North) and Ormiscodes amphinome as
far south as Ushuaia, Terra del Fuego (55
South). The group is most strongly represented
in the tropics, especially in the neotropics, but
with only two or three species in the West Indies.
Saturniidae are particularly abundant in the
mountains at moderate to relatively high eleva-
tions (highest record: 4600 meters for Roth-
schildia renatae, in Peru G. Lamas, pers.
comm. ); the richest area in the world is the rain-
forest on the east slopes of the Andes from Co-
lombia to Bolivia but there is a significant per-
centage (ca. 15 to 20%) of endemics in arid and
semi-arid areas. Attacus, Archaeoattacus and
Coscinocera, all inhabitants of the Indo-Austra-
lian region, are among the largest known Lepi-
doptera.
The family contains approximately 1480 spe-
cies in 165 genera. It is divided into nine subfam-
ilies, the adults of which can be distinguished
using the following key (in part from Michener
1952; however, the best diagnostic features lie in
the early stages, especially in the larvae):
Key to the subfamilies
1 Male with antennal flagellum dorsally scaled
to apex and bases of rami directed latero-
ventrad
- Male with flagellum unsealed, at least for
most of its length; bases of rami directed
laterad or latero-dorsad
2 Proboscis clearly present; apex of tibial
spurs not sharply pointed . . . Oxyteninae
- Proboscis absent or nearly so; tibial spurs
with a bare, hook-like apex
Hemileucinae, part (Perga)
3 Hindwing with well indicated crossvein (R)
between Sc and upper edge of discal cell
(Fig. 18.5 B) Cercophaninae
- Hindwing with crossvein nearly always ab-
sent or vestigial
4 Presence of strong bristles on pilifers or on
clypeal margin between pilifers Arsenurinae
- Pilifers and clypeal margin without bristles
5 Frons convex at sides, so that lateral sutures
are hidden in anterior view; antennal cones
(short ventral protuberances on flagello-
meres) simple Ceratocampinae
- Frons flat at sides or (if convex) antennal
cones multiple
6 Antennae, when quadripectinate, with bases
of rami invariably well separated; anterior
area of mesoscutum without middorsal pro-
jection; in forewing, when discal cell is
Claude Lemaire & Jol Minet
closed, base of Ml arising closer to M2 than
to Rs (Fig. 18.5 D) or about midway be-
tween M2 and Rs; labial palpi not fused to-
gether; hindwing sometimes with a long tail
Saturniinae
Antennae, when quadripectinate, with apical
rami of a segment usually adjacent to basal
rami of next segment; if rami separate, mes-
oscutum anteriorly provided with middorsal
projection or forewing with base of M1 dis-
tinctly closer to Rs than to M2 (or even
stalked with Rs); labial palpi occasionally
fused together; hindwing never with a long
tail 7
7 Antennal cones absent Salassinae
Antennal cones present 8
8 Fore- and hindwing with eyespot at apex of
discal cell (Fig. 18.4 J) 9
Forewing without eyespot (Figs. 18.41 and
L) 10
9 Antennal cones, reduced to small projec-
tions, present only on distal half of antenna
Hemileucinae: Polythysanini
- Antennal cones simple, not similarly re-
duced Agliinae
10 Frontal protuberance usually present, an-
tennal cones simple (American)
Hemileucinae: Hemileucini
Frontal protuberance absent; antennal cones
usually multiple (African) Ludiinae
Oxyteninae (Fig. 18.4 E). Neotropical sub-
family with about 35 species in 3 genera (Jordan
1924): Homoeopteryx, Oxytenis and Asthenidia.
2 Cercophaninae (Fig. 18.4 F). New World. 10
species in 4 genera (Jordan 1924): Janiodes,
Microdulia, Neocercophana and Cercophana.
3 Arsenurinae (Fig. 18.4G). Neotropical sub-
family divided in two tribes (Lemaire 1980):
Arsenurini (main genera: Arsenura, Paradae-
monia) and Almeidaiini (only one genus: Almei-
daia). 60 species in 10 genera. Hindwings pro-
vided with very long tails in Copiopteryx.
Ceratocampinae (Fig. 18.4 H). New World. 170
species in 27 genera (Lemaire 1988). Adults with
robust bodies, often lasiocampid-like or sphin-
gid-like. In all genera except Bathyphlebia, the
4 antenna of the male has a distal section entirely
devoid of rami. Main genera (alphabetically):
Adeloneivaia, Anisota, Citheronia, Eacles, Ptilo-
scola, Schausiella and Syssphinx.
5 Hemileucinae (Fig. 18.41). New World sub-
family divided into two tribes (Michener 1952):
Hemileucini and Polythysanini (with a single ge-
nus, Polythysana). 630 species in 51 genera
(Draudt 1929-1930, Ferguson 1971-1972,
Tuskes et al. 1996, Lemaire 1996). Main genera:
6 Automeris, Cerodirphia, Coloradla, Dirphia, Dir-
phiopsis, Gamelia, Hemileuca, Hylesia, Leuca-
nella, Lonomia, Meroleuca, Molippa, Ormiscodes,
Paradirphia, Periga, Periphoba, Pseudautomeris
and Pseudodirphia.
The Bombycoidea and their Relatives
Agliinae (Fig. 18.4 J). Palaearctic. Three spe-
cies in one genus: Aglia.
Salassinae (Fig. 18.4 K). Asiatic. Twelve spe-
cies in one genus: Salassa.
Ludiinae (Fig. 18.4L). African subfamily con-
taining two tribes (Ludiini, Goodiini; see Jordan
1922 b) and a genus that may deserve a separate
tribe (Micragone; see Rougeot 1962 and Minet
1994). 80 species in 8 genera (unconvincingly
transferred to Saturniinae, as tribe Ludiini, in
Oberprieler & Nssig 1994). Some populations
of Goodia kuntzei are parthenogenetic (Lemaire
1969). Main genera: Goodia, Ludia, Micragone
and Orthogonioptilum.
Saturniinae (Fig. 18.4 M). Practically world-
wide in distribution (see e. g. Michener 1952,
Rougeot 1962, Pinhey 1972, Lemaire 1978, Hol-
loway 1987, Peigler 1989, and Pinratana &
Lampe 1990). Divided into five tribes: Saturniini,
Attacini, Bunaeini, Decachordini and Pseuda-
pheliini (the latter three need reorganization).
480 species in 59 genera. Hindwing sometimes
with a long tail (e. g. Eudaemonia and Actias, at
least in males). Certain Saturniinae have had an
important place in sericulture in China and Ja-
pan (Peigler 1993). "Nonmulberry culture" is still
much practised in India where tussore and
muga-silk are produced by Antheraea and eria-
silk by Samia. Largest or best known genera: Ac-
tias, Antheraea, Attacus, Bunaea, Bunaeopsis,
Caligula, Copaxa, Cricula, Coscinocera, De-
cachorda, Epiphora, Gonimbrasia, Imbrasia, Lo-
bobunaea, Loepa, Maltagorea, Opodiphthera,
Pseudobunaea, Rothschildia, Samia and Saturnia.
Carthaeidae. Only one species, Carthaea saturni-
oides, is placed in the family. Among the apo-
morphies are the prominent eyespots of the larva
(which surround the abdominal spiracles), and
the dorsal transverse rows of deep pits on the
pupal abdomen.
Adults large, broad-winged, resembling cer-
tain members of Saturniidae (Fig. 18.6 B). An-
tennae tripectinate in male, serrate in female;
shaft dorsally scaled in both sexes. Compound
eyes naked. Ocelli and chaetosemata absent. Pili-
fere and proboscis well developed; maxillary
palpi small, 3-segmented (Common 1966: Fig. 4).
Labial palpi: second segment ascending, third
segment long, porrect and smooth-scaled. Epi-
physis well developed; tibial spurs: 0-2-4 (most
of them long); mid- and hindtibiae with long,
dorsal hair-scales; tarsi spinose, longer than the
corresponding tibiae. Pretarsus: claws simple
(i. e. without a medial tooth); arolium well devel-
oped; pulvilli with long, dorsal, setiform out-
growths. Male with subcostal retinaculum and
strong frenulum; female frenulum short,
multiple. Spinarea absent. Forewing venation
(Fig. 18.5 G): R free; Rsl and Rs2 long-stalked;
Rs3 and Rs4 short-stalked; M2 arising about
midway between Ml and M3; CuP absent; 1A
341
+ 2 A forked. Hindwing venation: subcosta and
upper edge of discal cell approximate for a short
distance, and connected by a crossvein (R); Rs
and Ml separate; CuP absent; 3 A rather short.
Base of abdomen: lateral bars of dorsum 1
strongly produced ventrad (posteriorly), coming
very close to sternum 2; venulae absent; sternum
2 with short apodemes and, in male, posterolat-
eral sclerotized arms, that are directed cephalad
and support well developed androconial tufts.
Male genitalia: valvae simple, rounded; gnathos
large, with a broad, scobinate apex; uncus trian-
gular; phallus with an anterior coecum. Female
genitalia: ductus bursae short and broad; corpus
bursae small and unarmed; apophyses of moder-
ate length, anterior pair connected both to
ventral and dorsal sclerites of A8.
Eggs ovoid, of the flat type; chorion smooth.
Last instar larva (e. g. Common 1970: Fig.
36.49 A): secondary setae minute; no scoli, but
dorsal hump on A8; abdominal spiracles sur-
rounded by ocellate markings; proleg crochets in
biordinal mesoseries. Penultimate instar (Com-
mon 1966: Figs. 9 and 10) with a number of
clubbed setae, mostly arising from short scoli; a
longer, middorsal, scolus on A8 (DI); scoli LI
and L2 separate on most abdominal segments.
Pupa (Common 1990: Fig. 119.4): antennae and
proboscis long; maxillary palpi very small; labial
palpi and forefemora concealed, except for a
minute, triangular area of the former; segments
Al to A8 with an anterodorsal row of deep pits;
cremaster with more than 20 apical hooklets.
The flimsy cocoons are spun in leaf litter and
perhaps also in the soil.
Erected by Common (1966), the family only
contains the Australian species, Carthaea satur-
nioides (Fig. 18.6B). The adults are nocturnal
and usually come to light after midnight. At rest,
they hide the eyelike markings of their hind-
wings, displaying them when disturbed. The lar-
vae feed externally on the foliage of several Pro-
teaceae.
Lemoniidae. As suggested by e. g. the hindwing
venation, Lemoniidae, Brahmaeidae and Sphin-
gidae are very likely to be close relatives (Minet
1994). Lemoniidae can be defined by the
following four autapomorphies: (a) no distinct
proboscis; (b) foreleg with a short tibia, and with
two very strong spines on the outer side of the
basitarsus (one spine or "tooth" lying dis-
tad of the other); (c) hindtibia without long me-
dial spurs; (d) frenular bristles absent from both
sexes.
Adults medium-sized to fairly large, with stout
bodies and broad wings (Figs. 18.20, R). Anten-
nae bipectinate to tip in both sexes, or even tri-
pectinate (i. e. with a ventral row of ramus-like
teeth); lateral rami directed latero-ventrad,
longer in males than in females; antennal shaft
scaled dorsally. Compound eyes naked, but often
342
"lashed" anteriorly (when lateral, frontal tufts
arise below the antennae). Ocelli and chaetose-
mata absent (or practically absent). Pilifer bris-
tles replaced by scales. Proboscis and maxillary
palpi absent. Labial palpi 3-segmented, moder-
ately developed to fairly short, and either decum-
bent, porrect or slightly ascending. Epiphysis
either well developed in both sexes (Sabalia), or
entirely absent (Lemonia); tibial spurs: 0-2-4, 0-
2-3 or, more often, 0-2-2. Forelegs modified for
burrowing: tibiae shortened (the tarsi may also
be very short); first tarsomere provided exter-
nally with two strong spines (particularly large,
tooth-like and remote from each other in Sab-
alia); in Lemonia, tarsomere 2 also with a strong
spine (tarsomere 3 similar occasionally), and pre-
tarsal claws conspicuously enlarged. Tarsi spined
ventrally. Arolium and pulvilli either well devel-
oped {Sabalia), or vestigial to absent (Lemonia);
claws simple, though sometimes with a serrulate
ventral edge. Wing-coupling amplexiform. Spi-
narea absent. Forewing venation: Rsl and Rs2
either entirely fused (Fig. 18.1 G) or stalked for
a long distance; R free or stalked for short length
with stem Rs (Ml and Rs may also be short-
stalked); M2 arising about midway between Ml
and M3, or nearer Ml than M3; discocellular
M 2 - M 3 strongly bent inwards; CuP replaced by
a fold (often well indicated). Hindwing venation:
humeral vein absent; an oblique crossvein (R)
between subcosta and upper edge of discal cell;
Se + R approximated to Rs beyond crossvein R,
CuP represented at most by strong fold; 3 A
rather well developed. Abdomen: dorsum 1 with
its median sclerite narrow (Lemonia) or absent
(Sabalia), and its lateral bars moderately en-
larged posteriorly; sternum 2 with short anterior
processes and short apodemes; venulae absent or
very short. Male tergum 8 large, markedly longer
than sternum 8; tegumen and vinculum synscleri-
tous; no distinct saccus; gnathos entire, well de-
veloped (e.g. Zolotuhin 1994a), sometimes
fused with the basal, dorsal processes of the val-
vae; uncus rather long; phallus without well de-
veloped coecum. Female venter 7 membranous
posteriorly; corpus bursae rather small, often un-
armed; ostium bursae on venter 8; apophyses
never very long (anterior pair short); papillae an-
ales approximate dorsally.
Egg of the upright type (Dring 1955), hemi-
spherical (Lemonia) to roughly spherical (Sa-
balia: Fontaine 1975), with a smooth chorion
(eggs are laid in groups). First instar larva of
Lemonia: many secondary setae arising from Ver-
rucae on the thorax and the abdomen; AlA7
sometimes with an extra verruca (or seta) caudad
of SD; seta L2 separate from verruca LI on
AlA8; Verrucae D l not fused together on A8.
Mature larva of Lemonia with a similar chaeto-
taxy; secondary setae more numerous (also pre-
sent between Verrucae and on the head capsule);
all setae devoid of barbs; prothoracic coxae
Claude Lemaire & Jol Minet
firmly fused; proleg crochets in biordinal meso-
series. In Sabalia (Fontaine 1975): last instar
larva without long setae, dorsally provided with
paired scoli (rather short, except on the protho-
rax where they are long and flexible in the mid-
dle); a middorsal scolus near the end of the abdo-
men (probably on A8). Pupa of Lemonia: labial
palpi sometimes distinctly exposed; maxillary
palpi concealed; proboscis well developed; meta-
notum variously sculptured; abdomen punctate,
devoid of transverse rows of teeth; AIO dorsum
produced posteriorly, bearing several acute teeth.
Pupation occurring in the soil (no genuine co-
coon).
The adults are usually nocturnal (though both
diurnal and nocturnal in certain species that
have small eyes). Males of Lemonia dumi fly very
rapidly by day, especially in bright sunshine, but
they are also attracted to light at night (Rougeot
1971). At rest, the wings are held roofwise (at
least in Lemonia), and hindwing costal margins
are frequently uncovered. The family contains
about 20 species, placed in two genera, which
have sometimes been referred to distinct subfam-
ilies (Lemoniinae and Sabaliinae: Berger 1958).
Lemonia (Fig. 18.20) occurs in the palaearctic
region, being better represented in the Mediter-
ranean area and the Middle East (Freina & Witt
1987). Its larvae feed essentially on certain Aste-
raceae, such as Hieracium, Sonchus and Taraxa-
cum (Rougeot 1971; Rougeot & Viette 1978).
Sabalia (Fig. 18.2 R) contains about 8 afrotropi-
cal species, mostly from East Africa. The larvae
of Sabalia tippelskirchi live gregariously on Eu-
phorbia granii (Euphorbiaceae) (Fontaine 1975).
Brahmaeidae. This family includes Spiramiopsis,
the true affinities of which have been ignored by
many authors. Despite uncommon plesiomor-
phies in its early stages (Oberprieler & Duke
1994), this genus shares at least eight imaginai
synapomorphies with Dactyloceras, Callipro-
gonos, Acanthobrahmaea and Brahmaea (Minet
1994). Among these traits are: (a) lower edge of
hindwing discal cell markedly shorter than vein
M3; (b) underside of the hindwing with a
number of transverse, parallel, wavy lines (such
lines may also be observable dorsally, and on
both surfaces of the forewings); (c) segments
A 4 - A 6 with the spiracles close to the edges of
the terga. Only Calliprogonos lacks apomorphy
(b), apparently due to secondary loss.
Moderately to very large moths with fairly ro-
bust bodies (Figs. 18.6 C F); wings broad, usu-
ally with several transverse wavy lines. Antennae
bipectinate to the tip in both sexes, frequently
with an additional ventral row of long teeth be-
tween the rami (the latter directed latero-
ventrad); "shaft" scaled dorsally. Antennal fo-
ramina more or less close together. Compound
eyes naked, sometimes "lashed" with frontal
tufts of scales arising near the antennae. Ocelli
The Bombycoidea and their Relatives 343

Fig. 18.6. Adults of Endromidae (A), Carthaeidae (B), Brahmaeidae ( C - F ) , Sphingidae ( G - L ) and Mirinidae
(M): A, Endrotnis versicolora; , Carthaea saturnioides; C, Spiramiopsis comma; D, Dactyloceras lucina; E, Calli-
progonos miraculosa; F, Brahmaea wallichii; G, Sphinx ligustri; , Akbesia davidi; I, Gynoeryx teteforti; J, Xeno-
sphingia jansei; , Cephonodes rufescens; L, Deiiephila porcellus; M, Mirina christophi. Scales A - L = 10 mm;
M = 5 mm (J. Minet).

absent or nearly so. No true chaetosemata. Pili-
fere present, sometimes weak. Proboscis un-
sealed, moderately developed to vestigial (Spira-
miopsis). Maxillary palpi small, 2-segmented
(.Dactyloceras), or vestigial. Labial palpi ascend-
ing or porrect, usually well developed; segment 1
without internal microtrichia; segment 3 short.
Epiphysis well developed; tibial spurs: 0-2-4 (Spi-
ramiopsis, Dactyloceras, Brahmaea) or 0-2-2
(Calliprogonos, Acanthobrahmaea). Tibiae with
344
strong spines in Acanthobrahmaea and Brah-
maea; tarsi spined ventrally (sometimes also with
dorsal spines). Arolium present; pulvilli some-
times very narrow and undivided (Dactyloceras),
ventral edge of the claws simple or serrulate.
Male wing-coupling retinaculo-frenate in Dacty-
loceras, amplexiform in the remaining genera.
Spinarea absent. Forewing venation (Fig.
18.5H): Rs branches all on a common stem; R
either free or stalked with Rs (Ml may also be
stalked with Rs); M2 about equidistant between
M l and M3, or closer to Ml than to M3; discal
cell often very short; CuP replaced by a fold.
Hindwing: no humeral vein; Se + R and Rs ap-
proximate beyond the (very short) discal cell; an
oblique crossvein (R) often present between Sc
and Rs, either before or beyond end of discal
cell (e. g. Holloway 1986: Fig. 32); stem of M
sometimes present within cell; tubular CuP lack-
ing; 3 A present or absent. Abdomen: lateral bars
of tergum 1 not connected with sternum 2,
though strongly produced laterally (posteriorly);
sternum 2 with short broad apodemes; venulae
present or absent; A2 sometimes with a pair of
lateral, eversible pouches that contain hair-scales
(males of Dactyloceras). Male sternum 8 often
very small; tegumen and vinculum synscleritous;
sacculus well sclerotized; uncus sometimes bilo-
bate (e. g. Meli 1937); gnathos large, often hard
and echinate mesally (sometimes divided or bilo-
bate). Female genitalia: corpus bursae with or
without signa; ostium bursae on venter 8; apoph-
yses anteriores short to moderately long; apoph-
yses posteriores sometimes vestigial {Dactylo-
ceras).
Egg hemispherical, of the upright type (e. g.
Okada 1968); chorion smooth. Larva, in all or
most instars, with a pair of scoli (D) on T2 and
T3, a middorsal scolus (Dl) on A8 (e. g. Bilek
1965), and sometimes various other scoli; last
larval instar of Brahmaea without scoli (e. g.
Paukstadt & Paukstadt 1986); scoli undulating
(e. g. Kozlov 1985) or rather straight (e. g.
Schultze 1931), the thoracic ones rarely being
short (Schultze 1914; Oberprieler & Duke 1994);
last instar larva with long ventral prolegs that
bear numerous secondary setae (crochets in
biordinal mesoseries). Pupa: maxillary palpi and
forefemora concealed; galeae long (Spiramiopsis)
to fairly short; metanotum with conspicuous
sculpturing (e.g. Meli 1930; Schultze 1931;
Freina 1985; Paukstadt & Ragus 1990; Ober-
prieler & Duke 1994); forewing base with a
strong, acute tooth in Acanthobrahmaea; abdo-
men of this genus provided with three dorsal
rows of strong teeth; dorsocaudal part of AIO
conical or subtriangular, with irregular surface.
Pupation in the soil or in leaf litter (no genuine
cocoon).
The adults are essentially nocturnal. At rest,
the wings are usually more or less outspread.
However, Spiramiopsis comma has a "bombycid-
Claude Lemaire & Jol Minet
like" resting posture (Holloway 1987; Ober-
prieler & Duke 1994: Fig. 1). The known larvae
feed externally on the foliage of the Asclepiada-
ceae (Spiramiopsis, Dactyloceras) and Oleaceae
(.Acanthobrahmaea, Brahmaea). The family con-
tains five genera and about 20 species. Sauter
(1986) divided it into two subfamilies: Dactylo-
ceratinae and Brahmaeinae. Since Sauter's classi-
fication does not take account of Spiramiopsis,
and because his Dactyloceratinae (Dactyloceras
+ Calliprogonos) are likely to be para- or poly-
phyletic, it seems premature to recognize sub-
families in Brahmaeidae. Spiramiopsis (Fig.
18.6 C) is a monotypic genus from southeastern
Africa. It has been placed in Brahmaeidae by a
few authors (Jordan 1923; Hering 1927; Vri &
Kroon 1986; Nssig & Paukstadt 1990; Scoble
1992; Minet 1994), but into other families by
many others (e. g. Gaede 1927; Forbes 1955;
Fletcher & Nye 1982). The afro tropical genus
Dactyloceras (Fig. 18.6D) includes about 8 spe-
cies. Calliprogonos miraculosa (Fig. 18.6E) is a
rare insect inhabiting the Chinese province of
Shaanxi. The monotypic genus Acanthobrahmaea
is restricted to Mount Vulture, in southern Italy
(e. g. Dufay 1970 and Rougeot 1971). Brahmaea
(Fig. 18.6 F) contains large moths distributed
from Caucasus and Turkey to Amur, Japan and
Sulawesi; Inoue's (1982) concept of this genus
seems preferable to that of Nssig & Paukstadt
(1990).
Sphingidae. The monophyly of Sphingidae is
supported by many autapomorphies (Minet
1994): e. g. (a) ventral arm of the laterocervicale
ending abruptly, mesad, in a very thin rod (Mad-
den 1944: Fig. 18); (b) vein M2 arising slightly
nearer M3 than M1 in the forewing; (c) hindwing
margin produced or angulate at the tip of 1 A
+ 2 A, the termen being somewhat emarginate
beyond this vein (Fig. 18.5 J, whereas Fig. 18.51
represents a case of secondary loss of this apo-
morphy); (d) segments Al A7 of the larva pro-
vided with a pair of lateral, oblique stripes (as-
cending caudad; those of A7 reaching the mid-
dorsal scolus on A8); (e) exposed parts of the
hindwings not reaching segment A4 in the pupa.
The lateral stripes of the larva have sometimes
been considered an autapomorphy of the non-
macroglossine Sphingidae (Fig. 18.7E), but they
actually also belong to the ground plan of the
Macroglossinae, being well indicated in several
of their members (Figs, in Bell & Scott 1937, Is-
siki et al. 1965, etc.).
Medium-sized (or even fairly small) to very
large moths, with stout bodies, rather long
subtriangular forewings and comparatively short
hindwings (Figs. 18.6G-L). Antennae often la-
mellate ventrally, and either roughly filiform,
davate or pectinate (bi-, tri- or quadripectinate);
apex tapering, usually upturned; shaft densely
scaled dorsally. Typically, a male flagellomere
The Bombycoidea and their Relatives
bears each side, ventrally, two rows of long
trichoid sensilla which meet dorsally (one ante-
rior, the other one posterior: e. g. Fig. 4.7 in Ea-
ton 1988); such sensilla are absent or shorter in
the female, which is usually also distinguished by
shorter and thinner antennae. Compound eyes
naked or, occasionally, hairy (e. g. Dolbina).
Ocelli and chaetosemata absent (or practically
absent). Pilifers often elongate, with numerous
long bristles, but sometimes more or less re-
duced. Proboscis usually long to extremely long
(its length then exceeding considerably that of
the body), but sometimes short or vestigial; pro-
boscis base occasionally scaled. Maxillary palpi
small, unsegmented. Labial palpi ascending or,
rarely, porrect, usually with robust second seg-
ments and short third segments (however, seg-
ment 3 is long in, e. g., Xenosphingia), second
segments usually meeting on the meson through
their dense vestiture, which may also merge into
similar vestiture of frontoclypeus; inner surface
of segment 1 frequently with a patch of micro-
trichia; inner surface of segment 2 sometimes
modified (especially when pilifers correspond to
phonoreceptors). Mesothorax: anepisternum and
meron large to very large, the latter having a
long, oblique line of junction with the epimeron
(e. g. Shepard 1930: Fig. 33); anterior area of the
epimeron provided with well defined "preepi-
meron" (called "protomerum" by Niculescu: e. g.
1989, Fig. 63). Tibiae sometimes spined, the fore
ones often ending in a strong point; epiphysis
present; tibial spurs (long to short): 0-2-4 or 0-2-
2. Tarsi with ventral spines, which may form a
comb-like structure at base of basitarsus (on
mid- and hindlegs). Male forecoxa sometimes
with an androconial tuft. Pretarsus: claws simple
(i. e. without distinct teeth); arolium occasionally
absent (this organ is sometimes wrongly called a
"pulvillus"); pulvilli, when present, either bilo-
bate or undivided. Wing-coupling usually reti-
naculo-frenate: frenulum long, simple, in male,
multiple in female (both sexes occasionally with-
out frenular bristles: if so, base of frenulum pre-
served as a tubular thickening). Spinarea absent.
Wings sometimes with large, transparent areas,
due to a labile scaling that disappears soon after
adult emergence (e. g. most species of Hemaris).
Forewing venation (Fig. 18.51, J): R free; Rsl
and Rs2 either long-stalked or entirely fused;
Rs3 and Rs4 stalked (their stem always sepa-
rated from Rsl 4- Rs2); M2 arising slightly
closer to M3 than to M l ; M3 and CuAl sepa-
rate, the latter arising far from CuA2; tubular
CuP absent; 1 A + 2 A forked basally. Hindwing:
upper edge of discal cell connected to Sc by a
distinct crossvein (R); postdiscal section of Rs
and Se + R more or less approximated or, occa-
sionally, shortly fused; M2 arising about midway
between Ml and M3; M3 and CuAl usually sep-
arate, seldom stalked; CuP replaced by a fold;
3 A well developed. Abdomen rather conical,
345
sometimes with lateral or terminal fan-shaped
scale-tufts; lanceolate or spine-like scales often
present, notably on terga (usually along their
caudal margin); dorsum 1 with a median sclerite
varying from narrow to fairly broad, and lateral
bars that form, in the rear, long lateroventral
projections (along which the anterolateral areas
of tergum 2 are produced ventrad, being some-
times fused with sternum 2); sternum 2 usually
with rather long and slender apodemes, and very
short anterolateral processes; venulae present or
absent; in most males, sternum 2 provided with
posterolateral arms that carry, distally, pleural
androconial brushes (hidden at rest in longitudi-
nal folds); male segment 8 variously shaped, its
tergum being occasionally bordered with modi-
fied scales. Male genitalia sometimes asymmetri-
cal; valva usually with well defined sacculus and
subtriangular dorsobasal process; modified
scales often present on outer surface of valva;
gnathos rarely absent, frequently complete and
broadly fused with tegumen; uncus apex simple
or bilobate; phallus not fused to juxta. Female
genitalia: corpus bursae often with long signum
formed by two rod-like dentate arms; signum
sometimes short (e. g. Agrius) or wholly absent
(many Smerinthinae); ostium bursae lying on
sternum 8 or just cephalad of it; both pairs of
apophyses distinctly present.
Egg of the flat type, ovoid or ellipsoidal, with
a rather smooth chorion (eggs usually laid sin-
gly). First instar larva with a globular head cap-
sule, and usually a long middorsal scolus on A8
(which may be distally bifid); scoli rarely present
on other trunk segments; LI and L2 remote from
each other on most abdominal segments. Last in-
star larva (Fig. 18.7E): head hypognathous,
either subtriangular (e. g. most Smerinthinae) or
fairly globular; thorax and abdomen densely
covered with minute secondary setae, sometimes
arising from domed swellings; most segments di-
vided into annulets; meso- and metathorax
rarely with a dorsal pair of scoli (e. g. Ceratomia
amyntor; Peterson 1948); thorax and abdomen
occasionally provided with numerous, well de-
veloped scoli (Lophostethus); A8 usually with a
fairly long middorsal scolus (Dl) (occasionally
absent or replaced by a button-like structure);
abdomen often ornamented with oblique lateral
stripes or conspicuous laterodorsal eye-spots
(sometimes also present on the thorax - see e. g.
Pelzer 1991); proleg crochets in biordinal me-
soseries; anal prolegs usually enlarged. Pupa:
maxillary and labial palpi concealed (e. g.
Mosher 1918, Nakamura 1977); ocular sutures
usually obliterated caudad; antennae and mid-
legs never reaching forewing apices; hindlegs
normally concealed; galeae sometimes short, but
ordinarily long, reaching ends of forewings, and
often strongly extended forward (they may also
bear, ventrally, an unpaired tubular projection:
Fig. 18.7 H); forefemora either exposed or con-
346
cealed; metathorax often with pair of dorsal pro-
tuberances (Pelzer 1996: Fig. 1); exposed por-
tions of hindwings never reaching A4 (see Figs,
in, e. g., Pitta way 1993); A5A7 (or only one
or two of these segments) often provided with
spiracular furrows, i. e. prespiracular grooves
parallel to intersegmental folds; AIO dorsum
usually subtriangular or conical (e. g. Zolotuhin
1994 b: Fig. 15), occasionally truncate or almost
rounded. Pupation usually occurs in the soil or
in leaf litter, rarely in a silken cocoon spun on
the food plant (e. g. Kendall 1976).
The adults are usually crepuscular or noctur-
nal, but sometimes exclusively diurnal (see e. g.
Li 1994). The species that have a well developed
proboscis use it for imbibing nectar from flow-
ers, hovering in front of these as do humming
birds; this behaviour may permit them to avoid
certain predators (Wasserthal 1994). In flight,
Sphingidae or "hawk-moths" are among
the fastest insects, and several are well-known
migrants (e. g. Reinhardt & Harz 1989). In sev-
eral genera, the larva rests with the anterior part
of the body slightly or widely remote from the
substrate.
The family occurs worldwide, but is better rep-
resented in tropical areas. It contains about 200
genera and at least 1200 species (I. J. Kitching,
pers. comm.). Rothschild & Jordan (1903) di-
vided the Sphingidae into two large groups, viz.
Asemanophorae and Semanophorae, containing
respectively two (Acherontiinae, "Ambulicinae")
and three ("Sesiinae", Philampelinae and Choe-
rocampinae) subfamilies. This classification has
been accepted by many authors, sometimes with
a different nomenclature. Thus, Zerny & Beier
(1936) rightly used Sphinginae instead of Ache-
rontiinae, and Smerinthinae instead of "Ambuli-
cinae" (correct spelling: Ambulycinae). More re-
cently, Asemanophorae and Semanophorae were
changed respectively to Sphinginae and Macro-
glossinae (Hodges 1971), the subfamily status of
these taxa having been proposed previously by
two authors (Janse 1932, Carcasson 1968), al-
though with an incorrect nomenclature. How-
ever, contrary to Macroglossinae, Sphinginae
(sensu Hodges) are probably paraphyletic, being
based mainly on a trait always found in the other
bombycoid families (viz. absence of microtrichia
from the inner surface of segment 1 of the labial
palpus). Following Minet (1994), the family is di-
vided into three subfamilies: Smerinthinae,
Sphinginae and Macroglossinae (the former two
sensu Zerny & Beier 1936, i. e. with the same
composition as the "Ambulicinae" and Acheron-
tiinae of Rothschild & Jordan 1903). For the
adults the following key can be used:
Key to the subfamilies
1 Labial palpus with patch of microtrichia on
inner surface of first segment; in male geni-
Claude Lemaire & Jol Minet
talia, outer surface of valva often provided
with row of large, lanceolate scales (some-
times with several rows of lanceolate scales)
Macroglossinae
Labial palpus without internal microtrichia
on first segment; valva never with row of
large, lanceolate scales (although sometimes
with modified scales) 2
2 Antenna with long terminal segment (i. e. at
least four times as long as basally high). . 3
Antenna with shorter terminal segment . . 4
3 Forewing without dark longitudinal streaks
in intervals M 3 - C u A l and C u A l - C u A 2 ;
its apex often truncate or falcate, and its in-
ner margin always markedly concave; fringe
evenly coloured along forewing termen;
sides of abdomen sometimes with oblique
stripes, but always devoid of conspicuous,
transverse, dark and light markings
Smerinthinae (in part)
Characters in a different combination; fore-
wing apex neither truncate nor f a l c a t e . . . .
Sphinginae (in part)
4 Forewing termen evenly (often weakly) con-
vex, with its fringe distinctly lighter between
apices of veins; inner margin of this wing
straight to very weakly concave; forewing
often with a dark longitudinal streak in each
of intervals M 3 - C u A l and C u A l - C u A 2 ;
tibial spurs usually short (long in Oligogra-
pha, a minor African genus)
Sphinginae (in part)
Characters in a different combination; fore-
wing without longitudinal streaks in in-
tervals M3CuAl and C u A l - C u A 2 ; tibial
spurs long to very short
Smerinthinae (in part)
Smerinthinae (Figs. 18.6HJ) often possess a
reduced proboscis. The forewing termen is fre-
quently wavy or irregular, though evenly convex
in a few taxa, e. g. Leucophlebia (Fig. 18.51) and
Xenosphingia (Fig. 18.6 J). Pectinate antennae
are occasionally present, sometimes only in cer-
tain species of a given genus (e. g. Ceridia). At
rest, the forewings are usually remote from the
abdomen (sometimes held in a bombycid-like
manner: Holloway 1987). The adults of most Sa-
taspes species mimic carpenter bees of the genus
Xylocopa (e. g. Riotte 1981, Li 1994). After its
first instar, the larva has a rather conical or tri-
angular head capsule (except in the final instar
of a few genera - see Rothschild & Jordan 1907:
PI. 3). The larval hostplants belong to various
families: Anacardiaceae, Bombacaceae, Faba-
ceae, Fagaceae, Lauraceae, Malvaceae, Mora-
ceae, Salicaceae, Tiliaceae, etc. (e. g. Bell & Scott
1937, Holloway 1987). Main smerinthine genera:
Protambulyx, Adhemarius, Ambulyx, Clanis, Po-
lyptychus, Phylloxiphia, Marumba and Smerin-
thus (D'Abrera 1987).
The Bombycoidea and their Relatives 347

Fig. 18.7. Second instar (C) / last instar (A, B, D, E) larvae and pupae ( F - H ) of Bombycidae (A, B, F, G),
Mirinidae (C), Saturniidae (D) and Sphingidae (, H): A, Phiditia minor-, B, Andraca bipunctata; C, Mirina christo-
phi; D, Citheronia phoronea; E, Acherontia lachesis; F and G, Gastridiota adoxima; H, Cocytius antaeus (A and H,
after Burmeister 1879; , after Kaishoven et al. 1951; C, after Kozlov 1985a; D, drawn from a photograph; E,
after Dupont & Roepke 1941; F and G, after Common & Edwards 1991). Scales A, B, D, , H = 10 mm; C =
5 mm; F, G = 2 mm (G. Hodebert).

Sphinginae (Fig. 18.6 G) usually possess a very
long proboscis, often several times as long as the
body (e. g. Xanthopan). In Acherontia, the imago
produces characteristic squeaks using its short,
stout proboscis. Closely related to this genus,
Agrius and Coelonia possess however a long pro-
boscis. At rest, the wings are held more or less
roofwise. The larva only rarely has a subtriangu-
lar head (Lapara). Among the main hostplant
families are the Asteraceae, Bignoniaceae, Con-
volvulaceae, Fabaceae, Lamiaceae, Oleaceae, So-
lanaceae and Verbenaceae (e. g. Moss 1920 and
Holloway 1987). Main sphingine genera: Cocy-
tius, Manduca and Sphinx (D'Abrera 1987).
Macroglossinae always have a well developed,
functional proboscis. Three tribes (Dilophono-
tini, Philampelini, Macroglossini) were recog-
nized in the subfamily by Hodges (1971), but at
least one of these is paraphyletic (I. J. Kitching,
pers. comm.) (a few subtribes have also been
proposed: see Carcasson 1968, Derzhavets 1984
and Pittaway 1993). Some of these moths are
day-flying (e. g. Hemaris, most Cephonodes and
many Macroglossum). At rest, the dorsal surface
of the abdomen is broadly to entirely exposed
and the inner margins of the forewings lie either
along the abdomen or at a short distance from
it (see e. g. Chang 1989). The male genitalia are
sometimes asymmetrical. Several genera (Choe-
rocampini) have pilifers transformed into ultra-
sonic receptors (e. g. Roeder et al. 1970; Cook &
Scoble 1992), for example Xylophones, Hyles,
Deilephila (Fig. 18.6L) and Hippotion. The larva
has a globular head and, frequently, conspicuous
thoracic or abdominal eye-spots. Occasionally,
the pupa is protruded before emergence (Brock
1990). Hostplants are very diverse: e. g. Apocy-
naceae, Araceae, Dilleniaceae, Onagraceae, Ru-
348
biaceae and Vitaceae (see Haxaire & Rasplus
1987 a and b, Lin 1987, Inoue et al. 1997, etc.).
Main macroglossine genera: Isognathus, Erin-
nyis, Callionima, Enyo, Nyceryx, Perigonio, He-
mar is, Cephonodes (Fig. 18.6K), Eumorpha,
Daphnis, Acosmeryx, Eupanacra, Nephele, Tem-
nora, Macroglossum, Xylophones, Hyles, Hippo-
tion, Theretra and Rhagastis (D'Abrera 1987).
Acknowledgements
We are indebted to many colleagues including
Mr J. Boudinot ( M N H N , Paris), Mr N . D u k e
(Mbabane), D r . Gustafsson ( N R , Stockholm),
M. J. Haxaire (Agen), Mr D . Herbin (Castanet-
Tolosan), D r J. D . H o l l o w a y (CABI, London),
Mr P. Kindl (Paris), D r W. A . N s s i g ( S N G ,
Frankfurt a m Main), Dr C. M . N a u m a n n
( M A K , Bonn), D r E. S. Nielsen (CSIRO, Can-
berra), D r R. G. Oberprieler (PPRI, Pretoria),
and D r R. S. Peigler ( D M N H , Denver, Co.) -
for providing valuable information and/or speci-
mens o f crucial importance. We also thank Mrs
M. Franey ( M N H N , Paris), w h o prepared the
photographic prints, and Mr G. Hodebert
( M N H N , Paris), the talented artist w h o executed
the wing venation drawings. We are especially
grateful to those w h o reviewed the manuscript,
in particular D r I. J. Kitching ( N H M , L o n d o n )
and D r M . J. Scoble ( N H M , London), and to
the v o l u m e organizer, D r . P. Kristensen
( Z M U C , Copenhagen), for valuable suggestions
and encouragement.
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 19. The Noctuoidea
Ian J. Kitching & John E. Rawlins
Moths of the superfamily Noctuoidea constitute
the most speciose and cosmopolitan lineage of
Lepidoptera, with adults and larvae exhibiting
an often bewildering diversity of sizes, color-
ation, behaviour and ecological habits. Histori-
cally classified using a small number of superfi-
cial characters (e. g. wing venation, eye vestiture,
leg spines), these moths comprise the traditional
large families Noctuidae (popularly known as
owlets, cutworms, underwings, deltoids), Noto-
dontidae (prominents, processionary moths),
Arctiidae (tiger-moths, handmaidens, footmen,
ermines) and Lymantriidae (gypsy moths, tus-
socks). Over 7,200 genera have been proposed
and nearly 70,000 species have been described.
The genera of all noctuoid families were cata-
logued by Nye (1975) and Watson, Fletcher &
Nye (1980). Species catalogues exist for Noctui-
dae, Nolidae and Pantheidae (Poole 1989), Aga-
nainae (Gaede 1932), Arctiidae (Bryk 1931, 1937;
Strand 1919, 1922; Watson & Goodger 1986;
Zerny 1912), Lymantriidae (Bryk 1934) and No-
todontidae (Bryk 1930; Gaede 1934).
The monophyly of Noctuoidea is unequivo-
cally established by the apomorphic presence of
metathoracic tympanal organs and associated
abdominal structures (Figs. 19.1 A - C , 19.2
- B ) . The tympanum perceives ultrasound over
a range of frequencies. Its primary function is
considered to be perception of the echolocation
signals of bats (Fenton & Fuller 1981; Spangler
1988). However, there is increasing evidence that
the tympanum may also be involved in reception
of mating signals (Sanderford & Conner 1990).
On each side, the metepimeron is modified poste-
riorly to form a cavity, within which lies the
tympanic membrane, which is usually separated
from the metepimeron by the nodular sclerite or
epaulette (Eggers 1919; Richards 1933). Anterior
to the nodular sclerite, the metepimeron may be
membranous ("accessory tympanum" of Rich-
ards 1933). The chordotonal organ comprises
one or two scolopidia. Dorso-medial to the tym-
panum and connected to it via the same air sac
is the counter-tympanic membrane, which prob-
ably acts as an accessory resonating organ (Eg-
gers 1919). The anterior face of abdominal seg-
ment 1 may be deeply invaginated to form a pair
of counter-tympanal cavities (antitympanal cavi-
ties of Minet 1986; abdominal bullae of Miller
1991). Laterally, there may be a sclerotized flap,
the counter-tympanal hood, which may be ante-
rior to (prespiracular; Fig. 19.1 C) or posterior to
(postspiracular; Fig. 19.1 B) the first abdominal
spiracle, and functions to localize incoming
sounds. In fact, precise interpretation of counter-
tympanal hood position relative to the spiracle is
problematic (e. g. Aganainae, many Herminii-
nae) as the spiracle may be below the hood, in
intermediate configurations, or the hood reduced
and its position unclear (Richards 1933). These
configurations are probably not homologous,
rendering dubious phylogenetic inferences based
on treating hood position as either prespiracular
or postspiracular (Speidel et al. 1996 a, notwith-
standing).
The presence of two M D setae on the larval
metathorax (Hinton 1946) is a consistent and re-
liable synapomorphy of all Noctuoidea except
the Australian Oenosandridae (Miller 1991)
(Fig. 19.14 A). But it is difficult to score in taxa
with numerous secondary setae. Other features
quoted as diagnostic for Noctuoidea are plesio-
morphic (relatively unmodified venulae; posteri-
orly complete ventral laminae of the metathora-
cic furca; forewing areole) (Brock 1971).
The basal noctuoid lineages. The fossil record
for Noctuoidea is too meagre and young (Middle
to Late Tertiary) to be phylogenetically informa-
tive. A few genera of Holarctic fossils have been
proposed: Arctiites, Stauropolia (Fig. 2.8) and
Oligamatites (Arctiidae); Cerurites (Notodonti-
dae); Noctuites and Phylledestes (Noctuidae). An
egg from the Cretaceous has been attributed to
Noctuidae (Gall & Tiffney 1983), but that deter-
mination is not based on a diagnostic apomor-
phy for noctuoid eggs and remains dubious.
Within Noctuoidea, there are two basic pat-
terns of forewing venation. In the plesiomorphic
state, MA2* arises midway between MAI and
MPI (Fig. 19.3 A), giving an apparent trifurca-
tion at the lower corner of the discal cell. This is
referred to as the "trifid" condition and occurs
in Oenosandra, Doidae and Notodontidae. In
contrast, the form in which MA2 arises very
close to, connate with or stalked with MPI, giv-
ing an apparent four-branched pattern, is re-
ferred to as the "quadrifid" condition and occurs
in all other noctuoids (Figs. 1 9 B - C , E - H ) ex-
cept derived lineages showing fewer branches
through fusion (Fig. 19.3 D).
The structure of the tympanum also shows
two basic forms. In Doidae, Notodontidae (Figs.
19.1 A, 19.2 A) and some Syntominae (Arctii-
dae), the metepimeron is strongly concave dor-
sally, causing the tympanic membrane to be ori-
ented ventrally, while the nodular sclerite and
counter-tympanal hood are absent (Kiriakoflf
* The vein notations M A I , M A 2 und MPI in this
chapter equal, respectively, M l , M 2 and M 3 used else-
where in this volume. Editor.
356 Ian J. Kitching & John E. Rawlins

Fig. 19.1. Metathoracic tympana and associated structures in Noctuoidea (external aspect, left lateral view). A,
Horizontal tympanum (Notodontidae, Heterocampinae, Schizura unicornis), B, Oblique tympanum with post-
spiracular counter-tympanal hood (Noctuidae, Catocalinae, Erebus ephesperis); C, Oblique tympanum with en-
larged, prespiracular counter-tympanal hood, (Arctiidae, Arctiinae, Ctenuchini, Ctenucha virginica). ac = acces-
sory tympanum, ct = counter-tympanic membrane, epm = metepimeron, h = counter-tympanal hood, sp = spira-
cle on first abdominal segment, t = tympanum.

1949; Miller 1991). In Oenosandra and the quad-
rifid families (Figs. 19.1 B; 19.2 B - C ) , the mete-
pimeron is only moderately concave, if at all, so
that the tympanum faces posteriorly. A counter-
tympanal hood and nodular sclerite are present.
The notodontid tympanum is innervated by a
single acoustical sensory cell, while the noctuid
tympanum is innervated by two such cells (Sur-
lykke 1984). However, the distribution of this
character is inadequately known.
We recognize three fundamental lineages of
Noctuoidea: Oenosandridae, Doidae + Noto-
dontidae, and the quadrifid families (here treated
as Arctiidae, Lymantriidae, Noctuidae, Nolidae
and Pantheidae). Miller (1991) favoured a clade
comprising Oenosandra and Doidae + Notodon-
tidae, on the basis of two synapomorphies: pres-
ence of a stipital lobe in the larvae (Godfrey
et al. 1989) (Fig. 19.13C) and presence of socii
(Fig. 19.11 D). However, both features are prob-
lematic. Oenosandra has a stipital structure that
is scarcely different from that of some noctuids
and lymantriids. It is also not clearly homolo-
gous with the prominent stipital lobes of many
other lymantriids, notodontids and some nolids.
In addition, although Miller (1991) stated that
The Noctuoidea 357

aepm
aepm /
epm eprn

Fig. 19.2. Internal view of the noctuoid tympanum. A, Antheua simplex, Notodontidae: Phalerinae; B, Psaphida
electilis, Noctuidae: Psaphidinae. 1 = pocket I, 2 = pocket II, 3 = pocket III, 4 = pocket IV, ac = accessory
tympanum, aepm = anapre-epimeron, ct = counter-tympanic membrane, epm = metepimeron, = nodular scle-
rite, ph = metascutal phragma, pn = metapostnotum, sb = subalare, sc = metascutum, sci = metascutellum, t =
tympanum, tps = tergopleural suture of first abdominal segment.

socii "do not occur in quadrifid noctuoid
groups", he did not address the homology of
similar structures in orgyiine Lymantriidae (Fer-
guson 1978). Furthermore, Tikhomirov (1979 b)
demonstrated that genitalic muscle M1 inserts on
the socii of notodontids rather than on the base
of the uncus as in many other Lepidoptera, sup-
porting the hypothesis that socii are not homolo-
gous in these groups. Socii in Oenosandra appear
similar to those in notodontids but the homolo-
gies in Doa remain problematic. Knowledge of
genitalic musculature in outgroup taxa is insuffi-
ciently documented. Other potential synapomor-
phies of Oenosandra, Doidae and Notodontidae,
such as the size of the saccus and the position of
larval head seta SI relative to stemma 3 and 4,
show reversal in some subfamilies of Notodon-
tidae.
In contrast, the presence of two MD setae on
the larval metathorax is a widely examined and
uniformly consistent feature supporting the mo-
nophyly of all noctuoids except Oenosandra. This
hypothesis, which is accepted here, implies that
the trifid venation and socii in Oenosandra, doids
and notodontids, and the tympanal configura-
tion in Oenosandra and the quadrifid lineages,
are symplesiomorphic.
Key to adult Noctuoidea at subfamily level
1 Forewing vein MA2 arising midway be-
tween MAI and MPI (trifid) 2 then valve broad, medially membranous,
1' Forewing vein MA2 arising very close to,
connate with or stalked with MPI (quadri-
fid) 10
2 Metepimeron strongly concave dorsally,
tympanic membrane facing ventrally; nod-
ular sclerite absent 3
2' Metepimeron only moderately concave or
not at all, tympanic membrane facing pos-
teriorly; nodular sclerite present
Oenosandridae
3 Sclerotized apices of tibial spurs smooth
Doidae
3' Sclerotized apices of tibial spurs serrate
4
4 In lateral view, junction between antennal
flagellar segments oblique; female T7
membranous with a tuft of deciduous
scales . . . Notodontidae: Thaumetopoeinae
4' In lateral view, junction between antennal
flagellar segments transverse; female T7
sclerotized without a tuft of deciduous
scales 5
5 A small, lightly sclerotized, setose patch
present in the pleuron above the first abdo-
minal spiracle; male abdomen without a
terminal scale tuft
Notodontidae: Dioptinae
5' Area above Al spiracle not setose; male
abdomen with a terminal scale t u f t . . . . 6
6 Manica heavily sclerotized, fused with
juxta to form a ring that tightly surrounds
the aedeagus . Notodontidae: Nystaleinae
6' Manica almost entirely m e m b r a n o u s . . . 7
7 Ocelli present and large; if rudimentary,
sclerotization restricted to costal and sac-
cular margins . . Notodontidae: Dudusiinae
7' Ocelli rudimentary or absent 8
358 Ian J. Kitching & J o h n E. Rawlins

Fig. 19.3. Wing venation in Noctuoidea (male wings except H). A, Trifid condition in b o t h wings, Anaphe (Noto-
dontidae, Thaumetopoeinae); B, Quadrifid condition in b o t h wings, Calidota (Arctiidae, Arctiinae); C, Quadrifid
f r o n t wing a n d trifid hind wing, Virbia (Arctiidae, Arctiinae); D, Reduced venation, Crambidia (Arctiidae, Lithosii-
nae); E, Enlarged costal region in front wing, Selepa (Nolidae); F, Quadrifine hind wing venation, Aon (Noctuidae,
Catocalinae); G , Quadrifine hind wing venation, Diloba (Noctuidae, Dilobinae); H, Trifine hind wing venation
(note multiple bristles in female frenulum), Tiracela (Noctuidae, Hadeninae); I, Absence of subcosta in hind wing,
Balacra (Arctiidae, Syntominae).

8 Uncus large, triangular, fused with socii; 11 A pair of glands invaginated dorsally just
female A8 fused into a heavily sclerotized anterior to the anal papillae 12
capsule Notodontidae: Phalerinae 11' Such glands absent 14
8' Uncus not fused with socii; pleuron of fe- 12 Branches of dorsal glands anterior to the
male A8 membranous, or if sclerotized, anal papillae short and broad, about as
then segment not capslate 9 long as wide Arctiidae: Lithosiinae
9 Female epiphysis absent 12' Branches long and narrow, more than
Notodontidae: Pygaerinae three times as long as wide 13
9' Female epiphysis present Notodontidae: 13 Hindwing veins Sc, Rs and MAI com-
Heterocampinae and Notodontinae pletely fused Arctiidae: Syntominae
10 Metascutal bulla present Notodontidae: 13' Hindwing veins Sc, Rs and MAI separate,
Platychasmatinae or only Sc and Rs fused, with a strong
10' Metascutal bulla absent 11 MAI Arctiidae: Arctiinae
The Noctuoidea 359

Fig. 19.4. Retinacula in male Noctuoidea. A, Catocala (Noctuidae, Catocalinae); B, Asola (Noctuidae, Aganai-
nae); C, Amastus (Arctiidae, Arctiinae); D, Earias (Nolidae, Eariadinae).

14 Male antenna pectinate, with 1 - 3 diver- 18' Ocelli present, large 20

gent setae apically on each ramus 19 Scaphium with lateral setose patches (lost
Lymantriidae in Nola) Nolidae: Nolinae
14' Male antenna simple; if pectinate, then api- 19' Scaphium without setose patches
cal setae not divergent 15 Nolidae: Afridinae
15 Male retinaculum elongate and narrow, 20 Frenulum and retinaculum absent
more than two times as long as wide ("bar- Nolidae: Eariadinae
shaped") 16 20' Frenulum and retinaculum present . . . 21
15' Male retinaculum wider than long . . . 24 21 Tegumen with coremata; sacculus with
16 Lower part of clypeofrons with a small but "bird's head"-shaped structures
distinct scaleless area Nolidae: Bleninae
Noctuidae: Aganainae 21' Tegumen without coremata; sacculus with-
16' Lower part of clypeofrons scaled . . . . 17 out such structures 22
17 Valves fused to vinculum 22 Distal margin of valve without a zone of
Nolidae: Eligminae basally-directed setae; basal abdominal
17' Valves not fused to vinculum 18 tymbal organs often present
18 Ocelli rudimentary or absent 19 Nolidae: Nolinae
360
22' Distal margin of valve with a zone of bas-
ally-directed setae; basal abdominal tymbal
organs never present 23
23 Uncus slender and sinuous; ovipositor
lobes not ring-like and highly setose . . . .
Nolidae: Risobinae
23' Uncus highly setose with a terminal hook,
or absent; ovipositor lobes ring-like and
highly setose . Nolidae: Westermanniinae
24 Lower part of clypeofrons with a distinct
scaleless area 25
24' Lower part of clypeofrons scaled . . . . 30
25 Proboscis heavily sclerotized and sharply
pointed apically, armed with tearing hooks
and with basiconic sensilla modified into
erectile barbs Noctuidae: Calpinae
25' Proboscis not heavily sclerotized and blunt
apically; basiconic sensilla unmodified 26
26 Counter-tympanal hood double; ductus
ejaculatorius with a small oval plate api-
cally; anal papillae modified so that the in-
terior surfaces face posteriorly 27
26' Counter tympanal hood single; ductus
ejaculatorius without a small oval plate;
anal papillae unmodified 28
27 Basal abdominal sternite without internal,
longitudinal flanges; female frenulum com-
prising a single seta; anal papillae apically
divided with numerous setae on elevated
bases Noctuidae: Stictopterinae
27' Basal abdominal sternite with a central
membranous area flanked by two internal,
longitudinal flanges; female frenulum with
more than 1 seta; anal papillae apically en-
tire Noctuidae: Euteliinae
28 Metepimeron strongly swollen ventral to
pocket IV of the tympanal organ; counter-
tympanal hood anterior to dorsal in posi-
tion relative to Al spiracle
Noctuidae: Herminiinae
28' Metepimeron not strongly swollen ventral
to pocket IV; counter-tympanal hood di-
rectly posterior to Al spiracle 29
29 Antenna distally widened so as to appear
somewhat clubbed; wings with large
transparent areas and metallic scaling . . .
Noctuidae: Cocytiinae
29' Antenna not distally widened; wings fully
scaled Noctuidae:
Catocalinae, Hypeninae, Strepsimaninae
30 Counter-tympanal membrane greatly en-
larged; base of abdomen with a paired vesi-
cle (bulla abdominalis)
Noctuidae: Agaristinae
30' Counter-tympanal membrane not en-
larged; bulla abdominalis absent . . . . 31
31 Counter-tympanal hood absent
Pantheidae
31 ' Counter-tympanal hood present . . . . 32
32 Antennae davate . Noctuidae: Eucocytiinae
32' Antennae various but never davate . . 33
33 Pocket IV of tympanum greatly enlarged
Ian J. K i t c h i n g & J o h n E. R a w l i n s
and fused to wall of the tympanal air-sac;
counter-tympanal hood with a pouch-like
secondary hood ventrally
Noctuidae: Plusiinae
33' Pocket IV not enlarged and fused to tym-
panal air-sac; ventral secondary hood ab-
sent 34
34 Female abdomen with a terminal tuft of
deciduous scales . . Noctuidae: Dilobinae
34' Female abdomen without such a tuft . 35
35 An enlarged, heavily-sclerotized alula pre-
sent overlying the tympanum
Noctuidae: Acontiinae
35' Heavily sclerotized alula absent 36
36 Base of uncus with a pair of small, well-
defined lateral lobes
Noctuidae: Condicinae
36' Base of uncus without lateral lobes . . 37
37 Resting posture with forewings overlap-
ping so that the costae are parallel to the
long axis of the body
Noctuidae: Noctuinae
37' Resting posture tectiform with dorsal
edges of forewings meeting along the mid-
line and the costae divergent posteriorly
. . . . Noctuidae: most tritine subfamilies
(Acronictinae, Bryophilinae, Raphiinae, Eu-
strotiinae, Amphipyrinae, Bagisarinae, Cu-
culliinae, Stiriinae, Heliothinae, Psaphidi-
nae, Glottulinae, Hadeninae, Ufeinae)
The Trifid Noctuoidea
Oenosandridae. The Australian genus, Oenosan-
dra, includes 1 - 3 species that have been mis-
placed in Thaumetopoeinae (Common 1990) due
to the presence in females of both taxa of a cor-
ethrogyne. However, Miller (1991) stated that
the two types of tuft were not homologous and
excluded Oenosandra from Notodontidae, plac-
ing it in a separate family. Oenosandra is consid-
ered the sister group of other Noctuoidea be-
cause it has only a single MD seta on larval seg-
ment T3 (Kitching & Rawlins pers. obs.).
Furthermore, the biserial and biordinal crotchets
are a unique configuration in Noctuoidea.
Adults are sexually dimorphic (Figs. 19.7 AB).
Eggs have faint longitudinal sculpture and are
laid in clusters covered with scales from the cor-
ethrogyne (McFarland 1972). The gregarious lar-
vae rest by day under loose bark of their Euca-
lyptus foodplants, emerging at night to feed on
mature leaves (McFarland 1979). The Australian
genus, Discophlebia, also belongs in this family
(Edwards 1996) although it differs greatly in sev-
eral features (especially oviposition and chori-
onic peculiarities of the eggs (McFarland 1972)).
Doidae. Characters shared by Doa and quadrifid
noctuoids (e. g. lack of a metascutal bulla) are
plesiomorphic (Miller 1991), but the tympanal
The Noctuoidea

Fig. 19.5. Head and associated structures of adult Noctuoidea. A, Lateral view of head showing naked frons,
Ogovia (Noctuidae: Catocalinae); B, Apex of haustellum modified for piercing mammalian skin, Calyptra eustri-
gata (Noctuidae, Calpinae, after Bnziger 1970); C, Unmodified apex of noctuid haustellum, Lygephila craccae
(Noctuidae, Catocalinae, after Bnziger 1970). D - G , Apices of antennal rami: D, Quadricalcarifera (Notodonti-
dae, Heterocampinae); E, Trisuloides (Pantheidae); F, Nyctemera (Arctiidae); G, Euproctis (Lymantriidae).

structure of Doa and Notodontidae is synapo-
morphic. These two taxa also share two apomor-
phies of larval head chaetotaxy: the distance be-
tween CI and PI is about 1 the distance be-
tween CI and Af2 (also found in Lithosiinae,
Nycteola and elsewhere), and seta P2 is further
from the epicranial suture than PI (although the
state in Oenosandra is problematic). In previous
classifications, Doa (5 spp.) has been included in
various other families, reflecting the uncertainty
of its phylogenetic relationships. Only recently
has Doa, and Leuculodes (Franclemont 1983),
been placed in a separate family (Donahue &
Brown 1987). Species are found from western
North America to northern South America in-
cluding Cuba. All doids are fragile moths with
thinly scaled wings (Fig. 19.7 C). The contrast-
ingly coloured larvae have biordinal crotchets
and feed communally in a silken web on leaves
of Euphorbiaceae (Brown 1990). Eggs are oblate,
without longitudinal sculpture and are laid di-
rectly on the hostplant. Pupation occurs in a spa-
cious, open-meshed cocoon amid debris. The
pupa is incompletely obtect with secondary setae
in ventral abdominal Verrucae. The terminal ab-
dominal segment is rounded, with numerous
hooked setae in some species, reduced in others.
Notodontidae. The monophyly of Notodontidae
is supported by several apomorphies (Miller
1991). In adults, the tibial spur apices have ser-
rate margins (also present in Lymantriidae, Pan-
theidae and others) and a metascutal bulla is pre-
sent (except Dudusiini). All larval notodontids
(except Dioptinae) can be recognized by the
presence of two M D setae on Al (Fig. 19.14 A);
other noctuoids have only one M D seta (Hinton
1946). All notodontids (except Dudusinae) have
seta X (Gerasimov 1935) in the anterolateral cor-
ner of the anal shield (Fig. 19.15 A). Early instars
of Doa lack seta X, but the condition is obscured
in mature larvae by secondary setae. In addition,
most notodontids have only one SV seta on T2
and T3, although reversal to the plesiomorphic
bisetose condition occurs in Heterocampinae
and other genera. Late instar notodontid larvae
have a smooth mandibular cutting edge, al-
though the ontogenetic change from a plesio-
morphic serrate mandible is suppressed in sev-
eral genera (Godfrey et al. 1989). The everted ad-
enosma (cervical gland of authors) is dorso-
ventrally compressed and bifid in all notodontids
except Dioptinae (Godfrey & Appleby 1987).
Uniquely among noctuoids, larvae of many No-
todontidae have the anal prolegs highly modified
as stemapods, often with eversible distal glands.
However, there seems to be little phylogenetic
pattern to the degree of modification. Closely re-
lated taxa often differ considerably (Miller 1991).
Similarly, while lateral flanges on the spinneret
362 Ian J. Kitching & John E. Rawlins

Fig. 19.6. Tymbal and sternal structures in the Noctuoidea. A, Subventral abdominal tymbal, subventral region
on left side of abdominal segment 3, external view, scales removed (Lymantriidae, Staetherinia); B, Basal abdomi-
nal tymbal, left side of abdominal segments 1 and 2, anterior view (Nolidae, Chloephorinae, Carea); C, Metathora-
cic tymbal on katepisternum, lateral external view (Arctiidae, Arctiinae, Bertholdia).

occur in m a n y notodontids, their occurrence is
inconsistent with other characters (Miller 1991).
Eggs of all groups of notodontids are surpris-
ingly u n i f o r m in shape, being hemispherical with
smooth chorions that lack conspicuous sculptur-
ing. The subfamilial classification presented here
is a preliminary one following Miller (1991), who
recognized eight subfamilies within Notodonti-
dae. However, he did not discuss the placement
for several Old World groups, such as Acrocteni-
nae, Ceirinae, Desmeocraerinae and Hylaeorinae.
Thaumetopoeinae. This strictly Old World sub-
family (Fig. 19.7 DE), the sister g r o u p of all
other Notodontidae, is diagnosed mainly by re-
duction characters. However, the female tergite
7 is m e m b r a n o u s with a corethrogyne, the hairs
of which are used to cover the egg masses. Al-
though this feature occurs in other noctuoid taxa
(notably Oenosandra, Diloba and Lymantriidae),
within N o t o d o n t i d a e it m a y be apomorphic for
Thaumetopoeinae. Larvae (Fig. 19.16 A) are
mostly gregarious, feeding on leaves of trees in
m a n y families, especially Apocynaceae, Faba-
ceae, Myrtaceae and Sterculiaceae. Some build
c o m m u n a l silk nests, f r o m which they sortie in
columns to feed, hence the c o m m o n name of
processionary m o t h s (Gmez 1986). The larval
setae of a number of species can cause urticaria
(Carter 1984). Pupation occurs in a dense co-
coon, which in Afrotropical species of Anaphe
are spun communally, with m a n y cocoons en-
closed in a tough outer silken envelope.
The Noctuoidea
All other notodontids (Fig. 19.1 A) have a
membranous cup surrounding the spiracle of A l .
This may be homologous with the counter-tym-
panal hood of other noctuoids. It is absent in
Dioptinae (Miller 1991). The apparent counter-
tympanal hood reported in Pterostoma by Minet
(1986) appears to be a particularly well-devel-
oped cup. Other synapomorphies include an ex-
panded larval anteclypeus, with several dorso-
ventral folds (Miller 1991), and males with a very
long A8A9 intersegmental membrane (except
Platychasma). This facilitates a distinctive mode
of copulation whereby males use segment 8 to
grasp the female (Miller 1988). Some members
of all notodontid subfamilies except Thaumeto-
poeinae also have a pleated sacculus enclosing
androconial organs. Although most authors have
considered this feature apomorphic for Noto-
dontidae (Holloway 1983; Weiler 1990), it has a
highly homoplastic distribution within the family
(Miller 1991).
Pygaerinae. Because most features used to
characterize Pygaerinae (hairy larvae, male ab-
domen ending in a tuft, Fig. 19.7 F) are plesio-
morphic, this subfamily is poorly delimited. Only
Clostera (= Melalopha, Ichthyura) and Pygaera
can be included with certainty. They have char-
acteristic accordion-like saccular pleats on the
male genitalia and setose tubercles on larval seg-
ments A l and A8 (Miller 1991). Micromelalopha,
and perhaps species of Rosema near R. deolis,
may also be included. Known larvae (Fig.
19.16 B) feed on woody plants including Salica -
ceae, Proteaceae and Fabaceae.
Larvae of taxa phylogenetically distal to Py-
gaerinae possess an extremely deep labral notch
that sometimes divides the labrum completely
(Fig. 19.13 E). Deciduous stellate cornuti on the
vesica only occur in members of this clade, al-
though they are by no means universal (Hollo-
way 1983; Miller 1991). These cornuti become
detached during copulation and remain in the
corpus bursae (Fig. 19.11 C). Their function is
unknown although it is usually thought that they
act to deter subsequent mating (Miller 1991).
Platychasmatinae. This subfamily was erected
by Nakamura (1956) for Platychasma (Fig.
19.7 G) and Cyphanta. Kiriakoff (1963 b) trans-
ferred both genera to Noctuidae on the basis of
quadrifid forewing venation and tympanal mor-
phology. Sugi (1982, 1987) and Holloway (1989)
disagreed, independently returning Platychasma
and Cyphanta respectively to Notodontidae.
Miller (1991) demonstrated that Kiriakoff had
misinterpreted the tympanal structure and de-
spite quadrifid venation and smooth apices to
the tibial spurs, Platychasma displays notodontid
apomorphies in both adults (metascutal bulla,
nodular sclerite absent, concave epimeron) and
larvae (developed stipital lobe in larvae, two M D
setae on larval Al). The known hosts are Acera-
ceae.
363
The remaining notodontids share several syna-
pomorphies, including a further modification of
the stipital lobe (reversed in the phalerines, Da-
tana and Phalera. Cteniophores (Jordan 1923), a
pair of wing-like, apically spinose flaps of un-
known function on sternum 4 (Fig. 19.12 B), are
known only from members of this clade (Miller
1991; Holloway 1983; Weiler 1989).
Notodontinae. This subfamily was expanded
by Miller (1991) to include Cerurinae (= Dicra-
nurini), Gluphisiini and Ptilophorinae. Synapo-
morphies include simple tarsal claws and four L
setae on larval segments A3 6, with the ventral
pair at approximately the same level. The sub-
family is divided into two tentatively monophy-
letic tribes, Dicranurini (Fig. 19.71) and Noto-
dontini (Fig. 19.7 H). The former is characterized
by loss of the pupal cremaster and with many
species having elongate, modified anal prolegs
(stemapods; Fig. 19.16 C). Hostplants for the
subfamily are mostly Fagaceae and Salicaceae,
but many temperate species feeding on Salica-
ceae have close tropical relatives feeding on Fla-
courtiaceae. Pupation occurs in loose cocoons
between leaves (Fig. 19.17M) or in very dense,
hardened cocoons partially imbedded in wood
(Fig. 19.17N).
Phalerinae (sensu stricto). This taxon, includ-
ing Phalera, Antheua and Datana (Fig. 19.7 J),
is supported by two genitalic synapomorphies:
uncus large and triangular and female segment 8
capsular. However, Miller's (1991) analysis also
associated other genera as Phalerinae (s. 1.), for
which no unambiguous synapomorphies were
found and the subfamily limits remain vague.
Typical phalerine larvae (Fig. 19.16D) have
long, scattered secondary setae and feed on both
woody and herbaceous plants including Anacar-
diaceae, Fabaceae, Fagaceae, Juglandaceae and
Poaceae.
Dudusiinae. This subfamily originally included
only large Asian species in Dudusa, Tarsolepis
and related genera (Fig. 19.7 K). Miller (1991)
expanded this concept as the tribe Dudusiini to
include the New World genera Crinodes, Goa-
campa and Cargida. A new tribe, Scranciini, was
established to accomodate narrow-winged Afri-
can and Asian moths in Scrancia, Gargetta and
related genera. Synapomorphies for Dudusiinae
include the presence of large ocelli (ocelli are ves-
tigial or absent in other notodontids) and the loss
of seta X from larval segment AIO. An interest-
ing possibility is that tear drinking (Bnziger
1988) is apomorphic for Dudusiini, although two
lachrymophagous genera, Ramesa (= Ponce ta)
and Pydnella have yet to be confirmed as dudusi-
ine (Miller 1991). Larvae of Dudusiini (Fig.
19.16 E) are variable in appearance, but often
have modified anal prolegs and feed predomi-
nantly on Aceraceae, Sapindaceae and Rhamna-
ceae. Larvae of Scranciini have greatly elongated
anal prolegs. In both Africa and Asia, they feed
364 Ian J. Kitching & John E. Rawlins

Fig. 19.7. Adult Noctuoidea. - B , Oenosandridae: A, Oenosandra boisduvalii, female, Australia (FW = 25 mm);
B, Oenosandra boisduvalii, male, Australia (FW = 2 2 m m ) . C, Doidae, Doa ampia, U . S . A . , Arizona (FW =
18 mm). D - P , Notodontidae: D, Thaumetopoeinae, Anaphe reticulata, Malawi (FW = 19 mm); E, Thaumetopoei-
nae, Marae melanospila, Australia (FW = 18 mm); F, Pygaerinae, Clostera albosigma, U. S. ., New York (FW =
15 mm); G, Platychasmatinae, Platychasma virgo, Japan ( F W = 17 mm); H, Notodontinae (Notodontini), Ptilodon
kuwayamae, China (FW = 17 mm); I, Notodontinae (Dicranurini), Furcula borealis, U. S. ., New York (FW =
18 mm); J, Phalerinae, Datana ministra, U. S. ., Pennsylvania (FW = 23 mm); , Dudusiinae, Tarsolepis taiwana,
Taiwan ( F W = 27 mm); L, Heterocampinae, Heterocampa astarte, U. S. ., Florida (FW = 22 mm); M, Nystalei-
nae, Calledema jocaste, Brazil (FW = 19 mm); N, Dioptinae, Myonia cingulina, Peru (FW = 15 mm); O, Diopti-
nae, Josia gigantea, Costa Rica (FW = 25 mm); P, Hemiceratinae, Hemiceras nigrigutta, Bolivia (FW = 24 mm).

on Antidesma (Euphorbiaceae). Some scranciine
larvae are semiloopers with reduced prolegs on
segments A3 and A4. The stout pupa of dudusi-
ines is in a slight cocoon on the ground, while the
slender, glossy pupa of scranciines is in a denser
cocoon among debris.
Heterocampinae. The composition of this sub-
family (Fig. 19.7 L) is uncertain but includes such
genera as Stauropus, Harpyia, Litodonta, Hypar-
pax, Schizura, Oligocentria and typical Rosema.
Confirmed apomorphies for the family are lar-
val: thoracic SV setal formula 2-2-2 (a reversal
to the plesiomorphic condition) and seta SV2 on
segment T1 short and spatulate (Miller 1991).
First instar larvae of many New World genera
have the bases of dorsal setae greatly enlarged
and extended into antler-like structures on the
prothorax and sometimes other body segments.
The Noctuoidea
Until an apomorphy is discovered in adult mor-
phology, the placement of numerous genera for
which the larvae are unknown remains unclear,
especially in the neotropics (Miller 1991). Larvae
(Fig. 19.16 F) feed on a wide range of woody
plants, especially Fagaceae, Juglandaceae, Rosa-
ceae and Ulmaceae. When disturbed, many eject
formic acid and ketones from the adenosma
(Kearby 1975).
Nystaleinae and Dioptinae are well-corrobo-
rated as sister groups (Weiler 1989), based on the
following synapomorphies: narrow adult tento-
rium; sensilla STI on the maxillary mesal lobe
shorter than the maxillary palp; larval tarsal seta
2 short; seta L2 on A8 located anterior to the
spiracle; and cremaster base fluted (Miller 1991;
1992 a). A unique proboscis structure, in which
the surface microspines on each annulet are re-
placed by longitudinal ridges, is tentative evi-
dence for a clade comprising these two subfamil-
ies and Heterocampinae (Miller 1991).
Nystaleinae. The classification and morphol-
ogy of this subfamily (Fig. 19.7 M), comprising
some 300 species restricted to the New World,
was revised by Weiler (1989). Numerous apo-
morphies support the monophyly of this subfam-
ily, including the fusion of juxta and sclerotized
manica and the presence of a callosum on the
aedeagus. Many species are characterized by
large tufts of scales on the antennal scape and
adopt characteristic resting postures resembling
sticks or twigs. Larvae are glossy (Fig. 19.16G),
often brightly coloured, and with medial pro-
cesses on A8. They have defensive secretions like
Heterocampinae. Hostplants are woody trees,
shrubs and vines, especially Fabaceae, Fagaceae
and Sapindales.
Dioptinae. This subfamily (Figs. 1 9 . 7 N - 0 )
comprises about 400 species, of which most are
neotropical and continental. Many are brightly
coloured and involved in mimicry complexes.
This distinctiveness had resulted in the group be-
ing awarded familial status but the work of Mi-
net (1983, 1986) and Weiler (1989) clearly de-
monstrated Dioptinae to be subordinate within
Notodontidae. The monophyly of Dioptinae is
beyond question. Miller (1991) listed seven un-
ambiguous apomorphies derived from larvae
and a further twelve from adults. Furthermore,
they feed on plants containing toxic secondary
chemicals (Violaceae, Passifloraceae, Aristoloch-
iaceae) and larvae (Fig. 19.16 H) are often
brightly or contrastingly coloured. This dietary
shift, together with derived diurnality and associ-
ated morphological changes (Weiler 1989) has re-
sulted in dioptine biology being more like that
of butterflies than moths. Indeed, although some
dioptines have shiny brown pupae in loose co-
coons, many have brightly coloured chrysalides
without cocoons (Fig. 19.170), anchored by
their cremasters in the exposed manner of many
Nymphalidae.
365
Miller (1991) placed Hemiceras (Fig. 19.7 P) as
sister group to Heterocampinae + (Dioptinae +
Nystaleinae), whereas Weiler (1989) considered
hemiceratines and dioptines to form a clade. We
retain Hemiceras and its relatives as Hemicera-
tini, a tribe of uncertain affinities. Pupal segment
AIO of Hemiceras has distinctive cuticular sculp-
turing (Rawlins pers. obs.), similar to many nys-
taleine pupae (Miller 1992 a).
Overall, there are more than 2,800 described
species of Notodontidae distributed worldwide,
with the exception of New Zealand (Dugdale
1988) and the Pacific islands. Old World genera
were treated by Kiriakoff (1960, 1964, 1967,
1968, 1970 b) and the Chinese fauna by Cai
(1979) and Schintlmeister (1991). Recent il-
lustrated works have been produced on the No-
todontidae of Borneo (Holloway 1983), Europe
(Freina & Witt 1987), Japan (Inoue et al. 1982),
Nepal (Haruta 1992, 1993), southern Africa (Pin-
hey 1975), Sumatra (Bender 1985) and Taiwan
(Chang 1989). Notodontidae are particularly di-
verse in the Neotropics (Forbes 1939; Godfrey &
Appleby 1987). The family has foodplants that
are predominantly woody and arborescent
(Miller 1992 b), although in Africa and tropical
Asia there are radiations (especially Phalerinae)
associated with herbs, both dicotyledons and
monocotyledons (Poaceae). This diet accounts
for the low number of economically important
species (Holloway 1983), although species of
Lochmaeus, Heterocampa, Symmerista and Schi-
zura are occasional defoliators of orchard and
timber trees (Holloway et al. 1987; Godfrey &
Appleby 1987). The larvae of many European
species are illustrated by Gmez (1988).
The Quadrifld Noctuoidea
The remaining families of noctuoids form a mo-
nophyletic group based upon the synapomorphic
possession of the quadrifld forewing venation
(Fig. 19.3 B - C , E - ) . Because they occur in the
trifid family Oenosandridae, several tympanal
features previously believed to be synapomor-
phies of the quadrifide (presence of the nodular
sclerite and a counter-tympanal hood), were
treated by Miller (1991) as apomorphies of Noc-
tuoidea as a whole. One of the most intractable
problems in Lepidoptera classification has been
the phylogeny of these quadrifld lineages of Noc-
tuoidea. This is due to great homogeneity in
adult morphology and widespread homoplasy in
virtually every character system. Features of the
immatures have been much more informative,
but resolution of phylogenetic problems has been
hindered by the absence of vouchered immatures
for the vast majority of world genera. Under-
standing of relationships between and among the
major subgroups awaits further research. With
366
these problems in mind, we recognize five fami-
lies of quadrifid noctuoids: Arctiidae, Lymantrii-
dae, Noctuidae, Nolidae and Pantheidae. Char-
acters supporting the monophyly of these fami-
lies are offered under their respective subhead-
ings and a discussion of their interrelationships
follows.
The monophyly of a group comprising Noli-
dae, Arctiidae, Pantheidae and Lymantriidae is
suggested by the presence in larvae of secondary
thoracic and abdominal setae that are restricted
to Verrucae at primary setal locations. However,
such setae also occur in Oenosandridae, Noto-
dontidae (Platychasma, Antheua) (Miller 1991)
and some species of Noctuidae (Acronicta) (Fig.
19.14 D), while reversal to the unisetose condi-
tion occurs in both Nolidae and Arctiidae.
Arctiidae, Pantheidae and Lymantriidae could
be united as a natural group based on three pu-
pal features: lack of a externally visible epicra-
nial suture; prothoracic femora not visible; and
presence of secondary setae, most of which arise
at the periphery of regions homologous with the
larval Verrucae (Mosher 1916) (Fig. 19.18 D).
Secondary setae on the larval head (Fig. 19.13 B)
are characteristic of all known pantheids, ly-
mantriids, and also many arctiids. However, all
these features are found elsewhere in Noctu-
oidea. Secondary setae occur in Nolidae (Noli-
nae) and many Acronictinae, and covered epicra-
nial sutures and prothoracic femora in Nolidae
and several groups of noctuids. The secondary
pupal setae of Arctiidae (Mosher 1916) and Pan-
theidae (Nakamura 1987) are usually micro-
scopic and invisible to the naked eye, while in
Lymantriidae they may be longer, giving the
pupa a hairy appearance (Fig. 19.18 D).
A sister group relationship between Panthei-
dae and Lymantriidae is supported by the pres-
ence of secondary setae in first instars, a condi-
tion lacking in other Noctuoidea. In addition,
the first instars of both families bear distinctive,
enlarged prespiracular Verrucae on the protho-
rax, making that segment the widest part of the
body. This condition is found elsewhere only in
some species of Acronicta. The pupal cremasters
of lymantriids and pantheids are well-developed,
often longitudinally rugulose, and bear eight to
many small, hooked cremastral setae. Pupae of
most nolids and arctiids lack a distinct cremaster.
Nolids and arctiids possess two SV setae on T2,
and elongate ("bar-shaped") male retinacula
(Fig. 19.4 C - D ) and heteroideous crotchets are
found in some lineages of those families.
Enigmatic but potentially informative struc-
tures are the sound-producing tymbal structures
in Arctiidae (metepisternum), Nolidae (second
abdominal sternite) and Lymantriidae (third ab-
dominal sternite). These structures bear exter-
nally similar rows of microtymbal grooves.
While the differences are probably autapomor-
phic for each family, the general structure of the
Ian J. Kitching & John E. Rawlins
abdominal organs may be homonomous (serial
homologues). Such structures are not known in
these positions in Pantheidae, nor in other Lepi-
doptera.
Although Arctiidae and Lymantriidae have
long been recognized as distinctive lineages, the
family Noctuidae has been notoriously difficult
to characterize (Kitching 1984). Only recently
have putative apomorphies been suggested. Mi-
net (1986) proposed two such features: the small
sclerotized band that links the anterolateral
elongations of St2 with the corresponding later-
otergal sclerite is distinctly widened dorsally, and
the counter-tympanal cavities are ventrally very
close to one another or even broadly contiguous.
However, the former character also occurs in
Arctiinae and is not found in many Noctuidae,
while the latter feature is present in the majority
of Nolidae, Lymantriidae and some Notodonti-
dae, and is not developed in several basal noctu-
ids (cf. Speidel et al. 1996 a).
Many larvae of Noctuidae are unique among
Noctuoidea in having a cylindrical galeal lobe on
the maxillary complex (Fig. 19.13D) (Grimes &
Neunzig 1986; Miller 1991), which is otherwise
recorded only in Nymphalidae (DeVries et al.
1985). The similar structure observed in Arctii-
dae (Grimes & Neunzig 1986) is an incomplete
ring rather than cylindrical and is non-homolo-
gous. However, because some noctuid larvae do
not have a cylindrical galeal lobe (e. g. Acronicti-
nae), the derivation may not be apomorphic for
that family and requires further study. In addi-
tion, noctuid larvae have only a single SV seta
on T2 and T3. However, the unisetose condition
occurs in many notodontid and nolid taxa, while
a few noctuids (Glottulinae, some Syngrapha, Li-
toprosopus) have the bisetose condition, proba-
bly as a secondary derivation.
Holloway (1988) suggested that the forewing
orbicular stigma might also be apomorphic for
Noctuidae, although it is lacking in many Her-
miniinae and present in some Notodontidae
(Phalera). Noctuidae also differ from Lymantrii-
dae and Pantheidae in that the retractor muscles
for the A9 apophyses originate on A7 rather
than A8 (Stekolnikov 1967) but the apomorphic
state and taxonomic distribution have yet to be
determined.
The widely accepted division of the Noctuidae
into "trifines" (in which hindwing vein MA2
arises nearer to MAI than to MPI and may be
reduced or absent; Fig. 19.3 H) and "quadri-
fines" (in which MA2 is a strong vein arising
close to MPI; Fig. 19.3FG) is rendered indis-
tinct by a failure to distinguish plesiomorphic ve-
nation from various independent derivations of
the trifine condition. Miller (1991) stressed two
aspects of hindwing venation in Notodontidae.
The first, the degree of development of MA2,
was highly homoplastic and is so in the quadrifid
families as well, especially Noctuidae (Kitching
The Noctuoidea
1984, 1987). The second aspect describes the po-
sition of MA2 (as in the forewing feature dis-
cussed above). In our view, hind wing venation is
plesiomorphic when MA2 is well-developed in
the trifine position, but may be apomorphic in
several non-homologous forms: MA2 greatly re-
duced or lost in any position (Fig. 19.3 H); MA2
adjacent to MPI in the quadrifine position; MA2
fused with MPI to secondarily assume a trifine
configuration (Fig. 19.3 C); CuAl fused with
MPI to form yet another trifine condition (some
nolids). The primitive trifine position of MA2
occurs in Oenosandridae, Doidae, Notodonti-
dae, Nolidae, some Arctiinae (Bertholdia) and
"trifine" Noctuidae (defined as Bagisarinae,
Condicinae, Eustrotiinae, some Acontiinae and
Plusiinae, Acronictinae, Agaristinae and Hamp-
son's (1903-1910a) trifine subfamilies, Noctui-
nae, Hadeninae, Cuculliinae and Heliothinae).
The Stiriinae, many plusiines and acontiines
have quadrifine venation, but the presence of
other features characteristic of trifines shows
that they are nevertheless part of that lineage.
Apomorphic reduction of MA2 occurs in the
Hemiceratinae and trifine Noctuidae, and apo-
morphic fusion occurs in many different groups
of Arctiidae, Nolidae and Noctuidae. The quad-
rifine position of MA2, if apomorphic, supports
the monophyly of two distinct clades: Arctiidae
+ Pantheidae + Lymantriidae; and "quadrifine"
Noctuidae. Enigmatically, a preliminary noctu-
oid analysis using molecular data (Weller et al.
1994) relates exemplars of quadrifine subfamilies
to different lineages of Arctiidae, placing a cato-
caline as sister-group to Arctiinae, but deriving
Nyctemera from Calpinae and Hypeninae.
In summary, there is evidence to suggest that
Arctiidae, Nolidae, Pantheidae and Lymantrii-
dae form a monophyletic group. Both Arctiidae
and Lymantriidae are defensibly monophyletic,
but Pantheidae, Nolidae and Noctuidae are not.
Several apomorphies support a sister group rela-
tionship between Arctiidae and part of a poten-
tially paraphyletic Nolidae. A similar relation-
ship is likely between Lymantriidae and a pa-
raphyletic Pantheidae. Finally, the number of
confusing similarities in eggs, larvae and pupae
between species in these families and those in lin-
eages currently contained within Acronicta (sensu
Forbes 1954) make uncertain the monophyly of
the quadrifid noctuoids here placed in Noctu-
idae.
Noctuidae. This family is the most speciose radi-
ation of Lepidoptera, containing more than
35,000 described species in more than 4,200 gen-
era. Most noctuid subfamilies erected by Hamp-
son (19031913) are now recognized as unnatu-
ral (Beck 1960, 1991, 1992; Kitching 1984; La-
fontaine & Poole 1991). Indeed, the composition
and monophyly of most is still open to question,
as is their number. The morphology of the imma-
367
ture stages, as well as the adults, is astonishingly
poorly known and such information will be criti-
cal to a defensible noctuid phylogeny. We have
chosen to discuss 28 subfamilies here, only some
of which are monophyletic.
There are a number of apomorphies that sup-
port subfamily groupings. Variation in hindwing
venation was discussed above and broadly di-
vides the subfamilies into two phenetic groups,
trifines and quadrifines. At the base of the abdo-
men, males of many trifine Noctuidae possess a
pair of brush organs (Fig. 19.12 A). When fully
developed, each consists of a brush of long scales
attached to a sclerotized lever that arises from
the posterior angle of the second sternite. At rest,
the brush is contained within a pocket. It is
everted by contraction of a muscle that origi-
nates on an apdeme produced from the anterior
angle of St3 and is inserted on the base of the
lever, further muscles fanning out the brush
scales (Birch 1979). Details of morphology and
operation are given by Birch (1970) and Clear-
water (1975). Pheromone is produced by Stob-
be's glands (Varley 1962), which are invaginated
from the membrane between St2 and the lever,
and are secretory in the pharate adult (Birch
1979). On emergence from the pupa, the glands
discharge their contents into the pockets, where
it is stored on the brushes. The glands then atro-
phy. Additional pheromone components are
secreted by glandular cells at the base of the
pocket scales (Birch 1970). The brush scales have
an intricate lattice structure (Birch 1979) that
provides a large surface area for storage and
evaporation of pheromone. The complexity of
this structure strongly suggests a single origin
that supports a clade comprising all trifine noc-
tuids except Bagisarinae, Condicinae, Acontii-
nae, Plusiinae, Acronictinae and Bryophilinae.
However, brush organs are lacking in most Noc-
tuinae (Birch 1979) and may also be fully present
or completely absent in otherwise very closely re-
lated species. In other taxa, individual elements
may be absent or in various stages of reduction.
These losses appear to follow a regular sequence:
glands first, followed by brushes, pockets, apo-
demes and finally levers (Birch 1972). Hence, al-
though the brush organs are doubtless a good
synapomorphy for a large subset of trifine Noc-
tuidae, their lability makes it difficult to assess
the limits of that clade.
In most noctuid larvae, seta SD on A9 has
the same thickness and attachment as D1 and
D2 on that segment (Fig. 19.15 D; Ripley 1923).
However, in Hadeninae, Noctuinae (Beck 1960;
Merzheevskaya 1988), Stiriinae, Condicinae,
some Plusiinae (Ichinos 1962), Glottulinae and
Ufeinae, SD is very fine, a tonosensillum (Fig.
19.15 B). Lafontaine & Poole (1991) considered
this condition to be synapomorphic for at least
Hadeninae and Noctuinae (as delimited here).
The feature has not been carefully surveyed in
368 Ian J. Kitching & John E. Rawlins

Fig. 19.8. Adult Noctuidae. A, Acronictinae, Acronicta lobeliae, U. S. ., Virginia ( F W = 22 mm); B, Bryophili-
nae, Cryphia domestica, England (FW = 12 mm); C, Raphiinae, Raphia pallida, U. S. ., Oregon (FW = 18 mm);
D, Herminiinae, Renia discoloralis, U. S. ., North Carolina (FW = 17 mm); E, Catocalinae, Lepidodes gallopavo,
Ecuador (FW = 29 mm); F, Catocalinae, Megacephalomana remaudi, Cameroon (FW = 26 mm); G, Catocalinae,
Sphingomorpha chlorea, Malawi (FW = 22 mm); H, Catocalinae, Dialithis gemmifera, Dominican Republic (FW =
19 mm); I, Catocalinae, Syllectra erycata, Bolivia (FW = 22 mm); J, Calpinae, Gonodonta correda, Bolivia (FW =
50 mm); K, Calpinae, Eudocima tyrannus, China (FW = 35 mm); L, Aganainae, Euplocia membliaria, Philippines
(FW = 37 mm); M, Cocytiinae, Cocytia durvillii, Indonesia, Irian Jaya (FW = 8 mm); N, Strepsimaninae, Dyspyr-
alispuncticosta, U. S. ., Pennsylvania (FW = 8 mm); O, Hypeninae, Hypena longipennis, Taiwan (FW = 19 mm);
P, Stictopterinae, Stictoptera griveaudi, Liberia (FW = 18 mm); Q, Euteliinae, Caligatus angasii, Malawi (FW =
23 mm); R, Dilobinae, Diloba caeruleocephala, Germany (FW = 18 mm); S, Plusiinae, Anagrapha falcifera,
U. S. ., Rhode Island (FW = 17 mm); T, Agaristinae, Episteme adulatrix, India ( F W = 34 mm).
The Noctuoidea
many problematic groups, especially tropical
ones, an effort complicated by intermediate setal
thickness in some groups.
In most noctuids the spinneret is a tubular
structure. However, in Hadeninae, Noctuinae
and a few other "higher" tritine noctuid groups,
the spinneret is dorsoventrally flattened, often
sinuous, with apical fringes (Beck 1960). This led
Beck (1989) to recognize just a single subfamily
with four tribes (Ipimorphini, Hadenini, Noctu-
ini and Euxoini). Fibiger & Hacker (1991) united
the last two of these and treated the resultant
three taxa as subfamilies. Beck (1991, 1992) dis-
cussed other alternatives for placement of trifine
genera with special reference to those originally
placed in Amphipyrinae, Acontiinae and Cuculli-
inae. Although resolution of this problem will
take comprehensive review of world taxa and
comparative study of all developmental stages,
we here adopt a classification that recognizes
Noctuinae and Ufeinae as separate from Hadeni-
nae, and use a broader concept of Hadeninae
that includes Apameini, Xylenini, Hadenini, Eri-
opygini and Ipimorphini of earlier authors, plus
isolated genus-groups, such as the Nedra-Acti-
notia group and the Elaphria-Spodoptera group.
The 'pleurite' (Forbes 1954), or 'paratergal
sclerite' (Tikhomirov 1979 a), is a small sclerite,
arising as a distal extension of the vinculum, that
articulates via a membrane with the base of the
tegumen (Fig. 19.11 A), which is thus separate
from the vinculum. From the pleurite originates
the valve flexor muscle, M.4 in the terminology
of Tikhomirov. In Lymantriidae, Arctiinae (Tik-
homirov 1979 c) and many Catocalinae and Cal-
pinae (Tikhomirov 1979 a, d) secondary fusion
of the pleurite with the tegumen has occurred
and muscle M.4 appears to originate on the tegu-
men. This condition is probably plesiomorphic at
the noctuoid level; it is similar to the state shown
by Notodontidae (Tikhomirov 1979 b, 1981), in
which muscle M.4 originates on the inner edge
of the tegumen, which is broadly contiguous
with the vinculum. The pleurite is absent in the
Nolidae and muscle M.4 usually originates near
the middle of the branches of the vinculum (Tik-
homirov 1979d; Speidel et al. 1996 a); Meganoia
[syn. Roeselia] strigula illustrated by Tikhomirov
1979 c is an exception). In Euteliinae, there is a
structure bearing the socius that may be homolo-
gous with the pleurite and in Stictopterinae, the
pleurite may be elongate and fused along the in-
ner margin of the tegumen (Kitching 1987).
Furthermore, in most families, there are complex
articulations between the tegumen and vinculum.
The detailed comparative morphology of the
pleurite needs further study to resolve these am-
biguities.
A muscle (M.2 of Tikhomirov 1979 a, d) be-
tween the tegumen and the transtilla is present
only in trifine Noctuidae, supporting their mono-
phyly (including Acontiinae and Plusiinae); its
369
presence in the Acronictinae is here considered
convergent. Speidel et al. (1996 a) consider this
muscle derived from a division of M.4, and in
their interpretation it supports the inclusion of
the Acronictinae in the trifines.
The number of prolegs has long been used in
noctuid classification. The primitive condition
(as in Raphiinae, Dilobinae, Psaphidinae and
Acronictinae) is a full complement in all larval
instars. Proleg reduction may have first evolved
as a dispersal adaptation in early instars of trifid
subfamilies and some Herminiinae. In these
groups, reduction in proleg size on A3 and A4 is
associated with "semi-looping", a locomotory
gait more rapid than normal crawling. Subse-
quent independent reductions extended semi-
looping into the later larval instars in several lin-
eages (quadrifid subfamilies, some Herminiinae,
Plusiinae, Acontiinae and Bagisarinae). Subfami-
lies predominantly feeding in herbaceous plants
and leading a terrestrial and relatively sedentary
existence (Noctuinae, Hadeninae) have semi-
looping in early instars, but develop a full com-
plement of prolegs in later instars. Some lineages
on woody plants have acquired greater speed
and mobility by retaining the semi-looping con-
dition through all instars, usually associated with
circadian cycles of activity, feeding on leaves at
night and withdrawing by day to hiding places
on the bark or ground. Less proleg reduction oc-
curs in arboreal lineages that do not withdraw
from the feeding site (Euteliinae, Stictopterinae,
Aganainae) and more in those that do (Catocali-
nae, Calpinae, Plusiinae). There is some evidence
that reversals in proleg reductions may be rare,
as that condition persists in semi-looping lin-
eages that have secondarily acquired sedentary
habits, such as algae grazing strepsimanines (Ni-
getia), fruit-boring acontiines (Tripudia) and
seed-eating eustrotiines (Eumicremma). Clearly,
the ontogeny of this feature requires further
study.
Three SV setae on the first abdominal segment
in larvae (Fig. 19.14C) have been considered ple-
siomorphic by many authors (Lafontaine &
Poole 1991) and occur widely in other families
of Noctuoidea. The monophyly of Plusiinae plus
most trifine noctuids (except Condicinae) ap-
pears supported by the loss of seta SV2 on A l .
However, Oenosandra has 2 SV seta on Al, as
do Acronictinae and many notodontids without
secondary setae. Although the vast majority of
species in the quadrifine subfamilies of Noctui-
dae have three SV setae, the bisetose condition
(Fig. 19.14 B) occurs in all Euteliinae (except An-
igraea), some Stictopterinae (Lophoptera, but
not Odontodes), most Acontiinae and Eustrotii-
nae (except Lithacodia, Eumicremma and Eu-
blemma), Dilobinae, Raphiinae and some Cato-
calinae (Donda, Rhesala; Gardner 1948 a). Her-
minia lacks SV2 on both Al and A2 but other
known herminiine larvae are trisetose on both
370
these segments. Furthermore, three SV setae oc-
cur in some trifine lineages, especially Condici-
nae (Condica (including Platysenta), Perigea,
Ogdoconta, Leuconycta, and Diastema). The
third SV seta on Al in these groups differs in
relative size, placement and degree of pinacular
fusion with other SV setae, and it is likely that
the trisetose condition is not homologous in all
lineages. Under such an interpretation, various
trisetose configurations are apomorphic and de-
limit several monophyletic groups including Bag-
isarinae, Condicinae, Herminiinae, the majority
of quadrifine subfamilies (Strepsimaninae, Cato-
calinae, Calpinae and Hypeninae) and one group
of Plusiinae (Eichlin & Cunningham 1978). In
Nolidae, the number of SV setae in Al varies:
Tympanistes and Selepa have three, Risobini and
Nycteola have two, and Carea and Earias have
one. Further study on the comparative morphol-
ogy of SV setae is required, especially in the
early instars.
Herminiinae, Hypeninae and Catocalinae may
form a clade based upon the use of polyenic hy-
drocarbons or their monoepoxide derivatives as
female sex pheromones (Arn et al. 1986; Renou
et al. 1988). All other subfamilies investigated
(together with Nolidae and Notodontidae) utilize
linear alkenes with even numbers of 1016 car-
bon atoms, one or two double bonds (Z- or E-
isomers) and an alcohol, aldehyde or acetate
functional group. However, because Arctiidae
and Lymantria (but not Euproctis, Calliteara and
Orgyia) also use polyenic hydrocarbons, and
many noctuid subfamilies remain to be exam-
ined, this hypothesis is highly tentative.
One overlooked feature of the quadrifine noc-
tuid subfamilies is the highly consistent absence
of scales on the lower part of the clypeofrons
(Fig. 19.5 A). This feature characterizes Strepsi-
maninae, Calpinae, Herminiinae, Hypeninae,
Cocytiinae, Euteliinae, Stictopterinae and Cato-
calinae. The scaleless area is smaller but distinct
in Aganainae, but the entire frons is scaled in the
trifine subfamilies and all other quadrifid Noctu-
oidea.
In conclusion, we tentatively assign the noc-
tuid subfamilies into three groups: 1. paraphy-
letic basal lineages including Acronictinae, Bryo-
philinae and Raphiinae; 2. quadrifine subfamilies
consisting of two subgroups, one paraphyletic
containing Herminiinae, Strepsimaninae, Cato-
calinae, Calpinae, Aganainae, Cocytiinae and
Hypeninae, and the other monophyletic includ-
ing Euteliinae and Stictopterinae; 3. chiefly 'tri-
fine' subfamilies, comprising Plusiinae, Agaristi-
nae, Eucocytiinae, Acontiinae, Eustrotiinae,
Bagisarinae, Cuculliinae, Psaphidinae, Dilobi-
nae, Condicinae, Stiriinae, Heliothinae, Glottuli-
nae, Hadeninae, Ufeinae and Noctuinae. This
preliminary grouping is not in agreement with a
phylogenetic alternative based primarily on adult
features hypothesized by Poole (1995), which
Ian J. Kitching & J o h n E. Rawlins
places Acronictinae, Bryophilinae, and Panthei-
nae as a derived lineage of trifine noctuids. It is
similarly incompatible with the proposal of
Speidel et al. (1996 a), which likewise has the
Acronictinae and Bryophilinae subordinate in
the trifines, excludes the Dilobinae from the lat-
ter and places our Nolidae inside the Noctuidae.
Several faunistic treatments of Noctuidae have
been published for eastern Canada (Rock-
burne & Lafontaine 1976), England (Skinner
1984), Europe (Fibiger 1990, 1993; Froster &
Wohlfahrt 1971; Skou 1991), Florida (Kimball
1965), Greece (Hacker 1989), Italy (Berio 1985,
1991), Japan (Inoue et al. 1982), Nepal (Haruta
1992, 1993), Ohio (Rings et al. 1992), southern
Africa (Pinhey 1975), Spain (Calle 1982), Taiwan
(Chang 1991) and the eastern United States (Co-
vell 1984). Illustrated works on larvae are less
common, but are available for Europe (Carter &
Hargreaves 1986), Japan (Sugi 1987), northeast-
ern North America (McCabe 1991), and Spain
(Gmez 1985, 1987).
Acronictinae. This subfamily, restricted to
Acronicta (s. 1.) (Fig. 19.8 A) and its close rela-
tives, is uniform in adult morphology. Many
have greyish, cryptic forewings with black basal
and subtornai streaks (Holloway 1989) but this
pattern is not unique to the group. Characteristic
features noted by early authors include a nar-
rower and paler segment A7 in first instar larvae
and a complex, stubby pupal cremaster with all
hooks similar in size and length (Mosher 1916;
Lorimer 1983). Both characters vary within the
subfamily as now delimited, as does degree of
development of secondary setae in larvae (Fig.
19.161). The Craniophora group (including Tha-
latha and Thalathoides) have primary setae only
(Holloway 1989). Acronicta alni has only primary
setae and displays two arctiid features: D1 and
D2 pinacula on T2 and T3 fused and L3 on
A36 bisetose. The SV group only of A. carbo-
naria, A. ovata, A. brumosa and related species
has additional secondary setae (Sugi 1979), while
in A. rumicis, A. impleta, A. impressa, Eulonche
and Simyra secondary setae are present on all
Verrucae. Finally, species related to A. leporina
and A. americana have numerous secondary se-
tae all over the body. The pupae of such hairy
larvae also possess secondary setae arranged
around the periphery of Verrucae as in Arctiidae.
This diversity of larval colour and pattern, to-
gether with genitalic differences, has led some
authors (Sugi 1987) to subdivide Acronicta into
several genera. The pupae of the Simyra group
differ greatly from those of the Acronicta group
and this confusing overlap in characters of all
stages of Acronictinae with other families of
quadrifid noctuids suggests that it may be pa-
raphyletic at the family level. The Acronictinae
lack a number of features (venation, brush or-
gans, proleg reduction) that would clearly associ-
ate it with a subgroup of Noctuidae, while the
The Noctuoidea 371

Fig. 19.9. Adult Noctuoidea. AS, Noctuidae: A, Eucocytiinae, Eucocytia meeki, Indonesia, Irian Jaya (FW =
25mm); B, Acontiinae, Acomia delecta, U . S . A . , New York (FW = 12mm); C, Eustrotiinae (Eublemmini),
Eumestleta recta, U. S. ., Florida (FW = 8 mm); D, Eustrotiinae, Zurobata vacillans, Taiwan (FW = 10 mm); E,
Eustrotiinae, Lithacodia bellicula, U. S. ., New Jersey (FW = 9 mm); F, Bagisarinae, Xanthodes transversa, Tai-
wan (FW = 19 mm); G, Cuculliinae, Cuculila argentea, China (FW = 15 mm); H, Stiriinae (Stiriini), Chrysoecia
atrolinea, U. S. ., Arizona (FW = 14 mm); I, Stiriinae (Grotellini), Groteila dis, U. S. ., Arizona (FW = 11 mm);
J, Condicinae, Condica confederata, U. S. ., Florida (FW 16 mm); , Heliothinae, Heliothis borealis, Canada,
Yukon (FW = 10 mm); L, Psaphidinae (Psaphidini), Psaphida thaxteriana, U. S. ., Indiana (FW = 16 mm); M,
Psaphidinae (Feraliini), Feralia comstocki, U. S. ., Pennsylvania ( F W = 16 mm); N, Amphipyrinae, Amphipyra
erebina, Japan (FW = 22 mm); O, Glottulinae, Diaphone eumela, Malawi (FW = 22 mm); P, Hadeninae (Hade-
nini), Persectania aversa, New Zealand (FW = 17 mm); Q, Hadeninae (Apameini), Papaipema eryngii, U. S. .,
Illinois (FW = 20 mm); R, Ufeinae, Ufeus satyricus, U. S. ., Vermont (FW = 21 mm); S, Noctuinae, A'estia
triangulum, Germany ( F W = 17 mm); T, Pantheidae, Panthea furcilla, U. S. ., West Virginia (FW = 18 mm).
372
male genitalia do have the 'trifine' trait of a tegu-
men-transtilla muscle. We tentatively consider it
to be a basal and probably paraphyletic taxon.
Craniophora group feeds largely on Oleaceae
(Holloway 1989), while North Temperate radia-
tions within Acronicta are variously restricted to
Ulmaceae, Fagaceae, Rosaceae, Betulaceae and
Salicaceae, with a few species being truly polyph-
agous. The Viminia-Eulonche-Simyra group feed
on low woody plants and herbs, especially Sali-
caceae, Polygonaceae and Typhaceae.
Bryophilinae. This subfamily comprises small,
lichen-coloured moths (Fig. 19.9 B) traditionally
included in Acronictinae (Franclemont & Todd
1983). Segment A7 of the first instar larvae is
unmodified and the cremaster is elongate with
the usual noctuid complement of hooks dimor-
phic in size and length (Giehsler 1975-1979).
Eggs are usually laid on rocks or walls near their
lichen foodplants, a unique oviposition behavi-
our for Noctuidae. The larva overwinters in a
silk-lined burrow, which it continues to inhabit
the following year. Pupation occurs in a blister-
like cocoon of silk and lichen fragments (Lori-
mer 1983).
Raphiinae. The aberrant genus Raphia (Fig.
19.8 C), the sole constituent of this subfamily,
has been variously placed in Ophiderinae, Pan-
theidae (Forbes 1954; Poole 1989) and Acronicti-
nae (Crumb 1956). Twelve species have been de-
scribed, all but one of which (from Nigeria) oc-
cur in the Holarctic Region. In contrast to pan-
theids and most Acronictinae, larvae of Raphia
lack secondary setae. However, they do have two
SV setae on the first abdominal segment, as do
Acronictinae and the trifine subfamilies. They
are characterized by seta D1 on T2 being borne
subapically on a conical process (Fig. 19.16 J).
The larval prolegs have enlarged plantae with a
large number of crotchets and foodplants re-
stricted to Salicaceae (Crumb 1956), features
that may be correlated (J. D. Lafontaine pers.
comm.; see also Ufeinae). The egg is spherical,
without prominent longitudinal sculpture and
deposited singly. The pupa is strongly rugulose
dorsally and bears two stout, divergent spines on
the broad cremaster, both features strongly remi-
niscent of agaristine pupae. The immature stages
of such genera as Thiacidas, Pteronycta and Pan-
thuama (currently Catocalinae s. 1.) should be ex-
amined as potential raphiines.
Herminiinae. This group (Fig. 19.8 D) has
been excluded from Noctuidae because the
counter-tympanal hood is prespiracular (Kiria-
koff 1963 a; Kitching 1984; Speidel et al. 1996 a).
However, the spiracle is only rarely posterior to
the hood, often situated directly ventral to the
lower edge of the hood, or even ventro-anterior
to it (Richards 1933; Forbes 1954). This condi-
tion is not homologous with the prespiracular
hoods of other noctuoids (Richards 1933;
Owada 1987) and the presence of other noctuid
Ian J. Kitching & John E. Rawlins
apomorphies demonstrates that Herminiinae are
true Noctuidae (Minet 1986). The subfamily
(which here excludes Litoprosopus and Meta-
lectra) is characterized by a number of apomor-
phies. In the male genitalia, muscle M.4 is di-
vided, the two parts originating from the apex
and margin of the pleurite respectively (Tikhomi-
rov 1979 a). The metepimeron is generally
strongly swollen ventral to pocket IV of the tym-
panal organ (Richards 1933), while the form of
the counter-tympanal hood is unique among
Noctuidae. Most Herminiinae lack the forewing
orbicular stigma, which may represent a loss.
Many Herminiinae have distinctive male second-
ary sexual organs, which can be found on virtu-
ally any part of the body (Owada 1987). There
are "knots" on the antennae, formed from a
broadened and flattened series of flagellomeres,
which may be olfactory or serve to clasp the an-
tennae of the female during mating (Polypogon).
Hair brushes occur on labial palps of some taxa.
Primitively, these are exposed (Adrapsa), but in
Trotosema they are concealed in grooves on the
vertex and thorax (Owada 1987). The male fore-
wing may have costal folds on the upperside
(Hydrillodes) or underside (Bocana). The foretib-
ial apex may be extended parallel to an elongate
basitarsus and contain a brush organ (Simplicia).
Other taxa have elongate trochanters (Polypo-
gon), while tarsal reduction is widespread, often
to a single segment without claws (Herminia).
Generally, abdominal brush organs are absent,
although a tuft of hairs on the vinculum occurs
in some genera (Paracolax). Most herminiine
larvae have three SV setae on those segments
(Crumb 1956). Many species have larvae with a
full complement of prolegs, but in several genera
(e. g. Idia) there is reduction in a manner con-
vergent with other quadrifine noctuids. In Asian
Gynaephila, early instars are "true loopers" as in
most Geometridae with prolegs present only on
A6 and AIO, the only known instance of this
configuration in Noctuoidea. Most known her-
miniine larvae (Fig. 19.16 K) feed on dead leaves
in various stages of fungal decomposition (Gard-
ner 1948 a, Crumb 1956), but a continuum of
diets exists from nearly totally fungal to living
leaves. Some species feed on dead insects, dried
fruits and vertebrate dung (Rawlins 1984;
Owada 1987). Oviposition may occur singly or
in large appressed masses. A number of species
drop eggs loose into debris. None is of economic
importance although they may be ecologically
significant as detritivores, probably augmenting
digestion from acquired enzymes obtained by in-
gesting fungi (Rawlins 1984). Pupation occurs in
cocoons often covered with debris or bits of the
host. The pupae are often covered with a waxy
bloom, as in Catocalinae, and the exposed por-
tion of the mesothoracic tibia contacts the eyes.
Strepsimaninae. First described as a family of
Microlepidoptera (Meyrick 1930), Strepsimani-
The Noctuoidea 373

Fig. 19.10. Adult Noctuoidea. AC, Lymantriidae: A, Aroa discalis, Malawi (FW = 17 mm); B, Dasychira mescal-
era, U. S. ., Colorado (FW = 17 mm); C, Mylantria xanthospila, Cameroon (FW = 23 mm). D - K , Nolidae: D,
Chloephorinae (Camptolomini), Camptoloma interiorata, China (FW = 15 mm); E, Nolinae, Nola minna, U. S. .,
California (FW = 8 mm); F, Westermanniinae, Westermannia anchorita, Cameroon (FW = 13 mm); G, Eariadinae,
Gabala argentata, Japan (FW = 12 mm); H, Bleninae, Blenina angulipennis, Taiwan (FW = 19 mm); I, Risobinae,
Baileya ophthalmica, U. S. ., Ohio (FW = 14 mm); J, Collomeninae, Iscadia aperta, Bolivia (FW = 20 mm); K,
Eligminae, Eligma narcissus, China (FW = 35 mm). L - T , Arctiidae: L, Lithosiinae, Cyana hamala, China (FW =
14 mm); M, Lithosiinae, Setina aurita, France (FW = 14 mm); N, Syntominae (Syntomini), Amata leucerythra,
Gabon (FW = 18 mm); O, Syntominae (Thyretini), Balacra rubrostriata, Malawi (FW = 23 mm); P, Arctiinae
(Arctiini), Arctia festiva, France (FW = 24 mm); Q, Arctiinae (Utetheisini), Amphicallia bellatrix, South Africa
(FW = 30 mm); R, Arctiinae (Pericopini), Chetone phyleis, Bolivia (FW = 34 mm); S, Arctiinae (Phaegopterini),
Halysidota ata, Dominican Republic (FW = 25 mm); T, Arctiinae (Euchromiini), Euchromia lethe, Cameroon
(FW = 23 mm).
374
nae (Fig. 19.8 N) were transferred to Noctuoidea
(Hodges 1978) and are here included within Noc-
tuidae. The subfamily includes the smallest noc-
tuids and is not economically important. Sub-
family Hypenodinae (Forbes 1954) syn. nov. is
here treated as a junior synonym following the
suggestion of Minet (1986). The latter taxon was
traditionally diagnosed by the absence of ocelli
(Forbes 1954), but the inclusion of Strepsimanes,
which has visible ocelli (Meyrick 1930), together
with research into other so-called anocellate
groups (Dickens & Eaton 1973), renders this
character untenable. The monophyly of Strepsi-
maninae remains undemonstrated and it is pos-
sible they constitute one or more lineages of re-
duced Catocalinae or Hypeninae. In Hypenodes
and Strepsimanes only, forewing vein Rs4 runs
to the costa rather than the apex or termen, a
condition unique in Noctuidae. The larvae are
primarily algivorous, lichenivorous, or fungivor-
ous, with prolegs reduced or absent on A3 and
A4. They lack tonosensillae on A9 and have
three SV setae on A l . Pupation occurs in fairly
dense cocoons that are heavily covered with
plant debris or bits of wood, and in many cases
are suspended on silken strands from the sub-
strate. The pupa has the exposed portion of the
mesothoracic leg touching the eye, an apomor-
phic configuration largely restricted to quadri-
fine noctuids.
Catocalinae. Berio (1959) demonstrated that
Hampson's (1903) character of "spined midtib-
iae" was a graded feature of minimal phyloge-
netic significance, which resulted in closely re-
lated species being separated into "Catocalinae"
and "Ophiderinae" (Kitching 1984; Holloway
et al. 1987). For the present, Catocalinae (Fig.
19.8 EI) remains a polyphyletic assemblage of
these quadrifine noctuids unassigned to other
subfamilies, including genera (e. g. Aon, Phypro-
sopus, Metalectra, Litoprosopus) removed from
those subfamilies to render them monophyletic.
There are doubtless a number of monophyletic
taxa in Catocalinae, some of which were treated
by Berio (1959) as "phyla". At present these are
not recognized as separate subfamilies because
the limits, composition and monophyly of most
are uncertain. They include groups centred upon
Catocala (with a distinctively sclerotized, elon-
gate ovipositor; Mitter & Silverfine 1988), Arete
(males with a dorsal, corrugated, sclerotized
structure of unknown function; Holloway et al.
1987, Aehaea (presence of a brush organ on the
valve; Berio 1959, Holloway et al. 1987), Dr ast-
eria (very long, spirally twisted ductus recepta-
culi (Speidel & Naumann 1995a), seta V on Al
and A2 highly modified to form a circular struc-
ture), and Mods and Caenurgina (loss of prolegs
on A3 and A4). The "Erebinae" of authors, in-
cluding Erebus and Ascalapha (forewing pattern
with the reniform stigma formed into a distinc-
tive eyespot), belong here. The subfamily in-
Ian J . Kitching & J o h n E. Rawlins
cludes Thysania agrippina, the world's largest
moth in terms of wingspan. Males of many gen-
era have a groove on the midtibia enclosing a
brush organ (Berio 1959). Pupae are often cov-
ered with a white waxy bloom (Kitching 1984),
especially in taxa related to Catocala and Mods
{Zale, Ischyja, Ercheia, Mods, Caenurgina, Par-
alalia). Foodplants are diverse, but there is a
special relationship in several subgroups with le-
gumes and grasses. The Arete group (Fig.
19.16L) feeds on Urticaceae; temperate species
of the Catocala group have lineages on Salica-
ceae, Myricaceae, Juglandaceae, Fagaceae, Rosa-
ceae, Fabaceae and some Zale on Pinaceae; trop-
ical lineages have species feeding on Euphor-
biaceae and Sapindaceae despite most species be-
ing on Fabaceae. A few are recorded as serious
agricultural pests. Anticarsia attacks soybean
(Holloway et al. 1987) and Mods is occasionally
a pest on sugar-cane and cereal crops (Common
1990). Wiltshire (1979) reviewed the Armadini
from desert regions in the Old World, but most
catocaline species remain unstudied. Berio (1992)
offered several alternative classifications of cato-
caline genera, grouping them into tribes, but the
monophyly of most of these is unconfirmed.
Calpinae. The three tribes of Calpinae
(= Ophiderinae sensu stricto of authors; Figs.
19.8 J - K ) (Calpini, Gonopterini and Anomiini)
form a clade based upon the specialized apical
armature of the proboscis (Speidel et al. 1996 b),
which is modified to pierce thick-skinned fruit or
mammalian skin (Figs. 19.5 and 19.5 C;
Bnziger 1975, 1979). Most highly developed in
Calyptra eustrigata (Bnziger 1980), the probos-
cis tip is heavily sclerotized and armed with tear-
ing hooks, basad of which are basiconic sensilla
modified as erectile barbs. Simultaneous, anti-
parallel movement of the two halves of the pro-
boscis, coupled with the erection by haemolytic
pressure of these hooks and barbs, enables the
proboscis to penetrate (Bnziger 1970, 1980).
Fruit-piercing is not restricted to Calpinae, but
most other species accredited with this behaviour
can only pierce soft-skinned fruit or need the
skin to have already been damaged. Bnziger
(1982) termed the latter "secondary piercers", in
contrast to "primary piercers", which make their
own entry into fruit. Some species, notably Eu-
docima (= Othreis) fullonia, can occur in large
numbers and cause extensive crop losses (White-
head & Rust 1972), much of which is attributable
to fungi and bacteria that enter through the hole
made by the moth or are introduced on the pro-
boscis. Many members of Calpini have a thin,
elongate labial palp segment 3, a feature also pre-
sent in Aganainae and suggestive that these
groups may be related. However, palps of Ca-
lyptra are porrect and beak-like. The forewing
dorsal margins of Calpini are sinuate, often with
a produced median lobe and tornai angle. The
former condition recalls that seen in Notodonti-
The Noctuoidea
dae, while the latter also occurs in Plusiinae. In
addition, the postmedian line is often extremely
oblique, running to the apex of the wing. Larvae
of Calpini (Fig. 19.16 M) are largely restricted to
Menispermaceae (Holloway et al. 1987), which
may be apomorphic for the tribe. Larvae are
often heterochromic, changing pattern and col-
oration abruptly in different instars. The green,
cryptic larvae of Anomiini and Gonopterini
{Scollopteryx) possess a subprimary seta verti-
cally below D2 on A l - 6 (Gardner 1947; Crumb
1956; Beck 1960). Scoliopteryx and Gonodonta
are unusual in having heteroideous crochets. In
Noctuidae, this feature is found only in some
genera of Euteliinae and Stictopterinae, and is
undoubtedly not homologous with similar heter-
oideous arrangements in Arctiidae and Nolidae
(Selepa). In Anomiini, the alula of the tympanal
organ is enlarged, although it is not sclerotized
as in Acontiinae (Richards 1933). Larvae of An-
omiini mostly feed on Malvales (Malvaceae, Tili-
aceae, Sterculiaceae), while Gonopterini feed on
Salicaceae (Scoliopteryx) (Carter & Hargreaves
1986), Annonaceae and Piperaceae (Gonodonta)
(Todd 1959). Pupation in Calpinae often occurs
between leaves of the host with minimal use of
silk (Fig. 19.17R), or in slight cocoons covered
with debris or bits of bark or wood. Several An-
omis species are cotton and jute pests and may
also defoliate agricultural and ornamental Hibis-
cus (Hollov/ay et al. 1987).
Aganainae. The relationships of this subfamily
(Fig. 19.8 L) have long been disputed (Kitching
1984). Generally, the group has been treated as
a separate family or subordinated within the
Arctiidae based on venation and the bar-shaped
retinaculum (Fig. 19.4 B), but the lack of both a
tymbal organ and pheromone glands associated
with the anal papillae (Holloway 1988) precludes
the latter position. A relationship with Noctui-
dae was first suggested by Gardner (1941) based
on larval characters. Kitching (1984) excluded
them from the Noctuidae because aganaines
have a prespiracular counter-tympanal hood, but
the spiracle is actually between two weak hood-
like structures (Holloway 1988). Minet (1986) in-
cluded aganaines in Noctuidae because they pos-
sess both his proposed noctuid apomorphies.
Additional features support placement of aga-
naines in Noctuidae. The forewing pattern in-
cludes an orbicular stigma which, although fre-
quently obscured by other pattern elements on
the upperside, is clearly visible on the ventral
side (Holloway 1988). The labial palps are up-
right, with an elongate, thin third segment. This
type of palp is also characteristic of Cocytiinae
and some Calpinae, although precise homologies
remain to be determined. Holloway (1988) also
considered that the single corema on St8 of the
male and the complex vesica lobes were also noc-
tuid-like. In contrast, the male retinaculum of
aganaines is elongate, as in Nolidae and Arctii-
375
nae but not Noctuidae. Moreover, Kuznetzov &
Stekolnikov (1989) noted a reduction in the ter-
gal extensor muscles of the valves in three aga-
naine genera (Asota, Neochera and Psephea),
suggesting a relationship with Arctiidae and Ly-
man triidae. Aganaine larvae (Fig. 19.16N) have
a single SV seta on the T2 and T3, but as dis-
cussed above, this character is a tentative noctuid
apomorphy, repeatedly found in other families.
Gardner (1941) reported two SV setae in Digama
hearseyana, but this seems to be a species-specific
reversal because Digama marmorea shows the
unisetose condition (Common 1990). The aden-
osma is absent in aganaines, as it is in Arctiidae
and most (but not all) Lymantriidae (Gardner
1941; Speidel et al. 1996a; Rawlins, pers. obs.).
However, if Miller (1919) is correct in concluding
that this structure is not homologous in noto-
dontids and noctuids, then absence in aganaines
may be plesiomorphic. The pupal prothoracic
femora are concealed (Fig. 19.18 B), a condition
found in quadrifine noctuids, Nolidae, Arctiidae,
Pantheidae and Lymantriidae. We postulate an
association between Aganainae and the quadrif-
ines because in both groups the pupal mesothor-
acic tibiae are contiguous with the eyes (Fig.
19.18 C). The metathorax of aganaine pupae
possesses a transverse row of deep impressions,
an unambiguous apomorphy of known pupae of
the subfamily (Fig. 19.18 A). Aganainae include
about 100 species placed in 15 genera (Munroe
1982), distributed throughout the tropical and
subtropical areas of the Old World. Adult aga-
naines are generally brightly coloured. Often said
to be diurnal, most fly at night and are readily
attracted to light (Common 1990). Several gen-
era (Peridrome, Euplocia) exhibit extreme sexual
dimorphism, males having large androconial
patches on the forewing upperside at the costal
base. Larval foodplants belong to Moraceae,
Apocynaceae and Asclepiadaceae (Holloway
1988; Common 1990), and thus the bright col-
ours of most adults are probably aposematic.
The larvae are sometimes aposematic, being pat-
terned variously in black and white, especially in
species that oviposit egg masses, which develop
into groups of gregarious larvae. Other species
oviposit singly and their larvae are usually cryp-
tic. Larvae appear hairy because primary setae
are very long and arise from chalazae (Holloway
1988). In some genera {Neochera), setae are plu-
mose. Secondary setae are absent. Larvae of
Neochera and Asota are "flaccid and rather slug-
gish" and "often rest with the body bent around
and the anal segments raised" (Common 1990).
The subfamily is not economically important.
Cocytiinae. The subfamily Cocytiinae (Roth-
schild 1915) stat. rev. comprises a single species,
Cocytia durvillii (Fig. 19.8 M), from the Moluc-
cas, New Guinea and neighbouring islands. It is
a remarkable diurnal hymenopteran mimic, with
transparent wings (but with dark margins, veins
376 Ian J. Kitching & John E. Rawlins

Fig. 19.11. Features of the male genitalia in Noctuoidea. A, Genitalia with paratergal sclerite or 'pleurite' (pi)
between tegumen and vinculum, anterodorsal view (Noctuidae, Hadeninae, Leucania dorsalis); B, Genitalia lack-
ing paratergal sclerite between tegumen and vinculum, anterodorsal view (Noctuidae, Catocalinae, Hypsoropha
adeona); C, Aedeagus with everted endophallus and enlarged detail of deciduous cornuti (Notodontidae, Nystalei-
nae, Pentobesa sp.); D, Genitalia with prominent socius (so) below anal tube (an), aedeagus removed (Notodonti-
dae, Dioptinae, Phryganidia californica).

and intervenulae and a bright orange base), met-
allic blue-green scaling on the head, thorax, legs
and abdomen, and long antennae that are dis-
tally widened but not clubbed (Boisduval 1828).
Labial palp segment 3 is long, thin and upright,
reminiscent of Calpinae and Aganainae. Cocytia
was described as a zygaenid but historically has
been placed in other families or in a family of its
own. However, Cocytia has a metathoracic tym-
panal organ that faces posteriorly and a postspir-
acular counter-tympanal hood and is clearly a
noctuid. The eyes are hairy. The quadrili d vena-
tion, naked clypeofrons, and labial palps suggest
an affinity with Aganainae or Catocalinae. The
immature stages are unknown. Consequently,
the placement of Cocytia within quadrifine Noc-
tuidae remains uncertain.
Hypeninae. This subfamily (Fig. 19.8 O), here
including Rivulinae, is poorly characterized. The
larval setae are usually borne on raised chalazae
(Gardner 1946 a; Forbes 1954; C o m m o n 1990),
although these are often not conspicuous in later
instars. Hindwing vein M A 2 is strong and arises
well above the lower angle of the cell, running
more-or-less parallel to M P I (Forbes 1954;
Kitching 1984; Holloway et al. 1987). Rivula and
Zebeeba have a unique proboscis structure
(Speidel et al. 1996 a, b). The Hypena g r o u p are
monophyletic on the basis of a spinose "cuff" on
the aedeagus (Lodi 1993). They also have long,
porrect labial palps, hence the c o m m o n name of
"snout moths", but these are not universal
within the subfamily. Three SV setae are present
on the first abdominal segment of known larvae,
including Rivula ( C r u m b 1956; Beck 1960). Lar-
vae are often semiloopers (Fig. 19.160), usually
green and without dark pigmentation, m a n y spe-
cies having restricted host preferences, especially
in Urticales, Malvales, Polygonaceae and Faba-
ceae. Larvae of Rivula and its relatives feed on
grasses and sedges, and lack an adenosma. Pupa-
tion occurs in loose cocoons on leaves (Fig.
19.17 Q), which are often covered with debris
(Fig. 19.17P); the pupal mesothoracic leg con-
tacts the eye. M a n y species are sexually dimor-
phic in colour and size, the males being larger
than females. Hypenines occur worldwide and
most are of little economic importance, but Pla-
thypena scabra can be m a j o r pest of legume crops
and a few Hypena having been recorded as mi-
n o r agricultural pests (Holloway et al. 1987). Hy-
pena laceratalis has been employed in Queens-
land, Norfolk Island and Hawaii to control Lon-
tana camara ( C o m m o n 1990).
Stictopterinae. Euteliinae and Stictopterinae
f o r m a monophyletic group, based on a large
The Noctuoidea
number of synapomorphies (Holloway 1985;
Kitching 1987) including: reduced female frenu-
lum; modified basiconic sensilla on the probos-
cis; presence of a small oval plate in the ductus
ejaculatorius; and anal papillae modified so that
their inner surfaces are directed posteriorly. The
counter-tympanal hood of Stictopterinae and
Euteliinae has a unique double structure (Rich-
ards 1933; Kitching 1987). In the former subfam-
ily, it is large, directed anteriorly over the tympa-
num and overlapped by a fan of narrow, lenticu-
lar scales arising from T3 dorsal of the tympa-
num, below which is a smaller flap forming a
shallow concavity into which the ventral part of
the counter-tympanal hood extends (Holloway
1985). In Euteliinae, the counter-tympanal hood
is shorter and the scales overlapping it are short
and rounded (Holloway 1985).
The monophyly of Stictopterinae (Fig. 19.8 P)
is supported by three apomorphies (Holloway
1985; Kitching 1987). The female frenulum is re-
duced to a single bristle and the anal papillae are
apically divided, with numerous setae on ele-
vated bases. The papillae are also generally semi-
circular, together forming a disc. The pupal
cremaster has a unique, obtuse, Y-shaped ar-
rangement of two slightly recurved conical
spines, which are fused medially into a broad
stalk and flanked by two or three small hooks
(Gardner 1948 b). Seta SV2 on Al in larvae may
be present or absent and crotchets may be homo-
ideous ur heteroideous (Gardner 1948 a). Many
stictopterines have highly variable forewing col-
our patterns, with similar recurrent themes,
while the hindwing often has a basal translucent
zone. Holloway (1985) divided the Bornean gen-
era into two monophyletic groups. All foodplant
records for Stictoptera group are from Clusia-
ceae (Guttiferae) and populations of Stictoptera
may occasionally reach levels where they damage
crop trees such as Garcinia (mangosteen). Food-
plant records for the Lophoptera group are pre-
dominantly from Dipterocarpaceae (occasionally
Tiliaceae and Euphorbiaceae).
Euteliinae. The monophyly of this subfamily
(Fig. 19.8 Q), excluding Aon, is supported by sev-
eral apomorphies (Kitching 1987), of which the
structure of the basal abdominal sternite is the
most definitive. The central membranous zone is
flanked by two internal longitudinal flanges aris-
ing from the bases of the anterior apophyses and
directed towards the midline (Holloway 1985).
This structure may stiffen the base of the abdo-
men and be associated with the characteristic
and unusual resting posture of the adults, in
which the wings are folded lengthwise and held
at an angle of 4590 to the abdomen, which
is curled dorsally or dorso-laterally. The overall
appearance is of a dried-up leaf or small cluster
of twigs. The antennae of most male euteliines
(except Anuga, Anigraea and Targalla) are
broadly and asymmetrically bipectinate over the
377
basal two-thirds and thence only narrowly so or
ciliate to the apex. This form is unique in Noctui-
dae but occurs convergently in Cossidae, Lima-
codidae and Notodontidae (Holloway 1985). The
"winged" structure of the apical basiconic sen-
silla of the proboscis may also prove apomorphic
(Kitching 1987). The anterior apophyses of fe-
male Euteliinae are often modified or lost (Hol-
loway 1985). Like Stictopterinae, larvae of Eutel-
iinae (Fig. 19.16 P) may have SV2 on Al present
or absent and the crotchets are homoideous or
heteroideous (Gardner 1948 a). Pupae lack a
cremaster (Gardner 1948 b), a feature con-
vergently derived in Nolidae. Meli (1943) recog-
nized four groups of genera within Euteliinae
based upon Penicillaria, Eutelia, Anuga and
Paedes. However, he examined very few species
and euteliine phylogeny remains unresolved. Eu-
teliinae are primarily tropical but are more char-
acteristic of savanna and semi-arid habitats than
Stictopterinae, which appear to be largely re-
stricted to forests. Foodplant records for euteli-
ines are heavily biassed towards Anacardiaceae,
and several species are pests of mango. However,
the range of foodplant families is quite extensive
(Holloway 1985).
Plusiinae. Following the studies of Eichlin &
Cunningham (1978), Kitching (1987) and Lafon-
taine & Poole (1991), the phylogeny of Plusiinae
(Fig. 19.8 S) is the best known of any noctuid
group. The inclusion of tribe Omorphinini has
yet to be confirmed (Lafontaine & Poole 1991),
but the monophyly of a clade comprising the re-
maining three monophyletic tribes (Abrostolini,
Argyrogrammatini and Plusiini) is well sup-
ported by the form of St8 and its associated hair
pencil (Kitching 1987), the detailed morphology
of the tympanal organ (Kitching 1987; Lafon-
taine & Poole 1991), biordinal crotchets (except
Plusia) and presence of a raduloid laterally on
the larval hypopharynx (Eichlin & Cunningham
1978). Although the last feature, which resembles
a sclerotized comb, is not universal within Plusii-
nae, it is not known to occur elsewhere in Noctu-
idae. Abrostoline larvae have the full comple-
ment of prolegs (although those on A3 and A4
are more slender than those on A5 and A6;
Gardner 1947), while Plusiini are semi-loopers
(Fig. 19.16 Q). In Argyrogrammatini, A3 and A4
have vestigial prolegs that lack crotchets (Eich-
lin & Cunningham 1978) (except Ctenoplusia
(Acanthoplusia); Ichinos 1962). The pupal wings
and proboscis project beyond the anterior mar-
gin of segment A5 (Mosher 1916). A sister group
relationship between Argyrogrammatini and
Plusiini is supported by four synapomorphies.
The forewing pattern often includes a character-
istic Y-, U- or V-shaped mark (giving rise to such
species names such as u-aureum and common
names as Silver-Y) and there is often a scale
tooth at the forewing tornai angle. The larval in-
tegument is covered in setose spinules (Lafon-
378 Ian J. Kitching & John E. Rawlins

Fig. 19.12. Secondary sexual features of Noctuoidea. A, Basal abdominal brush organ of male Noctuidae with
apex of brush still enclosed in pocket; ap = apdeme, br = brush, lv = lever, = pocket, Sg = Stobbe's gland,
St2 = sternite 2, St3 = sternite 3 (Noctuidae, Hadeninae, Oxythres splendens); B, Cteniophore (ct) on lateral edge
of sternite 4 (St4) (Notodontidae, Heterocampinae, Lochmaeus bilineata); C, Posterior abdominal coremata of
male arctiid in retracted position, ventral view; ac = antecosta of sternite 8, co = corema, St7 = sternite 7, St8 =
sternite 8 (Arctiidae, Arctiinae, Ctenucha virginica); D, Tubular pheromone gland of female arctiid, dorsal view
of posterior abdominal segments; A7 = segment 7, A8 = segment 8, gl = gland, os = ostium of gland (Arctiidae,
Arctiinae, Bertholdia detracta).

taine & Poole 1991). The pupae (Fig. 19.17 S)
have subdorsal grooves between tergi tes Al4
and intersegmental transverse ridges anterior to
each of the moveable abdominal segments (Na-
kamura 1974; Lafontaine & Poole 1991). Naka-
mura (1974, 1987) examined pupal chaetotaxy.
Within Plusiinae, seta SV2 is generally absent on
T2, T3 and A l . However, it has been secondarily
regained on Al in some Autographa and all Syn-
grapha, and on T2 and T3 in some Syngrapha
(Eichlin & Cunningham 1978). The distribution
of the 400 or so species of Plusiinae is distinctly
bimodal with most Abrostolini and Argyrogram-
matini being tropical and most Plusiini occurring
in temperate or subarctic biomes (Kitching
1987). Plusiines are more often associated with
open and disturbed habitats than with forests.
Many species are migratory and polyphagous on
herbaceous plants as larvae (Holloway et al.
1987). These traits combine to make a number
of plusiines economically important pests of a
wide variety of crops. Notable examples in the
Old World are Thysanoplusia orichalcea, Auto-
grapha nigrisigna, Trichoplusia ni and several spe-
cies of Chrysodeixis (Holloway et al. 1987; Com-
mon 1990). T. ni is also an important pest in the
New World, along with Pseudoplusia includens,
Autoplusia egena, Rachiplusia ou and Anagrapha
falcifera (Forbes 1954; Angulo & Weigert 1975).
However, not all are pests. Larvae of Exyra feed
on insectivorous pitcher plants (Sarracenia) and
several species of Syngrapha feed on coniferous
The Noctuoidea
trees. Many plusiines fly equally by day and
night and may be seen feeding at flowers in
bright sunshine.
Agaristinae. Members of this subfamily (Fig.
19.8 T) have long been recognized as merely spe-
cialized trifid noctuids (Jordan 1909), but they
are persistently and incorrectly referred to a dis-
tinct family (Forbes 1954; Kiriakoff 1977; Breth-
erton 1983). Mosher (1916), Gardner (1946 b,
1948 a, 1948 b) and Crumb (1956) all found no
differences in the immature stages that war-
ranted such a separation, while the presence of
basal abdominal brush organs supports inclusion
within the "higher" trifine noctuids. A number
of apomorphies support agaristine monophyly.
The counter-tympanal membrane is greatly en-
larged (Richards 1933) and associated with a re-
duced counter-tympanal hood (sometimes ab-
sent) and a paired vesicle (bulla abdominalis) at
the base of the abdomen (Kiriakoff 1977). The
ductus bursae is very long, narrow and unsclero-
tized, and leads into a small, unadorned or finely
scobinate corpus bursae (Holloway 1989). Lar-
vae (Fig. 19.16R) have a bisetose SV group on
A 7 - 9 (Fig. 19.15 C) (Crumb 1956; Rawlins
1992). Both these conditions may be apomorphic
for Agaristinae. Pupation usually occurs in a
chamber formed in soft or rotting wood. The
pupa bears a characteristic truncate cremaster
and is densely spinulose, especially dorsally
(Rawlins 1992). Correlated with a predominantly
diurnal habit, most agaristine adults are brightly
coloured and have davate antennae. Some spe-
cies indulge in territorial and hill-topping behavi-
our (McFarland 1976) while others produce
sounds in flight, either by rubbing a ridged
hindtarsus on swollen wing veins (Platagarista
tetrapleura) (Common 1990) or by a forewing
tymbal organ (Hecatesia spp.) (Bailey 1978).
Larvae are often brightly coloured and feed on
a variety of plant families, especially Onagraceae
and Vitaceae. Few are of economic importance,
although Phalaenoides glycinae damages grape-
vines in Australia (Common 1990).
Eucocytiinae. Eucocytia meeki (Fig. 19.9 A)
from Papua New Guinea, has traditionally been
included in Cocytiidae. However, it has little in
common with Cocytia other than metallic scaling
and davate antennae (which are apically blunt,
unlike Cocytia, in which they are pointed). The
counter-tympanal hood is postspiracular and the
male genitalia are phenetically similar to some
agaristines, as are the hairy eyes, clubbed anten-
nae and tuberculate but fully scaled clypeofrons.
However, the hindwing venation is quadrifine
and there are no brush organs at the base of the
male abdomen. It is here tentatively assigned to
its own subfamily, EUCOCYTIINAE stat. nov.
(Hampson 1918). Like Cocytia, nothing is
known of the immature stages of Eucocytia, and
although clearly noctuid, its affinities remain un-
known. It may be related to a Madagascan
379
group of genera comprising Epicausis, Daphoen-
ura, Adaphaenura, Eudaphaenura and perhaps
Mydrodoxa, which are currently placed in Pan-
theidae and Hadeninae.
Acontiinae. "Erastrianae" of Hampson
(1910b-1911) were divided into three sections
by Richards (1933) based upon characters of the
tympanal organ, and these were given tribal
status by Forbes (1954). Following Crumb
(1956), the first of these tribes is treated as Acon-
tiinae (Fig. 19.9 B), diagnosed by an enlarged,
heavily-sclerotized alula overlying the tympa-
num and a reduced counter-tympanal hood. The
male genitalia are often asymmetrical. Many spe-
cies are camouflaged as bird-droppings. In larvae
(Fig. 19.16 S), the spinneret is often reduced, SV2
is absent on Al and prolegs are absent on A34.
Acontiine larvae are frequently obligate feeders
on Malvales and Asteraceae, but are of no eco-
nomic importance. Cydosiini (Franclemont &
Todd 1983) (sole included genus, Cydosia) is
doubtfully included here; the male genitalia are
bizarrely asymmetrical and immature stages are
unknown.
Eustrotiinae. Two other tribes of earlier au-
thors, "Erastriini" and Eublemmini, are united
into this paraphyletic, perhaps polyphyletic, as-
semblage (Fig. 19.9 C - ) that lacks consistent
diagnostic features of adults and larvae. For ex-
ample, SV2 on Al of the larvae may be present
or absent (Crumb 1956). The Eublemma group
(Fig. 19.9 C) may be monophyletic based on en-
largement of M D and MSD setae on the larval
abdomen (Beck 1992). The east Asian species of
Deltote (which includes both Lithacodia and Eu-
strotia) and allied genera were revised by Ueda
(1984, 1987). Males of Amyna have a semitrans-
parent area on the forewing and a modified
hindwing anal area that may be involved in
sound production in flight (Common 1990).
Many feed on green plants (Figs. 19.16T) and
some are agricultural pests, such as Naranga dif-
fusa on rice (Holloway et al. 1987) and Eu-
blemma dimidialis on mung bean (Common
1990). Hostplants are frequently Poaceae, Aster-
aceae and Acanthaceae, although species are
known from Chenopodiaceae, Verbenaceae and
other unrelated families. Several Eublemma and
Catoblemma are predatory on coccids and may
be beneficial (Holloway et al. 1987); E. radda
feeds within the highly modified leaves of the in-
sectivorous plant, Nepenthes. Larvae of Enispa
feed on detritus in spiders' webs, where they are
apparently ignored by the spider (Common
1990). They even pupate in the web, suspended
without any cocoon by the cremastral hooks.
Revisionary and phylogenetic research in aconti-
ines and eustrotiines is badly needed.
Bagisarinae. This subfamily (Fig. 19.9 F) was
proposed by Crumb (1956) for the enigmatic am-
phipyrine Bagisara and the former chloephorine
genus Xanthodes. Both genera have "appendicu-
380 Ian J. Kitching & John E. Rawlins

Fig. 19.13. Head and mouthparts of larval Noctuoidea. A, Frontal view of head without secondary setae (Arctii-
dae, Arctiinae, Bertholdia sp.); B, Lateral view of head with secondary setae (Lymantriidae, Orgyia pseudotsugata):
C, Maxillolabial complex, ventral view; st = stipital lobe (Notodontidae, Heterocampinae, Lochmaeus bilineata);
D, Maxillolabial complex, ventral view; gl = cylindrical galeal lobe (Noctuidae, Catocalinae, Hypsoropha hormos),
E, External view of divided labrum; acl = anteclypeus, c = clypeus, lb = labrum (Notodontidae, Notodontinae,
Liparopsis postalbida).

late" crotchets, each bearing a large subapical
tooth (Crumb 1956; Holloway in prep.) and two
SV setae on A7 (Fig. 19.15 D; Rawlins 1992).
Appendiculate crotchets also occur in Chasmina,
Allocosmia, Oglasa, Androlymnia and Amyna
(Gardner 1947), some of which may also belong
in Bagisarinae (Holloway in prep.). Gardner
(1947) also noted similar crotchets in Anomis
(Cosmophila) and Martillada. However, in these
genera, the prolegs on A4 are well developed.
They are probably not related to Bagisarinae, in
which either the prolegs on A4 are reduced, with
minute crotchets (Fig. 19.16U), or are absent al-
together (Gardner 1947). Anomis (Cosmophila)
and Bagisarinae feed mostly on Malvales and it
is possible that the appendiculate crotchets rep-
resent a convergently derived adaptation to a
common problem involved in feeding on these
plants. The bisetose SV group on A7 in bagisar-
ines is not considered homologous with the biset-
ose SV group on A7 in Agaristinae (Rawlins
1992) or the noctuine Parabarrovia (J. D. Lafon-
taine pers. comm.).
Cuculliinae. Hampson (1906) defined Cucullii-
nae as comprising trifine noctuids with lashed
eyes but this taxon is polyphyletic (Forbes 1954;
Crumb 1956; Beck 1960; Kitching 1984). We fol-
low Fibiger & Hacker (1991) and restrict Cuculli-
inae to Cuculliini (Fig. 19.9G) and Oncocnemi-
dini (Forbes 1954), but place Amphipyra (Fig.
19.9 N) in its own subfamily. Poole (1995) ele-
vated these tribes to family status, separated Ox-
ycnemis and related genera as the psaphidine
tribe Triocnemidini, and revised the remaining
North American Cuculliini. The subspiracular
stripe of cuculliine larvae sweeps up and around
the anal plate instead of continuing down the
anal proleg as in Noctuinae and Hadeninae. This
pattern also occurs in Amphipyra and elsewhere
in Noctuidae (e. g. Hypeninae, Plathypena), and
needs more study. In Cuculliini and Oncocnemi-
dini (but not Amphipyra), larvae have biordinal
crochets, most distinctly so in Cucullia, Lathosea
and Oxycnemis (Crumb 1956; Merzheevskaya
1988). In addition, the pupal wings and probos-
cis project as a lobe onto A4 (Lafontaine &
The Noctuoidea
Poole 1991). A similar feature is seen in Plusii-
nae, where the projection is longer. Some genera
(e. g. Leucocnemis) have larvae with reduced
prolegs on A3 and A4, and are not easily distin-
guished from those of Acontiinae (Rawlins pers.
obs.). Larvae (Fig. 19.16 V) mostly feed on flow-
ers and fruits of herbaceous plants, with a de-
cided emphasis on Scrophulariaceae, Oleaceae
and Caprifoliaceae in Oncocnemidini, and Aster-
aceae in Cuculliini.
Amphipyrinae. Beck (1960) transferred Amphi-
pyra from "Amphipyrinae" (which he termed
Zenobiinae) to Cuculliinae because the larva has
a normal sized SD on A9, while almost all other
genera formerly placed in Amphipyrinae have
tonosensillae on A9 as in Hadeninae and Noctui-
nae. These genera are here considered to be ha-
denine. Pupae of Amphipyra lack the extended
wings and proboscis of Cuculliinae, the larvae
have uniordinal crochets, the male genitalia are
simple, and so we retain Amphipyra in Amphi-
pyrinae stat. rev. (Fig. 19.9N). Larvae of Amphi-
pyra feed on the leaves of deciduous trees and
shrubs (Carter & Hargreaves 1986).
Psaphidinae. This subfamily (Fig. 1 9 . 9 L - M )
comprises eight Nearctic genera, one of which,
Brachionycha, extends into the Palaearctic
(Franclemont & Todd 1983; Poole 1995). Psaphi-
dines are characterized by the presence of a
strong apical claw on the foretibia (Forbes 1954),
but this also occurs in various other noctuid
groups, including Oncocnemidini, Stiriinae and
Heliothinae. The valves of the male genitalia
bear a corona of stout setae as found in species
scattered through most trifine subfamilies, and
the harpe is near the ventral margin of the valve
as in cuculliines. The larvae have a tendency to
be humped on A8 (Forbes 1954) and known
hosts are restricted to forest trees, especially Fa-
gaceae, Betulaceae, Oleaceae, Rosaceae and Jug-
landaceae. The pupa is heavily sclerotized in an
underground cocoon, has deep pits dorsally be-
fore the cremaster, and the prothoracic femora
are concealed.
Feralia (Fig. 19.9 M) was included by Forbes
(1954) in his cuculliine tribe Psaphidini. Franc-
lemont & Todd (1983) placed it in a separate
tribe, Feraliini, formally described by Poole
(1994), which we treat as a tribe of Psaphidinae.
Poole (1995) also described Nocloini, Grotellini,
Azeniini, and Triocnemidini as tribes of Psaphi-
dinae, but we place those taxa elsewhere. Ferali-
ini lack the foretibial claw of Psaphidini and the
male genitalia are markedly different, similar to
those of Hadeninae, in that the valve has a digi-
tus with a free apex forming a pollex (Forbes
1954). However, the pupae of both tribes share
a distinctive transverse row of pits along the an-
terior margin of segment 10, a remarkable and
undoubtedly apomorphic feature of Psaphidi-
nae. Of the eight described species, six occur in
North America, one in Japan (Sugi 1968) and
381
one in the Ural Mountains (Kononenko 1984).
The known larvae (Fig. 1916 Z) all feed on coni-
fers, except F. februalis, which feeds on Fagaceae
and Myrtaceae (Crumb 1956). The crotchets are
uniordinal but otherwise larvae are similar to
those of Cuculliinae.
Dilobinae. Diloba caeruleocephala has had one
of the most chequered and varied taxonomic his-
tory of any lepidopteran (Kitching 1984). It has
been placed in Thyatirinae, Notodontidae, Ly-
mantriidae, Pantheidae, Acronictinae, Plusiinae
and its own family. A relationship with Psaphida
was first proposed by Tams (in Varley 1962),
while Minet (1983) suggested Diloba was cuculli-
ine. Kitching (1987) showed that Diloba shared
several features with Brachionycha, in particular,
unusually-shaped tegulae. However, Diloba lacks
the foreleg and pupal modifications of Psaphidi-
nae and cannot be included in that subfamily.
We allocate it to its own subfamily, Dilobinae
(Fig. 19.8 R). Females have a large, dark cor-
ethrogyne, used to conceal their eggs. Larvae
have D1 and D2 pinacula on T2 and T3 fused
and L3 on A 3 - 6 is bisetose (Merzheevskaya
1988), as in Acronictinae and Arctiidae. They
feed mainly on Rosaceae.
Condicinae. This subfamily (Fig. 19.9 J) com-
prises Condica, Ogdoconta and relatives, includ-
ing many species described in Platysenta and
Perigea. Condica, itself, contains a large number
of tropical species in need of extensive revision
(Holloway 1989). The concept of Condicinae
adopted here was first proposed by Berio (1981)
and subsequently expanded by Poole (1995). It
limits the subfamily to two tribes, Condicini with
the uncus base bearing a pair of small, well-de-
fined lateral lobes (Holloway 1989), and Leuco-
nyctini without lobes but associated on genitalic
and larval characters (Poole 1995). However,
basal lobes on the uncus occur elsewhere and
many condicines have little else in common
(M. R. Honey pers. comm.), so monophyly is
questionable. Basal abdominal brush organs and
a distinct pleurite are both absent, but the larva
bears uniordinal crochets and a tonosensillum
(piliform seta SD) on A9, a distinctive feature
of many Plusiinae, Stiriinae and most trifine spe-
cies. In addition, known larvae (Fig. 19.16X)
have the external portions of the spinneret al-
most completely reduced, similar to spinneret re-
duction in stiriine larvae. Larval foodplants are
mostly Asteraceae, in particular those containing
pyrrolizidine alkaloids (Eupatoriaeae and Sene-
cioneae), more rarely Acanthaceae (Holloway
1989).
Stiriinae and Heliothinae have long been con-
sidered to be closely related (Hardwick 1970).
The evidence was reviewed by Matthews (1991),
who noted several points of similarity, but found
no unambiguous apomorphies. Both subfamilies
feed on flowers and fruits as larvae, either ex-
posed or as borers, in contrast to a cutworm life-
382
style, but this habit also occurs in many Cucullii-
nae and some Condicinae, Hadeninae and Noc-
tuinae. A sister group relationship between Stirii-
nae and Heliothinae should be considered unde-
monstrated.
Stiriinae. This subfamily (Fig. 19.9 H I) was
restricted by Hogue (1963) to those species of
trifines with a stout, apical foretibial claw and a
sclerotized process on the frons consisting of a
raised ring surrounding a central protuberance.
Hardwick (1970) demonstrated the inadequacy
of this definition based on adult features, added
further genera including three closely related
ones grouped as Grotellini (Fig. 19.91), Groteila,
Hemigrotella and Neogrotella (Franclemont &
Todd 1983). Poole (1995) revised the Stiriinae
north of Mexico, but placed Grotellini as a tribe
of Psaphidinae. Matthews (1991) suggested that
Stiriinae are also represented in the Old World
(Panemeria, Aegle), but this requires further
study (Poole 1995, as "Panameria"). The group
is probably monophyletic and a diagnostic apo-
morphy, a greatly reduced, scale-like spinneret
(Crumb 1956), has been confirmed in a number
of stiriine genera (Rawlins pers. obs.) and Gro-
tella (Rawlins pers. obs. of larvae reared by N.
McFarland). We also include in Stiriinae those
genera grouped as Amphipyrinae: Nocloini
(Franclemont & Todd 1983), treated by Poole
(1995) as a tribe of Psaphidinae. Until larvae are
better known and the Old World fauna has been
investigated, the limits and affinities of Stiriinae
remain obscure. The group is most diverse in
seasonally dry habitats, a characteristic shared
with Heliothinae. No species is economically im-
portant. Larvae (Fig. 19.16W) of New World
species feed primarily on flowers and developing
fruits of herbaceous plants, especially composites
(Asteraceae), but Old World genera feed on flow-
ers of many families. The two known hosts of
Groteila are both in Nyctaginaceae (N. McFar-
land and R. Kendall pers. comm.). Several gen-
era have larvae that are highly specialized seed
feeders (Basilodes, Cirrhophanus, Stiria) and lar-
vae of Plagiomimicus bore into asteraceous inflo-
rescences to eat the developing achenes. The
weakly sclerotized pupa (Fig. 19.17 T) is formed
in a tight chamber constructed of fine dust or
clay particles cemented with labial secretions ap-
plied by the highly modified larval spinneret
(Rawlins pers. obs.).
Heliothinae. The monophyly of Heliothinae
(Fig. 19.9 K) is supported by two apomorphies.
The integument of heliothine larvae (Fig.
19.16 Y) is covered in conical granules with min-
ute apical spines. Spinose skin occurs in Hermin-
iinae, Cuculliinae and Plusiinae (Beck 1960;
Rawlins 1984) but these conditions are non-ho-
mologous. For example, the spinules in Plusiinae
are fine and hair-like (Lafontaine & Poole 1991).
Second, in most noctuid larvae, seta LI on T1 is
set vertically above seta L2, as it is in early in-
Ian J. P i t c h i n g & J o h n E. Rawlins
stars of Heliothinae. In later instar Heliothinae,
however, the orientation changes so that L2 lies
directly posterior to LI (Fig. 19.14 B; Hardwick
1958), although a few species exhibit intermedi-
ate states (Matthews 1991). There are no satis-
factory adult apomorphies, although the valves
are elongate and "strap-like" (Holloway 1989;
Matthews 1991). Many Heliothinae have a spiral
vesica and associated coiled appendix bursae but
this feature is homoplastic within the subfamily
(Matthews 1991). St8 in males possesses two
elongate sclerotized rods from the anterolateral
corners (Matthews 1991), but while these struc-
tures are absent in Stiriinae, they do occur in
Acronictinae and Hadeninae (Spodoptera). Some
Heliothinae have elongate, sclerotized oviposi-
tors, the precise form of which is related to ovi-
position site and is often, but not exclusively, as-
sociated with reduced fecundity (Matthews
1991). Many heliothine larvae have biordinal
crotchets (Gardner 1946 a; Crumb 1956). A phy-
logeny and classification of Heliothinae and dis-
cussion of additional characters is presented by
Matthews (1991) and Mitter, Poole & Matthews
(1993). Heliothinae also tend to be associated
with areas of lower rainfall but unlike Stiriinae,
many Heliothinae are economically important
and a number are extremely serious agricultural
pests. Their habit of feeding on flowers and
fruits, rather than leaves, results in great crop
losses. Perhaps the most destructive pests belong
to Helicoverpa, especially H. zea in the Americas
and H. armigera and H. assulta in the Old
World, all of which feed on a wide range of plant
families (Hardwick 1965; Common 1990). In the
Americas, members of the Heliothis virescens
complex (Poole, Mitter & Huettel 1993) are par-
ticularly damaging to tobacco, while Heliocheilus
albipunctella causes great losses to pearl millet in
the Sahel region of Africa (Matthews 1987).
Glottulinae. The subfamily Xanthopastinae
was erected by Biezanko (1971) for Xanthopastis
and Caridarctia, but Franclemont & Todd (1983)
resurrected Glottulidae of Guene (1852) as a
hadenine tribe for Xanthopastis and Brithys. The
two are here synonymized as Glottulinae (Fig.
19.9 O) stat. rev., and Diaphone, Polytela, Poly-
telodes, Capillamentum and Chlanidophora in-
cluded also. With the exception of Brithys, the
forewing pattern consists of transverse black stri-
ations on a pale, often pink, background, with
red and yellow elements. The larvae (Fig.
19.16AA) are also colourful, being black with
yellow stripes and spots. The chaetotaxy is unu-
sual in that the SV group is generally bisetose on
T2 and T3 as in Arctiidae (unisetose in Polytela),
but seta SD on A9 is a fine tonosensillum as in
Noctuinae and Hadeninae. Larvae feed by bor-
ing into the petioles, stems and bulbs of Amaryl-
lidaceae (especially Crinum, Hymenocallis, Pan-
cratium) (Gardner 1941; Crumb 1956; Biezanko
1971; Common 1990), and Diaphone eumelea
The Noctuoidea 383

Fig. 19.14. Semischematic configuration of setae on thorax and anterior segments of abdomen, T l A2, of larval
Noctuoidea. A, Two M D setae (MD1, MD2) on T3 and Al in Notodontidae, Heterocampinae (Heterocampa
umbrata); B, 2 SV setae (SVI, SV2) on Al and horizontal placement of L setae (LI, L2) on T l in Noctuidae,
Heliothinae (Heliothis virescens); C, 3 SV setae (SVI SV3) on A l in Noctuidae, Catocalinae (Phoberia atomaris);
D, Secondary setae in Verrucae in Noctuidae, Acronictinae (Acronicta hasta); E, Absence of seta M S D on Al and
A2 in Arctiidae, Syntominae (Syntomis flava); F, Unfused Verrucae and seta M S D on Al in Arctiidae, Arctiinae
( Virbia rosenbergi); G, Fusion of Verrucae in Arctiidae, Arctiinae, Nyctemerini (Nyctemera apicalis); H, Configura-
tion in Nolidae, Nolinae (Meganoia minuscula).

feeds on flowers and fruits of Liliaceae such as
Thuranthos, Ornithogalum, Scilla, Ornithoglos-
sum and Albuca (Pinhey 1975). The pupa is
strongly convergent on that of Arctiidae, often
covered with a pale waxy secretion and having
the wings meeting ventrally with the prothoracic
femora covered. The cremaster is stout with at
least one pair of enlarged posterolateral spines.
Hadeninae. This subfamily (Fig. 19.9 P - Q )
was defined by Hampson (1905) as comprising
trifine noctuids with hairy eyes. Usually, eyes are
classified as "hairy" only when the interfacetal
setae are long enough to be visible with a hand-
lens. But microscopic interfacetal setae occur in
most lepidopteran families (Yagi & Koyama
1963). In addition, long setae occur convergently
384
in Notodontidae (Miller 1991), Noctuidae (Noc-
tuinae, Catocalinae, Agaristinae), Pantheidae
and Nymphalidae (Satyrinae) to name but a few.
Recently, Fibiger & Hacker (1991), following
Crumb (1956), removed Xylenini and Apameini
(Fig. 19.9 Q) from Cuculliinae and Amphipyri-
nae respectively, uniting them as "Ipimorphi-
nae". However, no sound reasons exist for main-
taining this entity distinct from Hadeninae and
we include both taxa therein (as suggested by
Boursin 1964). A similar grouping, the "Hadapa-
meini", was proposed by Beck (1991, 1992) but
included Noctuinae (Beck 1960; 1992). Poole
(1995) conservatively placed most of these lin-
eages in the single subfamily Noctuinae. We pre-
fer to continue recognition of Noctuinae in the
traditional sense, excluding Ufeinae. This leaves
Hadeninae (excluding Glottulinae) as trifine noc-
tuids with a filiform SD seta on larval segment
A9, male genitalia with the harpe near the mid-
dle of the valve (Lafontaine & Poole 1991) and
a spinneret with its length greater than twice its
width. These characters are plesiomorphic and
hence Hadeninae is paraphyletic with respect to,
at least, Noctuinae, Ufeinae and Glottulinae. A
number of genera of trifine Noctuidae previously
included in Amphipyrinae are now transferred to
Hadeninae based on phenetic similarity in male
genitalia, an SD tonosensillum on larval seg-
ment A9, and plesiomorphic features of the
pupa. Amphipyrinae is restricted to only Amphi-
pyra. Most genera formerly considered amphi-
pyrine clearly belong to Hadeninae as here de-
fined, such as Nedra, Actinotia, Hyppa, Spodopt-
era, Elaphria, and Callopistria (the latter treated
as the subfamily Eriopinae by Poole 1995).
Many genera are misplaced in Hadeninae, or re-
main problematic such as Cropia and Phosphila
which were treated by Poole (1994) as the amphi-
pyrine tribe Phosphilini. As in the Catocalinae,
Hadeninae should be considered as a temporary,
paraphyletic taxon for genera of uncertain affini-
ties.
Hadeninae include numerous economically
important pests. The greatest losses are attribut-
able to species of Spodoptera. Known colloqui-
ally as armyworms because their larvae often oc-
cur in great numbers, these insects devastate
both dicotyledonous and monocotyledonous
crops (although a given species usually prefers
only one of these foodplant types) in all tropical
and subtropical areas of the world (Brown &
Dewhurst 1975; Todd & Poole 1980). Gramina-
ceous crops are also attacked by armyworms in
the genus Mythimna (including Pseudaletia and
Leucania, Franclemont 1951), borers in Sesamia
(Tams & Bowden 1953), or larvae of Mesapamea
and Cerapteryx (Carter 1984). In Europe, Pa-
nolis flammea can defoliate conifers, especially
lodgepole pines (Stoakley 1979). Many larvae
(Fig. 19.16 AB) are described and illustrated by
Godfrey (1972) and Merzheevskaya (1988). Pu-
Ian J. Kitching & John E. Rawlins
pation occurs in silk-lined chambers, usually un-
derground (Fig. 19.17 U). The Polia complex for
North America was revised by McCabe (1980),
but the limits of many genera are uncertain and
their identification difficult, particularly in Erio-
pygini, Apameini and the Mythimna-Leucania
group.
Ufeinae. This subfamily (Fig. 19.9 R) was
established (Crumb 1956) for feus, which had
previously been placed unsatisfactorily in Noctu-
inae (Forbes 1954). The larva (Fig. 19.17A) dif-
fers in having two L seta on A9 (L3 in addition
to the usual L2; Fig. 19.15 B). The proleg plantae
are exceptionally large and bear a relatively high
number of crochets. The known hosts are Salix
and Populus (Crumb 1956). Ufeus currently in-
cludes five species, found from Canada to north-
ern Mexico (a sixth species, U. carnea from
Kashmir, does not belong here and is probably a
hadenine; M. R. Honey pers. comm.). The ufeine
pupa is heavily sclerotized with a large protuber-
ant cremaster; the prothoracic femora are ex-
posed.
Noctuinae. Members of this subfamily (Fig.
19.9 S) are traditionally defined as trifine noctu-
ids with spined tibiae and unlashed, hairless eyes
(Hampson 1903). These characters are untenable
(Kitching 1984) but there are several potential
apomorphies for Noctuinae. At rest, most trifine
Noctuidae adopt a tectiform resting posture with
the posterior margins of the forewings meeting
on the midline or slightly overlapping at the tor-
nus (Tweedie & Emmet 1991). In contrast, Noc-
tuinae hold their wings flat in a horizontal plane
and with such a degree of overlap that the fore-
wing costae are almost parallel with the midline
of the body (Holloway 1989). Basal abdominal
brush organs are absent in the majority of male
Noctuinae (Birch 1979; J. D. Lafontaine pers.
comm.). First instar larvae have clubbed setae
borne on sclerotized pinacula (Common 1990).
Two tribes, Noctuini and Agrotini, are currently
recognized (Lafontaine 1987), of which the latter
is probably monophyletic. Lafontaine (1987)
considered Aniclini of Franclemont & Todd
(1983) to be a further paraphyletic group, con-
taining primitive elements of both Noctuini and
Agro tini. One lineage of Agro tini was recently
elevated to tribal status (Austrandesiini; An-
gulo & Olivares 1990). These species have a bul-
bous cornutus but the distribution of this charac-
ter is unknown. Noctuine larvae (Fig. 19.17 B)
are generally polyphagous on herbaceous angio-
sperms (Holloway 1989). Later instar larvae hide
in the soil by day or under broad-leaved weeds,
emerging after dark to cut off plant parts, which
they consume in their shelter (Forbes 1954; Com-
mon 1990). This "cutworm" behaviour causes
great damage to crops and many noctuines, espe-
cially in the genera Agrotis and Euxoa are serious
agricultural pests (Holloway et al., 1987). For ex-
ample, Agrotis ipsilon and A. segetum, which are
The Noctuoidea 385

Fig. 19.15. Semischematic configuration of setae on posterior abdominal segments, A 7 - A 1 0 , of larval Noctu-
oidea. A, Seta X on AIO in Notodontidae (Nadata gibbosa), , Two L setae (L2, L3) and filiform seta SD on
A9 in Noctuidae, Ufeinae ( i f f cus satyricus); C, Two SV setae (SVI, SV3) on A7 in Noctuidae, Agaristinae (Alypia
octomaculata); D, Two SV setae (SVI, SV2) on A7 and unmodified seta SD on A9 in Noctuidae, Bagisarinae
(Xanthodes transversa).

widespread throughout the tropics, subtropics
and warmer temperate regions, attack a wide
range of corps (Carter 1984), while A. munda
and A. infusa cause extensive damage in Aus-
tralia (Common 1990). This last species, the bo-
gong moth, is famous for its adult migrations
and aggregations in selected aestivation sites on
certain mountain tops in SE Australia (Common
1990), a behaviour duplicated by various Agrot-
ini in western North America. The European
fauna of Noctuinae has been reviewed by Fib-
inger (1990, 1993).
Pantheidae. Historically, members of this taxon
(Fig. 19.9T) have been characterized as quadri-
fine Noctuidae with hairy eyes (Hampson
1913 b), an unacceptable diagnosis (see Hadeni-
nae). Pantheids lack features thought to be char-
acteristic of noctuids, including a cylindrical ga-
leal lobe in larvae and features of the tympana
and genitalic musculature. Pantheids have sec-
ondary setae in the first larval instar and an en-
larged prespiracular Verrucae on the prothorax.
All larval instars lack middorsal glands on A6
and A7, the larval head bears secondary setae
and the LI verruca on A7 is below the level of
the spiracle. The adenosma is present in Tricho-
sea, Colocasia, Panthea and Gaujonia, but absent
in Charadra and Trisuloides. The pantheine pupa
bears a well-developed, often multisetose crem-
aster and the prothoracic femora are concealed
in many taxa (Panthea, Trisuloides (Nakamura
1987) and Charadra (Mosher 1916)). It rests in a
dense cocoon (Fig. 19.17W), similar to that of
Acronicta (Forbes 1954), which may or may not
include larval setae as in Lymantriidae (Mosher
1916). The adult eyes bear conspicuous setae and
the male genitalia have subuncal lobes.
The larvae, pupae and male genitalia of pan-
theids are strikingly similar to Acronicta, which
differ in hindwing venation and the consistently
exposed prothoracic femora of pupae. Gardner
(1946a) grouped Trichosea (as Diphthera) with
Acronicta on larval characters, while Kozhanchi-
kov (1950) considered Acronictini to be the sis-
ter-group to Pantheidae (as Momini). On the ba-
sis of the tympanum, Richards (1933) considered
the pantheids to form a relatively homogeneous
group, but one that was unplaceable within the
Noctuidae. They have an extremely modified
tympanum and, in particular, the counter-tym-
panal hood was generally completely missing.
The exact composition of the family remains un-
certain (Holloway 1985). Genera such as Pan-
thea, Trichosea, Gaujonia, Lichnoptera, Trisu-
loides, Charadra, Colocasia and Anacronicta have
similar biology, with larvae (Fig. 19.17 C) feeding
on woody plants, especially conifers, Podocarpa-
386
ceae, Fagaceae, Salicaceae, Aceraceae and Rosa-
ceae.
Lymantriidae. Although Lymantriidae (Figs.
19.10A-C) have long been regarded as mono-
phyletic, very little research has been carried out
on their phylogeny. Phylogenetic inference has
been confined to that of Kozhanchikov (1950)
who discussed relationships between lymantriids
and other Noctuoidea based primarily on fea-
tures of a few temperate genera, and placed Ly-
mantriidae sensu stricto as the subfamily Orgyi-
nae [sic], treating what we call Acronictinae and
Pantheidae as two tribes (Acronictini and Mom-
mi respectively) of the subfamily Acronictinae.
Kozhanchikov emphasized similarities in larval
chaetotaxy (D-group setae on the thorax, L-
group setae on A7), the presence of subunci
(= gnathos) in the male genitalia and a variety
of convergent reductions, including adult mouth-
parts, ocelli and life history strategies.
Many of the widely cited features of lymantri-
ids are plesiomorphies, such as the quadrifine
hindwing venation (Ferguson 1978), prespiracu-
lar counter-tympanal hood (Common 1990) and
larvae with paired Verrucae covered with dense
secondary setae (Godfrey 1987). Furthermore,
the presence in males of 13 divergent setae at
the apex of the ramus of each flagellomere (com-
pare Fig. 19.5G with Figs. 19.5 D - F ) , which is
frequently cited as being characteristic of Ly-
mantriidae, also occurs in a number of arctiine
genera related to Diacrisia (U. Dall'Asta pers.
comm.). The adenosma is present in larvae of
several tropical genera (e. g. Sarsina). The distri-
bution of this feature is complex, being absent
in Oenosandra, Doa, the "lower" Notodontidae
(Miller 1991), some Pantheidae (Charadra, Trisu-
loides), some Nolidae (Carea and most species of
Nolinae), and all known Arctiidae and Aganai-
nae. Many late-instar lymantriid larvae have
large and well-developed stipital lobes, equalling
or exceeding in size those of Notodontidae.
A number of possible apomorphies for Ly-
mantriidae have been suggested but their univer-
sality remains to be confirmed. All larvae (Fig.
19.17 D) examined so far possess a single, mid-
dorsal, eversible, often yellow to red gland on
segment A6 and usually A7 (Godfrey 1987; Hol-
loway et al. 1987). Only the gland on A6 occurs
in some Calliteara (e. g. C. pudibunda; Ferguson
1978), while externally developed gland openings
may occur on as many as six abdominal seg-
ments in other genera (Rawlins pers. obs.). Males
of many genera also possess a tymbal organ,
consisting of a pair of finely corrugated pockets
on sternite A3 (Dall'Asta 1988) (Fig. 19.6 A).
However, this organ may be both present or ab-
sent in a single genus (e. g. Lymantria; Dall'Asta
1988), and although it is likely that the feature is
apomorphic for a subset of genera, its signifi-
cance is obscured by multiple homoplastic losses.
Ian J. Kitching & J o h n E. Rawlins
The adult resting posture is also diagnostic, al-
though it is indistinguishable from that of lithosi-
ine arctiids related to Asura. The wings are held
in a broad triangle and pressed close to the sub-
strate, while the densely hairy forelegs are ex-
tended forward in front of the head (Ferguson
1978). Miller (1991) noted that female Lymantria
dispar and Dasychira obliquata possess a pair of
small, ventral setose lobes at the base of the anal
papillae and speculated that these might prove
to be an important synapomorphy for Lymantri-
idae, but such lobes occur in Aganainae (Hollo-
way 1988) and some Arctiidae (Nyctemera), al-
though apparently not in Nolidae, Pantheidae or
Noctuidae (Kitching pers. obs.).
There is currently no satisfactory suprageneric
classification of Lymantriidae. Kozhanchikov
(1950) and Ferguson (1978) both recognized only
a single subfamily for taxa recent authors have
considered to be lymantriid. Ferguson's Ly-
mantriinae grouped North American species into
two tribes, Orgyiini and Lymantriini, corre-
sponding to Kozhanchikov's Dasychira and Ar-
taxa groups. Holloway et al. (1987) thought
these tribes might merit subfamily status. How-
ever, much research still needs to be done before
monophyletic subtaxa can be confidently recog-
nized. Orgyiini (including Orgyia, Dasychira,
Calliteara and Gynaephora) are characterized
primarily by the placement and morphology of
larval setae. In most genera, Al4 have dense,
dorsal hair brushes. There are also a pair of di-
rected loose tufts on the T1 and sometimes A9,
with a similar mid-dorsal tuft on A8. The fore-
wing venation of this tribe usually incorporates
an areole (Ferguson 1978) but this feature may
comprise two or more non-homologous states
(Dall'Asta unpubl. obs.). It also varies within
genera (Arctornis) or even species (e. g. Pen-
thophera mor io) (Kozhanchikov 1950). The
valves of Orgyia and Dasychira bear a costal lobe
but this feature may simply indicate that these
two genera form a clade, as it is absent in Calli-
teara (Holloway 1982). Lymantriini (including
Lymantria, Euproctis and Leucoma) as currently
composed may be paraphyletic. The diagnostic
characters listed by Ferguson are generally ab-
sences (forewing areole absent in most species
and absence of a gnathos) or reductions (larval
hair tufts). Larvae of Lymantria and Leucoma
have closely adjacent, multisetose SD and L2
Verrucae, whereas these Verrucae in other ly-
mantriids (including Euproctis) are widely sepa-
rated. Lymantriini are thought to be restricted to
the Old World, although several Palaearctic pests
have been introduced into North America, and
some South American genera {Sarsina, Caviria)
have similar larvae. The arctic genus, Acsala,
placed by Ferguson (1978) in Lymantriidae, was
transferred to Arctiidae, Lithosiinae, by Lafon-
taine et al. (1982), and placed in its own tribe
(Acsalini) by Franclemont (1983). The neotropi-
The Noctuoidea 387

Fig. 19.16. Last instar larvae of N o c t u o i d e a . A - , N o t o d o n t i d a e : A. T h a u m e t o p o e i n a e , Anaphe pander, B, Py-

gaerinae, Clostera albosigma; C, N o t o d o n t i n a e , Furcula borealis; D, Phalerinae, Datana angusii; E, Dudusiinae,
Dudusa nobili.';; F. H e t e r o c a m p i n a e , Schizura ipomoeae; G , Nystaleinae, Dasylophia anguina; H, Dioptinae, Josia
fustula. I - A B . Noctuidae: I, Acronictinae, Acronicta ovala; J, Raphiinae, Raphia [rater; K, Herminiinae, Polypo-
gon inconspicuaiis; L, Catocalinae, Arete coerula; M, Calpinae, Eudocima homaena, penultimate instar; , Aganai-
nae. Asola plana; O . Hypeninae, Bomolocha madefactalis; P, Euteliinae, Marathyssa inficila; Q, Plusiinae, Exyra
semicrocea; R, Agaristinae, Euscirrhopterus gloveri; S, Acontiinae, Spragueia marmorea; , Eustrotiinae, Maliattha
concinnimacida; U, Bagisarinae, Xanthodes transversa; V, Cuculliinae, Calophasia lunula; W, Stiriinae, Basilodes
pepita; X, Condicinae, Condica vecors; Y, Heliothinae, Pyrrhia umbra; Z, Psaphidinae, Feralia comstocki; AA,
Glottulinae, Xanthopastis timis; AB, H a d e n i n a e , Morrisonia latex. All p h o t o g r a p h s by J. E. Rawlins.

cal genera Tepilia, Sorocaba, Rolepa and Phiditia
are bombycoids misplaced in Lymantriidae.
Lymantriidae include over 2500 described spe-
cies in nearly 360 genera, distributed primarily in
the Old World tropics. Only some 15 genera and
200 species occur in the New World (Ferguson
1978). The larvae of many genera feed on arbo-
real hosts and are frequently polyphagous (Hol-
loway et al. 1987). However, there are sizable ra-
diations specializing on flowers, fruits and low
herbs (Mylantria) and grasses (Laelia), and spe-
cies in both Asia and Africa that feed on detritus,
algae and fungi (Rawlins pers. obs.). Unlike the
arboreal Notodontidae, many lymantriids are
388
major forest pests, defoliating large areas. Such
economically important species include Ly-
mantria monacha, L. dispar, Leucoma salicis and
Euproctis chrysorrhoea (Europe, the last three
also introduced into North America; Ferguson
1978); Orgyia pseudotsugata in the western
United States (Wallner 1988); Dasychira men-
dosa, Indo-Australian tropics (Holloway et al.
1987); Leptocneria bino tata and L. reduca in
Australia (Common 1990); and species in several
genera in China (Sun 1988).
The larvae of a number of genera, most no-
tably M/?roc/-, have hollow, barbed, urticating
setae. In E. similis, these setae contain histamine,
while those of E. chrysorrhoea are more toxic,
also containing protease, esterase and phospholi-
pase A2 (Rothschild 1985). In many areas, such
larvae are sufficiently numerous to represent a
serious human health hazard (Holloway et al.
1987; Common 1990). These hairs are subse-
quently incorporated into the cocoon, from
which they are retrieved by the emerging female
in her corethrogyne and used to cover the egg
mass. In some species, the newly hatched larvae
feed in protected aggregations, their bodies cov-
ered with toxic scales deposited on the egg clus-
ter. Female scales and larval hairs thus provide
protection to all stages of the life cycle. In other
lineages, especially those feeding on detritus or
grass, the eggs may be dropped singly or in small
scale-covered clusters on the ground. There is a
tendency within Lymantriidae towards female
flightlessness (Sattler 1991). Female L. dispar are
unable to fly well, despite being fully winged.
Other Lymantria species (L. ampia), as well as
numerous other genera (Orgyia, Penthophera
and Iropoca), have females in which the wings
have become reduced to minute vestiges or com-
pletely lost and which are consequently com-
pletely flightless. Such species lay their eggs on
the outside of the cocoon (Fig. 19.17V) from
which the female emerged and the first instar lar-
vae disperse by wind on silk threads, or feed
communally forming aggregations in the early
instars.
Nolidae. This family includes those genera pre-
viously assigned to Nolinae, Sarrothripinae and
Chloephorinae. The last two have long been con-
sidered as subfamilies of Noctuidae (Hampson
1912). Nolinae (s. s.) were previously treated as
either an arctiid subfamily or a separate family
(Hampson 1900, 1914-1915), but were included
within the Noctuidae by Kitching (1984) and
Franclemont & Todd (1983). Most Nolidae do
have a dorsally broadened postspiracular bar at
the base of the abdomen, which was considered
by Minet (1986) to be a noctuid apomorphy, al-
though in many of the smaller species of Noli-
dae, the bar may be weakly sclerotized, with the
ventral part missing. A number of apomorphies
have been suggested to support the monophyly
Ian J. Kitching & John E. Rawlins
of the Nolidae. Holloway (in prep.) notes that
the tegulae and especially the patagia of nolids
are easily detached. This feature is not seen in
other noctuoids but the underlying mechanism
has yet to be investigated. Male Nolidae possess
an elongate ("bar-shaped") retinaculum (Fig.
19.4 D), similar to that of Arctiidae (Fig. 19.4 C).
Also, male genitalic muscle M.4 is directed
ventrally, rather than horizontally (Speidel &
Naumann 1995 b).
The Hampsonian taxa, Sarrothripinae and
Nolidae, were characterized in part by the pres-
ence of raised scales on the forewing upper sur-
face. However, such scales are not ubiquitous in
these taxa. Furthermore, their distribution con-
flicts with that of the basal abdominal tymbal
organs. These occur widely within the Nolidae
(Holloway in prep.), reaching their greatest com-
plexity in the Chloephorini (Fig. 19.6 B) and
Camptolomini. Within the former group, several
species have been recorded as producing sounds,
usually described as a "clicking" (Moore 1867;
Lorimer 1983). The precise function of the
tymbals remains to be elucidated. A modifica-
tion of the male genitalia, in which a flanged
structure is formed from interior extensions of
the dorsal margins of the sacculi fusing with the
juxta, may also be apomorphic for Nolidae (Hol-
loway in prep.). The larvae of most nolids (ex-
cept Earias, Selepa and Eligma) have only a sin-
gle SV seta on T2 and T3 (Gardner 1947), a
character shared with most Noctuidae but which
is also widespread in the Notodontidae (Miller
1991). In addition, the number of SV setae on
Al varies in nolids. Many have two, as found in
a large section of Noctuidae (mainly Plusiinae
and triflne subfamilies) (Gardner 1948 a). The
polarity of this character remains unconfirmed
but two SV setae on Al may be plesiomorphic
(see Noctuidae). The most characteristic nolid
apomorphy is the boat-shaped cocoon with a
vertical, anterior exit slit (Fig. 19.17X-Y) and
an unusual two-walled construction (McFarland
1980; Holloway in prep.). This feature serves not
only to unite the major groups of the family, but
also to confirm the membership of several other-
wise rather isolated taxa. The pupae of Nolidae
lack a cremaster (Gardner 1948 b; Kitching
1984), the terminal segment being rounded or
bluntly conical, often with beading or a series of
longitudinal ridges on the anterior margin (Hin-
ton 1948; Gardner 1948 b). These may function
as a defensive stridulatory organ within the co-
coon (Holloway in prep.). Meli (1943) united
Sarrothripinae and Chloephorinae into a single
subfamily, dividing this into eight tribes and a
species-group. Holloway (in prep.) proposes a
new classification for an expanded concept of
Nolidae and his treatment is employed in large
part here.
Nolinae. This subfamily (Fig. 19.10 E; Nolidae
of authors plus the Barasa group) is most clearly
The Noctuoidea
diagnosed by two apomorphies: presence of lat-
eral setose lobes on the scaphium (lost in Nola)
(Holloway in prep.) and loss of prolegs on A3
only, with the pair on A4 being fully developed
(Stehr 1987). Verrucae with secondary setae are
also present (Fig. 19.14H) but these may be ple-
siomorphic for nolids. The D1 and D2 Verrucae
are fused, as in Arctiidae. Ocelli are supposedly
absent (but see Dickens & Eaton 1973). Larvae
(Fig. 19.17 E) of several genera (McFarland
1980) retain the cast head capsules of earlier in-
stars stacked vertically above the thorax. Pupae
of Nolinae have short secondary setae arranged
around the scars of the larval Verrucae (Common
1990; Kitching & Rawlins pers. obs.) as in Arcti-
idae.
Chloephorinae. Five tribes with basal abdomi-
nal tymbal organs are here included in Chloeph-
orinae, although these structures are not univer-
sally present in all members. Sarrothripini are
primarily characterized by the position of the F
setae on the larval head (Gardner 1948 a). Some
species have the pupal prothoracic femora con-
cealed (Gardner 1948 b) as in Arctiidae, Panthei-
dae and Lymantriidae. Chloephorini have a par-
ticularly complex tymbal organ (Holloway in
prep.) and include the only genera in which
sound production has been recorded. Campto-
loma (Fig. 19.10D), the only included genus of
Camptolomini (Meli 1943; Holloway 1988), has
been variously considered to be arctiid, noctuid
or worthy of separate family status. Two SV se-
tae on T2 and T3 resemble Arctiidae and some
other nolids, but we follow Holloway in group-
ing in with lineages having complex abdominal
tymbal organs. Larvae of known members of
Careini (Fig. 19.17 F) have an apomorphic swell-
ing of thoracic segments (Gardner 1946 b), al-
though this feature also occurs in Lasiolopha of
Ariolicini (Holloway in prep.). This fifth tribe,
Ariolicini, is probably paraphyletic, there being
no unifying synapomorphies.
Westermanniinae, Eariadinae, Bleninae, Risobi-
nae, Collomeninae, Afridinae. The remaining nol-
ids considered by Holloway (in prep.) lack basal
abdominal tymbal organs and form a paraphy-
letic group consisting of seven subfamilies, an
unplaced genus-group based on Gelastocera, and
isolated genera such as Cacyparis. Three of these
subfamilies, Westermanniinae (Fig. 19.10 F),
Eariadinae (Figs. 19.10G, 19.17G)and Bleninae
(Fig. 19.10H), are strictly Old World, whereas
Risobinae includes the Nearctic Baileya (Fig.
19.101, 19.17H) and Neotropical Elaeognatha.
Holloway (in prep.) excludes the New World
Collomenini (including Iscadia) of Franclemont
and Todd (1983) based on their lack of supposed
nolid features, but this proposal requires further
study and they are treated as Collomeninae (Fig.
19.10 J). All the above groups, except Nolinae,
may form a clade based upon loss of secondary
setae. New World lineages related to Afrida lack
389
ocelli and were previously considered to be lith-
osiine arctiids (Franclemont & Todd 1983). They
do not have the distinctive synapomorphies of
Arctiidae and may be nolids of uncertain affinity,
here tentatively retained in Afridinae.
Genera such as Selepa and Eligma pose special
problems because they show both nolid and arc-
tiid features. In the larvae of both genera, there
are two SV setae on T2 and T3 and three L setae
on A 3 - 6 (Gardner 1947). However, both these
features may be plesiomorphic within Noctu-
oidea (Miller 1991) and thus provide no support
for a relationship with the Arctiidae and Noctui-
dae respectively. Females of Eligma possess a
pair of dorsal pheromone glands but these are
not homologous with those of Arctiidae (Jacob-
son & Weller in prep.). In Selepa, the prolegs
bear heteroideous crotchets indistinguishable
from those in Arctiinae. However, tymbal organs
are absent in both genera. Larvae of Selepa are
gregarious and have large eversible vesicles of
unknown function above seta SD on abdominal
segments A2 to A7; D and SD setae on the T2,
T3 and Al are mobile, particularly SD on Al.
Eligminae. We place Eligma (Fig. 19.10K) in
its own subfamily, based on the tubular phero-
mone gland in females. However, the presence of
stridulatory ridges on the inside of the cocoons
of Iscadia, Gadirtha and Plotheia suggest they
may also be related. These stridules, which rasp
against a transverse series of dorsal ridges on the
terminal abdominal segment as it wriggles or
"shivers" laterally (Hinton 1948) are absent in
known pupae of Nolinae, Chloephorinae, Risob-
inae, Bleninae, Eariadinae and Westermanniinae.
The cocoons of Eligma are elongate-ovoid, not
boat-shaped as in other nolids, and their mode
of construction appears different, but this may
be related to their large size (Meli 1943; Hem-
mingsen 1947).
There are an estimated 1400 described species
of Nolidae, in 308 genera. They have a world-
wide, but primarily palaeotropical, distribution.
While most are small moths of little or no eco-
nomic importance, several genera include a
number of agricultural pests, most notable of
which are the spiny bollworms of the subfamily
Eariadinae: Earias insulana and E. biplaga in
Africa, E. vittella in Indo-Australia and E. per-
huegeli in Australia. All are major pests of cot-
ton, causing damage by boring into the bolls
(Pinhey 1975). The communal larvae of Nola sor-
ghiella occasionally cause damage to sorghum in
the United States (Stehr 1987). Members of Ar-
cyophora visit the eyes of cattle and other domes-
tic stock at night to imbibe lachrymal juices and
thereby may transmit harmful viruses and other
infections (Reid 1954).
Arctiidae. Of the synapomorphies of adult Arctii-
dae, the most unambiguous is the presence of a
pair of dorsal, eversible pheromone glands asso-
390 Ian J. Kitching & J o h n E. Rawlins

Fig. 19.17. Last instar larvae, p u p a e , a n d c o c o o n s of N o c t u o i d e a . - B , Larvae of Noctuidae: A, Ufeinae, Ufeus

satyriats; B, Noctuinae, Agrotis malefida. C, Larva of Pantheidae, Panlhea pallescens. D, Larva of Lymantriidae,
Euproctis kanshireia. H, Larvae of Nolidae: E, Nolinae, Meganoia triangulalis; F, C h l o e p h o r i n a e , Carea varipes;
G , Eariadinae, Earias punclaria; H, Risobinae, Baileya doubledayi. I - L , Larvae of Arctiidae: I, Lithosiinae,
Chrysaeglia magnifica; J, Syntominae, Rhipidaretia danieli; K, Arctiinae, Spilosoma dubia; L, Arctiinae, Dinia sp.
M O, C o c o o n s a n d p u p a e of N o t o d o n t i d a e : M , N o t o d o n t i n a e , Liparopsis poslalbida; N, N o t o d o n t i n a e , Furcida
scolopendrina; O, Dioptinae, Phryganidia californica. P U, C o c o o n s a n d p u p a e of Noctuidae: P, Hypeninae,
Bomolocha bijugalis\ Q, Hypeninae, Rivula propinqualis; R, Calpinae, Eudocima homaena; S, Plusiinae, Exyra fax,
T, Stiriinae, Cirrhophanus triangulifer; U, H a d e n i n a e , Spodoptera mauritia. V, C o c o o n of Lymantriidae, Orgyia
postica. W, C o c o o n of Pantheidae, Panthea furcilla. XY, C o c o o n s of Nolidae: X, C h l o e p h o r i n a e , Tympanistes
fusimargo; Y, Nolinae, Meganoia sp. / A B, C o c o o n s a n d p u p a e of Arctiidae: Z, Lithosiinae, Schislophleps bi-
puncta; A A , S y n t o m i n a e , Rhipidaretia danieli; AB, Arctiinae, undescribed Arctiini, D o m i n i c a n Republic. All p h o -
t o g r a p h s by J. E. Rawlins.

dated with the anal papillae of females (Fig.
19.12D). These glands can be quite long (al-
though the branches are short and broad in
Lithosiinae; Holloway 1988), branched or un-
branched, and are apparently everted by haemo-
static pressure and retracted by longitudinal
muscles (MacFarlane & Earle 1970). In addition,
there are metathoracic tymbal organs, consisting
of a series of corrugations or grooves (micro-
tymbals) on the katepisternum (Fig. 19.6C).
The Noctuoidea
Tymbal organs occur in both sexes and all sub-
families, although they are reduced or absent in
many Syntominae, Euchromiini and Ctenuchini.
Sequential contraction of the basalare muscle
(Blest et al. 1963) deforms the ridges in rapid
succession to produce a burst of ultrasonic clicks
(Fenton & Roeder 1974). Sound production is
elicited by tactile stimulation or in response to
the hunting calls of bats (Watson 1975). Dunning
(1968) postulated that the sounds act as aural
aposematic signals, while Fullard et al. (1979)
suggested that they jammed chiropteran sonar.
Sanderford & Conner (1990) showed that in at
least one species, sounds are produced during
courtship. Watson (1975) noted the possibility of
Batesian mimicry. Lithosiinae, Arctiinae and
some Syntominae have prothoracic glands from
which they exude a liquid, sometimes foul-smell-
ing, when disturbed (Common 1990). This liquid
contains acetylcholine and histamine, and proba-
bly also pyrazines, the characteristic odour of
which is considered to signal distasteful or toxic
properties to predators (Rothschild 1985). Exu-
dation is often accompanied by reflex immobili-
zation and the display of aposematic coloration
(Blest 1964).
Two larval apomorphies support the mono-
phyly of Arctiidae: D1 and D2 on T2 and T3 are
fused into a single verruca and seta L3 on A3 6
is bisetose. The latter feature can be observed in
genera that have only primary setae, such as Ty-
ria, Utetheisa, Amerila and Nodozama, but not in
larvae with multisetose Verrucae. Both features
are independently derived in the noctuid genera
Acronicta and Diloba. The larvae of all known
Arctiidae lack an adenosma. The plantae of the
larval prolegs are often drawn out (Holloway
1988) but the significance of this has yet to be
evaluated. Most arctiids have heteroideous
crotchets, but there are genera in all subfamilies
with the homoideous condition, including all
Syntominae. Arctiidae have been traditionally
diagnosed by a basally swollen hindwing vein Sc
+ R but this is not universally present nor easy
to observe. The prespiracular counter-tympanal
hood is plesiomorphic. Three subfamilies are
here recognized: Lithosiinae, Syntominae and
Arctiinae. Lithosiinae and Syntominae form a
monophyletic group, as both lack seta MSD on
the abdomen, a synapomorphic condition un-
known elsewhere in the Noctuoidea (Fig.
19.14 E).
Lithosiinae. This subfamily (Fig. 19.10 LM)
has been characterized as arctiids that have lost
the ocelli (Pinhey 1975). However, this is incor-
rect, as ocelli are often extremely small and diffi-
cult to detect (Dickens & Eaton 1973). Their
presence in some genera (Macrobrochis) and ap-
parent absence in some non-lithosiines (e. g. Pa-
gara, Pygoctenucha, both Arctiinae) has caused
much confusion. The monophyly of Lithosiinae
is supported by larval mandibles with an en-
391
larged, basal molar area that is used to macerate
algae and lichens prior to ingestion (Rawlins
1984). Unlike the conical or spherical eggs of
other arctiids, which are covered with polygonal
networks of raised costulae, the eggs of lithosi-
ines are very smooth, without distinct costulae
and uniformly covered with shallow depressions.
Genitalia, venation and larval setal configura-
tions exhibit great variation. The Eilema group
is distinctive in Arctiidae by having Dl + D2 on
T2 and T3 located behind a large SD2 verruca
("horizontal" orientation of Forbes 1960). In
some groups of genera, SD2 is fused with D1
and D2, and others (Hypoprepia, Cisthene) lack
secondary setae. Both homoideous and heteroi-
deous crotchets occur. The form of the tymbal
organ, in which the microtymbals consist of fine,
shallow grooves, each with a single modified
scale (Forbes & Franclemont 1957), is distinc-
tive, but a number of variations and reductions
prevents this feature from being diagnostic for
all species. Finally, the thoracic spiracle of the
lithosiine pupa is fused shut, in many species ap-
pearing lost. Many species have elongate narrow
forewings with modified venation. Most com-
monly, Sc fuses for most of its length with R and
either Rs4 or MA2 may be absent (Fig. 19.3 D).
Where known, lithosiine larvae (Fig. 19.171) feed
on algae or the algal component of lichens (Raw-
lins 1984; Habeck 1987 b), occasionally liver-
worts (Forbes 1960) and mosses (Holloway
1988). Pupae vary greatly in size and shape, from
squat brown pupae in dense cocoons (Asura,
Crambidia), to brightly coloured ones in sparse
cocoons bearing complex arrays of larval setae
(Cyana, Bizone, Chamaita, Schistophleps, Fig.
19.17 Z). Tropical Lithosiinae have been very
poorly studied (but see Birket-Smith 1965).
There are numerous undescribed species in both
hemispheres, and the diversity of larval and pu-
pal structures exceeds that of all other arctiid lin-
eages. A number of genera are currently incor-
rectly placed in the subfamily and much remains
to be discovered about their biology and imma-
ture morphology. Lithosiinae are not economi-
cally important.
Syntominae. The monophyly of Syntominae
(Fig. 1 9 . 1 0 N - 0 ) is tentatively supported by the
loss of both the male frenulum and hindwing
vein A2 (except Meganaclia), and the complete
fusion of hindwing veins Sc, Rs and MAI (Fig.
19.31; Holloway 1988). Fusion of hindwing veins
Sc and Rs only, with a strong MAI, is a non-
homologous condition in Euchromiini and Cten-
uchini, while MA2 is considered to have been
lost independently in Syntomini and Euchromi-
ini (cf. Holloway 1988). We recognize two tribes
that have not been associated since the time of
Hampson (1898, 1914). Thyretini (Fig. 19.100)
have recently been treated as a separate but re-
lated family (KiriakofT 1970 a; Pinhey 1975) or
as a subfamily of Notodontidae (Minet 1983).
392
However, although the tympanal organ superfi-
cially appears to be directed ventrally (Kiriakoff
1949), the structural modifications are not ho-
mologous with those of Notodontidae. Thyretini
are unequivocally arctiid as many possess tymbal
organs and paired pheromone glands (Holloway
1988). The immature stages are very similar to
those of Syntomini. Syntomini (Fig. 19.ION) has
been grouped by many authors with Ctenuchini
and Euchromiini (Holloway 1988) in the hetero-
geneous family Ctenuchidae. But all known Syn-
tomini and Thyretini have homoideous crotchets
(Rawlins pers. obs.). Larval segments T2 and T3
have D l , D2 and SD2 fused into a large flat-
tened verruca that is only slightly separated or
narrowly connected to the pinaculum surround-
ing the tonosensillum SD. In contrast, Euch-
romiini and Ctenuchini have D l , D2 and SD2
fused into a prominent, circular verruca remote
from SD and always have heteroideous crotch-
ets. Lack of tymbal organs is interpreted as a
loss (contra Minet 1986) because these structures
occur in the sister group, Thyretini. Several gen-
era of Syntomini (e. g. Anapisa, Dysauxes) have
larval habits and morphology identical to Thyre-
tini and may be misplaced. Eggs, pupae and co-
coons (Fig. 19.17AA) are very similar to those
of Arctiinae (Fig. 19.17 AB). Many species are
diurnal and most mimic Hymenoptera, having
black or brown wings with translucent white or
yellow patches and an abdomen that is often
banded with red. Larvae of many species feed on
flowers (especially herbaceous composites) and a
few species are minor pests (Amata sperbius on
graminaceous crops; Holloway et al. 1987). Algi-
vory and associated lichenivory have been ob-
served in many genera of Syntominae, including
several genera of Thyretini (Rawlins pers. obs.).
In addition, detritivory and fungivory occur in
genera of both tribes. Many species have larvae
(Fig. 19.17 J) found on the ground, but feeding
on algae, sooty mould, lichens, detritus and
fungi has been often overlooked, and many spe-
cies in captivity develop on herbaceous plants
and flowers when preferred algae or fungi are
not available (Rawlins pers. obs.).
Arctiinae. This subfamily (Fig. 19.10P-T) is
tentatively considered to be monophyletic, based
upon an elongate male retinaculum (this is re-
duced or shortened in Lithosiinae, especially
small species, or absent in Syntominae, and sim-
ilar retinacula occur in Nolidae and Noctuidae:
Aganainae) and heteroideous crotchets (al-
though the homoideous condition occurs in Vir-
bia and Holomelina). Most subdivisions recog-
nized in the past are no more than genus-groups
based upon distinctive taxa. Ferguson (1985) dis-
cussed the relationships between world genera of
Arctiini (Figs. 19.10P, 19.17K), providing a key
and diagnoses to six groups centred on Holomel-
ina, Arctia, Spilosoma, Apantesis and Phragma-
tobia. In a study of the genitalia of Palaearctic
Ian J. K i t c h i n g & J o h n E. Rawlins
and Oriental Arctiinae, Kda (1987, 1988) recog-
nized six genus-groups, based on Amer ila, Uteth-
eisa, Callimorpha, Arctia, Rhyparioides and
Spilarctia. He concluded that each was mono-
phyletic but proposed no relationships between
them. Most species of Arctiinae are part of neo-
tropical radiations, traditionally placed by vari-
ous authors in artificial taxa, such as Ctenuchi-
dae, Pericopidae and Phaegopterinae. While
each of these groups contains numerous distinc-
tive genera, their interrelationships are unclear.
For convenience, these artificial groupings may
be treated as tribes (Arctiini, Ctenuchini, Euch-
romiini, Pericopini, Phaegopterini, Callimor-
phini, Nyctemerini, Utetheisini) and membership
largely follows Hampson (1898, 1901, 1914,
1920) and Forbes (1939, 1960). Attempts to char-
acterize the larvae and pupae of these tribes have
been misleading, as most available material is
from temperate regions, and is neither represen-
tative nor comprehensive enough to establish
meaningful generalizations about character dis-
tributions. By and large, Phaegopterini (Fig.
19.10S) and Pericopini (Fig. 19.10R) are pa-
raphyletic assemblages of strictly New World
taxa with relatively complete venation. From
these are derived several lineages with reduced
venation, assigned to the paraphyletic tribes
Euchromiini (Fig. 19.10T) and Ctenuchini.
Phaegopterines from a number of distinctive
genus-groups centred on Amastus, Carales,
Euchaetes, Halysidota, Hyperthaema, Idalus,
Melese, Neritos, Ormetica and Zatrephes. Larvae
feed primarily on woody plants (especially Ruta-
ceae, Myrtaceae, Asteraceae, Fabaceae, Euphor-
biaceae, Ulmaceae) but there are species that
feed on herbaceous dicotyledons, monocotyle-
dons and even algae. The Euchaetes group feed
on laticiferous plants (Euphorbiaceae, Apocy-
nales). Euchromiini and Ctenuchini are charac-
terized by homoplastic losses of vein Se + RI in
the hindwing. Euchromiini have branches of the
cubital vein stalked or fused, and Ctenuchini
have the cubital branches separate. Larvae of the
Palaeotropical genus, Euchromia, feed on Con-
volvulaceae. It is closely related to New World
genera feeding on that family, Asteraceae, Sapin-
daceae, Moraceae and many other plants. Cten-
uchini consists of several unrelated groups, in-
cluding a diverse one feeding on Poaceae and
Cyperaceae (Fig. 19.17 L; Ctenucha, Dinia,
Trichura, Cisseps, Philoros) and another, centred
on Theages and Eucereon, associated with laticif-
erous plants such as Moraceae and Apocynales.
This last group shares characters with the
Euchaetes group in the Phaegopterini and to-
gether may form a distinctive lineage. The Be-
lemniini of Forbes (1939) are simply one lineage
of Ctenuchini and do not merit tribal status
(Rawlins unpubl.). Finally, Pericopini are a pa-
raphyletic group characterized by plesiomorphic
hindwing venation with vein Se + RI only fused
The Noctuoidea 393

Fig. 19.18. Pupae of Noctuoidea. AC, Female pupa without secondary setae, Aso ta (Noctuidae, Aganainae); A,
Dorsal view; B, Ventral view; C, Lateral view; D, Pupa with secondary setae, lateral view, Euproctis (Lymantri-

for a short distance with Rs and by a divided or
deeply bifid uncus. Pericopine larvae feed on
plant groups known to contain pyrrolizidine
alkaloids, the Eupatoriaeae and Senecioneae of
the Asteraceae and various Boraginaceae. Lar-
vae, genitalia and hostplants suggest affinities
to generic groups in Phaegopterini and Ctenu-
chini.
In Arctiini and Callimorphini, larval segments
T2 and T3 have the primitive chaetotaxic config-
uration (Fig. 19.14 F) with six distinct and sepa-
rate Verrucae (Dl + D2, SD, SD2, LI + L2,
394
L3 and SV). Fusion of these Verrucae in various
ways occurs in all other tribes, confounding re-
peated attempts to use them as diagnostic fea-
tures (Forbes 1939; Habeck 1987 a, 1987 b,
1987 c; Common 1990). Most larvae of Arctiini
and Callimorphini also have a very large LI ver-
ruca located immediately behind the spiracle on
A7. Secondary cranial setae are usually absent,
but d o occur in species of both tribes. The Old
World Nyctemerini are characterized by fusion
of Verrucae D l and D2 on the abdominal seg-
ments, and fusion of D l , D2 and SD2 on seg-
ments T2 and T3 (Fig. 19.14G). Fusion of those
Verrucae on T2 and T3 has apparently occurred
many times in New World Phaegopterini and is
the usual (but not universal) condition in Peri-
copini, Ctenuchini and Euchromiini. Secondary
setae are missing in Utetheisini (Argina, Amphi-
callia, Utetheisa), a monophyletic lineage (Fig.
19.10 Q) with larvae feeding on plants containing
pyrrolizidine alkaloids (Crotalaria in Fabaceae
and various Boraginaceae).
Hostplants containing pyrrolizidine alkaloids
are widespread in Arctiidae and have been impli-
cated both in defence (Rothschild et al. 1979)
and in the synthesis of adult pheromones (Con-
nor et al. 1981). Krasnoff and Roelofs (1990) hy-
pothesized that ancestral Arctiinae utilized pher-
omones derived from pyrrolizidine alkaloids ob-
tained from larval hostplants. Subsequent lin-
eages may have evolved alternative reproductive
strategies, becoming less dependent on plants
containing such alkaloids and therefore more po-
lyphagous. Male arctiids often have specialized
coremata. These glandular organs are usually ab-
dominal and occur on almost any abdominal
segment, although they are most often located
on the eight sternite (Fig. 19.12C).
Hymenopteran mimicry is frequent within lin-
eages with reduced venation. The most extreme
forms are exhibited by ctenuchines such as Pseu-
dosphex, in which the basal abdominal segments
are constricted to form a "wasp-waist" and in
species where longitudinal wing folding resem-
bles the condition in vespid wasps (Danforth and
Michener 1988).
Recent illustrated works have been produced
on Arctiidae of Borneo (Holloway 1988), Europe
(Freina & Witt 1987), Japan (Inoue et al. 1982),
Nepal (Haruta 1992, 1993), southern Africa (Pin-
hey 1975), Taiwan (Chang 1989) and larvae in
Spain (Gmez 1986). Neotropical Arctiinae (ex-
cept for Ctenuchini) were catalogued by Wat-
son & Goodger (1986). Callimorpha dominula, a
species with highly variable wing pattern, played
a significant role in the early development of eco-
logical genetics (Kettlewell 1943). In some areas,
most notably the "Valley of the Butterflies" on
the island of Rhodes (Walker 1966), aestivating
adults of Euplagia quadripunctaria form vast ag-
gregations. Very few of the 4,400 currently valid
species of Arctiinae are of major economic im-
Ian J. Kitching & John E. Rawlins
portance, although some of the polyphagous
species will damage field crops. The fall web-
worm, Hyphantria cunea, causes considerable
damage to fruit and shade trees, both in its na-
tive North America and in Europe, where it was
accidentally introduced (Carter 1984). In con-
trast, Tyria jacobaeae was deliberately released in
western North America (Habeck 1987 b) and
New Zealand as a biological control agent for
Senecio (Asteraceae).
Acknowledgements
We thank the following specialists for advice, en-
couragement and unpublished information: D. J.
Carter, U. Dall'Asta, J. G. Franclemont, J. D.
Holloway, M. R. Honey, R. O. Kendall, J. D.
Lafontaine, . McFarland, J. S. Miller, M. J.
Scoble, S. J. Weiler, and C. W. Young. All illus-
trations were provided by M. A. Klingler, Carne-
gie Museum, and all photographs by J. E. Raw-
lins.
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 20. Evolution of Larval Food Preferences in Lepidoptera
Jerry A. Powell, Charles Mitter & Brian Farrell
Introduction
The Lepidoptera constitute probably the largest
radiation of phytophagous insects (Scoble 1992),
rivalled only by the coleopteran clade Phyto-
phaga (Chrysomeloidea + Curculionoidea;
Crowson 1981). The majority of lepidopteran
larvae live at the expense of living seed plants,
and virtually all orders of b o t h gymnosperms
and angiosperms, as well as ferns, liverworts and
mosses, are used. The Lepidoptera date at least
to the early Jurassic ( 1 - 2 ) , and their m a j o r radi-
ations probably began by at least the early Creta-
ceous, at about the same time as those of flower-
ing plants. Moreover, m a n y lepidopteran clades
are restricted primarily to particular higher taxa
of angiosperms. These facts suggest that the di-
versification of lepidopteran lineages might have
paralleled, perhaps even been promoted by, that
of their hostplants (Ehrlich and Raven 1964),
and that the present distribution of Lepidoptera
a m o n g host taxa might reflect such long-stand-
ing interactions.
Comprehensive review of larval foods within
m a j o r clades of Lepidoptera, however, raises
d o u b t that m a j o r features of lepidopteran evolu-
tion reflect interactive coevolution, or even par-
allel phylogenesis, with plants. Larvae of Lepi-
doptera feed on a vast array of substrates, in-
cluding not only all kinds of living plant materi-
als, but detritus of b o t h plant and animal origin
and occasionally living insects. Non-phytopha-
gous habits, while in the minority, are f o u n d pri-
marily in basal lineages of some m a j o r clades
(e. g. Tineoidea), suggesting that these might
have had non-phytophagous ancestors. We focus
this necessarily brief review on phytophages and
to a lesser extent on detritivores, and will only
mention carnivores in passing, where these occur
in each group. We refer the reader to the recent
comprehensive review by Pierce (1995). A strik-
ing generalization may be made concerning the
origins of lepidopteran carnivory. Of the more
than 200 species in eight superfamilies which are
known to be obligate carnivores or parasitoids,
nearly all attack only sedentary insects encoun-
tered on or near hostplants. With a notable ex-
ception in the geometrids, these caterpillars
largely feed on scale insects or other homopter-
ans and/or ant b r o o d or eggs (see Pierce 1995).
Considering just phytophagous lineages, there
is scant evidence for correspondence of lepidop-
teran phylogeny at higher levels to relationships
a m o n g m a j o r plant groups. Thus, analysis of a
general survey of larval food records more than
a decade ago (Powell 1980) showed that while a
few primitive taxa are associated with primitive
plants (bryophytes, gymnosperms), all the larger
superfamilies of both non-ditrysian and ditrysian
Lepidoptera feed on a wide variety of angio-
sperms. N o superfamily or higher taxon of Glos-
sata is primarily or even primitively associated
with non-flowering plants, though there are spo-
radic, clearly secondary transfers to such hosts
by unrelated species and genera. Moreover, the
differences a m o n g lepidopteran taxa in how
plants are used are often as pronounced as those
in which hosts are used. Larvae may bore in
roots, under bark, or in stems; feed on leaves or
within them as miners; induce galls or feed
within galls caused by other insects or patho-
gens; or feed in flowers, fruits or immature or
mature seeds. It is possible that the plant-re-
source "adaptive zones" in which lepidopteran
taxa have radiated are better described by such
ecological categories or "horizons" (Powell 1980)
than by plant taxonomy or chemistry.
In this chapter we summarize the evidence for
and against these contrasting views of feeding
habit evolution, focusing on the following ques-
tions:
1. Does specialized feeding on living plants have
a single origin in Lepidoptera, or has it arisen
multiple times independently? By what interven-
ing stages has it arisen f r o m , or given rise to,
non-phytophagous habits?
2. A m o n g phytophagous lineages, to what ex-
tent does evolution of host use reflect plant tax-
o n o m y or secondary chemistry, as opposed to
other ecological aspects of plant resources? To
what extent d o extant host associations reflect
parallel cladogenesis or reciprocally interactive
coevolution with hostplants?
We examine these questions separately for
non-ditrysian and ditrysian lineages, and briefly
contrast diet-evolutionary patterns in Lepidop-
tera to those in two comparably old phyto-
phage clades.
O u r review benefits f r o m m a j o r advances in
paleontology and phylogeny reconstruction, par-
ticularly for primitive Lepidoptera, during the
past 20 years. However, enormous gaps remain
in our knowledge, which impose limits on our
ability to d r a w conclusions. Lepidopteran fossils
are scarce compared with those for many hard
bodied insects (Labandeira and Sepkoski 1993),
larval fossils are virtually non existent, and it is
rarely possible to interpret the larval foods of
fossil taxa, though leaf miners are an important
exception (Labandeira et al. 1994). Hence, recon-
struction of the evolution of feeding habits de-
404 Jerry . Powell, Charles Mitter & Brian Farrell

NON-DITRYSIANS
approx. no. typical larval
descr. spp. feeding site/habit typical larval hosts chief distribution
Micropterigidae 100 humid forest floor bryophytes; herbs; detritus cosmopolitan
Agathiphagidae 2 seed borers Agalhis (Araucariaceae) Aus., S.W. Pacific
Heterobathmiidae 10 leaf miners Nothofagus S. So. Am.
Eriocraniidae 25 leaf miners 90 % on Fagales Holarctic
Acantherop- 5 [leaf miner] [Rhamnaceae]
W. No.Am.
leroctelidae
3 ?? miner S. Aus.
Lophocoronidae unknown
Neopseustidae 10 ?? miner unknown S. Asia, S. So. Am.
Mnesarchaeidae 6 humid forest floor spores; algae; bryophytes New Zealand
Hepialoidea 500+ underground or cosmop., esp.
in roots or stems generalists Australia
Nepticulidae 600+ leaf miners (a)_>40 fam., esp. Fagales, Rosales (a)cosmopolitan
(b)Myrtaceae (b) Australia
100 miners- leaves,bark, (a)Nothofagus (a)Chile
stems, fruits (b)Betulac., Saxifragac.,
Ranuncul., Polygon., others (b)cosmopolitan
100 leaf miners; pupate >17 fam., esp. Corn., Vit.(Holarc.),
in case Myrt. (Aus.) cosmopolitan
270 leaf miners; later instars >18 families (a)Afr., So. Am.
generally case bearers; (b) Holarctic
some in litter or detritus (a) Myrt., Proteac; Nothofagus (a)Aus., S. So. Am., Afr.
(b) > 12 fam.; some polyphag. (b)Holarctic
44 borers- stems,seeds (aJ?Myrtaceae (a)S. So. Am.
(b)Rosac., Saxifr., Eric.; (b) Holarctic
Prodoxinae on Agavaceae
Cecidosidae 7 gall formers Anacard iaceae S. So. Am; S. Afr.
Crinopterigidae 1 mine from portable case Cistaceae Europe
Tischeriidae 80 leaf miners Fagac., Rosac., Asterac..others mostly Holarctic
Palaephatidae 30 [twig or leaf tiers] [Verben.;Proteac.] S. So. Am., Aus,
DITRYSIA (Figure 2)

Fig. 20.1. Phylogenetic synopsis of larval feeding habits in primitive moths (non-Ditrysians). Cladogram follows
Kristensen and Nielsen (12, 5); sources for within-superfamily phylogeny, and all other information, in text, (a)
and (b) denote basal dichotomies within families. Square brackets denote habits inferred from very few observa-
tions or species. Habits of Lophocoronidae, Neopseustidae inferred from ovipositor morphology. Except as noted,
hosts indicated to family only.

pends largely on phylogenetic inference f r o m liv- nomically widespread or numerically predomi-

ing species. If changes in host are more frequent
than speciation, or if species with ancestral hab-
its are extinct, such reconstructions may be mis-
leading; for example, Van Nieukerken (1986) ar-
gued that in Nepticulidae, m a n y extant primitive
genera may have derived hostplant preferences.
A second limitation is the preliminary state of
knowledge of extant larval feeding habits, espe-
cially for tropical and Southern Hemisphere fau-
nas. Probably 75% of available larval food re-
cords for Lepidoptera come f r o m temperate
zones, while as many as 75% of the described
species of Lepidoptera may be tropical (Heppner
1991), with the true disparity undoubtedly still
greater. Given the floristic differences a m o n g re-
gions, this bias may seriously affect our recon-
structions of evolutionary patterns. F o r example,
plants in the relatively derived orders Fagales,
Rosales and Asterales appear to dominate lepi-
dopteran host associations (our plant classifica-
tion follows Cronquist [1981] unless otherwise
noted). But this may be in part an artifact of the
predominance of these taxa in the Holarctic (e. g.
Van Nieukerken 1986): Fagales and Rosales, at
least, are much less conspicuous elements in the
Southern Hemisphere flora.
A third problem is the rudimentary under-
standing of phylogeny in most groups, particu-
larly in the Ditrysia. Lacking detailed phyloge-
nies, we may be tempted to assume that taxo-
n a n t habits represent the groundplan, a poten-
tially misleading inference. Because uncertainty
about phylogeny often can be traced to igno-
rance about tropical and southern continent
forms, inferences about ancestral feeding habits
cannot be m a d e with confidence. For example,
the m a j o r obstacle to understanding of relation-
ships within the Tortricoidea ( H o r a k and Brown
1991) is insufficient knowledge of critical taxa
f r o m the M a l a y a n - P a p u a n region and southern
South America, areas for which we also have al-
most no larval food data.
Phylogeny of Lepidopteran Feeding
Habits: a Synopsis
O u r review of m a j o r trends in the evolution of
feeding habits emphasizes the basal lepidopteran
grades and is not exhaustive; detailed accounts
for individual groups are given elsewhere in this
volume. With exceptions to be noted, we follow
the phylogenetic arrangement and nomenclature
of Nielsen (1989), as updated by Kristensen
(15). Phylogenetic uncertainties are discussed
when these affect possible interpretations of
feeding habit evolution. The synopsis below is
mapped in summary f o r m on a cladogram (Figs.
2 0 . 1 , 2 , 3) in hopes of facilitating critical discus-
sion of evolutionary hypotheses.
Evolution o f L a r v a l F o o d Preferences in Lepidoptera 405

BASAL DITRYSIANS
typical larval
approx. no. feeding chief
TINEOIDEA descr. spp. site/habitat typical larval hosts distribution
Tineidae 2300 detritivory widespread, probably primitive cosmop., mostly tropical
J _ Erlocottidae 200 probably detritivorous Oriental, Ethiopian
^ ^ Acrolophidae 250 detritivorous New World, mostly tropical
Psychidae 1000 feed inside case polyphagous on living plants; some cosmop.; 90% O.W.
GRACILLARIOIDEA scavengers,lichenlvores
Gracillariidae 1600 leaf miners >80 fam., esp. Fagac., Fabaceae cosmop.

a Bucculatricidae 200 leaf miners >20 fam., esp. Asterac., Fagales cosmop., 75% N. Hemisph.
Roeslerstammidae >20 miners; extern, Epacrid.Proteac., Elaeoc., Paleare., Orient., Aus.
in later instars Fagac.; some polyphages
Oouglasiidae 25 miners herbs., esp. Rosac., Borag., mostly Holarctic
YPONOMEUTOIDEA Hydrophyll.
, Acrolepiidae <100 facultat. leaf miners, 5 fam., esp. Aster., Solan.,Liliac. Holarc., Neotrop., Hawaii
skeletonizers
Argyresthildae 160 leaf miners; later instars >13 fam.; 40% on conifers, 25% on mostly Holarctic
in silken shelters Rosaceae
Yponomeutidae 900 extern, folivores 50 fam., incl. ferns, gymnosp. mostly Holarctic
Ochsenheimeriidae miners, borers monocots, esp. Poac. Palaearctic
Heliodinidae 50 facultat. miners diverse dicots.; 50 % on mostly New World
Caryophyllales
Glyphipterigidae 400 stem & seed miners monocots, esp. Cyper., June. cosmop., primarily tropical
Lyonetiidae 250 leaf miners >27 fam.; a few polyphagous cosmop.
GELECHIOIDEA
Stenomatidae 1200 leaf tiers, 16 fam., mostly woody; esp. Myrt. cosmop.; 90% Neotropical
ext. feeders (S. Hemisph.), Fagales (Nearc).
Ethmiidae 250 within or on flowers, Boraginales (80%) cosmopolitan, esp. Neotrop
buds, or leaves
, Depressariidae 600 tiers, rollers, Ivs, > 17 fam., esp . Apiac., Asterac. cosmopolitan
flowers, seeds
, Elachistidae s.s. 250 leaf or stem miners monocots (89%); primitive cosmop., esp. New World
genera on Asterldae
. Agonoxenidae 95 borers, miners, Rosaceae, cosmopolitan
webbers Arecaceae, others
a Xylorictidae 500 leaf, bark feeders from >20 fam., esp. Proteac., Myrt. Oriental, Australian
stem tunnels, tied
leaves, etc.
Scythrididae 370 ext. feeders, leaf >20 fam, esp. Aster., Cistac. cosmop.; mostly Holarctic
tiers, miners In Holarctlc; some polyph.
Chimabachidae 6 tiers, rollers,Ivs, flwrs [polyphagous] Palaearctic
Oecophoridae 3000 case makers; most eat Myrtaceae In Australia cosmop., esp. Australia
dead plant material
Stathmopodidae 150 concealed feeders, in plant refuse or on living plants cosmop., esp. tropics
Lecithocerldae 500 feed on leaf litter most Austr., Oriental
Batrachedrldae 125 live plants; scavengers diverse cosmopolitan
Coleophorldae 1400 leaf miners; from portable >30 fam., woody & herbac., ind. cosmop.; 82 % Holarctic
case In later instars Aster., Betul., Rosac., Juncac.
Momphidae 60 miners, gallers Onagraceae (66%) mostly Holarctic
Blastobasidae 300 mostly detritivorous cosmop., esp. New World
Pterolonchidae 8 [root borer] [Asteraceae] Palaearctic; S. Africa
Symmocldae 170 mostly detritivorous cosmop., esp. Palaearctic
Peleopodidae 25 [leaf webber] [Verbenac., Malpighiac.,others] cosmopolitan
Cosmopterigidae 1200 miners, gallers, tiers, >35 fam., esp. Poac., Fabac. cosmopolitan
scavengers
Gelechiidae 4000 miners, borers, gallers, >80 fam., incl. Pinac., Myrt., cosmopolitan
ext. feeders & others morocots,mosses, ferns
APODITRYSIA (Figure 3)

Fig. 20.2. Phylogenetic synopsis o f larval feeding habits in primitive Ditrysians. Sources f o r phylogenetic hypothe-
ses in text. Sources f o r data in this and subsequent figure, when not referenced in text, include: H e p p n e r 1991;
H o d g e s 1974, 1978, 1 - 9 ; M u n r o e 1982; P o w e l l 1980; Scoble 1992; Stehr 1987; M . A . Solis, pers. c o m m . Square
brackets denote habits inferred f r o m very f e w observations or species. Except as noted, hosts indicated to family
only.

Editor's note: T h e names Z e u g l o p t e r a , Aglossata Smith 1984). R e c e n t r e p o r t s , h o w e v e r , indicate

and Heterobathmiina have purposely been retained in r e s t r i c t i o n o f t h e Sabatinca group to bryophytes
this chapter, though they are discarded in the preced-
b u t f e e d i n g o n a w i d e r a n g e o f f o o d s in Micro-
ing systematics accounts ( 1 2 , 13, 14).
pterix, including herbaceous angiosperms, gras-
ses, a n d p l a n t d e t r i t u s ( L o r e n z 1961, C a r t e r a n d
Dugdale 1982, Kristensen 1984, Davis 1987,
CLADE ZEUGLOPTERA C o m m o n 1990, N i e l s e n a n d C o m m o n 1991). T h e
MICROPTERIGOIDEA. Micropterigidae live in adults eat pollen and spores (Nielsen and Com-
humid forest floor habitats. Their distribution mon 1991).
appears relictual, c o s m o p o l i t a n w i t h highest ge-
neric and species diversity in the Palaearctic,
western Pacific, and Australian regions (Hepp- CLADE AGLOSSATA
n e r 1991). A G A T H I P H A G O I D E A . A g a t h i p h a g i d s are k n o w n
Traditionally micropterigid larvae have been f r o m just t w o species, in Q u e e n s l a n d a n d islands
characterized as specialists on bryophytes s. 1. o f the s o u t h w e s t P a c i f i c . T h e blind, a p o d o u s lar-
( e . g . I m m s et al. 1957, P o w e l l 1980, T u s k e s a n d v a e d e v e l o p w i t h i n t h e s e e d s o f Agathis (Araucar-
406 Jerry . Powell, Charles Mitter & Brian Farrell

APODITRYSIA
approx. no. typical
descr. spp. larval habit typical hosts chief distribution
Cossidae 670 borers, rootcrowns >17 fam. esp. Fabac.; cosmop., esp. tropics

& stems >20% polyph.
Dudgeonidae [stem borer] [Rubiaceae] Palaeotropical
Castniidae 200 borers monocots tropical S.A., Asia, Aus.
Sesiidae 1000 borers >32 fam., shrubs & vines cosmopolitan
Dalceridae 85 ext. folivores trees, shrubs; many polyph. Neotropical
Limacodidae 1000 ext. folivores trees & shrubs;most polyphag. cosmop., esp. tropics
Zygaenidae 800 ext. folivores many fam., esp. Vitaceae cosmop., esp. tropics
Megalopygidae 230 ext. folivores trees & shrubs, many fam. mostly Neotropical
Epipyropidae.Cyclotornidae - parasites of insects
Choreutidae 350 leaf tiers >18 fam., herbac. & woody cosmop., esp . O.W .tropics
Tortricidae 6600 leaf tiers; borers; herbs, shrubs, trees; cosmopolitan
detritiv.; others many fam.,incl. conifers
Urodidae 60 external folivores broadleaved trees mostly Neotropical
Schrecken- >5 web, skeletonize [Anacard., Rosac., Scroph.] cosmopolitan
steiniidae fruits, leaves
Epermeniidae 70 leaf miners; flower & Apiac., Santalac. cosmopolitan
fruit borers; ext. foliv.
Alucitoidea 140 leaf tiers; fruit & [woody Asteridae] cosmop.; mostly Indo-Aus.
seed borers
Pterophoridae 500 leaf miners; later instars [herbs & shrubs, incl. cosmopolitan
borers, leaf tiers, Asteridae]
OBTECTOMERA exposed feeders
Immidae 280 ext. folivores [Violac., Myrtac.] mainly Indo-Aus.
Copromorphidae 60 borers, tiers, esp. on [Eric.,Berberid.] tropics, mainly Indo-Aus.
flowers, fruits
Carposinidae 250 borers, leaf miners >7 fam., mostly woody pantropical
mm Hybiaeidae 20 leaf tiers [Verbenac., Bignoniac.] tropics, mainly O .W.
PYRALOIDEA
M Crambidae 11,700 diverse forms of subgroups specialized on cosmopolitan
concealed feeding diverse dicots and mono-
on living plants cots;some on lichens,
Pyralidae s.str.
mosses
Chrysauginae 400 leaf tiers; borers - cosmopolitan
seeds, fruits, roots, diverse fam., esp. Bignon.
stems; saprophages
Galleriinae 300 eating dry vegetable matter; some in nests of Hymenoptera cosmopolitan
Pyralinae 900 scavengers, esp . on dry plant material; some leaf feeders cosmopolitan
Phycitinae 4000 leaf rollers, borers; some on dry plant material, scavengers cosmopolitan
Epipaschiinae 700 leaf tiers, miners many woody fam., often pantropical
w/ secretory canals
Thyrididae 600 in stems or rolled leaves [many families] cosmop., esp. tropics
m MACROLEPIDOPTERA [12 superfamllies, 33 families, 87,000 species; most ext. folivores; many polyphagous]

Fig. 20.3. Phylogenetic synopsis of larval feeding habits in Apoditrysia. Sources for phylogenetic hypotheses in
text. Sources for other data see caption to previous Figure. Square brackets denote habits inferred from very
few observations or species. Except as noted, hosts indicated to family only.

iaceae). This association may be very ancient.
The earliest fossils ascribed to Lepidoptera are
lower Jurassic and are postulated, though with-
out rigorous cladistic analysis, to resemble Ag-
lossata more than Zeugloptera (Whalley 1986).
Fossil Araucariaceae are also known from the
early Jurassic, in both hemispheres (Townrow
1969). The placement in Figure 1 fits the mor-
phological evidence best and is corroborated by
new data from 18 S r D N A sequences (Wiegmann
1994). However, some characters would put
agathiphagids as the most basal lepidopteran lin-
eage, or as sister group to heterobathmiids
(1-2).
dibulate adults are believed to feed on host pol-
len. The fossil record of Nothofagus and of Lepi-
doptera of more derived lines than Heterobath-
mia extend well back into the Cretaceous, which
suggests that this strict larval host association is
the most primitive of a lepidopteran with an an-
giosperm (Kristensen and Nielsen 1983).
C L A D E GLOSSATA
The relationships among the remaining non-
Neolepidopteran families depicted in Fig. 20.1,
left unresolved by Nielsen (1989), follow Kris-
tensen (15).

ERIOCRANIOIDEA. Eriocraniidae are the basal

CLADE HETEROBATHMIINA
HETEROBATHMIOIDEA. This recently discov-
ered group is known from about 10 species in
southern South America. The larvae mine the
leaves of Nothofagus (Fagaceae), while the man-
family in this lineage. The 25 described species
are exclusively Holarctic (Heppner 1991). The
larvae are apodous leafminers of deciduous
woody angiosperms; 90% use Fagaceae or Betu-
laceae, while one species uses Rosaceae and Sali-
Evolution of Larval Food Preferences in Lepidoptera
caceae (Davis 1978; Powell 1980). The life cycle,
timed a r o u n d the early-spring availability of new
leaf tissue for larval development, resembles that
of heterobathmiids (Kristensen and Nielsen
1983).
ACANTHOPTEROCTETOIDEA. Acanthopteroc-
tetidae consist of five species in western N o r t h
America (Davis 1978), and Catapterix Zaguly-
aev & Sinev (1988) in Crimea, which has been
treated as a separate family (Scoble 1992) but
should be assigned here (N. P. Kristensen pers.
comm.). The larva of one Californian species has
been described, a leaf miner in Ceanothus
(Rhamnaceae) (Davis and Frack 1987).
Larval habits of Lophocoronidae and Neo-
pseustidae are not known, but possession by
these families of a piercing ovipositor, similar
(especially in Lophocoronidae) to that of Eri-
ocraniidae and Acanthopteroctetidae, suggests
that they are endophagous ( C o m m o n 1990,
Davis 1975, Kristensen, 15).
CLADE EXOPORIA
MNESARCHAEOIDEA. Mnesarchaeidae, one of
two sister lineages making u p the Exoporia, are
known only f r o m six New Zealand species found
in the periphyton layer of moist forest floors
characterized by mosses and liverworts. The lar-
vae are unspecialized, exophagous feeders that
appear to be generally phytophagous, feeding on
fern sporangia, fungal spores, and algae in addi-
tion to bryophytes (Gibbs 1979).
HEPIALOIDEA. Families in the other exoporian
lineage, Hepialoidea, need redefinition (Scoble
1992), as there are several genera, mostly South-
ern Hemisphere with u n k n o w n larval biologies,
whose placement is unclear. The Hepialidae
s. str. are the most diverse group of non-hetero-
neurans, with more t h a n 500 described species,
distributed in all faunal regions. Australia, with
more than 100 species, is the center of diversity
( C o m m o n 1990). The larvae, at least in late in-
stars, f o r m tunnels in sod or in soil and feed ex-
ternally on roots or surface vegetation, or tunnel
into roots or trunks of woody plants. A few spe-
cies appear to be specialists on mosses (Grehan
and Patrick 1984), ferns (McCabe and Wagner
1989), grasses (Pinhey 1975) or woody plants
such as conifers (Wagner et al. 1991) or Myrta-
ceae ( C o m m o n 1990), but these are exceptional.
Although hepialid species have been recorded in
older literature feeding on a wide variety of an-
giosperme, recent detailed studies have led to the
conclusion that most hepialids are generalists
(Grehan 1984, 1989; Wagner 1989), and often
detritivorous or mycophagous in early instars,
graduating to roots or stems of vascular plants
in later instars (Grehan 1981, 1987, 1989). Gen-
eralist feeding probably is ancestral (Grehan
1989).
407
CLADE HETERONEURA
The remaining superfamilies make u p the Heter-
oneura (Scoble 1992), relationships within which
are not definitively settled. Basal members of the
clade possess a piercing ovipositor by which the
eggs are inserted into plant tissue, where endoph-
agous larval feeding occurs, at least by early in-
stars.
NEPTICULOIDEA. This superfamily consists of
the sister families Opostegidae and Nepticulidae.
Larvae of Nepticulidae typically are miners in
mature leaves of woody angiosperms, though
some are stem miners or petiole gall formers. The
primitive Australian Pectinivalvinae, which feed
on Myrtaceae ( C o m m o n 1990), appear to be sis-
ter group to the cosmopolitan remainder of the
family (Van Nieukerken 1986). The latter are re-
corded f r o m over 40 angiosperm families, with a
preponderance of records for Fagales and Ro-
sales in the Holarctic (Powell 1980, Van Nieuker-
ken 1986, Fig. 1). The larger genera are not asso-
ciated with particular plant lineages, but several
smaller subgenera are almost completely limited
to one plant family, such as Anacardiaceae, Poly-
gonaceae, Fabaceae, or Cistaceae (Van Nieuker-
ken 1986).
A late Jurassic or early Cretaceous leaf mine
on a fossil seed fern in Australia was interpreted
by Rozefelds (1988) as lepidopteran and possibly
nepticulid. The mine f o r m is comparable to some
modern Nepticulidae, but this fossil may actually
be a dipteran rather than a lepidopteran; see also
1 2. Mid Cretaceous (97 mya) mines in leaves
of Magnoliidae (Lauraceae), Hamamelidae
(Platanaceae), and Rosidae have been identified
as representatives of two extant genera of Nepti-
culidae (Labandeira et al. 1994), and there are
late Cretaceous nepticulid mines in Fagales
(Skalski 1979, Van Nieukerken 1986).
The larvae of Opostegidae f o r m subcutaneous
mines, sometimes in bark, stems or fruit, on vari-
ous angiosperms (Powell 1980, Davis 1987). The
Chilean Notiopostega, proposed as sister group
to the bulk of this cosmopolitan family, mines
the cambium layer in trunks of Nothofagus
(Carey et al. 1978, Davis 1989). Other genera
mine leaves of Rutaceae (several Hawaiian spe-
cies), and stems of Betulaceae, Saxifragaceae,
Ranunculaceae and Polygonaceae in the H o -
larctic.
INCURVARIOIDEA. Relationships a m o n g the
six families of this superfamily were proposed by
Nielsen and Davis (1985). Foodplants are diverse
and larval biologies vary. Later instars of some
genera construct a portable case in which pupa-
tion (Heliozelidae) and larval feeding (some In-
curvariidae, Adelidae) take place. Strictly en-
d o p h a g o u s larvae that f o r m n o case (Cecidosi-
dae, derived Prodoxidae) are believed to be de-
rived (Nielsen and Davis 1985, Wagner and Pow-
ell 1988).
408
Heliozelidae, with about 100 described and
many described species, occur in all faunal re-
gions. They are leafminers in at least 17 angio-
sperm families, with numerical preponderance in
Vitaceae and Cornaceae in the Holarctic and
Myrtaceae in Australia (Common 1990; Powell
1980, unpublished data from Costa Rica).
Adelidae include 270 species (Heppner 1991).
The basal dichotomy appears to lie between a
mainly Southern Hemisphere Nematopogon
group of genera, and the primarily Holarctic Ad-
ela group, which are considered more derived
(Nielsen 1980). In Adela, larvae feed at first in
immature seeds, then in fallen leaves from a por-
table case. Oviposition by each species is specific
to one or two closely related plant genera (Pow-
ell 1969). Hosts include at least 18 angiosperm
families, with no strong numerical or geographi-
cal emphasis on one or a few, although Cauchas,
apparently the most basal lineage of the Adela
group, specializes on Brassicaceae (Powell 1980).
Nematopogon species evidently are non-specific
in oviposition preferences and feed mainly on
leaf litter, but the biologies are poorly known.
The few host records for other members of the
Nematopogon group are from Fabaceae in Africa
and Australia (Janse 1945, Common 1990).
The single, European species constituting Cri-
nopterygidae (Nielsen and Davis 1985) feeds on
Cistaceae, mining from a portable case in the
manner of coleophorid Gelechioidea (Petersen
1978).
Incurvariidae comprise a worldwide group of
about 135 species, 80% Palaearctic and Austra-
lian (Heppner 1991). Some genera mine through-
out larval life, while others leave the mine to feed
in later instars on fallen leaves (Davis 1987,
Common 1990). The basal, southern continent
groups specialize on Myrtaceae and Proteaceae,
while the more derived, Holarctic genera use
about 10 unrelated angiosperm families. A few
species are polyphagous (Nielsen 1981, 1982,
Nielsen and Davis 1981, Powell 1980, Scoble
1980).
Cecidosidae (including Ridiaschinidae, Niel-
sen and Davis 1985) are southern South Ameri-
can and southern African; the larvae cause galls
on Anacardiaceae, in which the apodous larvae
feed throughout growth (Powell 1980).
Prodoxidae are Holarctic, except for the
southern South American Prodoxoides, which
appears to be sister group to the rest of the fam-
ily (Nielsen and Davis 1985). The biology of Pro-
doxoides is unknown, but it is believed to be as-
sociated with Myrtaceae. Basal genera of the Ho-
larctic clade specialize on Rosaceae, Ericaceae,
and Saxifragaceae (Davis et al. 1992). The more
derived genera (yucca moths and allies = Pro-
doxinae of Davis 1987) are entirely endophagous
feeders on Agavaceae, the only non-ditrysians to
adapt to monocots (Powell 1992). Molecular
data indicate that this shift to monocots and the
Jerry A. Powell, Charles Mitter & Brian Farrell
correlated behavioral/morphological diversifica-
tion of the prodoxine genera occurred over a
brief interval during the Tertiary (Brown et al.
1994).
TISCHERIOIDEA. Tischeriidae are primarily Ho-
larctic, with about 70 described species (16).
The larvae make stout, silk-lined mines in which
pupation occurs. They are reported on seven
host plant families, primarily Fagaceae, Rosa-
ceae and Asteraceae (Braun 1972; Powell 1980);
a few species feed on annual herbs (Davis 1987).
PALAEPHATOIDEA. Palaephatidae consist of
five genera in southern South America and one
in Australia (Davis 1986; Nielsen 1987), totalling
about 30 species. A single larva observed in Ar-
gentina spun together the twigs of a verbena-
ceous shrub (Davis 1986) and may have fed on
dead leaves, while in Australia one larva was
found between leaves of Proteaceae (Common
1990).
DITRYSIAN CLADE
Phylogenetic relationships within Ditrysia, which
includes 98% of the described Lepidoptera spe-
cies, are not well resolved. Following Nielsen
(1989), and with exceptions to be noted, we
adopt the definitions of families, superfamilies
and several more inclusive groups proposed or
accepted by Minet (1986, 1990, 1991). The most
inclusive of these clades is the Apoditrysia
( 1 - 2 ) , and within it, the Obtectomera (Fig. 3),
consisting of those apoditrysians with both a
non-motile pupa at eclosion and a modified pul-
villus. While this scheme is tentative, there seems
little doubt that the non-apoditrysian groups, Ti-
neoidea, Gracillarioidea, Yponomeutoidea, and
Gelechioidea, are the most primitive (Fig. 20.2);
our arrangement of these follows Kristensen
(1 2). Because of the enormous diversity of Dit-
rysia, we attempt only a brief overview, particu-
larly in Apoditrysia, with focus on a few groups
in which recent phylogenetic studies alter the pic-
ture of feeding habit evolution, since Powell
(1980).
PRIMITIVE (NON-APODITRYSIAN)
SUPERFAMILIES
TINEOIDEA. We follow recent restrictions on the
definition of tineoids, including removal of Lyo-
netiidae (s. s.) and Ochsenheimeriidae to Ypono-
meutoidea (Kyrki 1984, 1990), and proposal of
the superfamily Gracillarioidea for the leaf-min-
ing gracillariid-bucculatricid set of families
(Davis 1988, cited in Nielsen 1989; Robinson
1988; Minet 1991). Relationships shown among
the remaining families (Fig. 20.2) follow Robin-
son (1988) and Robinson & Nielsen (1993).
Evolution of Larval Food Preferences in Lepidoptera
Tineidae include some 3,500 described species,
mostly tropical (Heppner 1991, Robinson and
Nielsen 1993). Feeding habits vary considerably
among the 15 subfamilies recognized by Robin-
son and Nielsen (1993), but generalized fun-
givores/detritivores occur in all the major sub-
families and probably represent the ancestral
condition (Robinson and Nielsen 1993). As sum-
marized by Robinson and Nielsen, Powell
(1980), and Davis (1987), Tineinae tend to be as-
sociated with mammals or social insects, and
their larvae are often keratophagous or coproph-
agous, although many are detritivores, and some
consume ant brood (Pierce 1995); Nemapogoni-
nae and Scardiinae mostly specialize on wood-
rot fungi, especially Polyporaceae (Lawrence and
Powell 1969); many or most Meessiinae prefer
lichens; Setomorphinae tend to feed on dead
plant matter; Hieroxestinae feed mostly on fungi
or dead plant materials but sometimes on living
plants, while members of one genus are coproph-
agous in caves (Robinson 1980, Zimmermann
1978); and many Myrmecozelinae are detriti-
vores and live in grass turf (Zagulyaev 1971),
caves, or mammal burrows (Davis et al. 1986,
Robinson 1980).
Larvae of Eriocottidae, a family of about 200
primarily Oriental and Ethiopian described spe-
cies (Heppner 1991), are thought to be detriti-
vores, perhaps endophagous in decaying wood
or leaf litter (Common 1990, Davis 1990 citing
Zagulyaev, Nielsen 1978).
There are 250 + described species of Acro-
lophidae in a single New World, mostly tropical
genus. The few known larvae are detritivorous,
feeding from tunnels in grass turf and sugarcane,
or in polypore fungi (Davis 1990); one species is
coprohagous and detritivorous in tortoise bur-
rows (Davis and Milstrey 1988).
Psychidae are cosmopolitan (90% Old World),
numbering about 800 described species (17).
Many species in the more primitive subfamily
Taleporiinae feed on lichens, some live in ant
nests where they are presumed to be scavengers,
and others are polyphagous on trees and shrubs.
Most Psychinae evidently are generalist herbi-
vores, some moving from initial herb feeding to
shrubs or trees in later instars (Common 1990,
Davis 1987, Powell 1980).
GRACILLARIOIDEA. Gracillariidae make up
the most species rich family of predominantly
leaf mining Lepidoptera: the 1,600 described spe-
cies undoubtedly represent fewer than half the
total. Most are specialists on one or a few closely
related plant species, usually woody angio-
sperms, and at least 80 families of hosts are re-
corded (Common 1990, Powell 1980). There is
no strong preference for particular plant taxa by
any of the three subfamilies. On a world basis,
Fagaceae and Fabaceae are most often recorded
(Powell 1980), but records are generally lacking
409
for tropical regions. Larval mines identified as
phyllocnistine Gracillariidae have been described
from mid Cretaceous (97 mya) Magnoliidae of
three genera representing two families (Laban-
deira et al. 1994). This is by far the earliest re-
corded instance of ditrysian leaf mining.
Bucculatricidae are a monogeneric leaf mining
family with about 175 described species (17),
occurring in all geographical regions, 75% in the
Northern Hemisphere (Heppner 1991). Host-
plant records that were summarized with Lyo-
netiidae (Powell 1980) represent one gymno-
sperm and 20 angiosperm families, predomi-
nantly Asteraceae (45%) and Fagales (23%), do-
minated by Nearctic records.
Roeslerstammiidae (= Amphitheridae) are
found in the Palaearctic, Oriental and Australian
regions. The larvae mine in leaves in early in-
stars, then feed externally from a slight web or
fully exposed, on Epacridaceae, Proteaceae,
Elaeocarpaceae and Fagaceae (Common 1990).
Roeslerstammia in Europe is polyphagous
(Kyrki 1983 a).
Douglasiidae comprise about 25 Holarctic
species plus one each in the Orient and Australia
(Heppner 1991). We follow Davis (1988) and Mi-
net (1991) in treating them as Gracillarioidea; ti-
neoid, yponomeutoid and other affinities also
have been proposed. The larvae mine herbaceous
angiosperms, primarily Rosaceae and Boragina-
ceae-Hydrophyllaceae (Powell 1980, Scoble
1992).
YPONOMEUTOIDEA. This superfamily has been
redefined recently. Kyrki (1983 b, 1984) recog-
nized lineages, though not defining families, that
represent Acrolepiidae, Argyresthiidae, Lyonetii-
dae (s. s.), Heliodinidae, Yponomeutidae (includ-
ing Plutellidae), and Ochsenheimeriidae. Minet
(1986) added to this list Glyphipterigidae (in-
cluding Orthotelidae, Kyrki and Itmies 1986),
which had been placed in Copromorphoidea, but
subordinated Acrolepiidae and Argyresthiidae to
Yponomeutidae. Kyrki (1990) again revised the
taxonomic status of the groups (18), but the
characters employed in his cladistic analysis need
rvaluation; the separation of Ypsolophidae
from Plutellidae in particular seems unwar-
ranted.
Acrolepiidae are a mostly Holarctic group of
80 + species, also occurring in the Neotropical
region and in Hawaii (Gaedike 1984, Heppner
1987, 1991, Zimmerman 1978). The larvae are
facultative leafminers or skeletonize the leaf sur-
face from slight webs, using plants of five un-
related families, with 30% of the records from
Asteraceae (Europe), 26% Solanaceae (Neotropi-
cal and endemic species in Hawaii), and 30%
Liliaceae (Holarctic) (Gaedike 1984, records in-
cluded in Yponomeutidae by Powell 1980, Zim-
merman 1978).
410
Argyresthiidae (Yponomeutidae, Argyresthii-
nae, 1 - 8 ) include about 160 species, primarily
Holarctic. The larvae are leafminers in early in-
stars, usually leaving the mines to form silken
shelters in new foliage. Some conifer-feeding spe-
cies remain in the mines until maturity. At least
13 gymnosperm and dicot families are used by
monophagous or oligophagous species of Argyr-
esthia; more than 40% of recorded species use
conifers, 26% Cupressaceae, the highest propor-
tional adaptation to conifers by any moth family.
Nearly 25% specialize on Rosaceae in the Pa-
laearctic (Moriuti 1977, Powell 1980).
Heliodinidae are mostly New World moths;
there are 75 + described species, more than half
Neotropical, with a few from other tropical areas
and Australia, and a considerably larger fauna is
likely (Common 1990, Heppner 1991, Y.-F. Hsu
unpubl. data, Powell 1991). The larvae often feed
on thick-leaved plants such as Abronia (Nyctagi-
naceae) in facultative mines. Knowledge of food-
plants has been increased greatly during recent
studies by Y.-F. Hsu. There are larval food re-
cords for 32 mainly Holarctic and American sub-
tropical species, from several unrelated dicotyle-
donous families, with pronounced specialization
on Caryophyllales (Aizoaceae, Chenopodiaceae,
Nyctaginaceae, Portulacaceae) (Powell 1980,
1991, Hsu and Powell unpubl. data).
Glyphipterigidae include nearly 400 described
species distributed in all faunal regions, primar-
ily at lower latitudes (Heppner 1982, 1991, Com-
mon 1990). The larvae of most species feed in
stems and seeds of Cyperaceae, Juncaceae and
other monocots (Heppner 1985, Powell 1980),
though a few use herbaceous Asteraceae.
Yponomeutidae (including Plutellidae and Yp-
solophidae of Kyrki 1990, 1 - 8 ) as defined by
Minet (1986) number nearly 900 species, and oc-
cur in all geographical regions. The larvae are
external feeders on leaves, living communally in
some genera. Plants in at least 50 families are
used, including a fern (Cyathaceae) and a gym-
nosperm (Podocarpaceae) in New Zealand, other
gymnosperms (Ephedraceae, Cupressaceae, Pi-
naceae) in the Holarctic, monocots (Cyperaceae,
Poaceae), and Proteaceae and Myrtaceae in the
Indo-Australian region. There is a predominance
of records from Brassicaceae, Cupressaceae, Fa-
gaceae, Celastraceae, and Rosaceae, especially
the last two, in the Holarctic (Moriuti 1977,
Powell 1980).
Lyonetiidae s. str. (18) are widely distributed
and include about 250 described species (Com-
mon 1990, Heppner 1991). The larvae are leaf-
miners in at least 27 families of angiosperms,
with no concentration on any one or a few re-
lated families. A few species use monocots; a few
in the Holarctic are polyphagous (Common
1990, lyonetiid records other than Phyllocnistis
and Bucculatrix from Powell 1980). About 17%
Jerry . Powell, Charles Mitter & Brian Farrell
of 75 records are Fabaceae, mainly in the western
Palaearctic, India, and Japan.
GELECHIOIDEA. This is the largest superfamily
of lower Ditrysia by far, and the species numbers
may rival those of Noctuidae and Pyralidae be-
cause vast numbers of species remain unde-
scribed. In better known temperate faunas (Aus-
tralia, Europe, and probably the Nearctic), Gel-
echioidea are more species rich than any other
superfamily. Although there is agreement on the
phylogenetic integrity of the superfamily (Com-
mon 1990; Hodges 1978 and 1 - 9 , Minet 1986,
1990, 1991), there have been wide differences of
opinion on the number and composition of in-
cluded families and their relationships (e. g. Kuz-
netsov and Stekol'nikov 1979, Minet 1990,1-9).
As a result, analysis of gelechioid larval substrate
evolution is tentative. Recent studies have begun
to agree on major lineages. Minet (1990) distin-
guished a clade consisting of Oecophoridae s. str.
and allies, and an unresolved residual assem-
blage including Elachistidae, Gelechiidae and
others, while Hodges (19) proposes a broader
oecophorid assemblage and a more restricted
elachistid assemblage. Our nomenclature follows
Minet (1990), except that we retain family status
for several biologically distinctive groups in-
cluded as subfamilies in Elachistidae, Xyloricti-
dae, or Coleophoridae by Minet or Hodges, such
as Stenomatidae, Ethmiidae, Agonoxenidae,
Scythrididae, and Blastobasidae. Our arrange-
ment (Fig. 20.2) follows Hodges (19).
In the subset of the oecophorid clade con-
sisting of Oecophoridae s. str. and its nearest rel-
atives (Fig. 20.2), detritivory is predominant
among the basal lineages and is possibly primi-
tive. Oecophoridae s. str. are a dominant group
in Australia (ca. 1,850 described species in 250 +
genera, Common 1994), a radiation that Com-
mon (1990) attributes to exploitation of the
highly diverse Myrtaceae flora, particularly the
persistent leaf litter; 95% of the larval food re-
cords are for living (28%) or fallen leaves of
Myrtaceae (Common 1990, 1994; Australian re-
cords in Powell 1980 from Common pers.
comm.). Elsewhere, at least 25 dicot families are
eaten, but nearly half the records represent det-
ritivory, on bark or wood of dead trees, plant
refuse, wood-rot fungi, ground litter, or stored
products (Powell 1980). Larvae of Stathmopodi-
nae live in fallen leaves, fruit or other decaying
vegetable matter, on living plants, usually in cat-
kins (including Araucariaceae), galls, fruits, seed
heads, bark, or as predators of scale insects
(Common 1990, Powell 1980). Lecithoceridae in
Australia resemble Oecophoridae in feeding on
leaf litter beneath Myrtaceae and grass tussocks
(Common 1990). Blastobasidae (Coleophoridae,
Blastobasinae, 19) and Symmocidae (Autos-
tichidae, Symmocinae, 19) as well as the re-
maining Autostichidae sensu Hodges (19) are
Evolution of Larval Food Preferences in Lepidoptera
also mostly detritivorous (Adamski 1989, Powell
1976 a, 1980), though some prey on scale insects
(Pierce 1995).
The remaining gelechioids, including the rest
of the oecophorid clade and the elachistid assem-
blage (Elachistidae of Hodges, 19), are primar-
ily living plant feeders. Scythrididae (Xyloricti-
dae, Scythridinae, 1 - 9 ) include 370 described
species mostly from the western Palaearctic, and
probably 400 + undescribed species in the Nearc-
tic (Landry 1991). Larvae of only a small frac-
tion of species are known, but at least 20 families
of angiosperms are used. Cistaceae and Astera-
ceae dominate Palaearctic records; a few species
use Poaceae, lichens, or mosses (Bengtsson 1984;
Landry 1991; Powell 1980). Polyphagy may be
more prevalent than the scattered records indi-
cate (Powell 1976 b). The cosmopolitan Coleoph-
oridae (Coleophorinae, 19), with about 1,400
described species, 82% in the Holarctic (Heppner
1991), and 5 0 0 + undescribed Nearctic species
(J.-F. Landry, pers. comm.), feed on more than
30 families of angiosperms. The major host taxa
are shared with those of the more primitive leaf
mining moths, especially Betulaceae, Rosaceae,
and Asteraceae. Many coleophorines use Junca-
ceae (Powell 1980), and some feed on scale in-
sects (Pierce 1995).
Gelechiidae, with more than 4,000 described
species (Heppner 1991) and probably at least as
many more known that are undescribed, feed on
an extremely diverse array of plants, with more
than 80 families recorded, especially Fabaceae
and Asteraceae, with appreciable numbers using
Pinaceae, Myrtaceae, monocots, and a few on
mosses and ferns (Powell 1980).
Stenomatidae (Elachistidae, Stenomatinae,
19) are richest in the Neotropical Region (90%
of 1,200 described species; Heppner 1991), and
foodplants are therefore poorly documented, but
larvae been recorded on at least 16 angiosperm
families, predominantly Myrtaceae (Southern
Hemisphere) and Fagales (Nearctic).
Lesser clades within Gelechioidea, probably
derived, specialize on derived kinds of angio-
sperms. Examples include Elachistidae (Elachis-
tinae, 1 - 9 ) on monocots (89%); Cosmopterigi-
dae (Cosmopteriginae, 19) on monocots;
Momphidae (Coleophoridae, Momphinae, 1 - 9 )
on Onagraceae (70% +); and Ethmiidae (Elachi-
stidae, Ethmiinae, 1 - 9 ) on Boraginales (Hydro-
phyllaceae, Ehretiaceae, Boraginaceae) (80%)
(Powell 1980, 1983, Janzen and Powell, unpubl.
data from USA and Costa Rica). Momphidae
and Cosmopterigidae also contain several preda-
tors of scale insects (Pierce 1995).
A P O D I T R Y S I A N C L A D E ( 1 - 2 ; Fig. 20.3)
Primitive (non-obtectomeran) apoditrysians are
nearly all living plant feeders. Many have
adopted leaf tying or other forms of partial con-
411
cealment, but there are also clades of both borers
and external leaf feeders (Fig. 20.3). We com-
ment only on selected major groups.
COSSOIDEA. Cossidae, often regarded as rela-
tively primitive, number about 670 described spe-
cies and are cosmopolitan but predominantly
tropical. Few life histories are known because the
larvae are borers in root crowns and stems, but
cossids appear to have broad host preferences.
More than 20% are known to be polyphagous,
and many others recorded from only one or two
hosts probably are generalists too. Recorded
hosts of specialist species span 17 families, in-
cluding one monocot, with woody legumes ac-
counting for 25% of recorded species (Common
1990; Powell 1980).
TORTRICOIDEA. Tortricidae, which constitute
the largest non-obtectomeran, apoditrysian su-
perfamily, exhibit a great range of larval habits
(Horak and Brown 1991, Powell 1980), and an-
cestral detritivory is possible. The family con-
tains more than 6,600 described species, and is
well represented in both temperate and tropical
regions (Heppner 1991). Phylogenetic relation-
ships among the three subfamilies are uncertain,
but the Tortricinae, probably not monophyletic,
appear to contain the most primitive tortricids.
Larval feeding in the Indo-Australian tortricine
tribe Epitymbiini is almost entirely restricted to
leaf litter, especially of Myrtaceae (Common
1990). Based on the Epitymbiini and unpub-
lished observations of other unrelated but primi-
tive species, Horak and Brown (1991) postulated
that detritus or mycelium feeding by a free-living
larva was the ancestral tortricid trait. However,
extant members of the morphologically primitive
tortricine tribes Phricanthini and Schoenotenini
are phytophagous (Powell and Common 1985),
as are nearly all Tortricini (Razowski 1966). No
Tortricini are known to be detritivores, but some
African species related to Accra, considered by
Razowski (1966, 1981) to be primitive, feed on
scale insects (Lamborn 1914, Ghesquiere 1940).
Hence, this hypothesis awaits rigorous phylo-
genetic testing.
Among phytophagous tortricids (the great
majority), external feeding probably is primitive,
with independent origin of root, stem, cone and
seed borers in at least three tribes in all three
subfamilies (Horak and Brown 1991). Among
external feeders there appears to be a trend to-
ward polyphagy, correlated with a transforma-
tion series in oviposition behavior (Powell 1964,
Powell and Common 1985). In Phricanthini,
Schoenotenini, Tortricini, and Cochylini, the
eggs are laid singly and most species are special-
ized feeders. The Phricanthini are recorded only
from Dilleniaceae, and in Tortricini and Cochyl-
ini only 17% and 11% of species, respectively,
use hosts in two or more plant orders (data from
Common 1990, Horak and Brown 1991, Powell
412
1980, Powell and Common 1985). In contrast, in
tribes that are believed to be more derived, eggs
are laid in imbricate masses, and larvae are more
often polyphagous. Thus, analysis of larval foods
for about 900 species of Tortricinae (Brown and
Powell 1991; Common 1990; Powell 1980, 1985,
unpubl. data) reveals that polyphagy is common
in Sparganothidini (70% + others recorded only
once) and Archipini (36%); Atteriini are proba-
bly also generalists, as species in three genera
have been reared on synthetic diet, which nor-
mally is not accepted by host-specific tortricids.
The proportions of generalists in these three
tribes are far greater than in any other phytopha-
gous non-obtectomeran Ditrysia.
Apart from Phricanthini and Epitymbiini, all
tortricid tribes use a wide diversity of angio-
sperme and gymnosperms (Horak and Brown
1991; Powell 1980; Powell and Common 1985),
with no predilection for one plant subclass or
order shown by any tribe or any species rich
genus.
OBTECTOMERAN CLADE
Within Obtectomera, we follow Minet's (1991)
provisional recognition of the clade Macrolepi-
doptera, following Scott (1986). The non-macro-
lepidopteran obtectomerans are primarily con-
cealed feeders in living plants. However, there is
extensive detritivory in the largest superfamily,
Pyraloidea.
PYRALOIDEA. Division of Pyralidae in the
traditional sense into Pyralidae and Crambidae
(Minet 1982, 1983, following Munroe 1972) has
been widely accepted. The Crambidae essentially
all feed on living plants, with diverse specializa-
tions among the 16 subfamilies, including for ex-
ample monocots (Crambinae), aquatic monocots
(Schoenobiinae), aquatic plants more generally
(Nymphulinae), mosses (Scopariinae), and Bras-
sicaceae (Evergestiinae) (Munroe and Solis
1 14). Although there are a few scavengers and
lichen feeders, e. g. some glaphyriines, no major
lineage is primarily detritivorous. By contrast,
detritivorous habits are widespread in the Pyrali-
dae s. str., concentrated especially in the Pyrali-
nae, Galleriinae, and Phycitinae. Phycitine habits
are especially diverse, including genera that feed
on foliage, bore into the cortex of succulents,
stems of cacti, conifer cones, or angiosperm seed
pods, even in an ascomycete fungus, while a
series of genera specialize on dry seeds, grain, or
nuts; some of these are detritivorous (Heinrich
1956, Powell 1967) and several genera contain
species predaceous on scale insects (Pierce 1995).
Further resolution is needed, but the current hy-
pothesis of subfamily relationships (Solis and
Mitter 1992) allows the possibility that feeding
on fallen or dead plant material is primitive for
the family.
Jerry . Powell, Charles Mitter & Brian Farrell
MACROLEPIDOPTERAN CLADE ( S c o b l e 1992;
1-2)
Caterpillars of Macrolepidoptera, which consti-
tute about half the described lepidopteran spe-
cies, are typically exposed feeders on foliage.
However, various forms of concealed feeding,
reminiscent of microlepidopterans, occur in sev-
eral groups, probably secondarily. Mimallonidae
(Bombycoidea s. I.), for example, construct por-
table cases, and Hesperioidea are typically leaf-
tiers or borers. Among Noctuidae (Godfrey 1987),
Apameini are often stem borers and include an
example of early-instar leaf mining, Heliothinae
often bore into reproductive structures; and leaf-
tying occurs within several subfamilies. A similar
array of concealed habits occurs in larentiine
Geometridae (Scoble 1992).
Although the great majority of Macrolepidop-
tera feed on living higher plants, adoption of
non-phytophagous habits has occurred repeat-
edly. Consumption of lower plants and/or detri-
tus has arisen in several groups, most notably
Noctuoidea. Thus, the lithosiine Arctiidae feed
on lichens and algae, while detritivory/mycoph-
agy is dominant in Herminiinae and recurs spo-
radically in other noctuid subfamilies (Rawlins
1984). Lichenivores also occur in Lycaenidae and
Geometridae (Scoble 1992).
As in microlepidopterans, derived members of
several macrolepidopteran families have become
facultative or obligate carnivores, again largely
of scale insects (Pierce 1995). Notable examples
occur in Noctuidae, and one group of larentiine
geometrids captures flies (Montgomery 1982).
Prdation is especially well developed in Lycaeni-
dae, many of which feed on the ants and/or ho-
mopterans with which they are associated (Sco-
ble 1992; Pierce 1995).
In phytophagous Macrolepidoptera, as in
non-ditrysians, the predominant and probably
ancestral hosts are woody dicots, but there have
been multiple radiations onto herbaceous plants.
Within Noctuoidea, for example, arboreal feed-
ing is probably primitive, being dominant in the
oldest lineages (Notodontidae s. 1.), in Lymantri-
idae, and in most "quadrifine" and some "trifine"
subfamilies of Noctuidae s. 1. (see Kitching and
Rawlins, 1 19). Among the trifines, Holloway
(1989) postulated a trend from tree feeding in
the basal groups, especially in tropical lowland
habitats, toward herb feeding, especially in open
habitats at higher elevations and latitudes, in de-
rived groups such as noctuine cutworms. Diver-
sification of the latter might have paralleled the
proliferation of herbaceous plant taxa as regions
of temperate climate expanded during the Terti-
ary, a hypothesis that needs testing. There have
been independent major radiation of herb-feed-
ing taxa in the Arctiidae, and within other super-
families (e. g. Geometridae).
Some of the most prominent departures from
arboreal feeding occur in the Rhopalocera, the
Evolution of Larval Food Preferences in Lepidoptera
habits of which have been intensively studied
(e.g. Ehrlich and Raven 1964; Ackery 1988;
Vane-Wright and Ackery 1984). Hesperioidea
and essentially all families of Papilionoidea con-
tain major elements associated with plant fami-
lies which are characteristically herbs (especially
in temperate regions), lianas, or shrubs. Selected
examples include: in the Pieridae, Pierinae on
Capparidaceae/Brassicaceae; in Papilionidae,
Troidini and Parnassiinae on Aristolochiaceae;
and in Nymphalidae, Satyrinae, Amathusiini and
Brassolini on monocots, Acraeini on Passiflora-
ceae and Urticaceae, Danainae on Apocynaceae/
Asclepiadaceae and Solanaceae. The contrast be-
tween such butterflies and more typical, tree-
feeding Macrolepidoptera has figured in the de-
velopment of theories relating plant growth and
defense to insect host choice and associated life
history traits including diurnality and aposema-
tism. We return to this topic below.
Discussion
In light of the foregoing review, we can address
the questions posed in the introduction.
Origins of higher-plant feeding
Several groups scattered through the primitive
lepidopteran lineages show habits reminiscent of
the panorpoid groundplan, living on the ground
and feeding on detritus, non-vascular plants or
fungi. Although further information is needed,
such habits appear to be primitive within Zeug-
loptera, Exoporia, Tineoidea, possibly the puta-
tive oecophoric clade of Gelechioidea, and possi-
bly, though doubtfully, in Tortricidae. This raises
a question introduced by Grehan (1989): Might
such generalized habits have persisted through
the early history of Lepidoptera, giving rise
multiple times to specialized higher-plant
feeding?
"Mapping" of the alternative conditions, "re-
striction to vascular plants" vs. "other habits",
on the cladograms of Figures 20.1-3 argues
against this idea. That is, if we allow that larval
phytophagy on higher plants might be a homolo-
gous trait, the most parsimonious hypothesis of
ancestral conditions is evolution of all Lepidop-
tera except Micropterigoidea from a higher
plant-feeding common ancestor. Similar reason-
ing suggests that all Lepidoptera except Micro-
pterigoidea and Agathiphagoidea derive from an
angiosperm-feeding common ancestor. These
hypotheses require independent reversion to
non-phytophagy in basal Exoporia and Ti-
neoidea, as well as within many more advanced
ditrysian groups. Conversely, a scenario of an-
cestral detritivory/fungivory in basal lineages up
413
through the common ancestor of Ditrysia re-
quires independent origins of higher plant phy-
tophagy in an additional seven or more lineages
(Fig. 1), i. e., Agathiphagoidea, Heterobathmi-
oidea, Eriocranioidea, Acanthopteroctetidae,
Neopseustoidea, Lophocoronidae, Tischerioidea
and/or Palaephatoidea, and basal non-tineoid
Ditrysia. Most of these groups, including Neo-
pseustidae and Lophocoronidae, whose habits
are inferred from the piercing ovipositor (1-5),
share a particular form of phytophagy, arboreal
leaf mining, although ovipositor form/function
and larval morphology vary markedly among
them.
Several problems are raised, however, by the
hypothesis that endophagous phytophagy origi-
nated only once, or twice (in Agathiphagoidea
and in Heterobathmioidea + Glossata), very
early in lepidopteran evolution (e. g. late Jurassic
or early Cretaceous). Why are there no extant
non-ditrysian lineages associated with pterido-
phytes, gymnosperms (other than agathipha-
gids), or primitive angiosperms (but see Laban-
deira et al. 1994)? An Upper Jurassic lepidop-
teran fossil (Kozlov 1989), inferred to have a well
developed maxillary siphon and large, three-seg-
mented labial palpi is provisionally assigned to
the Ditrysia (Labandeira et al. 1994). If the age
and morphology are correctly interpreted, the
major clades including Ditrysia must have devel-
oped prior to radiation of the angiosperms. Was
the larva of this Jurassic moth an exophagous
generalist with habits analogous to extant Micro-
pterigidae and Mnesarchaeidae, a detritivore
similar to modern tineids, or a specialist herbi-
vore on a primitive plant, an association that dis-
appeared following the radiation of the angio-
sperms? Labandeira et al. (1994) suggest that an-
giosperms in the Cretaceous offered a new food
resource to existing herbivores, but if so, what
did such herbivores eat and why did none of the
pre-angiosperm glossatan host associations sur-
vive?
The alternative hypothesis, that phytophagy
and endophagy arose in several lepidopteran lin-
eages independently during the early diversifica-
tion of the angiosperms, avoids these difficulties.
It also avoids the need to postulate the loss, in
both Exoporia and Tineoidea, of morphological
and behavioral specializations associated with
angiosperm leaf-mining. Although this alterna-
tive appears less parsimonious on the cladogram,
it should not be ruled out.
Whatever one's view of the earliest history of
phytophagy, secondary shifts to detritivory (and
thence often in stages to mycophagy, Rawlins
1984), are a recurrent feature of higher lepidop-
teran evolution, characterizing subgroups, for
example, of Pyralidae, Noctuoidea, Tortricoidea,
and Gelechioidea. Hypotheses of the selective
pressures favoring such shifts, including resource
414
availability (Rawlins 1984; Common 1990), war-
rant rigorous test.
How did typical, host-specific, higher plant
feeding initially arise in Lepidoptera? One model
is suggested by possible stepwise evolution
within several detritivorous/fungivorous lin-
eages. For example, in Hepialidae, early instars
appear to be detritivorous or mycophagous,
while later instars feed externally on the roots
of, or bore into, living plants and typically are
polyphagous. However, a few species, possibly
more derived, seem to be specific in their host-
plant selection (Grehan 1989). A sequence from
detritivory to generalized and thence to specific
phytophagy can similarly be envisioned in Ti-
neoidea (some Psychidae are host specific), sug-
gesting that lepidopteran phytophagy originally
arose in this way several times. However, the cla-
distic evidence for primitive moths suggests an
abrupt transition to arboreal, host-specific endo-
phagy; there are no phytophagous lineages that
are polyphagous among Agathiphagoidea, Het-
erobathmioidea, or basal glossatans. Indeed,
among strictly phytophagous clades, polyphagy
is relatively rare, seems clearly derived in many
instances, and is most prevalent among external
feeders (Gaston et al. 1992). It may also be of
relevance that the carnivorous lepidopterans are
often, if not invariably, in groups that also con-
tain detrivorous species (Pierce 1995).
The current hypothesis of phylogeny ( 1 - 2 )
suggests a trend among phytophagous Lepidop-
tera from internal to external feeding (Figs. 20.1,
2, 3; Scoble 1992) that is stronger than any corre-
lation with host plant phylogeny. Although
much further analysis is needed, the cladogram
(Fig. 20.1, 2) suggests there have been a number
of transformations through intermediate condi-
tions leading to external feeding. (Evolution in
the reverse direction may have occurred in some
groups, e. g. Tortricidae; Horak and Brown
1991.) Construction of a portable case from
which larvae feed either externally (Incurvari-
oidea) or by mining (Coleophoridae) has proba-
bly arisen several times (e. g. Incurvarioidea, Ti-
neoidea, Gelechioidea, Pyraloidea). Among in-
curvarioids there is a trend toward earlier con-
struction of the case, from just before pupation
(Heliozelidae), to an exophagous function after
one or more endophagous instars (Adelidae, In-
curvariidae). In several leaf mining groups, par-
ticularly among lower Ditrysia, there are transi-
tions from leaf mining to various forms of exter-
nal feeding. Such adaptations occur within indi-
vidual ontogenies, with exophagous instars
either concealed, as in Argyresthiidae and Grac-
illariidae, or exposed, as in Bucculatricidae and
Epermeniidae. Feeding exposed on foliage has
arisen a number of times, but it is most prevalent
in Macrolepidoptera.
Transition to exophagy might be viewed as an
adaptive "escape" from the constraints of endo-
Jerry A. Powell, Charles Mitter & Brian Farrell
phagy, and could have significant evolutionary
consequences. Disadvantages to endophagy
might include limits on body size and voltinism,
and in leafminers, vulnerability to excision. En-
dophagy might also limit the ability to colonize
alternative hosts (Gaston et al. 1992) and habi-
tats, and thereby restrict the potential for specia-
tion. Exophagy may be resisted by strong evolu-
tionary barriers among Holometabola (South-
wood 1973), and is found in relatively few holo-
metabolous phytophage lineages (Mitter et al.
1988), but may promote the evolutionary success
of groups adopting it. This hypothesis needs
testing.
Parallel diversification with hostplants?
If the predilection for a particular plant taxon
were faithfully passed from ancestral to descen-
dant species, lepidopteran clades should remain
associated with particular host clades over long
time spans, and might undergo divergence in
parallel with those hosts. Such a history should
be detectable by correspondence between lepi-
dopteran and hostplant phylogenies, unless ex-
tinction or within-species host switch obscure
early host associations. Different evolutionary
models predict different degrees of cladogram
match (Mitter and Farrell 1991), but the exis-
tence of any such pattern for a lepidopteran
group would suggest that its present-day host use
reflects long-term history, and that there has
been significant opportunity for reciprocal evo-
lutionary influence between these insects and
their hosts.
From comparison of host use among the basal
lineages, earlier reviews found little support for
parallel diversification between Lepidoptera as a
whole and their hostplants (Powell 1980; Mitter
and Brooks 1983). The question warrants re-ex-
amination, however, in light of new information
on phylogeny and host use, and development of
quantitative measures of cladogram correspon-
dence. To re-assess possible parallel cladogenesis
for the non-ditrysian lineages, we prune the cla-
dogram of Figure 20.1, leaving only phytopha-
gous lineages whose host associations can rea-
sonably be characterized to the plant family or
ordinal level. While our analysis is thus limited
by lack of phyletic resolution for groups with di-
verse hosts, its bias toward low-diversity and
biogeographically relictual taxa should maximize
the chance of detecting relictual host associa-
tions. This pruned tree, together with a recent
phylogeny estimate for the hostplants, is shown
in Figure 20.4. For evaluation of cladogram con-
cordance, we used component analysis (Nelson
and Platnick 1981) as implemented by Page
(1990).
If bryophytes are treated as the ancestral and
primary hosts of Micropterigoidea, the overall
concordance of lepidopteran and hostplant phy-
Evolution of Larval Food Preferences in Lepidoptera
, Micropterigidae ??Liverworts
, Agathiphagidae Coniferales
, Heterobathmiidae Nothofagus
Eriocraniidae Fagaceae,
Acanthop- Betulaceae
teroctetidae Rhamnaceae
, Crinopterigidae Cistaceae
Anacardiaceae
Cecidosidae
Nepticulidae:
Pectinivalvinae Myrtaceae
Opostegidae:
Notiopostega Nothofagus
Fig. 20.4. Phylogeny of primitive moths (non-ditry-
sians) (left), from Figure 1, excluding taxa not associ-
ated primarily with a single plant order, contrasted to
recent estimate of relationships among their host fami-
lies (right), based on rbcL gene sequences (Chase et al.
1993). Positions of Cistaceae, Myrtaceae, not studied
by Chase et al., inferred from those of confamilial or
con-ordinal species according to Cronquist (1981).
Cladogram concordance assessed using component
analysis under Assumption 1, as implemented in Com-
ponent 1.5 (Page 1990). For all taxa, items of error
(IOE) = 46, p. = 0.01 under Markovian null distribu-
tion of host phylogenies (1000 replications). With
Micropterigidae deleted, IOE = 46, = 0.08; with
Agathiphagidae also deleted, IOE = 46, > 0.50. Al-
ternative plant phylogeny of Hufford (1992) yields
very similar conclusions.
logenies is significantly greater than random
(Fig. 20.4). However, it is questionable whether
bryophyte feeding is primitive within micropteri-
gids. Even if it were, the habit is more likely to
reflect a retained, ancestral amphiesmenopteran
habitat preference (N. P. Kristensen, unpub-
lished) than plant phylogeny. That is, it seems
far-fetched to postulate that Micropterigidae fed
ancestrally on bryophytes because vascular
plants, host to nearly all subsequent lepidopteran
lineages and dominant since the Carboniferous
or before, were not yet available.
If Micropterigoidea are excluded, cladogram
concordance still approaches statistical signifi-
cance, due to the basal split between the conifer-
feeding Aglossata and the remaining lineages,
which primarily eat angiosperme. Whether or
not agathiphagids represent an independent ori-
gin of phytophagy, this is plausibly interpretable
as paralleling host phylogeny, perhaps the only
such instance in Lepidoptera (Scoble 1992); that
is, Aglossata may well retain an association
established before angiosperms or their more im-
mediate ancestors became available as hosts.
For the angiosperm-feeding lineages alone, in
contrast, there is no hint of concordance with
any recent host phylogeny estimate. It would not
be surprising for comparison across the major
non-ditrysian lineages to show no correspon-
dence to angiosperm phylogeny, if the suggestion
is accurate that divergence of these lineages pre-
ceded that of angiosperms. Match to host phy-
logeny might in that case be more apparent upon
detailed studies within the larger lineages. How-
415
ever, strikingly few species within any non-ditry-
sian group (Powell 1980) are associated with
hosts identified as early-diverging by recent cla-
distic analyses, such as the orders of Magnolii-
dae, "lower" Hamamelidae, or (primitive) mono-
cots (Chase et al. 1993; Hufford 1992). Even No-
thofagus and other Fagales, hosts for the two
oldest angiosperm-feeding lineages, appear rela-
tively derived in cladistic position, despite a long
fossil record. This association, ostensibly homol-
ogous between Heterobathmioidea and Eri-
ocranioidea, must be independently derived if
these lineages are both older than angiosperms.
This suggests that extinction and host transfer
may have obliterated most traces of early co-di-
versification with angiosperms if it occurred. (An
alternative explanation would be that lepidopter-
ans did not start feeding on angiosperms until
these plants were extensively diversified). The
possibility that primitive angiosperms were in
fact important hosts for early non-ditrysians is
supported by discovery in fossil magnoliid and
lower hamamelid (platanoid) leaves of mid Cre-
taceous age of mines attributed to two modern
nepticulid genera (Labandeira et al. 1994).
A conserved habit that merits further analysis
is the prevalent restriction of non-ditrysian lin-
eages to woody host plants. Association with
herbs, particularly those taxa that have prolifer-
ated in the Tertiary, occurs among non-ditrysians
only in some presumably derived clades, such as
Prodoxidae (Davis et al. 1992, Wagner & Powell
1988), a few nepticulid lineages (Van Nieukerken
1986), and Tischeriidae (Davis 1987). In con-
trast, the Ditrysia, including non-apoditrysian
groups, contain many lineages that appear to
have radiated on modern herbaceous host taxa.
Explicit phylogeny comparisons for lower
level lepidopteran taxa, e. g. genera or families,
and their hosts, have been few and mostly pre-
liminary. However, cladogram concordance be-
tween Lepidoptera and their host plants seems
rare at this level also (reviews in Miller 1987,
Mitter and Farrell 1991). The best candidate for
parallel cladogenesis is the subfamily Heliconii-
nae (Nymphalidae), which at the species level
provides evidence for coevolution in the sense of
reciprocal counter-adaptation with its passiflora-
ceous hosts (Gilbert 1984). However, even this
example is disputed (Benson et al. 1975; Mitter
and Brooks 1983; Mickevich and Weiler 1990;
. . . Venables, unpublished). Detailed study
of this and other intimate lepidopteran-host
plant associations is needed to determine
whether parallel phylogenesis, which is uncom-
mon but known in other phytophagous groups
(e. g. Chrysomeloidea; Farrell et al. 1992), can be
ruled out entirely in Lepidoptera.
The persistence of ancestral host associations
in Lepidoptera appears intermediate in compari-
son with that in two other recently-reviewed phy-
tophagous insect clades of comparable age. The
416
Fulgoroidea (19 families), which date from the
lower Cretaceous or earlier, show no convincing
instances of parallel phylogenesis with their host-
plants at any taxonomic level, despite prevailing
host specificity (Wilson et al. 1993). The only
trend reminiscent of Lepidoptera is a progression
from concealed (mostly underground) to ex-
posed sites of feeding.
In contrast, trends inferrable from morpholog-
ical classifications of the chrysomeloid Cole-
ptera (Chrysomelidae + Cerambycidae), sup-
ported by recent molecular studies (B. D. Farrell,
in prep.), suggest several parallels to Lepidoptera
in the evolution of host associations. In
chrysomeloids, which date from the upper Juras-
sic (Crowson 1981), saprophagy (dead wood
feeding) appears ancestral, and subsequent phy-
tophages have given rise to secondary detritivory
(Crowson 1981), though apparently much less
often than in Lepidoptera. Four morphologically
isolated, primitive, and species-poor subfamilies
of chrysomeloids are restricted to cycads or coni-
fers. An example is the Palophaginae (Chryso-
melidae), which consists of two monotypic gen-
era, one each in Chile and Australia, that feed
on Araucaria cones (Kuschel and May 1990),
analogous to Agathiphagidae. In the chrysomel-
oid sister group Curculionoidea, in which phy-
tophagy has arisen independently one or more
times (Anderson 1993), there are likewise several
small, Southern Hemisphere groups restricted to
gymnosperms, in the primitive families Nemo-
nychidae (Kuschel 1983), Brentidae and Belidae.
Chrysomeloid subfamilies of intermediate
morphological advancement feed characteristi-
cally either on monocots (e. g. Donaciinae;
Askevold 1991) or, like basal glossatane, on
woody Hamamelidae, Rosidae and Dilleniidae.
The most derived subfamilies, having the most
numerous generic radiations, feed on recent,
mostly herbaceous plant families, especially the
subclass Asteridae (Jolivet 1988). A further
chrysomeloid parallel to Lepidoptera is predomi-
nant and ancestral concealed feeding, which has
given rise multiple times to external larval herbi-
vory.
While similar trends are apparent, ancestral
host affiliations appear to be much better pre-
served in Chrysomeloidea than in Lepidoptera,
as reflected for example in the extent of the prim-
itive, gymnosperm-feeding element. Further-
more, two genera of advanced, herb-feeding
chrysomeloids provide the strongest examples
known of herbivore/plant cladogram correspon-
dence (Farrell and Mitter 1990; Farrell et al.
1992). Greater evolutionary lability of larval
food choice in Lepidoptera than in Chrysomel-
oidea is also indicated by the apparently much
more frequent origin of non-phytophagous hab-
its in lepidopterans. Apart from the basal wood-
feeding clades, departures from live-plant feed-
ing in chrysomeloids occur only in the case-bear-
Jerry A. Powell, Charles Mitter & Brian Farrell
ing clade of Chrysomelidae, which consists of
Lamprosomatinae, Chlamisinae, Cryptocephali-
nae and Clytrinae (Suzuki 1988). Chlamisinae
and Cryptocephalinae include some species with
detritivorous larvae, while Clytrinae are invaria-
bly myrmecophilic, and larvae may feed on ant
brood as well as detritus (Jolivet 1988), parallel-
ing some Lycaenidae (Pierce 1995).
This difference in evolutionary lability needs
explanation. Perhaps shifts of food preference in
living plant-feeding chrysomeloids are discour-
aged in part by the similar specificity of feeding
requirements between larvae and ovipositing
adults, both of which, particularly in Chrysomel-
idae, typically chew somatic tissue of the same
host plant. In contrast, adult Lepidoptera take
liquid food and usually are nutritionally inde-
pendent of the larval host. This presumably pro-
vides greater opportunity for host shifts through
oviposition on novel host plants (Chew and Rob-
bins 1984).
A final point of comparison between the
Chrysomeloidea and the Lepidoptera concerns
the evolution of carnivory. Feeding on ant brood
by species of Clytrinae is the only apparent de-
parture from phytophagous habits in Chryso-
meloidea, and it parallels lycaenid habits (Pierce
1995). Thus the Lepidoptera seem much more
labile overall in the evolution of non-phytopha-
gous habits, and prey items are typically insects
frequently encountered in the larval environ-
ment.
Evolutionary Patterns of Host Shift
Although parallel cladogenesis with host lineages
is evidently rare in Lepidoptera, the evolution of
lepidopteran host plant affiliations is not ran-
dom. It has long been noted, for example, that
in some groups, related species tend to feed on
related plants (e. g. Brues 1920). The prevalence
of this tendency across the order as a whole has
not been rigorously assessed. The preferred ap-
proach (Van Nieukerken 1986) would be to re-
construct the history of host shifts on clado-
grams inferred from other characters. Seven lepi-
dopteran groups, representing one macrolepi-
dopteran and three microlepidopteran superfam-
ilies, were included in a recent compilation of
species-level cladograms for phytophagous in-
sects (Mitter and Farrell 1991). The mean ratio
of host family shifts to speciation events inferred
from the lepidopteran cladograms was about
0.22, close to the value for all insect groups col-
lectively. This small sample can hardly character-
ize the order, but does illustrate the great varia-
tion among groups in the propensity for taxo-
nomically radical host shifts. For example, the
seven species in Yponomeuta "group A" (Ypono-
meutidae) have shifted among four host families,
with essentially all species using different host
genera (Menken 1982). In contrast, the ten
Evolution of Larval Food Preferences in Lepidoptera
congeneric species of Heterobathmioidea share a
single host genus, as do most of the 70 species
of the Heliconius group (Nymphalidae). The at-
tempt to classify and explain such heterogeneity
in host-use evolution has only begun. In the re-
mainder of this chapter we sketch several current
research directions, but a thorough treatment is
beyond our scope. Relevant recent reviews in-
clude Thompson and Pellmyr (1991), Stamp and
Casey (1993) and Renwick and Chew (1994).
Advances in lepidopteran and angiosperm
phylogenetics are increasing the detectability of
host use patterns reflecting plant relatedness
above the family level. Ehrlich and Raven (1964)
pointed out examples including papilionoid
groups broadly associated with monocots, with
the subclass Magnoliidae (Magnoliales, Laurales
etc.) and with Violales (Violaceae, Passifloraceae
etc.); Miller (1992) has described parallels in No-
todontidae. The disparate-seeming hosts of a
number of apoditrysian clades (e. g., Ackery
1988; Mitter and Farrell 1991) are united by
membership in an expanded concept of the sub-
class Asteridae (Asteridae of Cronquist 1981,
plus Apiales, Cornales, Ericales and several oth-
ers), monophyly of which is supported by recent
molecular studies (Olmstead et al. 1993).
Chemical similarities between host plants
probably underlie such conserved host-taxon as-
sociations, though the mechanisms are still
poorly understood. Lepidopteran host recogni-
tion is complex (Renwick and Chew 1994), and
host shift patterns no doubt reflect multivariate
similarity. In some groups, however, diet evolu-
tion appears strongly influenced by particular
secondary-chemical or other putatively defensive
plant attributes, which may even be "tracked"
across unrelated plant lineages (Ehrlich and Ra-
ven 1964). Recently-documented syndromes in-
clude associations with plants possessing com-
plex coumarins (Berenbaum 1983), and with
those containing iridoid glycosides (Bowers
1988). Iridoids are characteristic of, though not
universal in, Asteridae, and may in part account
for affiliations with that subclass. For example,
the host families of epiplemine Uraniidae as
compiled by Lees and Smith (1991) are Rubia-
ceae (Rubiales), Caprifoliaceae (Dipsacales),
Oleaceae and Bignoniaceae (both Scophulari-
ales), Verbenaceae (Lamales), and Daphniphyl-
laceae. All but the last are iridoid-bearing Asteri-
dae; Daphniphyllaceae (subclass Hamamelidae)
are one of the few non-asterid groups to contain
iridoids (Olmstead et al. 1993).
Numerous lepidopteran and other insect
clades have shifted among hosts, often unrelated
and chemically dissimilar, which have secretory
canals containing latex or resin (Peigler 1986;
Farrell and Mitter 1993). These repeatedly
evolved structures discourage herbivory by dis-
charging viscous, often toxic fluids when severed.
Caterpillars in a number of families overcome
417
this barrier by means such as severing the canal
and lowering its fluid pressure before feeding dis-
tal to the cut (Dussourd 1993). Canal-plant pat-
terns are best documented in Macrolepidoptera.
For example, the main hosts of danaine Nymph-
alidae are Asclepiadaceae/Apocynaceae; the
most reliable other records (Ackery and Vane-
Wright 1984) are from Moraceae, Convolvula-
ceae, Caricaceae, Euphorbiaceae and Rubiaceae.
All but the last bear latex. Euteliine Noctuidae
most commonly feed on Anacardiaceae; promi-
nent among their other hosts are Burseraceae,
Dipterocarpaceae, Moraceae and altingioid Ha-
mamelidaceae, all likewise canal-bearing (Hollo-
way 1985; Forbes 1954; Peigler 1986). The prob-
able euteliine sister group, Stictopterinae, is asso-
ciated primarily with Dipterocarpaceae and Gut-
tiferae ( = Clusiaceae) (1 19), which also bear
secretory canals. Further examples occur in epi-
paschiine Pyralidae (Solis 1993), aganaine Noc-
tuidae (Holloway 1988), Uraniidae (Lees and
Smith 1991), Saturniidae (Peigler 1986) and
doubtless many others.
The canal-plant syndrome, like that for com-
plex coumarins (Berenbaum 1983), illustrates a
way in which Lepidoptera may participate in
"diffuse" coevolution between plants and herbi-
vores. Canals seem to have evolved as broad-
spectrum defenses against herbivores and patho-
gens, presumably including lepidopterans, and
canal-bearing plant clades show consistently ele-
vated diversity as compared to their sister groups
(Farrell et al. 1991), as predicted by Ehrlich and
Raven's (1964) coevolutionary model. The mod-
el's converse prediction, that radiation should be
correspondingly accelerated in lepidopteran
clades able to colonize these or other sets of
plants bearing distinctive defenses, has yet to be
tested. Like that of other phytophagous insects,
lepidopteran diversity is elevated in comparison
to their sister group, Trichoptera (Mitter et al.
1988); but the question of whether this results
from radiation onto different plant resources,
however defined, is unexplored.
Several authors (e. g., Janzen 1984; Holloway
1987; Feeny 1991 b; Miller 1992) have contrasted
lepidopterans feeding on plants bearing distinc-
tive, "qualitative" or "mobile" secondary com-
pounds (Feeny 1991 a), typically toxic or repel-
lent to vertebrates as well as to non-adapted in-
sects, to species whose hosts typically lack such
features. Lepidoptera associated with "toxic"
plants often show marked specificity and conser-
vatism of host choice, an example being troidine
Papilionidae, restricted to Aristolochiaceae
(Feeny 1991b). Such conservatism has been at-
tributed in part to herbivore dependence on dis-
tinctive chemistry for host finding and accep-
tance, and possibly for defense as well: toxic
plant lepidopterans are often aposematic day-fli-
ers, and some sequester host toxins in the larval
stage for use in their own defense.
418
In contrast, plant in some taxonomic or eco-
logical assemblages may be less chemically dis-
tinctive, lowering the barriers to host transfer.
Thus, temperate deciduous forests are dominated
by tanniferous angiosperms in which resistance
to herbivores is manifested mainly by toughness
and low nutritional content of mature foliage.
A m o n g Lepidoptera that eat these plants the in-
cidence o f polyphagy is elevated, at least in
Macrolepidoptera (Futuyma 1976). Even conge-
neric species in host-specific groups often have
adapted to unrelated hosts (Powell 1980); exam-
ples include Yponomeuta, cited above, all the
larger genera of non-obtectomeran ditrysians
and o f Geometridae, and many genera o f arbo-
real N o c t u o i d e a (Miller 1992, Mitter and Farrell
1991). In such groups, feeding site and habitat
preference, particularly if these require special
adaptations, may be conserved more often than
associations with plant taxa.
Correlations of larval diet evolution with
other life history traits and with plant chemistry
and ecology have only begun to be documented,
and will prove much more complex than our
coarse d i c h o t o m y w o u l d suggest (see e. g. Janzen
1984). Phylogenetic study o f such patterns is a
promising avenue toward better understanding
o f the evolution of lepidopteran diet.
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 21. Biogeography of the Lepidoptera
Jeremy D. Holloway & Ebbe S. Nielsen
Introduction
Over half the world total of species consists of
insects (Barnes 1989). Lepidoptera are one of the
four major Orders of insects, though they are not
the most species rich of the four. Compared to
Coleoptera, Diptera and Hymenoptera, they of-
fer a better source of data in terms of their geo-
graphical coverage in world collections, develop-
ment of their taxonomy and knowledge of their
biology. The high representation of phytopha-
gous taxa means they do not exhibit the trophic
diversity of the other three Orders, but their spe-
cialization does offer opportunities for studies of
interdependence in insect and plant geography.
This combination of high diversity and a de-
tailed information base has meant that Lepidop-
tera studies are prominent in many areas of bio-
geographic research. Indeed, the seminal work
on island biogeography by MacArthur & Wilson
(1967) is now known to have been preceded in
the development of many of its concepts by a
largely unpublished doctoral thesis on Caribbean
Lepidoptera by E. G. Munroe (1948, 1953, as re-
lated by Brown & Lomolino (1989), and Wilkin-
son (1993)). We aim to focus this chapter as
much on this unique aspect of Lepidoptera bio-
geography as on a traditional, taxonomically
based catalogue of geographic patterns (e. g.
Heppner, 1991).
Concepts
Any account of the biogeography of a major tax-
onomic group will inevitably reflect the stand-
point of the authors in relation to current de-
bates on biogeographic method and philosophy
(e.g. Platnick 1976; Nelson & Platnick 1981;
Nelson & Rosen 1981; Myers & Giller (eds.),
1988; DSIR 1989; Cranston & Naumann 1991;
Morrone & Crisci 1995; Nelson & Ladiges 1996).
Wilson (1991) attempted to identify and catego-
rize three major biogeographic trends of thought
or models in relation to a number of criteria. The
trends included the more traditional Darwinian
centre of origin and dispersal philosophies (mi-
grationist), the vicariance model of the cladistic
biogeographers and the panbiogeographic model
progressing from the publications of Leon Croi-
zat. The criteria involved such topics as mapping
and analysis methodologies, assumptions about
dispersal and evolutionary process, attitudes to
fossil evidence and ecology, the interplay be-
tween taxonomic and biogeographic evidence,
and explanatory and predictive powers of the
models.
We would agree with the panbiogeographers
in emphasizing that modern patterns of distribu-
tion arise from the interaction of the geological
processes that are shaping our planet with the
processes of biological evolution, but would hold
that the mobilism of organisms is an important
part of the second process. We would also em-
phasize another physical factor, not disconnected
from the geophysical evolution of the planet, but
partly controlled by independent external
forces: climate.
Whilst not embracing the extreme dispersal-
from-centres-of-origin model attributed to the
migrationist school by Wilson and Cranston &
Naumann, we adopt the working hypothesis that
at any time in the past there has been a similar
mix of widespread v. localized taxa with similar
differences in diversity from place to place to
that seen today. The primeval cosmopolitanism
implicit in some biogeographic philosophies
seems at odds with the fossil record, both with
regard to evidence of past climates and evidence
of distribution patterns of organisms in ancient
areas, even though the changes in degree of geo-
graphical fragmentation from era to era must
be important.
In the absence of a well endowed fossil record
for the Lepidoptera (see 12), it is an open ques-
tion as to how far back in the Mesozoic tectonic
events have been influencing distribution pat-
terns in the order. Grehan (1991) considered that
Lepidoptera evolution has been closely associ-
ated with that of the angiosperms from well be-
fore the Cretaceous, and influenced by tectonic
events from then to today. The alleged noctuid
moth egg from the late (75 My) Cretaceous
(Gall & Tiffney 1983, but see 1 - 2 ) , may indicate
that most currently recognized ditrysian families
were extant at that time.
Such a long and complex history requires a
methodology for analysis of pattern that will
permit recognition of the respective influences of
mobilism in geology, mobilism of organisms and
the effects of climate on the development of that
pattern. Such interpretation will inevitably be
constrained by current hypotheses of the evolu-
tionary process. This analysis of patterns should
be independent from consideration of process
(Humphries & Parenti, 1986). A satisfactory
methodology has yet to be developed (Page
1989 b), but the Lepidoptera are an important
source of data that will also act as a stimulus for
further ideas.
424
Pattern Analysis: Q-mode v. R-mode
Biogeographic methodology is sometimes por-
trayed as having passed from a phenetic stage
to a cladistic one in much the same way as has
taxonomic methodology (e. g. Nelson & Platnick
1981). Cladistic information is of the utmost im-
portance in biogeographic analysis, but the anal-
ogy between systematic and biogeographic meth-
odology can be overstretched, obscuring the
more significant distinction between Q-mode (re-
lationships of areas of endemism in terms of
their biotas) and R-mode (relationships of taxa
in terms of their distributions amongst areas)
analyses. This distinction was made by Hollo-
way & Jardine (1968) in analyses of the geogra-
phy of butterflies and other taxa in the Indo-
Australian tropics. It is usually overlooked by
advocates of a cladistic approach to Q-mode
analysis that the phenetic approach can be linked
with non-metric multidimensional scaling meth-
ods that can be informative in terms of earth his-
tory (Holloway & Jardine 1968). Dennis et al.
(1991) have applied both cluster analysis and po-
lythetic division (TWINSPAN) in the Q-mode,
as well as non-metric multidimensional scaling,
to show regional structure in the distribution of
European butterflies. Phenetic analyses are still
performed where data are in the form of local
faunal records and do not permit clear recogni-
tion of areas of endemism, e. g. in Shapiro's
(1991) study of southern S. American pierine
butterflies and Gentili's (1988) study of Cossidae
in the same region.
More recently (Simberloff & Connor 1980;
Page 1987; Connor 1988) attention has again
been drawn to the distinction between the two.
Cladistic biogeography as practiced in recent
years is essentially Q-mode, seeking any unique,
underlying pattern of relationship between areas
of endemism, repeated in cladistic classifications
of those areas from biosystematic data, that re-
flects events in earth history. Examples of suc-
cessful application of this approach are very lim-
ited and the computational problems are great
(Humphries 1990; Page 1988, 1989 a, 1990; Hol-
loway 1991); methods of area classification using
taxa as 'characters' and applying parsimony
analysis appear to offer little more than do phe-
netic methods, and often yield very similar re-
sults (Holloway 1991; see critique of these meth-
ods in Page 1990). Figure 21.1 shows results for
the Indo-Australian tropics obtained by applying
phenetic and cladistic methods to data from the
Lepidoptera.
The challenge is to discover a method of R-
mode analysis that will incorporate cladistic in-
formation and allow recognition of statistically
significant groupings of taxa. Sets of clades with
high numbers of very widespread overlapping
taxa (e. g. possibly grouping together Helicov-
erpa, Spodoptera, Chrysodeixis, Anomis, Junonia,
Jeremy D. Holloway & Ebbe S. Nielsen
Danaus) will be identified at one extreme, and
sets of clades with many localized, allopatrically
distributed taxa at the other (e. g. Bracca com-
pared with Anomis subgenus Rusicada in Fig.
21.2). The former may hold information about
process in the mobilism of organisms (see also de
Jong 1990), and the latter, through the applica-
tion of Q-mode methods referred to above, may
provide information on earth history. Thus an
R-mode methodology does not foreclose on op-
tions for pattern recognition to the extent that a
Q-mode one does.
Advocacy of an R-mode approach to biogeo-
graphic analysis is an essential element of Croi-
zat's (1964) panbiogeography, in the form of der-
ivation of generalized tracks. Croizat performed
this subjectively, but recent attempts to translate
the approach into a statistically rigorous method
have had only limited success (Page 1987,
1989 b); Henderson 1989; Weston 1989). The fau-
nal element phenetic analysis of Holloway & Jar-
dine (1968) is similar but does not involve ex-
pression of the distribution of the taxon as a lin-
ear, usually branching track: distributions are
compared by measuring the extent of overlap
over primary areas of endemism.
Faunal element analyses have been applied ex-
tensively to data on Palaeotropical Lepidoptera
(Holloway 1973, 1974, 1979), and examples of
faunal elements for the Indo-Australian tropics
are given in Fig. 21.3. A more recent example,
applied to the European butterfly fauna, is by
Dennis et al. (1991). The faunal elements (units)
they recognized using cluster analysis and princi-
pal component analysis were characterized on
climatic (temperature) parameters prior to exam-
ining hypotheses for their historical development
(see also Dennis (1993)).
A related approach (Holloway 1969) includes
data on variation on species richness (or mass-
ing) within the range of a higher taxon: the deri-
vation of faunal centres (not to be confused with
centres of origin). Genera are classified as similar
on both overlap of geographical range and coin-
cidence of species richness in parts of that range.
Both these approaches were applied in an in-
vestigation of the butterfly fauna of India (Hol-
loway 1969, 1974). The Indian peninsula itself
contains no major centres of generic richness,
but the fauna is a composite one, with contribu-
tions of species from a variety of generic centres
(Fig. 21.4). The major contribution is from gen-
era with centres of richness coincident over much
of S. E. Asia, many with the greatest richness in
the N. E. Himalaya. Two centres include genera
of a more temperate range and character, one in
western China associated with the Oriental
Centre just mentioned and the other centred in
the N. W. Himalaya and Pamirs that is more
strongly Palaearctic in range and diversity. Mi-
nor contributions are also evident from genera
Biogeography of the Lepidoptera 425

MAINLAND ASIA
MALAYA
SUMATRA
BORNEO

MALAYA + SUMATRA
JAVA
BORNEO
LESSER SUNDAS
PHILIPPINES
PHILIPPINES
JAVA + BALI
SULAWESI
SULAWESI
AUSTRALIA
SULAIS. + S. MOLUCCAS
N. MOLUCCAS
NEW GUINEA+ N. MOLUCCAS
S. MOLUCCAS
NEW GUINEA
BISMARCKS
SOLOMONS

c c
SRI LANKA+ S. INDIA SRI LANKA+ S. INDIA
LESSER SUNDAS LESSER SUNDAS
MAINLAND ASIA MAINLAND ASIA
MALAYA MALAYA
BORNEO SUMATRA
SUMATRA BORNEO
JAVA JAVA

rC c
N. PHILIPPINES N. PHILIPPINES
S. PHILIPPINES S. PHILIPPINES
SULAWESI SULAWESI
AUSTRALIA S. MOLUCCAS
N. MOLUCCAS NEW GUINEA
S. MOLUCCAS N. MOLUCCAS

HZ NEW GUINEA
BISMARCKS
HZ AUSTRALIA
BISMARCKS

HZ SOLOMONS SOLOMONS

Fig. 21.1. Generalized statements of area relationships (Q-mode analyses) in the Indo-Australian tropics using
Lepidoptera data. A, phenetic - the butterfly analysis of Holloway & Jardine (1968). D, cladistic. B, Compo-
nent Analysis applied to cladograms for the genera Idea and Narosa (Humphries 1990). C and D, Brooks Parsi-
mony Analysis applied to a sample of nine cladograms (Holloway 1991), with areas in which a group is not
represented coded as plesiomorphic (absence; C) or unknown (D).

centred in the savannah region of Africa and in
the Ponto-Mediterranean area.
A number of other generic centres for butter-
flies are evident in Africa (Carcasson 1964) and
Australasia (data tabulated but not analysed;
Kitching 1981), but these do not contribute taxa
to the Indian fauna (an exception may be the
Madagascan-centred Papilio demodocus group).
In Africa there is a major group of 'sylvan'
genera with a coincident centre of richness in the
Congolese rainforests, and generic groups
centred on the Cape (Cottrell 1978) and in Mad-
agascar. In Australasia there is a major tropical
generic centre in New Guinea and a temperate
one in S. E. Australia.
Representatives of the various generic types
tend to segregate ecologically in India, with Orien-
tal tropical species well represented in the Himala-
yan lowland terai, the western Ghats and Sri
Lanka, the western Chinese centred genera repre-
sented more at middle Himalayan altitudes, giv-
ing way to species from the N. W. Himalayas and
Pamir centre above that. African savannah genera
are represented more in the seasonal savannah of
the plains, and Mediterranean taxa in semiarid
habitats of the north-west. For a similar approach
to the Lepidoptera of the Arabian Peninsula, see
Larsen (1984) and Wiltshire (1980,1990).
Though the Indian subregion is geologically
composite, the peninsula itself being a fragment
of Gondwanaland, there is no clear evidence of
surviving elements of Gondwanan biota (in but-
terflies at least), and the biogeographic pattern
appears to be climatically, and therefore ecologi-
426 Jeremy D. Holloway & Ebbe S. Nielsen

BRACCA
6 species
OUTGROUP endemic lo
New Guinea

georgiata BORNEO, SUMATRA, JAVA, PHILIPPINES, SULAWESI, S. MOLUCCAS

13757 . SC.SULAWESI

11539 VANUATU
teleleuca BISMARCKS
disrupts SOLOMONS

ribbei NEW GUINEA

lucida BISMARCKS
flavitaenia SULAWESI (SULA IS.)
bajularia S. MOLUCCAS
emolliens SULAWESI
14097 N S.E. & E. SULAWESI
14099 E. SULAWESI
maculosa MALAYA, SUMATRA, BORNEO
subfumosa SUMATRA, BORNEO
exul JAVA

monochrias PHILIPPINES (+ SANGIHE)

benguetana N. PHILIPPINES
xanthosoma N. & S. MOLUCCAS
matutinata AUSTRALIA (N.)
interrupta BISMARCKS
rotundata AUSTRALIA (N.), S. MOLUCCAS
barbara N. S S. MOLUCCAS
basiflava
rosenbergi SOUTH MOLUCCAS, NEW GUINEA, BISMARCKS
woodfordii SOLOMONS

- revocans BORNEO-S.W. PACIFIC

RUSICADA
- fulvida ORIENTAL TROPICS - SUMATRA, JAVA
- nigritarsis INDO-AUSTRALIAN TROPICS
- mafalui NEW GUINEA

- albipunctata NEW GUINEA-SOLOMONS

- combinans ORIENTAL TROPICS

- schislosema MOLUCCAS - QUEENSLAND & VANUATU
- privata CHINA, JAPAN
- metaxantha N.E. HIMALAYA
- lineosa INDIA
- fructusterebrans THAILAND

- sp. n. THAILAND
- prima N.E. HIMALAYA-SUNDALAND
- leucolopha ORIENTAL TROPICS - JAPAN, SUNDALAND
- (ulminans NEW GUINEA
- vulpina FUI, ? SAMOA
- albitibia INDO-AUSTRALIAN TROPICS
- leucosema AFRICA
- melanosema AFRICA

Fig. 21.2. Cladograms for two diverse Indo-Australian tropical genera. In the geometrid genus Bracca (Holloway
1991) the majority of species are extremely localized and show a high degree of allopatry. In the noctuid genus
Anomis, the subgenus Rusicada (Holloway, unpublished) shows a high degree of sympatry with two species, two
sister-pairs and one complex lineage spanning the whole Indo-Australian tropics. Characterization for the Rusicada
cladogram available from J D H on request.

cally, composite rather than tectonically deter-
mined.
A further modification of the R-mode ap-
proach, classifying natural generic and subgen-
eric groupings or clades in terms of the represen-
tation of their species amongst areas of ende-
mism and wider faunal elements is currently un-
der investigation for the Indo-Australian tropics
using data for Lepidoptera and cicadas (Hollo-
way, in press a).
Biogeography of the Lepidoptera 427

Fig. 21.3. Map of the Indo-Australian tropical archipelago showing some frequently encountered species distribu-
tion patterns (? generalized tracks) derived from R-mode analyses (Holloway 1973, 1979, 1982): A, mainland
Asian (can overlap B); B, Sundaland; C, Lesser Sunda; D, Wallacean; E, Melanesian (can overlap G and H); F,
tropical Australian (Torresian); G, New Caledonian; H, Fijian; I, Coral Sea; J, Indo-Australian. In addition, there
are species endemic to individual islands and to areas within islands. For an analysis of pattern within the Indian
Subregion, see Holloway (1974).

Pattern at an Ecological Level Transects from the more centrally placed is-
lands of the Indo-Australian tropics (Borneo, Su-
Similar Q-mode and R-mode methods of analy- lawesi, Seram) have revealed a humped profile
of diversity with altitude (Fig. 21.5), the greatest
sis can be utilized to investigate ecological pat-
diversity being recorded between 500 m and
tern in Lepidoptera, particularly moths. Light-
1000 m. Hypotheses attempting to address this
trapping provides a means of rapidly acquiring
phenomenon include: the possibility that though
large quantifiable samples. Indeed, the seminal
lowland site diversity is lower, species turnover
work on the measurement of biological diversity
(beta-diversity) between sites may be greater
(Fisher, Corbet & Williams, 1943) drew heavily
than at higher altitudes; vegetation zone depres-
on quantitative data for Lepidoptera. Conducted
sion during the glaciations decreasing extreme
over a transect it can provide a ready means of lowland diversity through extinction (Holloway,
investigating variations in diversity, assessing zo- Robinson & Tuck 1990). However, Colwell &
nation pattern or ecological continua, and recog- Hurtt (1994) have demonstrated that this effect
nizing associations of species that may be related could also arise as a sampling artefact.
to particular vegetation types. The most detailed
Within this humped profile are distinct but
transects have been undertaken in areas of rela-
overlapping associations of species: lowland,
tively uniform climate in the Indo-Australian
lower montane, upper montane and some at
tropics (Holloway 1970, 1977, 1979, 1984 a,
higher altitudes (Fig. 21.6). There is also an asso-
1987 a, 1993 b; Holloway, Robinson & Tuck
ciation characteristic of open, disturbed habitats.
1990; Robinson 1975), so there is a need for data
Diversity in S. W. Pacific islands is much lower
extending the approach into the more seasonal
(Fig. 21.5), and species associations recognized
and temperate parts of the world (e. g. Yoshida
are mostly characteristic of different lowland veg-
1981, 1983), as well as for comparisons with the
etation types, e. g. in Fiji (Robinson 1975) and
other major tropical regions.
New Caledonia (Holloway 1979,1993 a).
428 Jeremy D. Holloway & Ebbe S. Nielsen

Fig. 21.4. Map of the Old World showing the major coincident centres of butterfly generic species richness (mass-
ing centres), as derived numerically by Holloway (1969, 1974) and more subjectively by Carcasson (1964) and
Kitching (1981): 1, African sylvan; 2, African savannah; 3, Cape; 4, Madagascan; 5, Ponto-Mediterranean; 6,
N. W. Himalayan-Pamir; 7, W. Chinese; 8, Tropical Oriental; 9, New Guinea-Torresian; 10, Bassian; 11, New
Zealand. Tapering lines from 8, 7 and 6 indicate that species from genera of these centres replace each other with
altitude in the Himalaya. In peninsular India, species of centre 8 predominate in C but are increasingly replaced
by those from centres 2 and 5 in and A.

Holloway 1983 a; K. S. Brown 1991). The data

Altitude in metres
Fig. 21.5. Profiles with altitude of maximum macro-
lepidopteran diversity (the alpha statistic of Fisher
et al. (1943)) recorded in natural vegetation from local-
ities in the Indo-Australian tropics (from Holloway &
Barlow 1992).
Lepidoptera Biogeography in
Conservation: Critical Areas,
Vulnerable Groups and Bioindicators
Methods of pattern analysis can be combined
with the extensive information base available for
the Lepidoptera to provide data for conservation
planning and environmental monitoring (e. g.
presented earlier on faunal centres of Indian but-
terfly genera indicate that some parts of the
world support significantly higher species rich-
ness than others. Even data from phenetic bio-
geographic analyses can be used to identify areas
of significant endemism and richness (Holloway
1984). But considerably more sophisticated
methods are in the process of development.
Ackery & Vane-Wright (1984), followed by
Collins & Morris (1985), developed critical fau-
nas analysis, exemplified by butterfly data (Da-
nainae (Fig. 21.7) and Papilionidae respectively),
to rank areas of the world in order of conserva-
tion importance and identify the minimum set of
areas (or faunas) which would contain at least
one population of every species of the group con-
cerned. These analyses should lead to conserva-
tion action plans for the group (New & Collins
1991).
More recently a modified critical areas analy-
sis method (WORLDMAP) was presented by
Vane-Wright et al. (1991) and Williams et al.
(1991), combining a measure of taxic diversity
derived from the information content of avail-
able cladistic classifications (though any hierar-
Biogeography of the Lepidoptera
Geometroidea I
Zygaenoidea I
Arctlidae I
Sphingidaa [
Uranioidea I
Noctuidae
Nolodontidae I
Lymantriidae f
Drepanoidea I
Bonibycoidea |
{-Sphmgidae) L
Fig. 21.6. Summary diversity profile with altitude for
Borneo from Fig. 21.5 indicating how species associa-
tions recognized in numerical analyses replace each
other with altitude and contribute to overall diversity.
The bar charts below show proportions of species from
various macrolepidopteran families in samples from
different altitudes. The bar charts above compare fam-
ily proportions in a lowland forest sample in Sulawesi
(left) with proportions in a much less diverse sample
from a coconut and clove plantation at a similar alti-
tude (right). Partly after Holloway & Barlow (1992).
L M F lower montane forest; U M F - upper mon-
tane forest; RS Radio Sabah association; S sum-
mit association (the last two are virtually restricted to
Mt. Kinabalu).
chic system of classification can be used) with the
sort of distributional information used in critical
faunas analysis. Step-wise procedures using com-
plementarity between areas can identify opti-
mally efficient, single site sequences of priority
areas for a group. Danaines have again been
used to exemplify the method, and Lepidoptera
data promise to be particularly suitable for its
application: Ithomiinae, Papilionidae and Sphin-
gidae are currently under investigation (R. I.
Vane-Wright, pers. comm.), and an analysis of
Philippines Hesperiidae has been published by
Vane-Wright & Rahardja (1993). Figure 21.8
shows two measurements of biodiversity of the
Australian endemic primitive hepialid genus
Fraus using WORLDMAP based on data from
a taxonomic revision of that genus (Nielsen &
Kristensen, 1989). DIVERSITY, a software
package developed by Faith & Walker (1993,
1996), is another powerful tool estimating envi-
ronmental diversity.
429
A complementary concept is that of vulner-
able groups (Holloway & Barlow 1992; Hollo-
way 1994). Tables can be prepared for more di-
verse higher taxa in a given area showing pro-
portions of species represented in different bio-
geographic categories (endemic to widespread)
as against representation in different ecological
categories (lowland v. montane, forest v. open
habitats). This was exemplified by Holloway &
Barlow for a sample of about 1200 Bornean
moth species (Fig. 21.9). Obviously groups with
a high proportion of endemics restricted to low-
land forest are particularly vulnerable to logging
operations, and many of the groups studied are
in this category. Groups with high montane en-
demism will be more vulnerable to hill resort de-
velopments. Groups with a high proportion of
geographically widespread open habitat special-
ists, such as the trifine Noctuidae, will largely
benefit from human activity. Indeed, it is not sur-
prising that agricultural pests tend to be concen-
trated in such groups. See also Thomas (1991).
It is also possible to derive an index of biogeo-
graphic quality for different habitats by scoring
each species recorded on a scale of values from
low (widespread) to high (endemic) according to
its distribution type (e. g. the categories in
Fig. 21.9) and calculating the average score for
the species in the habitat. This can be combined
with a diversity index to give an assessment of
conservation value: high diversity is not neces-
sarily correlated with richness in endemics as is
seen in New Caledonian moth associations (Hol-
loway, 1993 a). Applications of this approach to
butterflies in forests of Vietnam and Indonesia
have been published by Spitzer et al. (1993), Hill
et al. (1995) and Hamer et al. (1997). These
studies observed a decline in biogeographic qual-
ity with increasing forest disturbance.
Lepidoptera data of this nature can provide a
rough guide to the biodiversity cost of various
types of human activity in modifying the envi-
ronment (e. g. Thomas 1991), and enable more
specific sampling programmes to be planned that
will measure the effects more precisely, e. g. of
logging (Holloway, Kirk-Spriggs & Chey 1992;
Hill et al. 1995; Intachat et al. 1997), shifting cul-
tivation or conversion to modern agricultural
systems (Holloway & Stork 1991; Holloway &
Barlow 1992; Holloway 1994, in press b; Chey
et al. 1997). Erhardt & Thomas (1991) have used
butterflies as indicators of the effects of different
regimes of pasture management in Europe.
More detailed assessments of the value of Lep-
idoptera as bioindicators, both in comparison
with other organisms and with comparisons of
different groups of Lepidoptera, are made with
respect to a range of criteria by K. S. Brown
(1991) and Holloway (1983 a, 1984 a, 1985 a).
The Geometroidea appear particularly suitable.
Rapid and predictable response to changing
environmental parameters is a criterion of a
430 Jeremy D. Holloway & Ebbe S. Nielsen

30 60 90 120" 150 180 150 120 90 60

Fig. 21.7. Map to show the distribution and composition of the minimum number of regional faunas need to be
conserved in order to ensure preservation of representatives of all danaine species; countries or islands listed in
italics include endemics. 1, Comoro Islands', 2, Seychelles; 3, Mauritius; 4, Zaire; 5, Sri Lanka; 6, southern India;
7, Nepal; 8, Burma; 9, Sumatra; 10, Java; 11, Borneo; 12, Luzon; 13, Negros; 14, Mindanao; 15, Sulawesi; 16,
Sumbawa; 17, Sumba; 18, Flores; 19, Timor; 20, Seram; 21, Biak; 22, New Guinea; 23, New Ireland; 24, New
Britain; 25, Guadalcanal; 26, San Cristobal; 27, New Caledonia; 28, Hispaniola; 29, Cuba; 30, Costa Rica; 31,
Bolivia. The New World has a danaine fauna of 14 species, the Afrotropical region has 21, and the Indo-Pacific
region has 125 (world total: 157). From Ackery & Vane-Wright (1984).

13

DO 03
, ,

T
mm^rni....;...
m 9s nf i .|
3
I I I 3 '6
R30

P P I l ' S

Fig. 21.8. Two measures of the biodiversity of the hepialid genus Fraus in Australia using W O R L D M A P 2.4;
species richness (left) and taxonomic dispersion (right). Note that the 'hotspot', the area with the highest diversity,
is in southwest Australia using the species richness measure and in southeast Australia applying the taxonomic
dispersion measure. Data from Nielsen & Kristensen (1989) compiled by R. I. Vane-Wright (unpublished).

good bioindicator group that will be increasingly
relevant in monitoring any climatic repercus-
sions of the greenhouse effect. The response of
an insular lepidopteran fauna to intensive
drought stress has been measured on Norfolk Is-
land over a period of several years (Holloway
1977, 1983 a, 1996), revealing both changes in
the equitability of species abundance and dif-
ferential effects with a biogeographic basis, sa-
vannah and humid tropical grass feeding species
switching abundance ranking in dry versus wet
periods. An important review and assessment of
butterfly ecology in relation to climate change
has recently been published by Dennis (1993):
the focus is predominantly European.
A range of computer software packages such
as BIOCLIM (Nix, 1986), CLIMEX (May-
wald & Sutherst 1991) and DOMAIN (Carpen-
ter et al. 1992) now allows the prediction of dis-
tributions of organisms using biological and cli-
matic information. Four steps are involved in de-
veloping predicted distribution: (1) a number of
high-quality real distribution records, (2) estima-
tion of the climate at each location where the
species has been recorded, (3) matching the bio-
climatic profile of the species with climate esti-
Biogeography of the Lepidoptera 431

RS
+ LMF
+ Low
+ RS
+ LMF
+ Low
+
2 Low LMF UMF Open 4 Low LMF UMF Open
S UMF Mont. S UMF Mont.

Endemic 22.58 4.30 6.45 1.08 1.08 Endemic 21.13 11.27 0.70 0.70

Borneo Borneo
+
1.08 1.41 1.41
Wallacea Wallacea

Sundaland 40.86 2.69 2.69 1.08 2.15 Sundaland 2.82 2.82 1.41

Sundaland Sundaland
+ + 4.23
Wallacea Wallacea
Himalaya Himalaya
+ 4.30 1.08
1.08
Sundaland Sundaland
Wide Wide
6.45 2.82 2.82 4.23 0.70 1.41 3.52
Oriental Oriental

Indo- Indo-
1.08 1.41 4.23 1.41 28.16
Australian Australian

Limacodidae Higher tritine Noctuidae

Similarity
30 40 50 60 70 80
I L _ L
Low Endemism
4 Higher Wines

7 Arctiinae

- 10 Sphingidao

5 Euteliinae

- 6 Stictoptennae
Montane
- 3 Lower trifines

- 12 Baptini

- 14 Boarmiini

- 15 Ennominae residue

- 1 Cossoidea

- 13 Cassyma group

- 11 Hypochrosini

- 8 Notodonlidae

- 2 Limacodidae

- 9 Lasiocampidae
Lowland
8 forest
High Endemism

Fig. 21.9. Single-link cluster analysis classification with a linkage diagram of two way tables (biogeography v.
ecology) for 15 moth groups in the fauna of Borneo to indicate vulnerability or potential for biodiversity loss.
Two example tables are given at the top. Limacodidae have a high proportion of endemics and relatively localized
species in lowland forest, hence are very vulnerable to forest clearance. Trifine Noctuidae have endemics in mon-
tane habitats and hence are vulnerable to hill resort development, but one third also are widespread in open
habitats and will respond positively to human disturbance (e. g. as pests). The biogeographic categories are based
on those illustrated in Fig. 21.3, and the ecological ones on the associations illustrated in Fig. 21.6. The similarity
measure is the summed percentage overlap over all the boxes in the tables for each pair of taxa being compared,
e. g. in the 'Sundaland' row for the two example tables, the overlap is 2.82 + 2.69 + 1.08. The linkage diagram indi-
cates pairs of taxa with percentage overlap of 60 or more (solid lines), and 50 to 59 (broken lines). Taxa 4 and 7 link
in at lower percentage levels as indicated by dotted lines with arrows. The classification indicates that high endemism
in lowland forest is the " n o r m " for Bornean moth groups, with specialism to open or montane habitats peripheral
and of irregular occurrence. See Holloway & Barlow ( 1992) and Holloway ( 1994) for a more detailed account.
432
mates in the wider area, and (4) a spatial repre-
sentation of the predicted potential distribution.
Such predicted distribution may allow optimal
identification of areas of high predicted species
richness and application of this technology may
also allow the prediction of future distribution
such as under various 'greenhouse' scenarios.
Lepidoptera Pattern and Process
In a short chapter it is not possible to give more
than a brief survey of the types of biogeographic
pattern seen across the world in the Lepidoptera,
or go deeply into the more contentious area of
the processes contributing to the development of
those patterns. The sample of modern cladistic
treatments of genera is still very small, though
those for the Lepidoptera are relatively nu-
merous.
One treatment of a worldwide group of but-
terfly genera (Craw 1989) can be used to il-
lustrate some of the problems involved in pre-
senting a concise account of pattern and process
in Lepidoptera biogeography. Craw provided a
cladistic analysis of Vanessa and Bassaris, with
the Cynthia complex as an outgroup. It is repro-
duced in Fig. 21.10, with additional input from
Shapiro (1992 a, b) and other sources. The
pattern in Vanessa and Bassaris is largely allopat-
ric though Vanessa indica and atalanta have some
overlap in Eurasia, and Bassaris itea is sympatric
with B. gonerilla in New Zealand. B. itea also oc-
curs on Norfolk Island and, even more remotely,
on the Polynesian island of Rapa. It and V. ata-
lanta are known migrants. V. tameamea is en-
demic to the Hawaiian archipelago, and there
are also localized taxa in the Indo-Australian ar-
chipelago. Craw interpreted the largely allopatric
pattern as one of progressive vicariance
following a period of mobilism in the absence of
barriers that established the ancestral range of
Bassaris + Vanessa. Subsequent limited mobil-
ism by the migratory taxa has resulted in the
only instances of breakdown of allopatry. The
genus Cynthia is predominantly New World,
with one subgroup, the carduHkershawi sister-
group (Africa, Eurasia v. Australasian allopatry),
occurring in the Old World. Migratory behavi-
our is seen in this subgroup and at least one
other subgroup in Cynthia (transatlantic dis-
persal of C. virginiensis).
Several process questions arise from this
pattern. Firstly, strong migratory behaviour is
exhibited by species in all branches of the clado-
gram. Is it therefore a plesiomorphic trait, with
homoplasious development of reduced mobility
in most lineages, particularly in extremely insular
situations? The presence or absence of barriers is
a relative concept dependent on the powers of
dispersal of the group compared to the distances
involved. The exponential model of dispersal of
Jeremy D. Holloway & Ebbe S. Nielsen
MacArthur & Wilson (1967) encapsulates this,
with distance, area of species range at source and
dispersal power being the variables in the equa-
tion. The model was tested and found to hold
for the biota of Norfolk Island (Holloway 1977,
1996) and was extrapolated to Sulawesi (Hollo-
way 1991) in an attempt to understand varia-
tions in species richness of various Lepidoptera
groups there. There appears to be a corollary
within the Vanessa group in an apparent correla-
tion of resident geographical range of taxa and
observations of migratory powers.
The carduHkershawi sister-pair has a similar
geography to the Vanessa/Bassaris sister-pair,
and hence the primary dichotomy might be at-
tributed to the same vicariance phenomenon. Is
the subsequent difference in the degree to which
fragmentation proceeded in each lineage due to
a different response of the taxa to the same vic-
ariance events, such as a different degree of mo-
bilism (and therefore retention of genetic conti-
nuity across the range) or to a failure in our abil-
ity to detect similar differences between popula-
tions in the range of cardui and kershawi
The presence of an endemic species, V. ta-
meamea, in the Hawaiian archipelago poses sim-
ilar questions. It is sister-taxon to the rest of
Vanessa except atalanta. Does this lead to a geo-
logical mobilism hypothesis of Hawaii as a de-
tached fragment of Eurasia or to the biological
mobilism hypothesis of a colonization event
from Eurasia? When one builds in more data,
the second hypothesis becomes more plausible,
particularly in face of evidence of the Hawaiian
chain having developed over a 'hot-spot' or man-
tle plume, counter to the detached fragment hy-
pothesis (see Wagner & Funk (1995) for a com-
prehensive treatment of Hawaiian biogeog-
raphy).
The subgenus Pseudaletia of Mythimna
(Franclemont, 1951; Holloway 1983 b) also has
endemic taxa in Hawaii. But the relationship of
these is to New World taxa (Fig. 21.10), one of
which, unipuncta, is migratory (McNeil 1987)
and occurs also in both Hawaii and Europe. The
sister-group consists of the widespread and mi-
gratory Indo-Australian species separata, and
more restricted taxa in Seram 4- New Guinea,
the Philippines, and Australia (with a subspecies
on Norfolk I.). The Australian species is also mi-
gratory (Farrow & McDonald 1987). Here again
migratory behaviour may well be plesiomorphic,
given the distribution of migratory taxa through-
out the cladogram, and there is also probably
some correlation between migratory powers and
geographical range within the subgenus.
The situation for Hawaiian Heliocoverpa is
more complex. The cladogram of Mitter et al.
(1993) has a Hawaiian species pair as sister-
group to a pan-tropical, but mostly Old World
complex that includes three further Hawaiian en-
demics.
Biogeography of the Lepidoptera 433

MYTHIMNA SUBGEN. PSEUDALETIA

narrowed sacculus converta AUSTRALIA
asymmetric cucullus
(costa concave) nobbsi NORFOLK IS.
quadrate harpe
sp. n. NEW GUINEA, SERAM

idlsana LUZON

separata INDO-AUSTRALIAN TROPICS & SUBTROPICS

sp. n. TAHITI

unlpuncta NEW WORLD + S.W. EUROPE + HAWAII

cunyada COLOMBIA

apically roraimae N. BRAZIL

bifid juxta
adultera S. BRAZIL, . ARGENTINA, URUGUAY, PARAGUAY

punctulata CHILE

sequax C. & S. AMERICA

cucullus slender,
dasuta HAWAII
apically reduced
macrosaris HAWAII

amblycasis HAWAII

VANESSA GROUP
tameamea HAWAII
amerindica
COLORADO, USA
- ? vulcania CANARY IS.
(Oligocene fossil)
saman SUMATRA

dejeanii JAVA, BALI, LOMBOK, SUMBAWA

I
L VANESSA
..?
mounseyi

buana

Indica
MINDANAO

S. SULAWESI

INDIA, CHINA, KOREA, TAIWAN, LUZON

atalanta HOLARCTIC

tea AUSTRALIA, NEW ZEALAND, NORFOLK IS., RAPA

gonerilla NEW ZEALAND

BASSARIS
ida CHATHAM IS.

CYNTHIA - 3 subgroups

> mainly AMERICAS

except cardui and kershawi

Fig. 2 1 . 1 0 . C l a d o g r a m s for: the Vanessa generic complex m o d i f i e d from Craw ( 1 9 8 9 ) in view of the discussion by
Shapiro ( 1 9 9 2 a, b) and personal c o m m u n i c a t i o n from R . I . Vane-Wright o n the status o f buana; subgenus Pseuda-
letia of Mythimna as reviewed by H o l l o w a y ( 1 9 8 3 b) from a study o f Franclemont ( 1 9 5 1 ) and Z i m m e r m a n ( 1 9 5 8 ) .
See the text for further details. The position of the fossil amerindica is uncertain; Shapiro stated it t o be phenetically
m o s t similar t o indica.

If o n e t h e n a d d s in o t h e r c o m p o n e n t s o f the Continents and Islands

H a w a i i a n f a u n a s u c h a s t h e Anomis species with
m o r e southerly tropical relationships, then main- The Lepidoptera faunas of oceanic islands s h o w
tenance of any hypothesis of geological mobilism a s u r p r i s i n g h o m o g e n e i t y w o r l d w i d e (e. g. F e r g u -
requires c o m p l e x c o n c e p t s o f H a w a i i as a c o m - s o n et al. 1 9 9 1 ) t h a t a g a i n a c c o r d s w i t h h y p o t h e -
posite area through 'island integration' (Ro- ses o f biological rather t h a n g e o l o g i c a l m o b i l i s m .
t o n d o et al. 1981). We have collated data for proportions of macro-
434 Jeremy D. Holloway & Ebbe S. Nielsen

1 2 3 4 5 6 7 8 9 10 11 12
Cossoidea KEY
A World 73582
Geometroidea
1 Japan 2555 1

Zygaenoidea M 2 Borneo c.4500

3 Sulawesi c.3000 I

Arctiidae i
4 Seram c.2000
Sphingidae 5 Solomons 600+
Uranioidea 6 Vanuatu 293
7 Hawaii 140
Noctuidae
8 Fiji 344
Notodontidae 9 Samoa 140
Lymantriidae
10 North America 5159
Drepanoidea
Bombycoidea 11 Galapagos 103
(-Sphingidae) 12 Bermuda

13 Ascension 16
14 Saudi Arabia 622
15 South Africa 3400
16 Reunion 178
17 Taiwan 2374
18 Cocos-Keeling 28
19 Australia 5674
20 New Zealand 365
21 Norfolk resident 58
22 Norfolk migrant 40
23 New Caledonia 345
24 Tonga 73
25 Polynesia 63
26 Henderson 14

13 14 15 16 17 18 19 20 21 22 23 24 25 26

Fig. 21.11. Proportions of species in different macrolepidopteran families from various parts of the world as
discussed in the text.

lepidoptera families represented in the faunas of
N. America (Hodges et al. ), Bermuda (Ferguson
et al. 1991), Ascension I. (Robinson & Kirke
1990), the Galapagos (Hayes 1975), Saudi Arabia
(Wiltshire 1990), South Africa (Vari & Kroon
1986), Reunion I, (Guillermet & Guillermet
1986) Cocos-Keeling Is. (Holloway 1983 c), Ja-
pan (Inoue et al. 1982), Taiwan (Heppner & In-
oue 1992), Borneo, Sulawesi, Seram, Solomons,
Vanuatu, New Caledonia, Fiji, Samoa, Tonga,
Norfolk I., an average high volcanic Polynesian
island (Holloway 1987 a, 1993 b), Henderson I.
(Holloway 1990; Benton 1995), New Zealand
(Dugdale 1988), Australia (Nielsen et al. 1996)
and Hawaii (Zimmerman 1958). These data are
presented in Fig. 21.11, together with some indi-
cation of the number of species in each fauna. A
column for the world fauna is also included,
Biogeography of the Lepidoptera
Ascension Bermuda Reunion Cocos -
Keeling
Agrius
Utetheisa
Agrotis
Heliothis
+ Helicoverpa
Mythimna #
Spodoptera
Condica
Chrysodeixis
w
+ Pseudoplusia
Mocis "
Anomis #
Anticarsia
Hypena
Table I. Distribution of widespread genera on remote islands.
based on Heppner (1991) but with composition
of families and superfamilies as in other chapters
of this Handbook. Heppner's figures probably
require further checking (e. g. 75 Oriental Helio-
thinae compared with perhaps 1520 recognized
by Matthews (1991), and see also Herbulot
(1992) and Scoble, Gaston & Crook (1995) for
criticism of the Geometridae figures).
Several trends are apparent. Tropical faunas
are much more diverse than temperate ones both
in terms of absolute richness and in terms of the
representation of families, with smaller families
relatively better represented in the tropics. In
tropical latitudes insular faunas show a relation-
ship between island area and number of species.
Robinson (1975) demonstrated that the classic
log./log. linear species/area relationship (Preston
1962; MacArthur & Wilson 1967) is valid for
macrolepidoptera faunas of islands in the S. W.
Pacific. Latitudinal changes of family diversity
are reflected in similar changes with altitude in
the tropics (Fig. 21.2; Holloway 1986).
The Australian fauna is unique in the extraor-
dinary radiation of the oecophorid subfamily
Oecophorinae: with about 6000 species in about
270 genera the Oecophorinae are the second
largest family group of fauna in Australia and
comprise over 20% of the Australian Lepidop-
tera. The group has undoubtedly radiated with
the Myrtaceae, particularly the large genus Euca-
lyptus; a large percentage of the Australian Oeco-
phorinae feed as larvae on dead Eucalyptus
leaves and are important to the leaf litter nutri-
ent cycle (Common 1981, 1994, 1997). Other as-
pects of the Australian Lepidoptera fauna are
discussed by Common (1990).
435
Henderson Norfolk Norfolk Galapagos Texas oil
resident migrant platforms
Family diversity declines with isolation in is-
lands such that in the most remote faunas only
the Noctuidae, Arctiidae, Geometridae and
Sphingidae are represented. This phenomenon is
common to remote islands in all the major
oceans, such as Ascension and Bermuda in the
Atlantic, Reunion and Cocos-Keeling in the In-
dian Ocean, and Polynesian islands (e. g. Hen-
derson) and the Galapagos in the Pacific.
Furthermore, for these and for Norfolk I. (Tas-
man Sea) a dozen genera have representatives re-
corded from almost all of them as residents or
migrants (Table 1). Ferguson et al. (1991) have
presented a more extensive account of this ge-
neric selectivity in island faunas. A high propor-
tion of these genera are rich in open habitat spe-
cialist species, often agricultural pests. Many are
recorded offshore on oil platforms (Sparks et al.
1986) and weather ships (Yoshimatsu 1991), or
in unsuitable localities such as deserts (Pedgley &
Yathom 1993). They contribute to the rich array
of migrant species recorded on Norfolk I., most
with resident species also (Holloway 1977,
1982 b). Migrants are also well represented in the
Bermuda fauna (Ferguson et al. 1991).
These observations can probably be extrapo-
lated to the Ditrysia generally, though compre-
hensive data are not available. Amongst the
microlepidopteran ditrysian families the Cos-
mopterigidae, Gelechiidae, Pterophoridae, Pyra-
lidae, Tortricidae and Tineidae are usually repre-
sented on even the most remote islands such as
Easter I., Henderson or Ascension, and these are
also the most diverse of the microlepidopteran
families on other oceanic islands with larger fau-
nas such as Bermuda.
436
Fig. 21.12. Dynamics of Spodoptera exempta, the African army worm, a migratory pest of pasture [from Haggis
(1984)]. The total recorded African distribution is shown top left, and the three other maps show the recorded
incidence of outbreaks from 1940-1982 for the months of January, June and August.
In contrast, non-ditrysian groups show an
increase in family diversity towards temperate
latitudes, are rarely found on oceanic islands (the
Opostegidae on Hawaii are exceptional), and
their presence might be taken as indicative for
land areas with a geologically long and con-
tinuous subaerial history. This difference is de-
scribed in more detail in the final section.
Grehan (1991) has highlighted the transoceanic
relationship of most of these non-Ditrysian
groups, but within the oceanic tectonic plates
themselves, the archipelagos support faunas that
are highly biased towards the more mobile com-
ponents of the ditrysian families. It is therefore
plausible that, through the evolution of the Lepi-
doptera, biological mobilism has increasingly re-
placed geological mobilism as the predominant
influence on development of biogeographic
pattern in the Order.
Migration
Migration in the Lepidoptera is now a well
known phenomenon. Since the classic review by
Jeremy D. Holloway & Ebbe S. Nielsen
Williams (1930), it has been increasingly well
documented, e. g. for the Tasman Sea as re-
viewed by Holloway (1977, 1982 b) and Dugdale
(1989), with particular reference to the records
made by K. J. Fox (Fox, 1978). The most dra-
matic instance is the recording of Utetheisa
pulchelloides of presumably Australian origin in
Vanuatu, Norfolk I. and New Zealand within the
space of a few days. The literature is too exten-
sive to review here, but the unravelling of the
Danaus plexippus story in the U. S. A. (e. g. Ur-
quhart 1987; Brower 1986) and Australia (Smith-
ers 1977) and its range expansion within histori-
cal times (Vane-Wright 1993 a) is worthy of men-
tion, as is the detailed analysis of the regular
Spodoptera exempta movements in Africa (Fig.
21.12) by Haggis (1984). Radar (Fig. 21.13) is
increasingly being used to track migration (e. g.
Drake 1985; Drake & Farrow 1985), and sophis-
ticated climatic back-tracking is often possible
(e. g. Larsen & Pedgley (1985), discussing the ar-
rival of Indian butterflies in the Arabian Penin-
sula). Migrant taxa recorded (Sparks et al. 1986)
on unmanned oil platforms at 32-160 km off
Biogeography of the Lepidoptera
Fig. 21.13. Radar scan showing two layers of migrat-
ing insects with a strong orientation pattern (i. e. the
bands); the insects are migrating towards the south-
east. Each individual dot represents one insect. Insects
are identified by their wingbeat frequency and through
collecting by nets from kites. (Photograph courtesy of
R. A. Farrow)
Fig. 21.14. Migration of bogong moth, Agrotis infusa,
in Australia. Open arrows indicate spring migration
from the breeding areas and solid arrows late summer/
autumn migration from the aestivation areas.
the coast of Texas include many of those listed
for remote islands in Table 21.1. In mainland
areas, light-trapping on high towers can provide
good indications of migrant movements (Gregg
et al. 1993).
The Australian bogong moth, Agrotis infusa,
is a particularly interesting example of lepidop-
teran migration where the same specimens mi-
grate from the breeding areas to the mountains
in southeast Australia where they aestivate dur-
ing the summer and then move back to the
breeding areas (Fig. 21.14). The bogong moth
occurs commonly in non-tropical Australia and
only the populations in eastern Australia are
known to undertake this two-way annual migra-
tion (Common 1954). Strays from this migration,
437
particularly the movement to aestivation sites,
are regularly recorded on Norfolk Island (Hollo-
way 1977, 1982 b).
Refugia! Patterns
It is possible to recognize several broad cate-
gories of biogeographic pattern in the Lepidop-
tera, though these are by no means restricted to
the Order, and mostly very general.
The first might broadly be termed rfugiai,
and related to Quaternary climatic changes, par-
ticularly the glaciations. With the advance and
retreat of the ice in the northern hemisphere, the
trend for both plant and insect species was to
track their climatic optima rather than to adapt
to changing conditions. This is well documented
for beetles (Coope 1975, 1991), and there is no
reason to expect the Lepidoptera to have be-
haved differently. Vegetational changes during
these times were marked not by block movement
of plant communities but by significant disasso-
ciation, independent movement of taxa and reas-
sociation (Huntley & Webb 1989). The phytoph-
agous specificity of the Lepidoptera may have
led to a similar effect (Dennis et al. 1991) but, in
the absence of a fossil record such as exists for
beetles, this is mere speculation.
The patterns of Lepidoptera distribution in
the Palaearctic today are usually attributed to re-
colonization of glaciated areas from an array of
refugia in middle latitudes (De Lattin 1967; Den-
nis et al. 1991, 1995 a, b): the east central and
west Mediterranean, Middle East and east of the
Himalayas. In many butterfly groups there are
instances where the greatest diversity and ende-
micity is still observed in 'rfugiai' latitudes, with
only one or two species widespread at higher lat-
itudes, e. g. the Parnassiinae, Gonepteryx, Antho-
charis and Melanargia. The initial development
of this diversity was probably influenced by
earlier geological events as discussed for the Me-
diterranean on p. 444. The analysis of Dennis
et al. establishes climatic constraints for groups
of taxa with similar modern ranges, and these
could be projected onto maps under glacial cli-
matic regimes (as in Huntley & Webb) to assess
the contribution of glacial (climatic) versus pre-
glacial (tectonic) events towards development of
modern butterfly geography in the Mediterra-
nean area. Dennis et al. have proposed a more
dynamic model of species turnover to explain
higher endemicity in the Mediterranean area,
with the events of the glaciations serving to ac-
celerate a more general process of speciation and
species turnover.
In North America there is a similar rfugiai
pattern that is shown by a wide range of plant
and animal groups, and is also evident in human
culture (Rogers et al. 1992). The distribution
438
Fig. 21.15. Pleistocene refugia for North America (left, from Rogers et al. (1992)) during the Wisconsinan Glacial
Maximum (map 1), with possible segregation of Munroe's (1952) typical ranges of Canadian insects (maps 213)
amongst them.
categories for Canadian Lepidoptera recognized
by Munroe (1956) and those for the noctuid ge-
nus Euxoa (Lafontaine 1982) appear generally to
fit this pattern also (Fig. 21.15). Mikkola et al.
(1991) proposed the Beringian area as the most
important refugium for naturally Holarctic noc-
tuids but suggested the origins of many such taxa
could be traced further back in the Tertiary to
diversification in the mountains or steppes of
Central Asia (e. g. Xestia, and compare Parnassi-
inae on p. 455).
In tropical latitudes rfugiai patterns are also
thought to occur. They have their greatest com-
plexity in South America, where it is thought
that drier periods during the glaciations caused
the retreat of the moist tropical forests into a
considerable number of pockets where suitable
conditions persisted. Much of the evidence for
this has been gained from studies of butterfly dis-
tributions (Fig. 21.16), particularly the Ithomiini
and Heliconiini (e. g. K. S. Brown 1982, 1987,
1991), but other Lepidoptera groups have also
been studied in this context, such as the Sphingi-
dae (Schreiber 1978). For groups having taxa en-
demic to a high number of putative rfugiai
areas, application of the methods of cladistic
biogeography would be appropriate to test
hypotheses of the sequence of fragmentation of
the rain forest extent into rfugiai pockets, or to
generate such hypotheses for assessment against
Jeremy D. Holloway & Ebbe S. Nielsen
other evidence. Turner (1983) made some prelim-
inary observations of this nature on two helicon-
iine groups.
However, the rfugiai hypotheses for tropical
America have been criticized on a number of
counts, e. g. by Humphries (1989), particularly
for failing to consider that any pattern due to
events during the glaciations is overlain on a
template of patterns originating from earlier
periods in geological time, and by Bush (1994),
who suggested that observed centres of ende-
mism are in areas of maximal disturbance rather
than stability.
Carcasson (1964) related pattern in African
butterfly geography to alternating pluvial and
arid periods in the Quaternary, again with refer-
ence to forest refugia, though these did not ap-
pear to be as complex or numerous as those in
the Neotropics.
The Indo-Australian tropics differ from the
other major tropical areas in being archipelagic,
hence the fluctuations in extent of habitat types
thought to have occurred in continental masses
would here be further complicated by the frag-
mented geography. Sea-level falls of 100 m or
more during glacial episodes exposed major
areas of the Sunda and Sahul continental shelves
such that Java, Sumatra and Borneo were united
with the Asian mainland, and New Guinea was
united with Australia. The butterfly and bird
Biogeography of the Lepidoptera 439

Fig. 21.16. Subspecies endemic centres for three Neotropical butterfly groups (top), with an indication of the
correspondence those for Heliconiini and Ithomiinae with regions thought to have seen the persistence of tropical
forest during long cold, dry spells during the glaciations. From K. S. Brown (1991).

faunas of the Sundaland areas show a greater
uniformity than would be expected from present
geography, suggestive of this past union (Hollo-
way & Jardine 1968). The glaciations appear to
have caused a sequence of alternating episodes
of mobilism and vicariance in the Lepidoptera
(Holloway 1973, 1982 a, 1985 b), with mobilism
from west to east, from Asia into the archipel-
ago, being predominant.
Montane Pattern
In all continents at tropical latitudes, montane
habitats tend to be archipelagic. During the gla-
ciations, vegetation zones fluctuated over alti-
tude of up to 1000 m (Flenley 1979). The Lepi-
doptera faunas of tropical mountains tend to be
made up of representatives of groups with
centres of species richness in temperate latitudes
mingling with intrinsic tropical montane special-
ist taxa and species belonging to predominantly
lowland genera that have given rise to montane
taxa. For example, these categories are repre-
sented amongst Bornean montane geometrids
(Holloway 1986, in press c) as follows: Dys-
stroma, Pasiphila subgenus Gymnodisca (north
temperate), Poecilasthena (south temperate),
Trichopterygini (mainly intrinsic but see Dug-
dale (1980)), Synegia (tropical centre of diversity
with an even spread of lowland to montane spe-
cialist taxa). Lepidoptera diversity decreases with
440
altitude from above 1000 m, and family and sub-
family proportions change in macrolepidoptera
in a manner similar to that with latitude (Hollo-
way 1986 b, 1987 a). Thus larentiine geometrids
are more diverse at high altitudes in the tropics
relative to other subfamilies, as they are at high
latitudes. The butterflies are somewhat of an ex-
ception to this: tropical montane diversity is low
relative to that in the Holarctic (Descimon 1986;
Congdon & de Jong 1992). The latter speculated
that absence of suitable host-plants in tropical
mountains (e. g. in herb-rich meadow habitats)
might be the cause.
Analyses of pattern in the distribution of mon-
tane Lepidoptera are few. The works just men-
tioned are restricted to the butterflies of the An-
des and mountains of eastern Africa respectively.
Rawlins (pers. comm.), in cladistic analyses of
largely allopatric arrays of species in Andean
arctiid groups, found no evidence of an underly-
ing pattern of area relationships, and Cong-
don & de Jong (1992) reached similar conclu-
sions for E. African montane butterfly patterns.
Montane moth distribution patterns in the Indo-
Australian tropics show some correlation with
generalized tracks identified for the flora (Fig.
21.17): from E. China through Taiwan and the
Philippines south to Borneo and Sulawesi; from
the E. Himalaya down through Burma to the
mountains of Sumatra, Java and the Lesser Sun-
das; from New Guinea west to the S. Moluccas,
Sulawesi and Borneo (Holloway 1970, 1986).
The situation in the Himalaya has been dis-
cussed earlier.
Speciation Process
Enrichment of tropical montane faunas by in-
trinsic speciation rather than by colonization
from temperate latitudes has been studied partic-
ularly with respect to montane Andean butter-
flies (Adams 1973, 1977; Adams & Bernard
1977). The fluctuation in vegetation zones during
the glaciations was suggested to have caused al-
ternating episodes of mobilism and vicariance,
particularly of the lower altitude taxa, such that
the products of earlier events are progressively
displaced upwards by competition from below,
partitioned into parapatric altitudinal bands.
Holloway (1970) proposed a similar process for
the evolution of species pairs on Mt. Kinabalu
in Borneo.
Though most episodes of speciation in mon-
tane and actual archipelagos appear to be pre-
dominantly allopatric, this may not always be
the case. Species-rich groups endemic to Su-
lawesi and New Guinea show a high degree of
sympatry across these islands despite evidence
for areas of endemism within them in other
groups. Several instances of this were docu-
Jeremy D. Holloway & Ebbe S. Nielsen
mented for Sulawesi by Holloway (1991). In all
cases the sister taxon to the Sulawesi group was
a lowland Sundanian species or species pair (Fig.
21.18). In Sulawesi the majority of species in
these groups extended over a much greater range
of altitude, through both lowland and lower
montane zones and occasionally into the upper
montane zone. In New Guinea such speciation
has also been accompanied by the evolution of
strikingly colourful wing facies in many lineages
that elsewhere are cryptic, and in many instances
specific strong colour patterns manifest them-
selves across a number of unrelated groups, sug-
gestive of the sort of mimicry rings that are
found in the Neotropics (Holloway 1984 c).
There are Lepidoptera species complexes en-
demic to some of the larger Caribbean islands
(Miller & Miller 1989; Vane-Wright & Ackery
1992; Brower et al. 1992 (see p. 451); J. E. Raw-
lins, pers. comm.) that would similarly repay
analysis, as would the even more dramatic exam-
ples on Hawaii. In each of these cases the leading
research on speciation process has involved other
groups of insects (Liebherr 1988; Liebherr & Ha-
jek 1990; Carson 1984, 1987; Otte 1989).
Process in Island Biogeography
The island biogeographic theory of MacAr-
thur & Wilson (1967) is primarily concerned with
process. Mention has already been made of the
early work of Munroe (1948) that anticipated
many of the concepts (Brown & Lomolino 1989),
of the study of Pacific species/area relationships
by Robinson (1975) and of the assessment of the
exponential model of dispersal in relation to
Norfolk I., extrapolated to Sulawesi, by Hollo-
way (1977, 1991). Lepidoptera studies have also
been used in assessing the validity of the equilib-
rium theory, that species number on an island
is determined by a balance of immigration and
extinction processes.
The equilibrium theory has been used to ex-
plain the observed decline of species richness
from base to apex of a peninsula (Simpson 1964)
as an equilibrium between colonization from the
basal mainland source and extinction along the
peninsula, i. e. a result of the peculiarities of pen-
insula geometry. This has been examined with re-
gard to the butterfly faunas of two major N.
American peninsulas (Fig. 21.19): Baja Cali-
fornia (J. W. Brown 1987) and Florida (Brown &
Opler 1990).
In Baja California species richness was lowest
centrally rather than apically. Vegetation com-
plexity, proximity to the mainland, geological
and climatic factors have obscured any peninsula
effect if it occurs. Though Nearctic species de-
cline in richness from base to apex, Neotropical
ones decline in the reverse direction, having orig-
Biogeography of the Lepidoptera 441

Fig. 21.17. Probable geography of the Indo-Australian tropics during Pleistocene glaciation. The dashed line indi-
cates the margins of the Sahul and Sunda Shelves exposed by sea level depression. Stippled areas indicate zones
of montane habitat. The generalized tracks of van Steenis (1964) for the montane flora are indicated: 1, Luzon
track; 2, Sumatran track; 3, Papuan track. Montane Macrolepidoptera taxa (mostly Geometridae) concordant to
some degree with these tracks are: 1, Dysstroma fumata group, Papilio xuthus group, Cosmorhoe spp.; 2, Habrosyne
spp., Anacronicta spp., Dysstroma cuneifera group, Loxofldonia spp.; 3, Poecilasthena spp., Tympanota spp., Cyclo-
phora flavissima, Tripteridia spp., Ruttellerona lithina.

i n a t e d either vicariously t h r o u g h geological p r o - suggested t o p r e d o m i n a t e : presence of a suitable

cesses o r t h r o u g h dispersal across the sea f r o m
sources t o t h e s o u t h .
In F l o r i d a a similar p a t t e r n w a s observed,
with a t e m p e r a t e c o m p o n e n t exhibiting a p e n i n -
sula effect a n d a t r o p i c a l c o m p o n e n t exhibiting
t h e reverse. Again, ecological c o n s t r a i n t s were
h o s t - p l a n t within a general c o r r e l a t i o n of b u t t e r -
fly richness a n d w o o d y p l a n t richness (Fig.
21.19); t h e ability, f o r tropical species a t least, t o
survive low w i n t e r t e m p e r a t u r e .
In c o n t r a s t , M a r t i n & G u r r e a (1990) f o u n d a
significant p e n i n s u l a effect in I b e r i a n butterflies,
442
ASTYGISA
9170
m
12695
metaspila
vexillaria
12349
14024
14060
furva ? C
14026
14016
14020
12376
14025
circulara
O 00
500
Fig. 21.18. Representation of species o f the e n n o m i n a e geometrici genus Astygisa at different altitudes in B o r n e o
and Sulawesi. The cladogram illustrates relationships within a swarm of interrelated species in Sulawesi and their
probable sister-relationship to A. vexillaria (India t o B o r n e o and Java). Species 9170 and 12695 (genitalia slide
numbers) have subsequently been described as A. waterstradti
Sulawesi is indicated at the right. F r o m H o l l o w a y (1991).
diversity correlating more strongly with distance
f r o m the Pyrenees than with ecological factors
such as topography, soil and climate. African
contribution to the f a u n a via the straits of Gi-
braltar is very small relative to the European
contribution. This peninsula effect in fact ex-
tends over the whole of Europe and is exhibited
particularly strongly by the most widespread
faunal element, the extent unit, in the analysis of
Dennis et al. (1991: 16).
The equilibrium theory model not only pre-
dicts that there is a relationship between number
of species (which remains constant within nar-
row bounds) and area, but that the dynamic pro-
cesses of immigration and extinction be manifest
in a turnover of species on each island (Gilbert
1980; S. E. Miller 1984). Miller f o u n d no clear
indications of turnover in the butterfly f a u n a s of
the California Channel Is. O n the Cocos-Keeling
Is. in the Indian Ocean (Holloway 1983 c), seven
species recorded in the earliest survey ( 1 9 0 5 - 6 )
were not taken in more recent surveys (1952,
1964, 1978), and the former did not include
seven recorded in the latter, though again the
evidence is not unequivocal. During twelve
years' intensive light-trap recording of Lepidop-
tera on Norfolk Is. (Holloway 1977, 1982 b,
1984 d) extensive immigration was noted but
Jeremy D . H o l l o w a y & Ebbe S. Nielsen
Borneo
C
C
'
a>
C,S*
C,S*
=3
CO
, C
, C
Sulawesi
o o
1000 1500 2000
Altitude in metres
and A. stueningi respectively. Localization within
none of the species involved established for more
than a generation, nor did any of the resident
f a u n a go extinct.
Amongst the Norfolk I. butterflies, Holloway
(1977) reviewed literature on the occurrence of
Belenois java, a pierine Capparis-feeder, b o t h on
Norfolk I. and on Lord Howe I. Capparis occurs
on the former but not on the latter. The New
Caledonian race, peristhene, was recorded on
both during the 19th century but then was ap-
parently absent until it reestablished on Norfolk
in the early 1970s. It did not displace the en-
demic Cepora perimale subspecies that also feeds
on the Capparis, and the two taxa thrive and co-
exist together at the present (1996), possibly be-
cause perimale oviposits singly and java in egg
masses. Indeed, the situation has been further
complicated by the apparent introgression of
Australian B. j. teutonia characteristics, presuma-
bly arising f r o m immigrants of this race (as
noted on Lord Howe in 1969), such that now
a range of intermediates between peristhene and
teutonia occurs on the island (Holloway, unpub-
lished observations, February 1992, and material
f r o m the subsequent season reared by Mrs. M.
Jowett).
Bush & Whittaker (1991) have used Lepidop-
tera d a t a f r o m K r a k a t a u to question the validity
Biogeography of the Lepidoptera 443

Y/\ Temperate species

140 f ~ l Tropical species

Total species density

Y / \ Temperate species
IH Tropical species

Total species density

I,,.,,.,,.,.,,,,,.,,.
3 4 5 6 7

Y / \ Temperate species

Tropical species

-- Total species density

2 3 4 5 6 7 8 9 10

peninsular subdivisions

Fig. 21.19. Peninsula effects in tropical v. temperate components of butterfly faunas in Baja California (A) and
Florida (B) with data for woody plants for the latter (C) as discussed in the text. From J. W. Brown (1987) and
Brown & Opler (1990).

of the equilibrium model when applied to com-
plex forest communities in the lowland tropics.
The millenial time scale they suggested from con-
sideration of generation times of the dominant
organisms (trees) to be necessary for attainment
of equilibrium was probably much greater than
the periodicity of natural disturbance cycles in
such environments and certainly greater than the
eruptive periodicity of Krakatau. Most compo-
nents of the fauna show dependence of some sort
on the vegetation, hence fluctuations in their di-
versity are deterministic rather than stochastic
(see also Whittaker 1993). This is particularly
true for phytophagous animals exhibiting host
specificity such as butterflies.
The majority of butterfly taxa that have colo-
nized the Krakatau group since the 1883 erup-
tion are mostly geographically very widespread,
444 Jeremy D. Holloway & Ebbe S. Nielsen

MELANARGIA

C. S E . ASIAN OUTGROUP TAXA PALAEARCTIC

occitanica IBERIA, . AFRICA, S. FRANCE

tz pherusa
ines
arge
SICILY
IBERIA, . AFRICA
ITALY
IBERIA

ITALY
larissa BALKANS, TURKEY
hylata TURKEY - C. ASIA ASIA MINOR
syiaca . SYRIA
lachesis IBERIA W. MEDITERRANEAN
galathea S. & C. EUROPE & CAUCASUS
C. & E. MEDITERRANEAN
russiae ITALY - IRAN & C. ASIA

ZERYNTHIINI (part)
Zerynthia rumina W. MEDITERRANEAN
. polyxena C. & E. MEDITERRANEAN
Allancastria spp. ASIA MINOR
Sericinus spp. CHINA, KOREA (PALAEARTIC)

Fig. 21.20. A biogeographic hypothesis (Oosterbroek & Arntzen, 1992) of general area relationships in the Medi-
terranean area (right) is compared with a phylogenetic hypothesis (Mensi et al. 1992) for the genus Melanargia
based on electrophoretic data. The position of M. russiae, excluded from the analysis, is speculative, based on
comments in Mensi et al. A major component of the general area cladogram is also represented in a lineage of
the Zerynthiini (bottom, partly after Hancock (1983)).

and many are known to be migratory. However,
there are many butterfly species in this ecological
category that have failed to establish, probably
because their plant hosts have also failed to colo-
nize.
Holloway (1977, 1987 a, in press) has also con-
sidered stochastic and deterministic components
in the development of the moth fauna of Norfolk
I. It has already been mentioned that the expo-
nential model of dispersal of MacArthur & Wil-
son had been assessed in this context, propor-
tions of taxa with affinities to Australia, New
Zealand and New Caledonia in groups of vary-
ing dispersal powers appearing to fulfil the pre-
dictions of the model. But the proportions of
macro-lepidoptera families present in the resis-
tent fauna of the island do not reflect particu-
larly closely to proportions of these families in
the migrant influx of non-resistent taxa recorded
over the twelve years of the survey. Many of the
migrants that do not establish are dependent on
host plants absent from Norfolk, and the mi-
grants generally show higher host specificity
than the resident fauna. But the determinism of
establishment does not completely mask the
stochasticity of arrival.
Lepidoptera and Earth History
The contribution of Lepidoptera studies to inves-
tigation of earth history, often stated as the pri-
mary goal of cladistic biogeographers, is grow-
ing. The premise of Croizat (1964), reiterated in
cladistic biogeographic methodology (e. g.
Humphries & Parenti 1986), that congruent dis-
tribution patterns common to a diverse range of
plant and animal groups may provide the strong-
est pointers to events in earth history should not
be ignored, but for systematists specializing in a
single group it may be difficult to live up to. Any
contribution to the data set of modern cladistic
analyses is welcome, but those featuring groups
with a high proportion of endemic species, yet
distributed over a wide geographical range, are
probably the most useful.
An example is given by Oosterbroek & Arnt-
zen (1992) where a phylogenetic hypothesis for
the Procris statices group (Tarmann 1979) is
combined with others for groups of crane flies,
scorpions and amphibia to generate a general
area statement for the Mediterranean. This is
compared with hypotheses of the tectonic history
of the region from the Oligocene and found to
show correlation with the sequence of events
postulated. Dating of some of the biological lin-
eages using biochemical and molecular data gen-
erally supports the correlation. The authors are
now seeking further groups, particularly from
the crane flies, beetles and reptiles, to add to
their data set. An electrophoretically derived hy-
pothesis of relationships amongst Mediterranean
Melanargia taxa (Mensi et al. 1992) shows some
consistency with the general area cladogram of
Oosterbroek & Arntzen (Fig. 21.20), but possibly
with the cladogenetic events occurring over a
Biogeography of the Lepidoptera
much more recent time sequence. Relationships
within the Zerynthiini also show some consis-
tency, but the Italy and Iberia terms are lacking.
The model of Oosterbroek & Arntzen should be
compared with that of Dennis et al. (1991,
1995 a, b), discussed earlier, suggesting a much
more rapid and recent turnover in speciation.
The criticism by Humphries (1989) and Bush
(1994) of glacial refuge hypotheses as applied to
Neotropical biogeography (p. 438) should be
borne in mind here also. Bush has suggested that
the observed centres of endemism are in areas
with a history of maximal disturbance rather
than stability.
Tropical archipelagos provide the severest test
of biogeographic method in unravelling earth
history. Miller & Miller (1989) have attempted to
segregate the effects of dispersal and geological
vicariance in the development of biogeographic
patterns in West Indian butterflies but without
the hard cladistic information used by Liebherr
(1988) to relate carabid beetle data to alternative
dispersal and geological vicariance hypotheses
within the Caribbean: the data showed a better
fit to the latter. Vane-Wright, Ackery & Turner
(1992) and Brower et al. (1992) consider the da-
naine genus Anetia in this context, and Johnson
(1991) discusses two lycaenid genera that indi-
cate a better fit with the vicariance hypothesis.
Geological data on the history of the Caribbean
are an essential adjunct to such an approach.
Michaux (1991) advised biogeographers at-
tempting to relate geographic distributions to
tectonic events to include in their studies thor-
ough discussions of pertinent geological evi-
dence, but Cranston & Naumann (1992) have
admonished biogeographers not to be con-
strainted by current dogma of earth history (see
also Hall & Holloway, in press).
In the Indo-Australian tropics also, biogeo-
graphic data have been used more in assessment
of current geological hypotheses than to develop
hypotheses anew. Caribbean hypotheses suggest
the major islands may be fragments detached
from what is now Central America, hence con-
gruent vicariance patterns might be expected in
their biota. The geology of the Indo-Australian
archipelago primarily involves emergence of is-
land arcs from a marine environment with subse-
quent convergence and coalescence with each
other and with continental fragments at, or de-
tached from, the Australian margin (Holloway
1979; Burrett et al. 1991). For such islands to
acquire biotas requires a significant amount of
biological mobilism. Yet even there information
on past geography many be provided in the form
of biogeographic snapshots produced by what
Duffels (1986) termed dispersive vicariance (see
also Holloway 1982 a): the spread of an ancestral
organism through an archipelago followed by
speciation, the patterns of which will include in-
formation on the geography of the area soon af-
445
ter the dispersal event. The cicadas studied by
Duffels are particularly suitable for such studies,
and both they and the Lepidoptera have thrown
light on a number of questions about Indo-Aus-
tralian past geography. The problem, as always,
is how to segregate the influence of dispersal and
vicariance (earth history) events in current distri-
bution patterns, particularly in one of the more
mobile insect groups such as the Lepidoptera
(e. g. Page, 1989; Holloway 1992).
Sulawesi has long been suggested to be geolog-
ically composite, with eastern Australasian and
western Oriental components, but the interaction
of these fragments with neighbouring land areas
and their relative contributions to the Sulawesi
biota, indeed their subaerial history, have been
unclear. Data from Lepidoptera suggest no
strong relationships with Borneo, despite its
modern proximity, and some interaction with the
Philippines to the north and the Lesser Sundas to
the south. Relationships with Australasian biotas
appear to be relatively recent through the S. Mo-
luccas, suggesting the Australasian geological
component was submerged until fusion and thus
contributed no early biotic elements to Sulawesi
(Holloway & Jardine 1968; Holloway 1973,
1987b, 1991, 1997; Kitching et al. 1987; Duffels
1990; Vane-Wright 1991).
New Guinea is also geologically composite,
formed by the fusion of a complex of Melanesian
archipelagos at the approach of the northing
Australian continent; the outer archipelagos ex-
tend eastwards as the Bismarcks, Solomons, Va-
nuatu and Fiji. This composite nature is reflected
most strongly in the biogeography of cicadas
(Duffels 1986; Duffels & de Boer 1990; de Boer
1995). Some Lepidoptera patterns (Fig. 21.21)
also show consistency (Holloway 1979, 1984 c)
though patterns in the heteropteran family Miri-
dae do not (Schuh & Stonedahl 1986). See also
Polhemus (1996).
The changing position of archipelagos in the
S. W. Pacific, particularly Vanuatu relative to
Fiji and New Caledonia, and the role of small
islands such as Rotuma as a link from Vanuatu
to Samoa and the Pacific has also left an imprint
in biogeographic pattern (Robinson 1975; Hollo-
way 1979, 1983 b; Duffels 1988).
Major events such as the dispersal of Gondwa-
naland will have left their trace in modern Lepi-
doptera geography, particularly amongst the
more primitive groups (see Cranston & Nau-
mann (1992) for a general review). Grehan
(1991) and the next section review pattern in
these groups, and more localized studies are pro-
viding indications of complex patterns, such as
the groupings of Sabatinca discussed by Gibbs
(1983, 1989) that occur in Australia, New
Zealand and New Caledonia: different lineages
have different biogeographic patterns that may
be indicative of a composite geological origin for
some of the lands involved, a concept considered
446 Jeremy D. Holloway & Ebbe S. Nielsen

Fig. 21.21. Distribution of species in the Melanesian complex of the genus Cyana suggested by Holloway (1984c)
to represent two-arc sister-groups. Above, the Outer Arc group: A, croceizona; B, thoracica; C, bicolor; D, nigro-
marginata; E, basiflava\ F, scintillons, G, miles', H, securizonis; I, guizonis; J, omissa. Below, the Inner Arc group:
K, tricolor, L,fulvia; M, postdivisa; , metamelas; O, dinawa, , meyricki; Q, undescr. sp.
Biogeography of the Lepidoptera
for the New Zealand Lepidoptera more generally
by Dugdale (1989).
Some ditrysian groups have been identified as
largely Gondwanan in their distributions, such
as the Chlidanotini (e. g. Tuck 1981; J. W. Brown
1990) and some Tortricinae (Horak 1984). The
geometrid tribe Trichopterygini might also yield
interesting biogeographic pattern (Dugdale 1980;
Holloway 1986).
Amphipolar patterns such as in the archiea-
rine geometrids would repay further study
(McQuillan 1986). McQuillan also drew atten-
tion to the related arctiine genera Phaos and
Metacrias in S. E. Australia and New Zealand
respectively. These each contain three essentially
montane species that have close relationships
with S. American taxa, the Patagonian Phragma-
tobia thursbyi and the high Andean P. flavata,
and perhaps other Andean taxa (D. Ferguson,
in litt.). Ferguson notes that all have extremely
brachypterous females, making dispersal by
flight unlikely. He suggests the group is related
to the north temperate Phragmatobia and Ocno-
gyna group, with the montane South African
P. parvula also involved. The enigmatic presence
of Lycaena in New Zealand, only distantly re-
lated to the Lycaenini of montane New Guinea,
and the affinities of the New Zealand satyrines
also present major challenges to biogeographers
(Gibbs 1980). Holloway (in press c) associated
the Palaearctic and montane tropical larentiine
taxon Gymnodisca (= Rhinoprora) as a subgenus
of the New Zealand Pasiphila. Dugdale (1989)
has reviewed the relationships of the New
Zealand Lepidoptera in relation to current pan-
biogeographic hypotheses of the composite de-
velopment of the New Zealand biota through the
coalescence of geological terranes.
Biogeography in Relation to Host
Plants
Coincidence of distribution pattern in ecologi-
cally disparate taxonomic groups has already
been mentioned as likely to provide the strongest
evidence for past episodes of earth history. It
might therefore be logical, in specialist plant-
feeding animals such as Lepidoptera where a de-
gree of coevolution is likely, to except a more
precise biogeographic relationship between the
systematics of animal and host taxa.
In the next section examples are given (Agath-
iphagidae, Heterobathmiidae) where primitive
Lepidoptera groups appear to have failed to
track the dispersal of their host plant. The one
tentative attempt to relate lepidopteran (Hetero-
bathmia) taxonomy to host (Nothofagus) taxon-
omy did not result in clear congruence of biogeo-
graphic pattern (Humphries et al. 1986).
Wider considerations of the interrelationships
of plant diversity and lepidopteran herbivore di-
447
versity may present clues to why precise relation-
ships are uncommon. Vane-Wright (1978) sug-
gested, from observations on the frequency of
lepidopteran exploitation of plant families, that
this was preferential towards the larger, more di-
verse plant families. These present greater 'target
areas' for host-shift. He suggested plant diversifi-
cation may precede herbivore occupation rather
than being driven by it in a strict coevolutionary
fashion. This may be particularly true for the
Dipterocarpaceae in S. E. Asia, a diverse, domi-
nant family where the Lepidoptera herbivore
load is low and primarily non-specialist (Hollo-
way 1989). The relationship of papilionid diver-
sity to larval host plant diversity over latitudinal
gradients is discussed in the final section.
It is currently the case that localized higher
plant taxa of low and sometimes moderate, di-
versity lack specialist lepidopteran herbivores
(Cottrell 1978, 1985; Ackery 1991). Cottrell, with
reference to this phenomenon in the relationship
of Cape-centred butterfly groups to the diverse
and highly endemic Cape flora of southern
Africa, suggested that much of this flora is pro-
tected from butterfly exploitation in its adaption
to oligotrophic soils in the form of sclerophyll
foliage with low nitrogen content. A similar situ-
ation occurs in New Caledonia where vegetation
associations of high endemism, antiquity and di-
versity grow on ultrabasic rock types and sup-
port minimal lepidopteran diversity. Here an ad-
ditional factor is the accumulation by some of
the plants of the heavy metals of the ultrabasic
soils in their sap (Holloway 1979, 1989, 1993 a).
Hence there may be many process factors that
have acted through time against the development
and preservation of precisely congruent Lepi-
doptera/plant biogeographic patterns, apart
from the factor of differential dispersal powers
of insect and host taxa.
Gross Pattern in Higher Taxa
Non-Glossatans and Non-Heteroneuran Glossata
The non-glossatans and the Glossata basal to the
Heteroneura exhibit a high proportion of dis-
junct and localized distribution patterns. Most of
these patterns and those of the more widespread
families such as Eriocraniidae, Hepialidae and
Micropterigidae are restricted to, or show more
species diversity in, temperate latitudes (Fig.
21.22).
The order as a whole shows a strong ecologi-
cal relationship with the angiosperms ( 1 - 2 0 )
that may well date back to the Jurassic (Grehan
1991). While larvae of the two most primitive
suborders feed primarily on liverworts (Micro-
pterigidae) or gymnosperms (Agathiphagidae),
adults of at least the former visit the flowers of
angiosperms for resources (Micropterix), though
448 Jeremy D. Holloway & Ebbe S. Nielsen

^ Micropterix S] Agathiphagidae
^ O t h e r Micropterigidae H Heterobathmiidae

^ Eriocraniidae O Lophocoronidae
Acanthopteroctetidae Neopseustidae

ES Neotheoridae
^Mnesarchaeidae ] Anomosetidae
^ Palaeosetidae HD] Prototheoridae

Fig. 21.22

there is some suggestion of association with ferns
in the Sabatinca lineage. In both instances adult
feeding is on the spores or their angiosperm
equivalent pollen (Nielsen 1985).
The current distributions of these more primi-
tive lepidopteran groups may therefore have re-
sulted from a combination of two processes: the
tectonic and possibly associated climatic changes
that have occurred from the Jurassic to the pre-
sent day, affecting both angiosperms and their
dependent fauna alike; competitive interaction
with more modern groups, particularly in the
more continuous continental land masses. Both
processes could have contributed to the general
Biogeography of the Lepidoptera
Fig. 21.22. Generalized distribution of some groups of non-ditrysian ('primitive') Lepidoptera.
temperate latitude diversity exhibited by the
groups as a whole, though biogeographers will
no doubt continue to dispute the importance of
each process. Grehan (1991) has presented an as-
sessment of Lepidoptera geography, collated
with Croizat's panbiogeographic synthesis of an-
giosperm evolution and geography. He stresses
the importance of the Pacific Basin.
As our concepts of the interrelationships of
the Lepidoptera are refined and our knowledge
of their biology and interdependence with other
organisms increases, the historical biogeographic
picture may become clearer.
Micropterigidae. The Micropterix lineage of the
single family, Micropterigidae, is purely Pa-
laearctic, but the Sabatinca lineage has a local-
ized but more widespread and disjunct distribu-
tion (mostly round the Pacific rim) in eastern
and western N. America, Ecuador, temperate S.
America, southern Africa, Madagascar extreme
eastern Asia (mainly Japan) and Australia (E.
Australia, New Caledonia, New Zealand). The
449
Australasian taxa and their affinities have been
studied by Gibbs (1983, 1989), and the distribu-
tion of the family as a whole has been mapped
by Grehan (1991). The S. American Sabatinca-
group genera may well be most closely related to
Australasian taxa (Nielsen 1985).
Agathiphagidae. Only two species are included,
distributed in Queensland, Vanuatu, the south-
ern Solomons and Fiji, feeding as larvae in the
seeds in the cones of Agathis. The distribution of
the group is very much more restricted than that
of its host genus which ranges from Malaysia to
Fiji and New Zealand. Agathiphaga has not yet
been positively recorded from New Caledonia,
the centre of species richness of Agathis.
Heterobathmiidae. The single genus is restricted
to temperate S. America where the larva mines
leaves of Nothofagus (Kristensen & Nielsen 1983;
Nielsen 1985). Heterobathmia is unknown from
South Patagonia and Tierra del Fuego and, as
with Agathiphaga, the distribution is very much
more restricted than that of its host genus which
is also widely represented in Australasia.
450
Eriocraniidae are a small holarctic and widely
distributed family (Davis 1978).
Acanthopteroctetidae occur in western North
America, Crimea and Peru (D. R. Davis & O.
Karsholt, pers. comm.); the phylogeny of the
taxa is not resolved.
Lophocoronidae occur across southern temperate
Australia in mallee and dry sclerophyll wood-
lands (Nielsen & Kristensen, in press).
Neopseustidae. This small family is known only
from subtropical eastern Asia and from Chile
and Argentina (Davis 1975; Davis & Nielsen
1980, 1985), an unusual distribution that Grehan
(1991) notes to have a very strong resemblance
to that of the angiosperm family Lardizabala-
ceae. The biology of the immature stages is un-
known.
Mnesarchaeoidea. The single small exoporian
family Mnesarchaeidae is restricted to New
Zealand. It is regarded as the sister group to the
Hepialoidea (Kristensen 1984). Gibbs (1989) has
proposed a phylogeny for the fourteen species,
discussed possible external relationships, and
mapped their distribution within New Zealand.
Both species richness and taxic diversity in-
creases towards the north in New Zealand.
Hepialoidea. The major family, Hepialidae, is
found virtually worldwide, but is particularly di-
verse in temperate latitudes. The superfamily in-
cludes eleven other lineages (Nielsen & Scoble
1986), many of which have more restricted distri-
butions. These include the Prototheoridae
(southern Africa), Neotheoridae (S. Brazil), Ano-
mosetidae (Australia: S. Queensland and N. New
South Wales), and Palaeosetidae (Colombia, As-
sam, Burma, Thailand and Taiwan, Queensland),
and are mapped by Grehan (1991). The so-called
primitive Hepialidae also occur in the temperate
latitudes or elevated altitudes in the northern
(Gazoryctra) and southern Hemisphere (Fraus,
Antihepialus, Afrotheora).
Heteroneura Excluding Ditrysia
The trend of temperate diversity and hemisphere
localization continues with the non-ditrysian
Heteroneura.
Nepticuloidea. The Nepticulidae include two sub-
families, one widespread but predominantly tem-
perate in diversity and the other restricted to
Australia (Scoble 1983). The second family in-
cluded, Opostegidae, is much smaller but also
widely distributed (Davis 1989), and is excep-
tional in its occurrence on Hawaii, the only rep-
resentation of a non-ditrysian group on a remote
oceanic archipelago.
Incurvarioidea. Of the six families included, the
Incurvariidae, Heliozelidae and Adelidae are
found more or less worldwide but the other fami-
lies are more restricted. The Prodoxidae are
mainly Holarctic but also have a single species
Jeremy D. Holloway & Ebbe S. Nielsen
in temperate S. America, New Zealand (Dugdale
1989) and South Africa. The Cecidosidae occur
in temperate S. America. The Crinopterygidae
are restricted to the Mediterranean.
Palaephatoidea. The small family Palaephatidae
is only known from Australia and temperate S.
America (Nielsen 1987).
Tischerioidea. The single family Tischeriidae is
predominantly Holarctic but extends to the trop-
ics and is absent from Australia and New
Zealand.
The Ditrysia
Only amongst the Ditrysia does one begin to find
higher taxa that are strongly or exclusively tropi-
cal in their diversity. Where diversification in
temperate latitudes occurs, it is often a derived
feature amongst the groups concerned, such as
in the zygaenine Zygaenidae, the trifine Noctui-
dae, the majority of the larentiine Geometridae
(perhaps excluding the Trichopterygini (Dugdale
1980; Holloway 1986, in press c)) the thyatirine
Drepanoidea and parnassiine Papilionidae and
other sub-groups in the Nymphalidae, Pieridae
and Lycaenidae.
Strong tropical diversity amongst the larger
ditrysian families is seen in Choreutidae, Castnii-
dae, Immidae, Thyrididae, Megalopygidae, Li-
macodidae, Lasiocampidae, Eupterotidae, Sa-
turniidae, Mimallonidae, Lymantriidae, Uranii-
dae and Callidulidae. In most other families di-
versity increases towards the tropics.
Exceptions to this tropical diversity are seen
in some of the smaller family group taxa, such
as the Douglasiidae (Holarctic, Australia), Och-
senheimeriinae (Palaearctic, Himalayan), the gel-
echioid families Elachistinae, Scythridinae, Col-
eophorinae (mainly Holarctic) and Pterolonchi-
nae (Mediterranean, N. America, S. Africa), Het-
erogynidae (W. Europe, S. Africa), Lemoniidae,
Endromidae and Mirinidae (Palaearctic), and
Axiidae (Mediterranean).
There are a number of family-group taxa of
interest that have ranges that are much more re-
stricted geographically within the tropics, such
as the Neotropical Arrhenophanidae, Dalceri-
dae, Megalopygidae, Oxytenidae, Cercophani-
dae, Apatelodidae, Mimallonidae and Hedyli-
dae, the Palaeotropical Eriocottidae, Eupteroti-
dae (a few questionable Neotropical taxa), Bom-
bycidae, Brahmaeidae (also temperate) and Cal-
lidulidae. The Hibrildinae and Apoprogoninae
are purely African, the Ratardinae and Epicopei-
idae are restricted to the Oriental tropics, and
the Carthaeidae, Cyclotornidae are endemic to
Australia and the Hypertrophinae and Antheli-
dae to Australia and New Guinea. A number of
families are much more diverse in, or more or
less restricted to the Oriental and Australasian
tropics: Hyblaeidae, Tineodidae (including Ox-
ychirotidae). The Castniidae show great diversity
Biogeography of the Lepidoptera
in the Neotropics, but there is a lineage endemic
to Australia and a few exceedingly rare taxa in
the Oriental tropics. The Agonoxeninae include
five Indo-Australian and Pacific species associ-
ated with coconut palms.
Biogeographic Aspects of Phylogenetic
Hypotheses for Two Family-Level
Butterfly Groups
In this final section we attempt to place a few of
the concepts discussed earlier within the context
of current best estimates of phylogeny within
two butterfly higher taxa, the subfamily Danai-
nae of the Nymphalidae, and the Papilionidae.
These are the only diverse higher taxa where
phylogenetic analyses have been performed ex-
tensively down to the level of species, and these
analyses are inevitably continually subject to
criticism, disagreement and further refinement.
Danainae
The trio of tribes in the Danainae has been ex-
tensively researched, with phylogenetic hypothe-
ses available down to the species level except for
the Ithomiini (Ackery & Vane-Wright 1984; Ack-
ery 1987; Kitching, Vane-Wright & Ackery 1987;
Vane-Wright, Schulz & Boppr 1992; Vane-
Wright, Ackery & Turner 1992; Kitching, Ack-
ery & Vane-Wright 1993; Schulz, Boppr &
Vane-Wright, in press). Some cladograms have
been reproduced by Hollo way (1984, 1991) with
geographical information added, and some are
shown similarly in Fig. 21.23. The subfamily is
considered a natural group but the interrelation-
ship of the tribes Danaini, Ithomiini and Teller-
vini within it is still unresolved.
All three groups are virtually restricted to the
tropics and subtropics, as are their larval host
families, the Asclepiadaceae, Apocynaceae and
Moraceae for the Danaini, Solanaceae and Apo-
cynaceae for the Ithomiini and Apocynaceae for
the Tellervini. The Danaini are found in all tropi-
cal regions but the Ithomiini are strictly Neo-
tropical and the Tellervini are found only in the
Australasian tropics, an uncommon restriction
for a lepidopteran higher taxon (the Hesperiidae:
Trapezitinae are more broadly Australasian as
are the Hypocystini in the Nymphalidae).
The Danaini show their greatest diversity in
the Indo-Australian tropics (see Fig. 21.7). Ack-
ery & Vane-Wright recognized two approxi-
mately equally diverse lineages. Strongly migra-
tory taxa occur in each. The nominate subspecies
of Danaus (Danaus) plexippus has already been
mentioned on p. 436, showing major range ex-
pansion to Europe and across the Pacific to Aus-
tralasia (Vane-Wright 1993 a) from N. America
as well as regular migratory behaviour in its N.
451
American home and in its colonization of Aus-
tralia. Danaus (Anosia) chrysippus has been ob-
served to disperse over long distances at both the
Mediterranean and Australasian extremes of its
range. Parantica sita is migratory in Japan. In
the other lineage, the Australasian member (cor-
ima) of the rather disjunctly distributed Indo-
Australian superspecies Euploea core (Vane-
Wright 1993 b) has been recorded as an adven-
tive on Cocos-Keeling I. in the Indian Ocean,
and on Norfolk I. and Lord Howe I. in the Tas-
man Sea.
This second lineage (currently under review
and possibly paraphyletic with regard to the first
lineage, with the New World group as the basal
branch (R. I. Vane-Wright, pers. comm. )) divides
further into New World and Indo-Australian
groups of genera and is surprisingly absent from
Africa and Madagascar. Two lineages of Euploea
are represented in the Indian Ocean islands, one
on the Seychelles, the other with a species dis-
tributed over Mauritius, Reunion and Rodriguez
Is. Euploea is also the most successful and speci-
ose danaine genus in terms of penetration of Pa-
cific archipelagos, but it has proved difficult to
derive a satisfactory phylogeny for it (Ackery &
Vane-Wright 1984).
Euploea is grouped with Idea and the New
Guinea endemic genus Protoploea. Idea is pri-
marily Oriental with much sympatry, but there is
a distinctive endemic on Sulawesi and a sequence
of allopatric species from Sundaland to the Phil-
ippines, Sulawesi, Moluccas and New Guinea
that has been interpreted (Kitching et al. 1987;
Vane-Wright 1990; see also Holloway 1991) as
arising through progressive spread eastwards
with speciation rather than through vicariance
from west to east of a widespread ancestral
taxon.
Vicariance is a plausible explanation for the
distribution of Anetia in the New World group
of the sister lineage to Euploea and allies (though
Brower et al. (1992) argue for a more basal posi-
tion in the cladogram for Anetia). The five spe-
cies (Fig. 21.23) are found in Central America
and the larger Caribbean islands. The distribu-
tion pattern is compatible with hypotheses for
vicariance through geological changes in the
Caribbean, but sympatry and cladistic ambiguity
(A.jaegeri is a paraspecies with no autapomor-
phy) render the group somewhat uninformative
(Vane-Wright, Ackery & Turner 1992). The sis-
ter-genus to Anetia, Lycorea, is represented by a
few much more widespread Neotropical species.
The lineage that includes Danaus also contains
two groups of genera. The first has the Indo-
Australian Parantica and Ideopsis in a sister-rela-
tionship to the African and Malagasy Amauris.
Vane-Wright, Schulz & Boppr (1992) and
Schulz et al. (1993) provide the most rigorous
phylogenetic hypothesis for Amauris. The major-
ity of taxa are, however, widely distributed in
452 Jeremy D. Holloway & Ebbe S. Nielsen

Parantica INDO-AUSTRALIAN TROPICS

C Ideopsis

Amauris

Tirumala
INDO-AUSTRALIAN TROPICS

AFRICA, MALAGASY

AFRICA

Tirumala AFRICA & ORIENTAL

Danaus AMERICAS

Salatura INDO-AUSTRALIAN

Anosia PAN-TROPICAL
DANAINI Tiradelphe SOLOMON IS.

Euploea SULAWESI

Euploea MAURITIUS, RODRIGUEZ, REUNION (1 species)

Euploea INDO-AUSTRALIAN TROPICS (diverse)

Euploea INDO-AUSTRALIAN & 1 species in SEYCHELLES

Idea ORIENTAL & INDO-AUSTRALIAN GROUPS

Protoploea NEW GUINEA

Lycorea NEOTROPICAL

Anetia C. AMERICA & CARIBBEAN

ITHOMIINI NEOTROPICAL

TELLERVINI AUSTRALASIAN TROPICS

thirza S. MEXICO - PANAMA

cubana CUBA
ANETIA jaegeri HISPANIOLA, JAMAICA

briarea CUBA & HISPANIOLA

pantheratus CUBA & HISPANIOLA

niavius WIDESPREAD, AFRICA

tartarea WIDESPREAD, AFRICA

AMAURIS
ellioti WIDESPREAD, CENTRAL & EASTERN AFRICA

comorana COMOROS

phoedon MAURITIUS

echeria WIDESPREAD, AFRICA

vashti CENTRAL AFRICAN RAIN FORESTS

crawshayi WIDESPREAD, AFRICA

hecate WIDESPREAD, AFRICA EXCEPT EAST & SOUTH

albimaculata EASTERN & SOUTHERN AFRICA

damocles WESTERN AFRICA

EAST AFRICA TO SOUTH AMERICA, COMOROS

MADAGASCAR, COMOROS

ZAIRE, ZAMBIA, ANGOLA

CENTRAL AFRICAN RAIN FORESTS

Fig. 21.23. Cladograms, with distributions added, for D a n a i n a e groups, the Central American and Caribbean
genus Anetia and the Afrotropical genus Amauris (from literature sources cited in the text).

tropical Africa, hence the cladogram is relatively
uninformative. It is interesting to note that,
whilst the Comoros (A. comorana) and Mauri-
tius (A. phoedon) endemics arise independently
from a relatively basal position in the cladogram,
the Madagascar species (A. nossima; also on
Mayotte in Comoros) is in the terminal group,
sister-species to A. ochlea, a species widespread
in E. Africa and also in the more westerly Com-
oros.
The cladogram presented by Ackery & Vane-
Wright for Parantica is detailed, with numerous
island endemics, some of which are on relatively
basal branches, e. g. P. weiskei in New Guinea,
Biogeography of the Lepidoptera
P. menadensis in Sulawesi and P. crowleyi in
Borneo, but in most instances occur in small
groups of largely allopatrically distributed spe-
cies with at least some indication of pattern repe-
tition. There are two small clades of species dis-
tributed over islands of the outer Melanesian ar-
chipelagos (see p. 445). One, a trio including
P. schenkii, falls within a clade adding three fur-
ther allopatric species (Sundaland; Philippines;
Sulawesi and Moluccas), with two mainland
Asian species on more basal branches. The se-
cond contains five species, extends to New Cale-
donia (P. puntila), but is more isolated within the
cladogram, having a sister-relationship to a ma-
jor Oriental clade that extends no further east
than Timor or Sulawesi.
This major Oriental clade contains two exam-
ples of speciation within an island (the P. toxopei
group of four in Sulawesi) or restricted to an ar-
chipelago (the P. phyle group of four in the Phil-
ippines) as well as the P. sita group of seven spe-
cies that are more widespread and sympatric
within the Oriental tropics. A trio of species re-
lated to the sita and phyle groups is distributed
in southern Sulawesi (Mt. Lompobattang) and
the Lesser Sunda Is., and the toxopei quartet in
Sulawesi belongs to a much larger group with
the other species distributed over the Banda Arc
Islands from Sumatra and Java to Timor.
Firm indications of repeated patterns in the
Indo-Australian tropics are, however, elusive. It
appears more a case of variations on many
themes. Only when an effective method of R-
mode pattern recognition is applied to a much
larger sample of cladograms that contain a high
level of endemism and allopatry, will it be pos-
sible to gauge whether statistically significant
pattern of area relationships is present (see Hol-
loway (in press a)). The phylogenetic hypotheses
of Paralitica and Idea discussed in this section
and of Graphium in the next contribute to that
sample.
In contrast, the sister-group to the Parantical
Amauris group contains a high proportion of
geographically very widespread, dispersive spe-
cies in Tirumala and Danaus joined, paradoxi-
cally, by Tiradelphe, a monobasic genus only
known from a mountain-top on a single island
of the Solomons group. Subgenus Anosia of Da-
naus is pan-tropical, with an intriguing trans-At-
lantic relationship that unites the Old World
D. chrysippus with a number of Neotropical spe-
cies (Ackery & Vane-Wright 1984).
Papilionidae
Phylogenetic hypotheses for the family Papilioni-
dae as a whole, with some biogeographic discus-
sion, have been published by Hancock (1983)
and J. S. Miller (1987 a). The more recent analy-
ses in Tyler, Brown & Wilson (1994) and by
Brown et al. (1995) came to hand too late for
453
detailed consideration here, but they indicate al-
ternative relationships among the tribes of the
Papilioninae. We present in Fig. 21.24 the gene-
ral structure of Miller's cladogram, with geo-
graphical data added, and with the relationships
of the Parnassiinae suggested by Hancock added
on. This hypothesis for the Parnassiinae has
achieved more general acceptance than that of
Hiura (1980), but is not immune from criticism.
Huser (1993) indicates that there is no firm evi-
dence for the monophyly of the subfamily, and
that the genus Hypermnestra is misplaced within
it. The host specialism of Hypermnestra on Zy-
gophyllaceae is anomalous within the general
Aristolochiaceae specialism of all of the rest of
the subfamily (except Parnassius itself)
Hancock (1983) provided hypotheses for phy-
logeny (with extensive biogeographic interpreta-
tion) within the Papilionini, but these were criti-
cized by Miller (1987 a). Nevertheless, this phy-
logeny provides a basis for biogeographic discus-
sion. Modern phylogenetic hypotheses are also
available for the Neotropical genus Battus
(Racheli & Pariset, 1992) and for the nominate
subgenus of Graphium (Saigusa et al. 1982; see
also Holloway 1984 c).
Features of biogeographic interest in the ge-
neric-level cladogram of Fig. 21.24 include two
Gondwanan pairings between S. America and
Australasia (EurytideslProtographium in Graphi-
ini and EuryadeslCressida in Troidini). Some
early branches in the cladogram consist of
monotypic or only weakly speciose genera that
are geographically very restricted, e. g. the Baro-
niinae in Central America, Teinopalpus in the Hi-
malaya and Pharmacophagus in Madagascar.
Miller noted that a number of these occurred in
the N. E. Himalaya and S. E. Asia, each sister to
a morphologically rather uniform, speciose
clade.
Except for the Parnassiinae, all major lineages
are predominantly tropical. The Parnassiinae are
subtropical to temperate, found mostly in the
Oriental and Palaearctic Regions except for a
few Parnassius in N. America. The Graphiini,
with the exception of Eurytides, are exclusively
Old World, with only Graphium subgenus Arisbe
in the Afrotropics, and one predominantly Pa-
laearctic genus, Iphiclides. The Troidini are more
generally tropical, with a greater degree of allop-
atry within the generic groupings. Parsons (1906)
has suggested this has arisen through Gondwa-
nan vicariance. Speciation within Troides itself is
thought to have been allopatric, with successive
episodes of range expansion and fragmentation
within the Indo-Australian archipelago (Zeuner
1943, reviewed by Holloway 1973).
The non-cladistic phylogeny for the Papilio-
nini of Hancock (1983) recognizes two major lin-
eages, only one of which occurs in the tropical
New World. The American Pterourus group has
a sister-relationship to another American group
454 Jeremy D. Holloway & Ebbe S. Nielsen

BARONIINAE
Baronia C. AMERICA

Parnassius HOLARCTIC

Hypermnestra IRAN, AFGHANISTAN, USBEKISTAN, KIRGHIZIA

Archon PONTO-MEDITERRANEAN

Zerynthia MEDITERRANEAN

Allancastria PONTO-MEDITERRANEAN

Sericinus CHINA, KOREA

Luehdorlia CHINA, KOREA, JAPAN

Bhutanitls . . HIMALAYA

Eurytides AMERICAS

Protographium AUSTRALIA

Iphiclides PALAEARCTIC

Lamproptera ORIENTAL

Pazala HIMALAYA, TAIWAN

Graphium INDO-AUSTRALIAN

Pathysa INDO-AUSTRALIAN

Arisbe AFRICA, MADAGASCAR

Teinopalpus HIMALAYA

Paplllo COSMOPOLITAN

Meandrusa ORIENTAL

Battus AMERICAS

Pharmacophagus MADAGASCAR

Cresslda AUSTRALIA, NEW GUINEA

Euryades PATAGONIA

Losara ORIENTAL

Pachliopta INDO-AUSTRALIAN

Trogonoptera SUNDALAND

Troides ORIENTAL - NEW GUINEA

TROPICAL AUSTRALASIA

Atrophaneura ORIENTAL

Parides NEOTROPICAL

Panosmia ORIENTAL MAINLAND

Fig. 21.24. Full area cladogram for genera of Papilionidae compiled from Miller (1987), with Parnassiinae after
Hancock (1983). See also Hiura (1980) for alternative phylogeny of Parnassiinae, and Parsons (1996) for distinc-
tion of Ornithoptera from Troides and association with Pharmacophagus.

(Heraclides) plus an Oriental (Chilasa), Austral-
asian (Eleppone) sister-pair. The second lineage
has the Holarctic Papilio sister to a predomi-
nantly tropical Old World assemblage of groups,
Princeps. Princeps divides into E. Palaearctic (Si-
noprinceps), Indo-Australian (Menelaides), Afro-
tropical (Druryia) lineages and a more wide-
spread lineage (Princeps) that itself comprises
distinct Afrotropical and Indo-Australian lin-
eages.
The detailed phylogeny for Battus (Racheli &
Pariset 1992) was established using Euryades as
outgroup. Two main lineages were recognized,
one with many rather widespread taxa and much
allopatry, the other with most species much more
restricted and allopatric, including two taxa en-
demic to Caribbean islands. This second lineage
therefore provides more data for historical bio-
geographic analysis of the Caribbean (see
p. 445). The two endemics are among the more
basal branches of the lineage, and are suggested
by Racheli & Pariset to show concordance with
the general vicariance pattern for the region.
The analysis of subgenus Graphium by Saigusa
et al. (1982) suggested three lineages could be
recognized. One consists of a number of exten-
sively sympatric Oriental species and provides
limited information for historical biogeographic
analysis. The other two contain a much higher
proportion of localized, allopatric species, with
one distributed over the inner Melanesian archi-
pelagos (Moluccas, New Guinea, Australia, New
Biogeography of the Lepidoptera
north
Percent "Generalist" species per 10%
Fig. 21.25. Numbers of papilionid species decrease with latitude, and the percentage with "generalist" feeding
larvae increases. After Scriber (1984).
Caledonia), and the other distributed over the
outer Melanesian archipelagos, particularly the
Solomons. But these lineages do not bear a sis-
ter-relationship to each other, and each contains
one or more very widespread species. Hence they
do not provide such clear support for the geolo-
gical hypothesis of two convergent archipelagos
as do some cicada groups (see p. 445).
Unlike the Danainae groupings, the Papilioni-
dae are also moderately represented in northern
temperate latitudes. In most instances this has
been facilitated by a host plant shift away from
the families Aristolochiaceae, Rutaceae, Lau-
raceae and Annonaceae favoured by the tropical
papilionid fauna. Scriber (1984) and Miller
(1987 b) have surveyed papilionid diversity and
distribution in relation to host-plant specializa-
tion, and there are a number of chapters on papi-
lionid host-plant relationships in Scriber, Tsu-
baki & Lederhouse (1995). Scriber showed that
papilionid species richness declines from the
equator polewards, but there is a corresponding
increase in the proportion of species showing ge-
neralism in host-plant selection (Fig. 21.25).
There are even latitudinal bands in host-plant
specialism within single species, demonstrated
clearly for the N. American Papilio glaucus by
Scriber. The related glaucus and troilus groups
are most diverse in N. American temperate lati-
455
Total no.
of species
tudes, the larvae feeding generally on compo-
nents of the eastern broadleaved forest systems.
The other major group in Papilio to have suc-
cessfully established in temperate latitudes con-
sists of P. machaon and related species, most of
which feed on Umbelliferae rather than the trop-
ical Rutaceae favoured by the majority of Papi-
lio. The Umbelliferae are diverse in open habitats
of temperate latitudes. Similarly, in Graphiini,
the most temperate genus is Iphiclides with lar-
vae feeding on Rosaceae.
In the Parnassiinae the majority of genera are
found in Mediterranean climate, subtropical and
subtropical montane localities in the Palaearctic
and Oriental Regions (mapped by Holloway
(1974)). All except Parnassius and the dubiously
parnassiine Hypermnestra feed on Aristolochia-
ceae (Miller (1987 b)) suggests this specialism is
homoplasious relative to that of the Troidini).
The genus Parnassius has its centre of richness in
the mountain systems to the west of the Tibetan
Plateau along with a number of other speciose
Palaearctic genera such as Lycaena sensu lato
and Colias (Holloway 1974). For all these gen-
era, preadaptation to open habitats at altitude in
a region of mountain building on the fringes of
the tropics appears to have facilitated their suc-
cess, probably conjointly with that of their host-
plants, in extending into the vast areas of open,
456
temperate habitat developing in the northern
hemisphere as the climate cooled through the
Tertiary.
The m o s t widespread and successful species
group in Parnassius extending to the Nearctic,
has the widest range o f host-plants, including
families such as Crassulaceae, Saxifragaceae and
Caryphyllaceae, as well as the family favoured
by many o f the more restricted species groups,
Fumariaceae.
The more westerly parnassiine taxa, compris-
ing the Zerynthiini, s h o w concordance in their
systematics and area relationships with the gene-
ral pattern for Mediterranean areas discussed o n
p. 444 (Fig. 21.20).
Acknowledgements
We are grateful to the following w h o have helped
in various ways such as commenting o n earlier
drafts o f the text or providing information: Tim
Benton, D o u g l a s Ferguson, Christoph Huser,
Scott Miller, G a d e n Robinson, Gary Stonedahl,
and Dick Vane-Wright; Mr. G. Davies prepared
the figures and Phillipa H o l l o w a y key-boarded
the bulk o f the text and Ms. Judith Hull finalized
the editing o f the manuscript.
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 Index of Scientific Lepidoptera Names

Principal entries of suprageneric taxa recognized in the classification of this work are indexed by boldface page
numbers. Subspecific names are not indexed.

Abantiades, 62 Acronictini, 385, 386

Abantis, 273 Acropolitis xuthobapta, 202
Abegesta reluctalis, 248 Acropteris, 309
Abisara, 284 Acsala, 386
Abraxas grossulariata, 314 Acsalini, 386
Abraxini, 315 Actios, 338, 341
Abrostolini, 377-378 Actinotia, 369, 384
Acalyphes, 313, 314 Adaphaenura, 379
Acalyptris, 18, 69 Adela, 73, 75, 77, 408; A. caerulella, 82; A. reawnurella,
Acanthobrahmaea, 342, 343, 344 70; A. trigrapha, 66, 74, 76, 78
Acanthoplusia, 377 Adelidae, 31, 66, 70, 72, 73-75, 76, 78, 80, 81, 82, 84,
Acanthopsyche, 105, 107 404, 407, 408, 414, 450
Acanthopteroctetes, 52, 54; A. bifascia, 56; A. unifas- Adelinae, 71, 75
cia, 54 Adeloneivaia, 340; A. subangulata, 339
Acanthopteroctetidae, 11, 34, 51, 52, 53-54, 55, 56, Adelpha, 289; A. ixia, 277
404, 407,413,415, 448, 450 Adhemarius, 346
Acanthopteroctetoidea, 10, 11, 28, 29, 41, 53-54, 407 Adoxophyes, 207
Accra, 411 Adrapsa, 372
Acentria, 246; A. ephemerella, 248 Adscita, 174; A. subtristis, 171
Achaea, 374 Aegle, 382
Acharia, 167 Aenetus, 61
Acherontia, 347; A. lachesis, 347 Aeolanthes, 139; A. callidora, 137
Acherontiinae, 346 Aeolanthinae, 133, 134, 139
Achroia, 242 Aepytus, 62
Achyra, 250 Aethes, 206; A. smeathmanniana, 200
Acidaliini, 316 Aethiopsestini, 305
Acleris, 205, 209; A. rhombana, 200 Afrida, 389
Acontia delecta, 371 Afridinae, 359, 389
Acontiinae, 360, 367, 369, 370, 371, 375, 379, 381, 387 Afrotheora, 58, 61, 450
Acosmeryx, 348 Agalope, 174, 291
Acousmaticus, 189, 194 Agamopsyche threnodes, 160
Acraea, 278, 286, 288, 331; A horta, 266 Aganainae, 38, 227, 355, 359, 368, 369, 370, 374, 375,
Acraeinae, 288 376, 386, 387, 392, 393
Acraeini, 279, 288, 413 Agaristidae, 174
Acraga, 169 Agaristinae, 184, 360, 367, 368, 370, 379, 380, 384,
Acria, 146 385, 387
Acriinae, 157 Agathiphaga, 44, 45, 449; A. vitiensis, 45, 46
Acridotarsa, 95 Agathiphagidae, 34,41, 43-46, 404,415, 447,448,449
Acritocera, 195; A. negligens, 192 Agathiphagoidea, 9, 10, 11, 27, 28, 29, 41, 43-46, 405,
Acrobasis, 243 413, 414
Acrocercops, 114; A. astaurota, 96; A. brongniardella, Agathodes, 250
96 Agavenema, 11, 79; A. barberella, 66
Acrocteninae, 362 Agdistinae, 224
Acrojana, 330 Agdistis, 218, 224; A. bennettii, 221
Acrolepia, 119, 124 Agdistopis, 223, 224
Acrolepiidae, 35, 119, 120, 121, 123, 124,125, 405, 409 Agais, 288; A. karaganica, 19; A. urticae, 19
Acrolepiopsis, 121, 125; A. assectella, 123, 125 Aglaope, 170, 174; A. infausta, 171
Acrolophidae, 18, 91, 92, 93, 98, 101, 102, 103, 106, Aglaopus, 229
405, 409 Aglia, 341; A. tau, 335
Acrolophus, 102, 103; A. arboreus, 106; A. arcanellus, Agliinae, 337, 338, 340, 341
103; A. pholeter, 101, 103; A. popeanellus, 93, 98, 101 Aglossa, 243
Acronicta, 366, 367, 370, 372, 385, 391; A. alni, 370; A. Aglossata, 27, 405, 415
americana, 370; A. brumosa, 370; A. carbonaria, 370; Agnidra, 305
A. hasta, 383; A. impleta, 370; A. impressa, 370; A. Agonoplerix 138; A. umbellana, 137
leporina, 370; A. lobeliae, 368; A. ovata, 370, 387; A, Agonoxena, 139
rumicis, 370 Agonoxenidae, 405, 410
Acronictinae, 360, 366, 367, 368, 369, 370-372, 381, Agonoxeninae, 35, 133, 134, 139, 155, 156, 157, 451
382, 383, 386, 387 Agrias, 289
464 Index of Scientific Lepidoptera Names

Agrionympha, 43 Amyna, 379, 380

Agriothera, 110; A. elaeocarpophaga, 110; A. issikii, Anacampsis, 136
96, 110 Anacampsidae, 158
Agriphila, 244; A. vulgivagella, 245 Anacampsinae, 158
Agrius, 345, 347, 435 Anacronicta, 385, 441
Agrotini, 384 Anacrusis, 199, 208; A. stapiana, 200
Agrotis, 384, 435; A. infusa, 385, 437; A. ipsilon, 384; Anadasmus, 136
A. maleflda, 390; A. munda, 385; A. segetum, 384 Anadiasa, 329
Agrumenia, 174 Anaea, 289
Aididae, 159, 164, 165-166 Anaeini, 289
Aldus amanda, 166 Anagasta kuehniella, 243
Akbesia davidi, 343 Anagogini, 315
Akesina, 164 Anagrapha falcifera, 368, 378
Alabonia geoffrella, 137 Anania, 250
Alaena, 284 Anaphe, 358, 362; A. panda, 387; A. reticulata, 364
Alatuncusiinae, 234, 249 Anapisa, 392
Alcides, 307, 308, 310 Anarmodia, 250
Aleimma, 205 Anarsia, 136, 147
Alheita, 323, 324 Anarsiidae, 158
Allancastria, 454 Anartia, 288
Alloclemensia, 81, 83 Anathamna, 200
Allocosmia, 380 Anathyrsa, 144
Allotinus, 285 Anchimompha, 144
Almeidaia, 340 Ancylis, 212
Almeidaiini, 340 Ancyluris, 284
Aloeides, 285 Andraca, 332, 333, 334; A. bipunctata, 347
Alophogaster, 174 Androlymnia, 380
Alsophila, 315; A. aescularia, 315; A. pometaria, 315 Anetia, 292, 445, 451, 452; A. briarea, 452; A. cubana,
Alsophilinae, 315 452; A. jaegeri, 452; A. pentheratus, 452; A. thirza,
Alucita, 1, 218, 223; A. coffeina, 223; A. hexadactyla, 452
221 Angeronini, 315
Alucitidae, 12, 37, 217, 221, 222-223, 233 Aniclini, 384
Alucitoidea, 10, 12, 13, 14, 28, 30, 219, 222-223, Anigraea, 369, 377
234, 406 Anisodes, 316; A. rufulata, 311
Alypia octomaculata, 385 Anisota, 338, 340
Amana, 302 Anisynta, 273
Amastus, 359, 392 Anomiini, 374375
Amala leucerythra, 373; A. sperbius, 392 Anomis, 375, 380, 424, 426, 433, 435
Amathusia, 290; A. phidippus, 268 Anomoeotes, 164
Amathusiini, 290, 413 Anomoeotidae, 159, 160, 161, 163-164, 175
Amauris, 286, 292, 451, 452, 453; A. albimaculata, 452; Anomoeotis felderi, 175
A. comorana, 452; A. crawshayi, 452; A. damocles, Anomologidae, 158
452; A. dannfelti, 452; A. echeria, 452; A. ellioti, 452; Anomoses hylecoetes, 56, 60
A. hecate, 452; A. inferna, 452; A. niavius, 452; A. Anomosetidae, 56, 6 0 - 6 1 , 448, 450
nossima, 452; A. ochlea, 452; A. phoedon, 452; A. Anoncia, 147; A. brunneipes, 140
tartarea, 452; A. vashti, 452 Anopina, 208
Ambia, 246 Anosia, 451, 452, 453
Amblyscirtes, 274 Antaeotricha, 136
Ambulicinae, 346 Antequera, 147
Ambulycinae, 346 Antequerinae, 135, 136, 147, 155, 156, 157
Ambulyx, 346 Anteros, 284
Amerila, 391, 392 Anthela, 326; A. connexa, 328; A. ocellata, 325
Ametridini, 315 Anthelidae, 39, 321, 322, 324-326, 328, 450
Ametroptila, 305 Anthelinae, 325, 326
Amida, 107; A. quadrangularis, 106 Anthene, 285; A. emolus, 270
Amnosia, 287 Antheraea, 338, 341; A. larissa, 335
Amorbia, 208 Antheua, 363, 366; A. simplex, 357
Amphicallia, 394; A. bellatrix, 373 Anthocharis, 281
Amphiesmenoptera, 7, 9, 16 Anthocaris, 437
Amphipyra, 380, 381, 384; A. erebina, 371 Anticarsia, 374, 435
Amphipyrinae, 360, 369, 371, 381, 382, 384 Anticrates, 170
Amphisbatidae, 135, 136, 138, 146, 155, 156, 157 Antihepialus, 61, 449, 450
Amphisbatis, 146 Antirrhea, 287, 289, 290
Amphithera, 110 Antirrheini, 290
Amphitheridae, 409 Antispila, 73; A. nysaefoliella, 66, 85
Amphixystis, 94, 99 Antoloea, 146
Amselina, 146 Anuga, 377
Amydria, 103; A. effrentella, 93, 102, 103 Aon, 358, 374, 377
Index of Scientific Lepidoptera Names 465

Aoraia, 58, 61, 62 Agyreus, 288

Apameini, 369, 371, 384, 412 Aorista, 307
Apantesis, 392 Aretornis, 386
Apanthesis leuce, 19 Arguda, 329
Apatelodes, 332, 333, 334; A. singularis, 335 Argyria, 244
Apatelodidae, 321, 331, 450 Argyriini, 236
Apatelodinae, 38, 333, 334 Argyrogrammatini, 377378
Apatetrinae, 158, 157 Argyrophorus, 290
Apatetris, 136, 147 Argyrotypini, 229
Apatura, 289; A. iris, 266, 289 Arhopala, 285
Apaturinae, 267, 287, 288, 289, 290 Ariadne, 289, 290
Apethistis, 146 Aridomeria, 124
Aphelosetiidae, 157 Ariolicini, 389
Aphiaris, 211 Arisbe, 453, 454
Aphnaeinae, 285 Aristaea, 114
Aphnaeini, 279, 285 Aristotelia, 136, 147
Aphnaeus, 285 Aristoteliinae, 158
Aphomia, 242 Arniocera, 229; A. cyanoxantha, 228
Aplotinae, 157 Aroa, 373
Apoda, 167 Aroga, 147
Apoditrysia, 10, 13, 28, 127, 181, 188, 217, 405, 406, Arotrophora, 203, 206
408 Arotura, 139, 140
Apophatus, 83, 84, 86 Arragonia, 144
Apoplania, 54, 56 Arrhenophanes, 104; A. perspicilla, 93, 98, 101, 103, 106
Apoprogones, 307 Arrhenophanidae, 91, 92, 93, 98, 101, 103, 104, 106,
Apoprogoninae, 307, 450 450
Aporia, 302 Arsenura, 340
Apostibes, 140 Arsenurinae, 337, 338, 340
Apotomis, 211 Arsenurini, 340
Appias, 281; A. nero 279 Artace, 329
Aproaerema, 147 Artaxa, 386
Aprominta, 146; A. desigmatella, 138 Artifodina, 113
Apterona, 105, 107; A. helix, 106 Artogeia, 281
Aracimini, 315 Artona, 172, 174
Araschnia, 288 Asaphocrita, 144
Arbela, 193 Ascalapha, 374
Arbudas, 174 Ascalenia, 146
Archaeoaltacus, 340 Ascia, 281
Archaeopacha, 329; A. obsoleta, 328 Asemanophorae, 346
Archaeopachinae, 327, 328 Ashinagidae, 157
Archaeoses, 195 Aslauga, 285
Archeolepis, 28; A. mane, 16, 17 Asota, 359, 375, 393; A. plana, 387
Archepiolus, 56 Asterivora, 219
Archiearides, 313 Asteroeampa, 289
Archiearinae, 310, 311, 3 1 3 - 3 1 4 Asterope, 289
Archiearis, 313, 314; A. notha, 314; A. parthenias, 311 Asthenidia, 340; A. buckleyi, 335; A. diffissa, 339
Archipini, 199, 200, 201, 202, 203, 206, 207, 412 Asthenini, 316
Archips, 207; A. podana, 200 Asthenoptycha, 207
Archon, 454 Astraptes, 273
Arete, 374; A. coerula, 387 Astygisa, 442; A. stueningi, 442; A. vexillaria, 442; A.
Arctia, 392; A. festiva, 373 waterstradti, 442
Arctiidae, 19, 20, 38, 174, 292, 302, 356, 358, 359, 361, Asura, 386, 391
362, 366, 367, 370, 373, 375, 378, 380, 383, 386, 388, Asymphorodes, 147
389-394, 412, 429, 434, 435 Asymplecta, 100
Arctiinae, 356, 358, 359, 362, 366, 367, 369, 373, 378, Ategumia, 250
380, 383, 390, 391, 392-394, 431 A this inca, 185
Arctiini, 373, 390, 392, 393, 394 Atrichozancla, 143
Arctiites, 355 Atrophaneura, 279, 454
Areyophora, 389 Attacidae, 321
Areniscythris, 140 Attacini, 337, 341
Areticulata, 69 Attacus, 338, 340, 341; A. atlas, 338
Argina, 394 Atteria, 203, 208
Argynninae, 288 Atteriini, 199, 200, 201, 203, 208
Argynnis, 288; A. paphia, 266 Atteva, 122; A. punctella, 123
Argyractini, 247 Attevinae, 120, 122
Argyresthia, 121, 123, 410; A. brockella, 123 Atychia, 181
Argyresthiidae, 405, 409, 410, 414 Audre, 284
Argyresthiinae, 36, 120, 122-123, 410 Augasma, 144
466 Index of Scientific Lepidoptera Names

Augasmidae, 157 Belenois, 19, 279, 281; B. java, 442

Aulacodes, 247 Beralade, 329
Auliepterix, 17, 28 Bertholdia, 362, 367, 380; B. detracta, 378
Auratonota, 210 Bhutanitis, 19, 278, 454; lidderdalii, 264
Austrandesiini, 384 Bibasis, 272; B. sena, 271
Austrocossus, 193, 195 Biblidini, 288
Autochthonus, 100 Biblini, 289
Autographa, 378; A. nigrisigna, 378 Biblis, 282, 289; B. hyperia, 266
Automerina, 337, 338 Bicyclus, 286, 289, 290
Automeris, 242, 340 Bijugis, 107
Autoplusia egena, 378 Bipectilus, 61, 62
Autosticha, 146; A. pelodes, 148 Birthana, 227; . saturata, 228
Autostichidae, 134, 135, 138, 144-146, 157 Biselachista, 139
Autostichinae, 135,146, 155, 156, 157 Bissetia, 244
Auximobasis, 144 Biston, 310; B. betularia, 311, 314
Auzea, 308; A. rufifrontata, 306 Bistonini, 315
Auzeinae, 309 Bizone, 391
Axia, 258; A. margarita, 257, 258; A. napoleona, 257; Blastobasidae, 405, 410
A. nesiota, 260 Blastobasinae, 36, 134, 135, 144, 155, 156, 157
Axiidae, 257-258, 260, 450 Blastobasis, 144; B. glandulella, 137, 142, 145, 148
Axioidea, 10, 14, 28, 30, 2 5 7 - 2 5 8 Blastodacna, 139
Azaleodes, 86; A. micronipha, 67 Blastodacninae, 157
Azamora, 242; A. sororia, 241 Blenina angulipennis, 373
Azelinini, 315 Bleninae, 359, 373, 389
Azeniini, 381
Blepharomastix, 250
Azinidae, 156
Boarmiini, 310, 312, 315, 431
Azochis, 250
Bocana, 372
Azygophleps, 191, 193
Bohemannia, 69
Boisduvalodes, 165
Bac tra, 211; . verutana, 200 Boloria, 288
Bactrini, 200, 203, 210, 211 Bombycidae, 38, 39, 321, 322, 331-334, 335, 336,
Badamia exclamationis, 272 347, 450
Baeophylla, 97 Bombycinae, 332, 333, 334
Bagisara, 379
Bombycini, 332, 334
Bagisarinae, 360, 367, 369, 370, 371, 379-380, 385,
Bombycoidea, 10, 14, 19, 28, 193, 321, 322, 329-348,
387
412, 429, 434
Baileya, 389; B. doubledayi, 390; B. ophthalmica, 373
Bombycopsis, 329
Balacra, 358; B. rubrostriata, 373
Bombyx, 1, 3, 4, 334; B. huttoni, 332; . mandarina,
Balataea octomaculata, 171
334; . mori, 332, 333, 334
Baliochila, 284
Bomolocha bijugalis, 390; B. madefactalis, 387
Banisia, 229
Bondia, 225, 226; B. comonana, 226
Bankesia, 107
Borkhausenia, 141
Baptini, 315, 431
Borocera, 329; B. marginepunctata, 328
Barasa, 388
Bracca, 424, 426
Barbarus, 1
Barca, 141 Brachionycha, 381
Baronia, 265, 275, 277, 278, 454; B. brevicornis, 264, Brachmia, 147
278 Brachmiinae, 158
Baroniinae, 265, 278, 454 Brachodes, 181, 182; B. appendiculata, 182
Barygnathella, 205 Brachodidae, 13, 36, 181-182, 183, 191
Basilarchia, 289 Brachycodilla, 166
Basileura, 81, 83 Brachycyttarus, 107
Basilodes, 382; B. pepita, 387 Brodino, 250
Basonga, 235 Bradypodicola, 242
Bassaris, 432; B. gonerilla, 432, 433; B. ida, 433; . Bradypophila, 242
ilea, 432, 433 Brahmaea, 342, 344; B. wallichii, 343
Bathyphlebia, 340 Brahmaeidae, 38, 321, 322, 339, 341, 342-344, 450
Bathypluta, 209 Brassolinae, 287, 289
Batrachedra, 143; B. enormis, 140, 145 Brassolini, 290, 413
Batrachedridae, 135, 137, 143, 157, 405 Brassolis, 290; B. sophorae, 268
Batrachedrinae, 135, 143, 155, 156, 157 Brenthia, 219; B. albimaculana, 221
Battaristis, 147 Brephinae, 313
Battus, 277, 453, 454 Brithys, 382
Bebearia, 289 Bruandia, 107
Bedellia, 126, 127; B. somnulentella, 127 Bryophilinae, 360, 367, 368, 370, 372
Bedelliidae, 119, 120, 123, 124, 126, 127, 128 Bryotropha, 136, 147
Bedosia, 324 Bucculatricidae, 18, 35, 96, 106, 107, 108, 109,
Belemniini, 392 110-112, 405, 409, 414
Index of Scientific Lepidoptera Names 467

Bucculatrix, 106, 112, 410; . magnellalla, 96; 110; . Candalides, 285

solidaginiella, 108; . thurberiella, 111; . tridenti- Capillamentum, 382
cola, 96 Capnoptycha ipnitis, 203
Buckleria, 224 Caprona, 273
Bunaea, 341 Capys, 285
Bunaeini, 341 Carales, 392
Bunaeopsis, 341 Carea, 362, 370, 386; C. van>es, 390
Bupalus piniaria, 314 Careini, 389
Bursadella, 227 Cargida, 363
Butalidae, 157 Caridarctia, 382
Carmenta, 183
Caberini, 315 Carposina, 225, 226
Cacographis, 246; C. osteolalis, 246 Carposinidae, 37, 225, 226, 228, 406
Cacotherapia leucocope, 245 Carterocephalus, 269, 273
Cacotherapiini, 236 Carthaea saturnioides, 339, 341, 343
Cacozelia basiochrealis, 241 Carthaeidae, 39, 321, 322, 339, 341, 343, 450
Cactoblastis cactorum, 243 Carthara abrupta, 245
Cacyparis, 389 Caryocolum, 147
Cadmogenes, 119 Casana, 107
Cadra cautella, 243 Cassus, 290
Caenurgina, 374 Cassyma, 431
Caerois, 289 Castrila, 274, 282; C. eudesmia, 185, 186
Caw, 338 Castniidae, 13, 19, 20, 34, 35, 181, 184, 406, 450
Caladium, 246 Castniinae, 185, 187, 188, 450
Calamotropha, 244 Castniini, 185, 186, 188
Calephelis, 284 Castniomera atymnius, 186; C. humboldti, 186
Calidota, 358 Casuario catocalis, 245
Caligatus angasii, 368 Catacroptera cloanthe, 291
Caligo, 286, 290; C. fe/irao, 277 Catapterix, 54, 407
Caligula, 341 Catasticta, 279, 281
Calinaga, 278, 286, 287, 291; C. W i f a , 268 Catharia, 249; C. pyrenaealis, 248
Calinaginae, 267, 285, 287, 290, 291, 292 Cathariinae, 234, 240, 249
Calisto, 290 Catoblemma, 379
Calledema jocaste, 364 Catocala, 359, 374
Callerebia, 290 Catocalinae, 356, 358, 359, 360, 361, 368, 369, 370,
Callicore, 289 372, 374, 376, 380, 383, 384, 387
Callidrepana, 305 Catonephele, 286, 289
Callidula, 259, 274 Catopsilia, 281
Callidulidae, 37, 258-259, 260, 331, 450 Catopta, 192
Callidulinae, 258, 259 Catoptria, 244
Calliduloidea, 10, 14, 28, 30, 193, 257, 2 5 8 - 2 5 9 Cauchas, 73, 75, 408
Callimorpha, 392; C. dominula, 394 Caviria, 386
Callimorphini, 392, 393, 394 Cecidoses, 79, 80, 82; C. eremita, 66, 74, 76, 78, 85
Callionima, 348 Cecidosidae, 34, 66, 67, 70, 71, 74, 76, 78, 7 9 - 8 1 , 82,
Callipielus, 62 85, 404, 407, 408, 415, 449, 450
Calliprogonos, 342, 344; C. miraculosa, 343, 344 Ceirinae, 362
Callisto, 114 Celaenorrhinus, 269, 272, 273; C. eligius, 271; C. fritz-
Calliteara, 370, 386; C. pudibunda, 386 gaertneri, 269
Callithrinca, 122 Celastrina, 285
Callizygaena, 174; C. durata, 171 Cemiostominae, 127, 128
Callizygaeninae, 170, 171, 172, 174 Cenoloba, 222; C. obliteralis, 221
Callophrys, 285 Cephimallota, 97
Callopistria, 384 Cephonodes, 347, 348; C. rufescens, 343
Callosiope, 193 Cepora, 281 ; C. perimale, 442
Calophasia lunula, 387 Cerace, 209; C. xanthocosma, 200
Caloptilia, 114; C. azaleella, 96, 111; C. stigmatella, Ceracini, 199, 200, 201, 203, 209
108, 110; C. syringella, 96 Ceranemotini, 305
Calosaturnia, 337 Cerapteryx, 384
Calosima, 144 Ceratocampinae, 337, 338, 340
Calothysanini, 316 Ceratomia amyntor, 345
Calpinae, 360, 361, 367, 368, 369, 370, 374-375, 376, Ceratophaga vastella, 93, 106
387, 390 Cerconota anonella, 136, 137, 148; C. palliata, 1361
Calpini, 3 7 4 - 3 7 5 Cercophana, 338, 340
Calyptra eustrigata, 361, 374 Cercophanidae, 321, 336, 450
Cameraria, 114; C. guttifnitella, 113 Cercophaninae, 39, 337, 338, 340
Campaeini, 315 Ceridia, 346
Camptoloma, 389; C. interiorata, 373 Cerodirphia, 340
Camptolomini, 373, 388, 389 Ceromitia, 73, 75; C. fasciolata, 80; C. iolampra, 66
468 Index of Scientific Lepidoptera Names

Cerurinae, 363 Chrysoecia atrolinea, 371

Cerurites, 355 Chrysoestiidae, 158
Cethosia, 288 Chrysonoma, 141
Cethosiini, 288 Chrysopeleiinae, 135, 136, 146, 155, 156, 157
Ceuthomadarinae, 143, 157 Chrysopoloma, 166, 168; C. venata, 161
Ceuthomadarus, 143 Chrysopolomidae, 188
Chaeopsestis, 303 Chrysopolominae, 159, 160, 161, 1 6 7 - 1 6 8
Chaetocneme, 269, 273 Chrysopsyche, 329
Chalcoela pegasalis, 248 Chrysozephyrus, 285
Chalcosia, 174 Cicinnus, 324; C. anysia, 325
Chalcosiidae, 163 Cilix, 305
Chalcosiinae, 159, 170, 171, 172, 173, 174 Cingiliini, 315
Chalcoteuches, 110 Cirrhochrista, 250
Chamaita, 391 Cirrhophanus, 382; C. triangulifer, 390
Charadra, 385, 386 Cisseps, 392
Charadraula, 144 Cissia, 290
Charaxes, 3, 278, 286, 289; C. jasius, 266, 289 Cissuvorini, 184
Charaxinae, 267, 287, 288, 289, 290 Cisthene, 391
Charaxini, 289 Citheronia, 340; C. phoronea, 347
Charideinae, 228, 229 Clania, 106, 107; C. peyeri, 96
Charltona, 244 Clanis, 346
Chasmina, 380 Cledeobia moldvico, 245
Chedra, 143 Cleis, 259; C. evander, 260
Cheimophila, 141; C. solicello, 137 Cleorini, 315
Chelariinae, 158 Clepsis, 207
Chelepteryx, 326; C. chalepteryx, 324; C. collesi, 328 Cliniodes, 235, 247; C. vinacea, 248
Chenuala, 326 Clostera, 363; C. albosigma, 364, 387
Chetone phyleis, 373 Closterothrix, 329
Chezala, 141 Clydonopteron, 242
Chilecomadia, 188; C. moorei, 190; C. valdiviano, 191, Cnephasia, 206
192 Cnephasiini, 201, 203, 204, 205, 2 0 6 - 2 0 7
Chilo, 244 Cnissostages, 104; C. oleagina, 98
Chimabachidae, 15, 36, 133, 134, 136, 137, 141, 405 Cochlidion, 167
Chinodes, 147; C. retiniella, 138 Cochylini, 199, 200, 201, 202, 203, 204, 206, 208, 411
Chionopsyche, 326, 327, 329; C. montana, 328 Cochylis, 206
Chionopsychinae, 326, 327, 329 Cocytia, 376, 379; C. durvillii, 368, 375
Chlamydastis, 136 Cocytiidae, 379
Chlanidophora, 382 Cocytiinae, 360, 368, 370, 375
Chlenias, 310 Cocytius, 347; C. antaeus, 347
Chlidanotinae, 199, 200, 201, 202, 203, 204, 205, 206, Coeliades, 272; C. forestan, 272
209-210 Coeliadinae, 263, 270, 272
Chlidanotini, 200, 201, 202, 204, 209-210, 447 Coelolepida, 10, 11, 28, 53
Chloephorinae, 362, 373, 388, 389, 390 CoeIonia, 347
Chloephorini, 388, 389 Coenonympha, 290
Chloroclystis rectangulata, 316 Coenotephria incultaria, 316
Chlosyne, 288 Colabata, 334
Choaspes, 272, 276 Coladenia, 273
Choerocampinae, 346 Coleophora, 144; C. atromarginata, 142; C. eleagnisella,
Choerocampini, 347 137; C. sacramenta, 140
Cholo tis, 146 Coleophoridae, 33, 36, 37, 132, 133, 134, 135, 137, 138,
Chondrostega, 326, 327, 329; C. tingitana, 325, 328 144, 157, 405, 410, 411, 414
Chondrosteginae, 326, 327, 329 Coleophorinae, 33, 131, 134, 135, 144, 155, 156, 157,
Chondrostegoides, 329 411, 450
Choreutidae, 36, 218, 219-220, 221, 406, 450 Coelopoetinae, 157
Choreutinae, 2 1 9 - 2 2 0 Coleotechnites, 147; C. edulicola, 142
Choreutis, 218, 219; C. pariana, 219; C. tigroides, 219 Coliadinae, 265, 280, 281, 291
Choreutoidea, 10, 14, 28, 29, 131, 219-220, 226 Colias, 278, 279, 280, 281, 291, 455; C. hyale, 264
Choristoneura, 207 Colla, 334
Choristostigma, 235 Collicularia, 213
Choropleca, 18 Collinsa, 229
Chrysaeglia magnifica, 390 Collomeninae, 373, 389
Chrysamma purpuripulcra, 161 Collomenini, 389
Chrysaster, 113, 114 Colobura dirce, 266
Chrysauginae, 234, 236, 238, 242, 406 Coloburini, 289
Chrysendeton, 247 Colocasia, 385
Chrysiridia, 308, 309, 310; C. rhipheus, 307 Coloradia, 337, 340
Chrysoclista, 139; C. linneella, 140 Colons, 279, 281
Chrysodeixis, 378, 424, 435 Comadia, 192
Index of Scientific Lepidoptera Names 469

Cornelia, 258, 259 Crocidolomia, 249

Comibaenini, 315 Crocidosema, 212; C. plebejana, 200, 202
Comodica, 94, 100 Cropia, 384
Comostolini, 315 Cryptaspasma, 210; C. sordida, 200, 201
Compsoctena, 100, 102; C. thwaitesii, 93, 98 Cryptolechiidae, 156
Compsocteninae, 102 Cryptophasa, 139; C. balteata, 139
Compsolechia, 147 Cryptophasidae, 157
Condica, 370, 381, 435; C. confederata, 371; C. vec- Cryphia domestica, 368
ors, 387 Cryphiomystis, 114
Condicinae, 360, 367, 369, 370, 371, 381-382, 387 Crypsiphona ocultara, 311
Condicini, 381 Crypsithyris, 94
Conogethes, 250 Cryptoblabes, 243
Conquassata, 146 Cryptophasa setiotricha, 131
Consul, 286, 289 Cryptophlebia, 215
Copaxa, 341; C. decrescens, 339 Cryptoptila, 202
Copiopteryx, 340 Cryptoses, 242
Copromorpha, 225, 226; C. efflorescens, 228 Cryptothelea, 107
Copromorphidae, 37, 225, 226, 228, 406 Ctenoplusia, 377
Copromorphoidea, 10, 13, 14, 28, 30, 125, 222, Ctenopseustis, 207
225-226, 409 Ctenucha, 392; C. virginica, 356, 378
Coptodisca, 71, 72, 73; C. arbutiella, 84; C. splendorif- Ctenuchidae, 174, 391
erella, 74 Ctenuchinae, 38
Corades, 290 Ctenuchini, 356, 391, 392, 394
Corcyra cephalonica, 242 Cuculila, 380; C. argentea, 371
Cornuticlava, 206 Cuculliinae, 360, 367, 369, 370, 371, 380-381, 382,
Coronidia, 305, 307 384, 387
Corrachia, 284; C. leucoplaga, 270, 284 Cuculliini, 380-381
Corsocasis, 220 Culama, 189, 191, 192, 193
Corybantes pylades, 185 Culladia, 244
Coryptilum, 94; C. klugii, 93 Cupha, 288
Coscinocera, 340, 341 Cupido, 285
Coscinoptycha, 226 Curetidae, 281
Cosmiotes, 139 Curetinae, 265, 283, 284, 285
Cosmophila, 380 Curetis, 283, 285; C. brunnea, 282; C. thetis, 270
Cosmopterigidae, 18, 36, 131, 135, 136, 138, 146-147, Curvicubitus triassicus, 16
157, 405, 411, 435 Cyana, 391, 446; C. basiava, 446; C. bicolor, 446; C.
Cosmopteriginae, 131, 135, 136, 146-147, 155, 156, croceizona, 446; C. dinawa, 446; C. fulvia, 446; C.
158, 411 guizonis, 446; C. hamata, 373; C. metamelas, 446; C.
Cosmopterix, 147; C. montisella, 142, 145 meyricki, 446; C. miles, 446; C. nigromarginata, 446;
Cosmorhoe, 441 C. omissa, 446; C. postdivisa, 446; C. scintillons, 446;
Cossidae, 36, 184, 188, 189-194, 195, 329, 377, 406, C. securizonis, 446; C. thoracica, 446; C. tricolor, 446
411,424 Cybalomia, 246; C. pentadalis, 245
Cossiformia, 159 Cybalomiinae, 234, 239, 246
Cossinae, 36, 188, 189, 190, 191,192 Cyclidia, 305; C. substigmaria, 305, 306
Cossoidea, 10, 13, 18, 28, 30, 31, 159, 181, 188-195, Cyclidiidae, 302
411,429, 431,434 Cyclidiinae, 304, 305
Cossula, 189, 193 Cyclophora, 316; C. flavissima, 441
Cossulinae, 188, 189, 190, 191, 193, 195 Cyclophorini, 316
Cossus, 3, 188; C. cossus, 192 Cyclophragma, 329
Cosymbiini, 316 Cyclosia, 174
Cotana, 331 Cyclotorna diplocentra, 161; C. monocentra, 162
Crambidae, 233, 234, 235, 238, 241, 243-250, 406, 412 Cyclotornidae, 37, 159, 160, 161, 162, 406, 450
Crambidia, 358, 391 Cycnodiidae, 157
Crambiformes, 234 Cydia, 213; C. pomonella, 200, 201
Crambinae, 233, 234, 235, 236, 239, 244-245, 412 Cydosia, 379
Crambini, 235, 236 Cydosiini, 379
Crambus, 244 Cyllopoda, 316; C. claudicla, 311
Craniophora, 370, 372 Cyllopodini, 316
Cremastobombycia, 114; C. ignota, 113 Cymolomia, 211
Crenidomimas concordia, 291 Cymoriza, 246
Crepidochares, 91, 100, 102; C. neblinae, 98, 101 Cymothoe, 289
Cressida, 453, 454 Cynaeda, 247
Cricula, 341 Cynthia, 288, 432; C. cardui, 432, 433; C. kershawi,
Crinodes, 363 432, 433; C. virginiensis, 432
Crinopterygidae, 67, 70, 71, 74, 76, 78, 80, 83, 84, 404, Cyphanta, 363
408, 415, 449, 450 Cyphura, 309
Crinopteryx familiella, 67, 70, 74, 76, 78, 80, 83, 84 Cyrestini, 287, 289
Crocanthes, 143 Cyrestis, 289
470 Index of Scientific Lepidoptera Names

Dacnonypha, 51 Dichomeridinae, 133, 135, 136, 147, 155, 156, 158

Dactyloceras, 342, 344; D. lucina, 339, 343 Dichomeris, 147; D. margine lia, 142, 145; D. mulsa,
Dahlica, 105, 107; D. triquetrella, 96, 107 138; D. punctipennella, 148; D. simpliciella, 148
Daiopterix, 16, 28 Dichordophorini, 315
Dalaca, 62 Dichrorampha, 212, 213
Dalailama, 336 Dicranoctetes, 139
Dalcera, 169; D. abrasa, 161 Dicranoses, 71, 79; D. congregateli, 66, 70, 74, 76, 85
Dalceridae, 34, 159, 160, 161, 164, 167, 168, 175, 188, Dicranurini, 363, 364
406, 450 Dicroplema, 308
Dalcerides, 169; D. ingenita, 169; D. ingenitus, 175 Dicymolomia, 249
Dalla, 273 Digama hearseyana, 375; D. marmorea, 375
Danainae, 267, 278, 286, 287, 292, 413, 428, 430, 451, Digglesia, 329
455 Digitivalva, 125
Danaini, 292, 451 Diloba, 358, 362, 391; D. caeruleocephala, 368, 381
Danaus, 1, 3, 4, 278, 292, 331, 424, 452, 453; D. chrys- Dilobinae, 37, 358, 360, 368, 369, 370, 381
ippus, 451, 453; D. plexippus, 268, 277, 286, 292, Dilophonotini, 347
436, 451 Dinia, 390, 392
Daphnis, 348 Diocosma, 141
Daphoenura, 379 Dione, 288
Daraba, 250 Dioptinae, 357, 361, 363, 364, 365, 376, 387, 390
Dasy carea, 110 Dioryctria, 235, 243; D. rossi, 245
Dasyceridae, 157 Diphthera, 385
Dasychira, 386; D. mendosa, 388; D. mescoler, 373 Diplodoma, 107
Dasylophia anguina, 387 Diplosaridae, 158
Datana, 363; D. angusii, 387; D. ministra, 364 Diptychophorini, 235
Decachorda, 341 Dirades, 309
Decachordini, 341 Dirce, 313, 314
Deceda, 309 Dirphia, 340; D. lichyi, 335
Deilephila, 347; D. porcellus, 343 Dirphiopsis, 340
Delias, 279, 281; D. aglaia, 264 Discophlebis, 360
Deloneura sheppardi, 282 Dismidila, 246
Deltophora, 147 Dismorphia, 279, 280; D. laia, 264
Deltoplastis figurata, 140; D. ocreata, 142 Dismorphiinae, 265, 275, 276, 280
Deltote, 379 Dissoctena, 107
Demopsestini, 305 Dissoctenioides, 107
Denaeantha nivigera, 205 Dissoprumna erycinaria, 306
Dendrolimus, 329 Ditrigona, 305
Dendrorycter, 113 Ditrysia, 10, 11, 12, 14, 16, 27, 28, 29, 57, 65, 230, 301,
Deoclona yuccasella, 137 404, 408, 412, 413, 415, 435, 450
Deoclonidae, 134, 135, 137, 143-144, 157 Diurnea fagella, 140
Deocloninae, 134, 135, 143, 155, 156, 157 Diurneinae, 157
Depressaria, 138; D. cinereocostella, 145; D. juliella, Doa, 37, 357, 361, 386; D. ampia, 364
140 Dodona, 284
Depressariidae, 405 Doidae, 37, 355, 356, 357, 3 6 0 - 3 6 1 , 364, 367
Depressariinae, 133, 134,138, 155, 156, 158 Doina, 146
Deramas, 284 Doleschallia, 286, 288; D. bisaltide, 282
Desmeocraerinae, 362 Dolbina, 345
Desmia, 250; D. vulcanalis, 241 Dolichomia, 243
Desmobathrinae, 315 Donacaula, 246
Desmobathrini, 315 Donda, 369
Deudorix, 285 Doratopteryx, 163
Deuterocopinae, 224 Doritites, 19, 20; D. bosniaskii, 16
Deuterocopus, 224; D. planeta, 221 Douglasia, 110
Deuterogonia, 139; D. chionoxantha, 137, 145 Douglasiidae, 34, 96, 107, 108, 109, 110, 405, 409, 450
Deuterogoniinae, 133, 134, 139 155, 156, 157 Doxocopa, 289
Deuterollyta conspicualis, 237 Drachmobola, 207, 208
Deuterotinea, 100, 102 Dragmatucha, 143
Diacrisia, 386 Drapetodes, 305
Dialithis gemmifera, 368 Drastea, 103
Dianeura, 163, 164 Drasteria, 374
Diaphania, 250 Dreata, 330, 331
Diaphone, 382; D. eumela, 371, 382 Drepana, 305; D. curvatula, 306
Diastema, 370 Drepanidae, 37, 257, 301, 3 0 2 - 3 0 5 , 306
Diatraea, 244 Drepaninae, 301, 303, 304, 305
Dicallaneura pulchra, 291 Drepanini, 304, 305
Dichocrocis, 250 Drepanoidea, 10, 14, 28, 30, 257, 3 0 1 - 3 0 5 , 429, 434,
Dichogama, 249 450
Dichogaminae, 234, 249 Druentica, 324
Index of Scientific Lepidoptera Names 471

Druryia, 454 Endromidae, 39, 321, 322, 334-336, 339, 343, 450
Dryadaula, 94, 99 Endromis versicolora, 336, 339, 343
Dryadaulinae, 9 7 - 9 9 Endrosis, 141
Dryocampa, 338 Enicostominae, 156
Dudgeonea, 189, 190, 192, 195; D. actinias, 194 Enispa, 379
Dudgeoneidae, 188, 189, 190, 192, 194-195, 233, 406 Ennominae, 311, 312, 313, 314-315, 431
Dudua, 211 Ennomini, 315
Dudusa, 363; D. nobilis, 387 Ennomos autumnaria, 314; E. subsignaria, 314
Dudusiinae, 357, 363-364, 387 Enteucha, 69
Dudusiini, 361, 363 Enyo, 348
Durbania, 284 Eocorona iani, 16
Durrantia, 146; D. amabilis, 138 Eolepidopterigidae, 16
Dynamine, 289 Eolepidopterigina, 16
Dysauxes, 392 Eolepidopterix, 16, 28
Dyseriocrania, 52, 53, 54 Eoophyla, 247
Dysodia, 229; D. rajah, 228 Eoparargyractis plevie, 241
Dysoptus tantalota, 93 Eoreuma, 244
Dyspessa, 192 Eoses trissica, 16
Dyspyralis puncticosta, 368 Eosopostega, 69, 71
Dysspastus, 146 Epermenia, 218, 222; E. chaerophylella, 221, 222
Dysstroma, 439; D. cuneifera, 441; D. fumata, 441 Epermeniidae, 35, 218, 220-222, 406, 414
Epermeniinae, 222
Eacles, 340; E. masoni, 335 Epermenioidea, 10, 13, 14, 28, 30, 2 2 0 - 2 2 2
Eana, 206 Epia, 334; E. muscosa, 332, 335
Eariadinae, 359, 373, 389, 390 Epiblema, 212
Earias, 38, 359, 370, 388; E. biplaga, 389; E. insulana, Epicampoptera, 305
389; E. perhuegeli, 389; E. punctaria, 390; E Vit- Epicausis, 379
iello, 389 Epicephala, 114
Eccritothrix trimaculella, 106 Epichnopterix, 107
Eclipsiodini, 236 Epichnopteryginae, 107
Ecpathophanes, 104 Epichoristodes, 207
Ecpyrrhorrhoe, 250 Epicimelia, 258; E theresiae, 260
Ectoedemia, 18, 68, 69, 87; E. phleophaga, 70, 74, 76, Epicopeia, 302; E. mencia, 306
78, 84; E. sericopeza, 67 Epicopeiidae, 301-302, 303, 306, 307, 450
Ectomyelois ceratoniae, 243 Ephigraphiinae, 156, 158
Ectropa, 168 Epiini, 334
Edosa, 99 Epimarptinae, 135, 143, 155, 156, 157
Eidophasia, 125; E. messingiella, 123 Epimarptis, 137, 143; E. philocoma, 145
Eilema, 391 Epimariyria, 41, 42, 43, 44; E. pardella, 45
Elachista, 139 Epimecis hortaria, 311
Elachistidae, 18, 35, 36, 131, 132, 133, 134, 136-139, Epinotia, 212
137, 156, 158,405,410,411 Epipaschia, 243; E. superatalis, 245
Elachistinae, 18, 35, 133, 134, 139, 155, 156, 411, 450 Epipaschiinae, 234, 235, 236, 238, 242, 243, 406
Elaeognatha, 389 Epiphora, 341; E. atbarina, 338
Elaphria, 369, 384 Epiphyas, 207; E. postvittana, 201, 202
Elbella, 273 Epiplema, 309
Elcysma, 174 Epipleminae, 308, 309
Eldana, 242 Epipomponia, 162; E. nawai, 160, 161
Electrocrania immensipalpia, 17 Epipsestis, 305
Eleppone, 454 Epipyropidae, 35, 159, 160-162, 168, 175, 406
Eligma, 388, 389; E. narcissus, 373 Epipyrops, 160, 162
Eligminae, 359, 373, 389 Epirranthini, 315
Elophila, 247 Episteme adulatrix, 368
Elymnias, 290, 291 Epitola, 284
Embryonopsis, 125 Epitrichosma, 206; E. neurobapta, 200
Emesis, 284; E. tenedia, 270 Epitymbia, 207; E. alaudana, 200
Emmelina, 218 Epitymbiini, 199, 200, 201, 203, 207, 411, 412
Enanlia, 280 Eporidia, 235
Enarmonia, 212 Eques, 1
Enarmoniini, 200, 211212 Erannis defoliaria, 312, 314; E. tiliaria, 314
Enchoptila, 110 Erastrianae, 379
Endoclita, 61 Erastriini, 379
Endopiza, 211 Ercheia, 374
Endothenia, 210, 211 Erebia, 290
Endotricha, 243 Erebinae, 374
Endotrichini, 235, 236 Erebus, 374; E. ephesperis, 356
Endoxyla, 189, 190, 193, 195; E. encalypti, 191; E. litur- Erechthias, 94, 100
atus, 192 Erechthiinae, 92, 94, 100
472 Index of Scientific Lepidoptera Names

Eresia, 288 Eumorpha, 348

Eretmocera, 140 Eupalamides, 186; E. cyparissias, 186
Eretris, 290 Eupanacra, 348
Eridachtha, 143; E. prolocha, 140 Euparyphasmini, 305
Erinnyis, 348 Eupastranaia fenestrata, 246
Eriocottidae, 91, 92, 93, 98, 100, 101, 405, 409, 450 Eupistidae, 157
Eriocottinae, 100-102 Euphaedra, 289
Eriocotds, 100, 102; E. fuscanella, 93, 101 Euphydryas, 288
Eriocrania, 51, 52, 53, 54; E. sparrmannella, 56 Eupithecia, 316; E. millefoliata, 312
Eriocraniella, 53 Eupitheciini, 316
Eriocraniidae, 11, 17, 34, 41, 5 1 - 5 3 , 54, 56, 404, 406, Euplagia quadripunctaria, 394
407, 415, 447, 448, 450 Euplocaminae, 100
Eriocranioidea, 10, 11, 13, 28, 29, 5 1 - 5 3 , 4 0 6 , 4 1 3 , 415 Euplocamus, 100
Eriogaster, 327, 329; E. lanestris, 325; E. rimicola, 328 Euplocia, 375; E. membliaria, 368
Erionota, 269; E. thrax, 271 Euploea, 292, 452; E. core, 451
Eriopinae, 384 Eupoecilia, 206
Eriopygini, 369, 384 Eupragia, 146
Erites, 290 Euproctis, 361, 370, 386, 393; E. chrysorrhoea, 388; E.
Erosia, 309; E. incendiata, 303, 306 kanshireia, 390; E. similis, 388
Erupa, 244 Eupselia, 139
Erynnis, 269, 273 Eupterote, 331
Erythrina, 250 Eupterotidae, 39, 321, 322, 325, 328, 329-331, 450,
Eschata, 244 450
Eterusia aedea, 171 Eupterotinae, 331
Ethmia assamensis, 138; E. nigroapicella, 138; E. semi- Euptychia, 290, 291
lugens, 137; E. sphaerosticha, 138 Eurema, 279, 280, 281, 282; E. daira, 264; E. hecabe,
Ethmiidae, 405, 410, 411 291
Ethmiinae, 133, 134, 136-138, 411, 155, 156 Euripus, 289
Etiella zinckenella, 243 Euryades, 453, 454
Eublemma, 379; E. dimidialis, 379; E. radda, 379 Eurybia, 284
Eublemmini, 371, 379 Eurydoxa, 209
Eucecidoses, 79; E. minutanus, 85 Eurytides, 453, 454
Eucereon, 392 Euschemon, 272, 273, 274; E. rafflesia, 271, 273, 286
Euchaetes, 392 Euscirrhopterus gloveri, 387
Eucheira socialis, 280 Euselasia, 284; E. gelon, 270
Euchloe, 281; E. ausonia, 264
Eustrotia, 379
Euchromia, 392; E. lethe, 373
Eustrotiinae, 360, 367, 369, 370, 371, 379, 387
Euchromiini, 373, 391, 392, 394
Eutelia, 377
Euchromius, 244; E. ocelleus, 245
Euteliinae, 360, 368, 369, 370, 375, 376, 377, 387, 431
Euclea, 167
Euthalia, 289
Euclemensia, 147; E. bassettella, 138
Euthrix, 327, 329
Eucleoidea, 159
Euxanthini, 289
Eucocytia meeki, 371, 379
Eucocytiinae, 360, 370, 371, 379 Euxanthe, 289
Eucosmini, 199, 200, 201, 202, 203, 210, 212 Euxoa, 384, 438
Eucosmomorpha, 212 Euxoini, 369
Eucryptogona, 102 E veres, 285
Eudaemonia, 337, 341 Evergestinae, 234, 236, 240, 249, 412
Eudalaca, 62 Evergestis, 249; E. limbata, 248
Eudaphaenura, 379 Exaeretia, 138
Eudarcia, 94, 97; E. simulatricella, 101 Exapate, 206
Eudia, 340 Excurvaria, 81
Eudocima fullonia, 374; E. homaena, 387, 390; E. tyran- Exoncotis, 103
nus, 368 Exoporia, 9, 11, 17, 28, 55, 57, 407, 413
Eudonia, 235, 236, 244; E. torniplagalis, 245 Exoteleia, 147
Eudulini, 316 Exyra fax, 390; E. semicrocea, 387
Eueides, 288 Falkovitshiinae, 157
Euglyphis, 327, 329; E. mama, 328
Eulechria, 141 Feniseca, 285
Eulia, 208; E. ministrana, 200, 208 Feralia, 381; F. comstocki, 371, 387; F. februalis, 381
Euliini, 199, 200, 201, 206, 2 0 8 - 2 0 9 Feraliini, 371, 381
Eulonche, 370, 372 Fernandocrambus, 244
Eulophonotus, 193 Filatima, 147
Eumaeus, 284, 285; E. minijas, 271 Fissicrambus, 244
Eumeleini, 315 Fraus, 54, 61, 429, 430, 449, 450
Eumestleta recta, 371 Frechinia, 247
Eumeta, 107 Frisilia, 143
Eumicremma, 369 Fulgoraecia exigua, 160
Index of Scientific Lepidoptera Names 473

Fumea, 107 Gnophini, 312

Furcula borealis, 364, 387; F. scolopendrina, 390 Gnorimoschema, 147
Goacampa, 363
Gabala argentata, 373 Gonepteryx, 281, 437
Gadirtha, 389 Gonimbrasia, 341
Galactica, 217, 220 Goniodoma, 144
Galacticidae, 35, 217, 221 Gonodonta, 375; G. correda, 368
Galacticoidea, 10, 28, 31, 217 Gonodontini, 315
Galasa, 242 Gonometa, 329
Galleria mellonella, 242 Gonometinae, 327, 328
Galleriinae, 234, 235, 236, 238, 242, 406, 412 Gonopterini, 374375
Galleriini, 242 Gonoretodes timea, 306
Gamelia, 337, 340 Goodia, 341; G. kuntzei, 341
Gargetta, 363 Goodiini, 341
Garrha, 143 Gorgopis, 62
Gastridiota adoxima, 347 Gracilepterix, 17, 28
Gastropacha, 327, 329 Gracillariidae, 16, 18, 34, 35, 96, 107, 108, 109, 110,
Gastropachinae, 327, 328 111, 112-114, 405, 409, 414
Gastropachini, 329 Gracillariinae, 112, 114
Gastroplakaeis, 329 Gracillarioidea, 10, 12, 18, 28, 91, 107-114, 405, 408,
Gatesclarkeana, 200, 211 409
Gatesclarkeanini, 200, 210, 211 Graphiini, 453, 454, 455
Gathynia, 309 Graphium, 274, 277, 279, 291, 453, 454; G. agamemnon,
Gaujonia, 385 277; G. sarpedon, 264
Gaurena, 305 Grapholita, 213
Gaurenini, 305 Grapholitini, 200, 203, 210, 212, 215
Gazoryctra, 61, 449, 450 Greya, 77, 79; G. pectinifera, 75; G. punctiferella, 66, 76
Gelastocera, 389 Griveaudia, 259; G. charlesi, 260; G. vieui, 258, 260
Gelechia, 147; G. rhombelliformis, 142 Griveaudiinae, 258, 259
Gelechiidae, 18, 34, 35, 36, 131, 132, 135, 136, 138, Groteila, 382; G. dis, 371
147-149, 158, 405, 410, 411, 435, Grotellini, 371, 381, 382
Gelechiinae, 133, 135, 147, 155, 156, 158 Gunda, 334
Gelechioidea, 9, 10, 12, 13, 18, 28, 29, 127, 131-149, Gymnodisca, 439, 447
155, 156, 158,219, 2 2 2 , 4 0 5 , 4 0 8 , 4 1 0 , 4 1 1 , 4 1 3 , 4 1 4 Gymnogramma, 170; G. rufiventris, 161
Genduara, 329 Gynaephila, 372
Genus tes, 60 Gynaephora, 386
Geometra, 1, 233 Gynoeryx teteforti, 343
Geometridae, 37, 301, 302, 305, 3 1 0 - 3 1 6 , 412, 435,
441, 450 Habrosyne, 305, 441; H. pyritoides, 306
Geometrinae, 311, 312, 313, 314, 3 1 5 - 3 1 6 Habrosynini, 305
Geometrini, 315 Hadapameini, 384
Geometroidea, 10, 14, 19, 28, 30, 257, 301, 3 0 5 - 3 1 6 , Hadeninae, 358, 360, 367, 369, 370, 371, 376, 378, 379,
429, 434 380, 381, 382, 383, 387, 390
Gephyroneura, 324, 326 Hadenini, 369, 371
Gerontha, 94; G. captiosella, 93 Haemilidae, 156
Gesneria centuriella, 245 Haetera, 290, 291; H. piera, 268
Girdharia, 244 Haeterini, 267
Givira, 194; G. nais, 190 Hahncappsia, 250
Glanycus, 229; G. tricolor, 229 Haimbachia, 244
Glaphyria, 249 Halpe, 274
Glaphyriinae, 234, 235, 236, 240, 249 Halysidota, 392; H. ata, 373
Glaucacna iridea, 139 Hamadryas, 289
Glaucocharis, 244 Hamanumida daedalus, 282
Glaucopsyche, 285 Hamearis, 284; H. lucina, 270
Glossata, 9, 10, 11, 16, 27, 28, 41, 43, 51, 403, 404, Haplotineinae, 97
406,413, 447 Haploptilidae, 157
Glottulinae, 360, 366, 367, 370, 3 7 1 , 3 8 2 - 3 8 3 , 384, 387 Hapsifera, 99
Gloveria, 329 Hapsiferinae, 92, 99
Gluphisiini, 363 Harmaclona, 94, 97
Glyphidocera, 137, 141; G. juniperella, 141 Harmocloninae, 91
Glyphidoceridae, 136, 137, 141 Harpyia, 364
Glyphipterigidae, 34, 119, 120, 121, 123, 124,125, 127, Harrisina, 172, 174
134,217, 226,405, 409,410 Hasora, 269, 272
Glyphipteriginae, 125126, 127 Hecatesia, 379
Glyphipterix, 121, 126; G. haworthana, 123 Hednota, 244
Glyphodes, 250; G. extorris, 248 Hedya, 211
Glypiphteriginae, 120 Hedylidae, 37, 265, 271, 293, 450
Gnathifera, 222 Hedyloidea, 10, 14, 28, 257, 263, 267, 268, 293, 315
474 Index of Scientific Lepidoptera Names

Helcystogramma, 147 Heterocrossa, 226

Helicomitra, 259 Heterogymna, 225
Heliconiinae, 267, 279, 287, 288, 292, 415 Heterogynidae, 37, 159, 160, 161, 170, 450
Heliconiini, 286, 288, 438, 439 Heterogynis, 170; H. penella, 161
Heliconius, 174, 278, 279, 286, 288; IL charitonia, 266 Heteroneura, 9, 10, 11, 12, 18, 28, 51, 53, 59, 65, 86,
Helicopis, 284 404, 407, 447, 450
Helicoverpa, 382, 424, 432, 435; H. armigera, 382; H. Heteropan, 174
assulta, 382; H. zea, 382 Heteropsyche, 162, 175
Heliocheilits albipunctella, 382 Heteropterinae, 265, 272, 273
Heliodines, 126, 127 Heteropterus, 273; H. morpheus, 271
Heliodinidae, 18, 36, 119, 123, 124, 126,127, 128, 405, Hexeretmis, 223; H. argo, 221
409, 410 Hexeriiis, 19
Heliophorus, 285 Hibrildes, 328, 329, 330, 331; H. norax, 325
Heliothela, 239 Hibrildidae, 321
Heliothelinae, 234, 244 Hibrildinae, 331, 450
Heliothelini, 236 Hidari irava, 271
Heliothinae, 360, 367, 370, 371, 381, 382, 383, 387, Hieromantis, 143
412, 435 Hierophanes, 127
Heliothis, 435; H. borealis, 371; H. virescens, 382, 383 Hieroxestinae, 92, 97, 9 9 - 1 0 0 , 409
Heliozela, 66, 72, 73; H. aesella, 70, 74, 76, 78, 82 Hilarographa, 210
Heliozelidae, 34, 65, 66, 70, 7 1 - 7 3 , 74, 76, 78, 82, 84, Hilarographini, 200, 201, 203, 204, 209, 210
85, 404, 407, 408, 414, 450 Himantopteridae, 159, 161, 1 6 2 - 1 6 3
Hellinsia, 224 Himantopterus dohertyi, 163; H. fuscinervis, 163
Hellula, 249 Hipparchia, 290
Hemaris, 345, 347, 348; H. fuciformis, 339 Hippotion, 347, 348
Hemiceras, 365; H. nigrigutta, 364 Historis, 289
Hemiceratinae, 364, 367 Hofmannophila, 143
Hemiceratini, 365 Holcocera, 144
Hemigrotella, 382 Holcocerinae, 157
Hemileuca, 337, 338, 340 Holcopogon bubulcella, 138, 144
Hemileucinae, 337, 338, 339, 340 Holcopogoninae, 135, 144-146, 155, 156, 157
Hemileucini, 340 Holocerina rhodesiensis, 335
Hemiostolini, 315 Holomelina, 392
Hemipecten niveogrisea, 190 Homadaula, 217; H. anisocentra, 217, 221
Hemipectrona nigripennata, 190 Homaledra, 143; H. heptathalama, 137
Hemitheini, 315 Homaloxestis, 143
Henotesia, 290 Homidiana, 305, 307; H. japet, 303, 306
Heortia, 247 Homoeopteryx, 340
Hepialidae, 35, 54, 58, 59, 6 1 - 6 2 , 407, 414, 447, 450 Homoeosoma, 243
Hepialoidea, 10, 11, 28, 29, 5 9 - 6 2 , 404, 407, 450 Homona, 207, 208
Hepialus, 54, 61; H. humuli, 61 Homosetia, 97
Heraclides, 454 Hoplojana, 331
Herculia, 243 Hoplophanes chlorochrysa, 66
Her ingita, 144 Hoplophractis, 181, 182
Herminia, 369, 372 Hoploscopini, 236
Herminiinae, 355, 360, 366, 368, 369, 370, 372, 382, Hositea, 246
387,412 Hulstia, 243
Herpetogramma, 250 Hyalea, 250
Herrichiinae, 157 Hyblaea, 225, 227; H. flavifasciata, 228; H. puera, 227
Hesperia, 269, 274; H. comma, 271, 286 Hyblaeidae, 37, 225, 227-228, 233, 406, 450
Hesperiidae, 19, 35, 174, 263, 269-274, 275, 276, 282, Hyblaeoidea, 10, 14, 28, 30, 2 2 7 - 2 2 8
286, 287, 291, 429, 451 Hydrillodes, 372
Hesperiinae, 265, 272, 2 7 3 - 2 7 4 Hydriomeninae, 316
Hesperilla, 273 Hydriomenini, 316
Hesperioidea, 10, 14, 19, 28, 29, 257, 263, 267, 268, Hylaeorinae, 362
269-274, 276, 278, 412, 413 Hyles, 347, 348
Hestiochora, 172, 173 Hylesia, 338, 339, 340; H. lineata, 338
Heterobalhmia, 44, 45, 447, 449; H. pseuderiocrania, Hymenia, 250
45, 47; H. valvifer, 46, 47 Hyparpax, 364
Heterobathmiidae, 9, 34, 41, 44, 45, 4 6 - 4 7 , 404, 415, Hypatima, 147
447, 448, 449 Hypatiminae, 158
Heterobathmiina, 27, 405, 406 Hypatopa, 144
Heterobathmioidea, 10, 11, 27, 28, 29, 41, 43, 45, Hypena, 376, 435; H. longipennis, 368
46-47,406,413,414,415 Hypeninae, 360, 367, 368, 370, 374, 376, 380, 387, 390
Heterocampa, 365; H. astarte, 364; H umbrata, 383 Hypenodes, 374
Heterocampinae, 356, 358, 361, 364-365, 378, 380, Hypenodinae, 374
383, 387 Hyperchiria, 338
Heterochorista, 208 Hypermnestra, 19, 278, 453, 454, 455
Index of Scientific Lepidoptera Names 475

Hyperthaema, 392 Ithomia, 293

Hypertropha, 139; H. tortriciformis, 137 Ithomiinae, 287, 429, 439
Hypertrophinae, 133, 134, 139, 155, 157, 158, 450 Ithomiini, 287, 292-293, 438, 451, 452
Hyphantria cunea, 394 Ixias, 281
Hypochrosini, 315, 431
Hypochrosis, 315 Jalmenus, 285
Hypochrysops, 285 Jamides, 285
Hypocysta, 290 Jana, 331
Hypocystini, 451 Janinae, 331
Hypolamprus, 229 Janiodes, 337, 340
Hypoleria, 293 Janomima karschi, 328
Hypolimnas, 286, 288 Jemadia, 269, 273
Hypolycaena, 285 Jodiini, 315
Hypomartyria, 44 Jordanita, 174
Hypomecis, 310, 315 Josia fustula, 387; J. gigantea, 364
Hypoprepia, 391 Junonia, 286, 288, 289, 424
Hypopta, 189, 194; H. watsoni, 190
Hypoptinae, 188, 189, 190, 191, 193-194 Kallima, 286, 288
Hyposcada, 293 Kallimini, 288
Hyposmocoma, 147 Karataunia lapidaria, 16
Hyposmocomidae, 158 Kearfottia, 103, 105, 107; K. albifasciella, 93, 98, 101
Hypothyris, 293 Kedestes, 274
Hypotiinae, 235 Keiferia, 147
Hypotiini, 236 Kessleria, 122
Hyppa, 384 Kiwaia, 147
Hypsidia, 303, 304, 305; H. erythropsalis, 306 Klimeschia, 110
Hypsoropha adeona, 376; H. hormos, 380 Korscheltellus, 62
Kunugia, 329
Ichthyura, 363 Kurtomesis, 146
Idaea, 316; I. sericeata, 314
Idaeini, 316 Labdia, 147
Idalus, 392 Lachneini, 329
Idea, 292, 425, 451, 452, 453 Lachnocephala, 313
Ideopsis, 451, 452 Lachnocnema, 285
Idia, 372 Lacosoma, 324
Idioblasta, 235 Lacosomatinae, 324
Idiopteryx, 143 Lacosominae, 324
Mioses, 189, 192 Lactistica, 143; L. longimana, 140
Idrusia nyctelius, 289 Lactura, 170, 175
Ilionarsis, 144 Lacturidae, 160, 161, 169, 175
Illiberis, 174 Laelia, 387
Imbrasia, 341 Lambdina fiscellaria, 314
Imma, 226, 227; I. albo/asciata, 228 Lambertiodes, 208
Immidae, 37, 226, 227, 228, 406, 450 Lampides, 285; L. boeticus, 283
Immoidea, 10, 14, 28, 31, 226-227 Lampronia, 77; L. capitella, 80; L. fuscatella, 81; L.
Incurvarla, 71, 81; I. masculella, 66, 70, 74, 76, 78, 85; luzella, 70, 74; L. russatella, 66, 76, 78
I. vetulella, 80 Lamproniinae, 35, 77
Incurvariidae, 33, 66, 67, 70, 71, 74, 76, 78, 80, 81-83, Lamproptera, 279, 454
85, 86, 404, 407, 408, 414, 450 Lampropterini, 454
Incurvariina, 65 Lamprosema, 250
Incurvarioidea, 9, 10, 12,18, 28, 29, 65, 66, 71-83, 404, Lamyristis, 105
407, 414, 450 Langsdorfs, 189, 194
Indarbela, 193 Lapara, 347
Infurcitinea, 95, 97 Larentia clavaria, 314
Inga, 143 Larentiinae, 312, 313, 314, 316
Iolaus, 285 Lasiocampa, 327, 329; L. quercus, 326
Iphiclides, 453, 454, 455 Lasiocampidae, 38, 321, 322, 324, 325, 326-329, 337,
Ipimorphinae, 384 431, 450
Ipimorphini, 369 Lasiocampinae, 326, 327, 328, 329
Iropoca, 388 Lasiocampini, 329
Iscadia, 389; I. aperta, 373 Lasiocampoidea, 10, 14, 28, 321, 322, 324-329
Ischyja, 374 Lasiolopha, 389
Isognathus, 348 Lathosea, 380
Isonomeutis, 13, 226 Lavernidae, 157
Isophrictis, 147 Lebeda, 329
I so trias, 209 Lebedodes, 189
Issoria, 288 Lechriolepis, 329
Ithome, 146 Lecithocera, 143
476 Index of Scientific Lepidoptera Names

Lecithoceridae, 35, 131, 134, 137, 143, 155, 156, 157, Lithocolletinae, 112, 113, 114
405, 410 Lithocolletis, 114
Lecithocerinae, 143, 157 Lithopleur a monochopis, 106
Leioptilus, 218, 224 Lithosiinae, 358, 361, 373, 386, 390, 391
Leipoxais, 329 Litidae, 158
Lemaireia, 338 Litobrenthia, 219
Lemonia, 342; L. dumi, 342; L. sardanapalus, 325, 328 Litodonta, 364
Lemonias, 284 Litoprosopus, 366, 372, 374
Lemoniidae, 38, 321, 322, 325, 328, 341-342, 450 Lobesia, 211
Lemoniinae, 342 Lobobunaea, 341
Lepidochrysops, 285 Lobophorin/, 316
Lepidodes gallopavo, 368 Lochmaeus, 365; L. bilineata, 378, 380
Leptidea, 280 Loepa, 341
Leplocneria binotata, 388; L. reduca, 388 Lonomia, 337, 339, 340
Leptodeuterocopus, 224 Lophochoristini, 315
Leptosia, 281 Lophocorona, 52, 55; L. pediasia, 56
Leptosteges, 246 Lophocoronidae, 11, 51, 52, 55, 56, 404, 407, 413,
Leptotes, 285 448, 450
Leptozestis, 146, 147 Lophocoronoidea, 10, 11, 17, 28, 29, 41, 53, 54, 55, 57
Lethe, 290 Lophoptera, 369, 377
Leto, 62 Lophostethus, 345
Letogenes, 146 Lopinga achine, 291
Leucanella, 340 Losara, 454
Leucania, 384; L. dorsalis, 376 Loxofdonia, 441
Leucanthiza, 114 Loxostege, 250
Leucinodes, 250 Loxura, 285
Leucobrephos, 313 Lucia, 285; L. timbara, 270
Leucocnemis, 381 Ludia, 341
Leucoedemia, 110, 112; L. ingens, 96 Ludiinae, 337, 338, 340, 341
Leucoma, 386; L. salicis, 388 Ludiini, 341
Leuconycta, 370 Luehdorfia, 19, 278, 454
Leuconyctini, 381 Luffia, 105, 107
Leucophlebia, 346; L. afra, 339 Lycaena, 285, 447, 455; L. phlaeas, 270
Leucoptera, 126, 128; L. malifoliella, 128; L. psopho- Lycaenidae, 19, 39, 174, 265, 270, 271, 274, 275, 276,
carpella, 128 279, 281-285, 286, 291, 412, 416, 450
Leuculodes, 361 Lycaeninae, 265, 283, 285
Leuroperna, 125 Lycaenini, 285, 447
Levuana iridescens, 175 Lycorea, 451, 452
Libythea, 285, 286, 288; L. cuvierii, 288; L. geoffroy, Lygephila craccae, 361
266, 288 Lygropia, 250
Libytheana, 288 Lymanopoda, 290
Libytheidae, 274 Lymantria, 370, 386; L. dispar, 388; L. monacha, 388
Libytheinae, 19, 267, 269, 285, 287-288, 291 Lymantriidae, 38, 170, 326, 334, 355, 356, 357, 359,
Lichenaula, 139 361, 362, 366, 367, 369, 373, 375, 380, 381, 385,
Lichnoptera, 385 386-388, 389, 390, 393, 412, 429, 434, 450
Limacodes, 167 Lymantriinae, 386
Limacodidae, 34, 159, 161, 162, 164, 165, 166-168, Lymantriini, 386
169, 172, 175, 188, 377, 406, 431, 450 Lyonetia, 126, 128; L. clerckella, 128
Limacodinae, 161, 166-167, 168 Lyonetiidae, 18, 119, 120, 123, 124, 126, 127-128, 405,
Limenitinae, 267, 287, 288, 289, 291 408, 409, 410
Limentini, 288, 289 Lyonetiinae, 128
Limenitis, 289; L. populi, 266 Lypothora, 209
Limnaecia, 147 Lypusa maurella, 102
Lindera, 99 Lypusidae, 91, 92, 102
Lineodes, 250 Lyssa, 309
Linggana, 107
Linosta, 246; L. sinceralis, 245 Macalla, 243
Linostinae, 234, 236, 240, 246 Macarangela, 110
Liopasia andrealis, 248 Macariini, 312, 315
Liparopsis postalbida, 380, 390 Macarostola pontificalis, 96
Liphyra, 283, 284, 285; L. brassolis, 271, 282 Machaeropteris, 92
Liphyrini, 266, 284, 285 Machimia, 146
Lipocosma, 249 Macrobathra, 147
Liptena, 283, 284; L. undularis, 270 Macrobrochis, 391
Liptenini, 284 Macrocilix, 305; M. mysticata, 306
Lita, 147 Macrocyttara, 191, 192
Lithacodia, 369, 379; L. bellicula, 371 Macroglossinae, 344, 346, 3 4 7 - 3 4 8
Lithinini, 315 Macroglossini, 347
Index of Scientific Lepidoptera Names 477

Macroglossum, 347, 348 Melittiini, 184

Macrolepidoptera, 10, 14, 28, 224, 230, 233, 321, 406, Memphis, 289
412,413,414,417,418 Mendesia, 139
Macromphalia, 329 Menelaides, 454
Macromphaliinae, 326, 327, 329 Menevia, 324
Macropiratinae, 224 Meridarchis, 226
Macropiratis, 224 Meritastis, 207
Macrosoma, 268, 282, 293; M. tipulata, 271 Meroleuca, 336, 337, 340
Macrostola, 114 Mesapamea, 384
Macrothyatirini, 305 Mesembrynus, 173
Maculinea, 285; M. alcon, 285; M. arion, 285; M. reb- Mesene, 284
eli, 285 Mesepiola, 77, 79
Mahasena polytricha, 106 Mesodina, 273
Malacosoma, 327, 329; M. castrensis, 328 Mesolia, 244
Malacosomatinae, 327 Mesoscia, 164
Malacosomatini, 327, 329 Mesosemia, 284
Maliattha concinnimacula, 387 Mesoses optata, 16; M magna, 16
Maltagorea, 341 Mestolobes, 235, 244
Manataria, 279 Mestra, 290
Manduca, 347 Metachandidae, 157
Manila, 290 Metacrias, 447
Mapsidius, 140 Metalectra, 372, 374
Marae melanospila, 364 Metanomeuta, 121
Marasmia, 250 Metaphatus, 83, 85, 86; M. cirrhus, 82; M. ochraceous,
Marathyssa inficita, 387 82; M ochraceus, 84
Marcillada, 380 Metapherna, 92
Marmara, 113 Metaprotus, 219
Marmax, 229 Metarbela, 193
Marpesia petreus, 277 Metarbelianae, 36
Marshalliana, 193; M. bivittata, 190 Metarbelidae, 188
Maruca, 250 Metarbelinae, 188, 189, 190, 191, 193, 329
Marumba, 346 Metarbelodes umtaliana, 190
Meandrusa, 454 Metardaris, 272; M. cosinga, 271
Mechanitis, 293
Metaselena, 212
Mecyna submedialis, 237
Metathrinca, 139
Mediavia discalis, 237
Metaxmeste, 249
Meessia, 97
Metisella, 273
Meessiinae, 92, 95, 409 Metrea, 235
Megacephalomana remaudi, 368 Metriochroa, 114; M. psychotriella, 113
Megacraspedus, 147 Metriotes, 144
Megalopalpus, 285 Metriotinae, 157
Megalopyge, 164; M. lanata, 161 Metzneria, 136, 147
Megalopygidae, 38, 159, 160, 161, 164, 165, 166, 168, Metzneriinae, 158
169, 175, 188, 406, 450 Micraglossa, 244
Megalopyginae, 164,450 Micragone, 341
Megalorhipida defectalis, 224 Micrerethista, 94, 97
Megalura coresia, 266 Microblepsis, 305
Meganoia, 369, 390; M. minuscula, 383; M. triangu- Microcolona, 139
lis, 390 Microcorsini, 200, 201, 210-211
Megarthridiini, 242 Microcrambus, 244
Megastes, 250 Microdulia, 340
Megathymus, 269, 270-271, 274; M. yuccae, 271 Micromelalopha, 363
Melalopha, 363 Micronia, 309
Melanargia, 290, 437, 444; M. arge, 444; M. galathea, Microniinae, 308, 309
444; M. hylata, 444; M. ines, 444; M. lachesis, 444; Micropsychidae, 107
M. larissa, 444; M. occitanica, 444; M. pherusa, 444; Micropsychiniidae, 107
M. russiae, 444; M. syiaca, 444 Micropterigidae, 9, 33, 4 1 - 4 3 , 44, 45, 53, 404, 405,
Melanis, 284 413, 415, 447, 448, 449
Melanitini, 279 Micropterigoidea, 8, 9, 10, 11, 17, 27, 28, 29, 4 1 - 4 3 ,
Melanitis, 290 45, 46, 405, 413, 414
Melanolophini, 315 Micropterix, 41, 42, 43, 44, 45, 405, 447, 448, 449; M.
Melanothrix, 330 aureatella, 45; M. pervetus, 17
Melasina, 107 Midila, 246; M. daphne, 245
Melese, 392 Midiliformes, 234
Melinaea, 293 Midilinae, 234, 236, 239, 246
Melitaea, 288; M athalia, 291; M cinxia, 266 Miletinae, 266, 283, 2 8 4 - 2 8 5
Melitaeini, 286, 287, 288 Miletini, 266, 285
Melittia, 183, 184 Miletus, 283, 284, 285; M. symethus, 270
478 Index of Scientific Lepidoptera Names

Millieria, 219; M. dolosana, 221 Nacophorini, 315

Millieriinae, 219 Nadata gibbosa, 385
Mimacraea, 284 Napeogenes, 293
Mimallo, 324; M. amilia, 324 Napta Serratilinea, 325
Mimallonidae, 34, 321, 322, 323-324, 325, 328, 412, Naranga diffusa, 379
450 Narosa, 167, 425
Mimalloninae, 324 Narycia, 107; N. monilifera, 93
Mimallonoidea, 10, 14, 28, 30, 321, 322, 3 2 3 - 3 2 4 Naryciinae, 107
Mimoniades, 269, 273 Nataxa, 326
Mimopsestis, 304 Nathalis, 281
Mimozethes argentilinearia, 305, 306 Nedra, 369, 384
Miresa, 167 Nelcynda, 305
Mirificarma, 147 Nemapogon, 95, 99; N. granella, 93, 101, 103
Mirino, 321, 336; M. christophi, 339, 343, 347 Nemapogoninae, 97, 99, 409
Mirinidae, 321, 323, 336, 339, 343, 347, 450 Nemoria, 316
Miscera, 182 Nemoriini, 315
Mnesarchaea, 54, 58, 59; M. acuta, 56 Nematocentropus, 56
Mnesarchaeidae, 17, 54, 56, 5 7 - 5 9 , 58, 404, 407, 413, Nematopogon, 73, 75, 408; N. metallicus, 66; N. metax-
448, 450 ella, 74
Mnesarchaeoidea, 10, 11, 28, 29, 5 7 - 5 9 , 407, 450 Nematopogoninae, 75
Moca, 227 Nemaxera, 99
Mods, 374, 435 Nemophora, li, 75; N. amatella, 66; N. aursparselia,
Moerarchis, 94; M. australasiella, 93 66; N. raddei, 74, 84
Molippa, 340 Neoanathamna, 212
Momini, 385, 386 Neocastnia, 187
Mompha, 144; M. definitella, 137; M. franclemonti, Neocastniinae, 187
140, 142 Neocercophana, 340; N. philippii, 335, 339
Momphidae, 405, 411 Neochera, 375
Momphinae, 36, 37, 131, 134, 135, 144, 155, 156, 157, Neocoenyra, 290
411 Neoepiscardia, 99
Monochroa, 136 Neogrotella, 382
Monopis, 94, 95, 99; M. monachella, 93 Neohesperilla, 273
Monotrysia, 12, 65 Neohipparchini, 315
Moriloma, 144 Neolepidoptera, 10, 11, 28, 41, 55, 5 6 - 5 7 , 404
Morophaga, 99; M. bucephala, 93 Neomicropteryx, 42, 43
Morphinae, 267, 272, 287, 289-290, 292 Neope, 290
Morphini, 290 Neophasia, 279, 281
Morpho, 287, 290; M. achillei, 268; M. amathonte, 282 Neopreptos, 331
Morphopsis, 290 Neopseustidae, 11, 52, 54, 5 5 - 5 6 , 56, 404, 407, 448,
Morrisonia latex, 387 450
Munychryia, 324, 325, 326; M. senicula, 328 Neopseustis, 52; N. meyricki, 56
Munychryiinae, 325, 326 Neopseustoidea, 10, 11, 28, 29, 5 5 - 5 6 , 413
Musotima, 246; M. acclaralis, 245 Neoscythris, 140
Musotiminae, 234, 236, 240, 246 Neosphaleroptera, 206
Mustilia, 274, 334; M. falcipennis, 335 Neotheora chiloides, 56, 60
Mycalesini, 290 Neotheoridae, 56, 59, 60, 448, 450
Mycalesis, 290 Neorinops sepulta, 19
Mycelia, 288 Nephele, 348
Mydrodoxa, 379 Nephopterix, 243
Myelobia smerintha, 244 Nepticulidae, 34, 65, 67, 6 8 - 6 9 , 70, 71, 74, 76, 78, 80,
Mylantria, 387; M. xanthospila, 373 82, 84, 87, 404, 407, 415, 450
Mylothia, 146 Nepticulina, 65
Mylothris, 279, 281 Nepticulinae, 69
Myoglossata, 10, 11, 28, 53, 54, 55 Nepticuloidea, 10, 11, 12, 18, 28, 29, 41, 65, 6 7 - 7 1 ,
Myonia cingulina, 364 407, 450
Myrmecozela, 97 Neptis, 288, 289
Myrmecozelinae, 97, 409 Neritos, 392
Myscelus, 273 Nesara, 329
Mysoria, 273 Netrobalane, 273
Mythimna, 384, 432, 433, 435; M. adultera, 433; M. Neurophyseta, 246; N. completalis, 245
amblycasis, 433; M. convecta, 433; M. cunyada, 433; Neurosymploca, 174
M. dasuta, 433; M. idisana, 433; M. macrosaris, 433; Niasoma, 207, 208
M. nobbsi, 433; M. punctulata, 433; M. roraimae, Nidarini, 305
433; M. separata, 432, 433; M. sequax, 433; M. uni- Niepeltia, 18
puncta, 432, 433 Nigetia, 369
Niphograpta albiguttalis, 250
Nacaduba, 285 Niphopyralis, 249; N. sufftdalis, 248
Nacoleia, 250 Nisaga, 331; N. simplex, 330
Index of Scientific Lepidoptera Names 479

Nocloini, 381, 382 Odontiinae, 234, 235, 236, 239, 247, 249
Noctua, 1 Odontodes, 369
Noctueliopsis, 247 Oecia, 137, 141
Noctuidae, 19, 20, 21 ,37, 38, 355, 356, 357, 358, 359, Oeciinae, 134, 135, 141, 155, 156, 157
360, 361, 363, 366, 367-385, 387, 388, 390, 392, 393, Oecophoridae, 18, 36, 131, 133, 134, 137, 141-143,
410, 417, 429, 431, 434, 435, 450 157, 405, 410
Noctuinae, 360, 367, 369, 370, 371, 380, 382, Oecophorinae, 131, 133, 134, 141-143,155, 156, 157,
384-385, 390 435
Noctuini, 369, 384 Oedematopoda, 143
Noctuites, 355 Oegoconia, 146
Noctuoidea, 10, 14, 21, 28, 30, 355-394, 413, 418 Oeneis, 290
Nodozama, 391 Oenochroma, 315; O. vinaria, 311
Nola, 389; . minna, 373; . sorghiella, 389 Oenochrominae, 311, 312, 313, 315
Nolidae, 355, 356, 358, 359, 360, 362, 366, 367, 369, Oenochromini, 315
370, 373, 375, 383, 386, 388-389, 390, 392 Oenosandra, 355, 356, 357, 360, 361, 362, 369, 386; O.
Nolinae, 38, 359, 366, 383, 386, 388-389, 390 boisduvalii, 364
Nomophila, 250; N. noctuella, 235, 250 Oenosandridae, 38, 355, 356, 357, 360, 364, 365, 366,
Noorda, 247; N. blitealis, 248 367
Noordinae, 234, 240, 247 Ogdoconta, 370, 381
Norape, 164; N. ovina, 175 Oglasa, 380
Nordstromia, 305 Ogmograpis scribula, 112
Nossa, 302 Ogovia, 361
Notiopostega, 69, 71, 407, 415; N. atraa, 67, 78, 85 Ogygioses, 59, 60
Notodontidae, 19, 37, 38, 355, 356, 357, 358, 361-365, Ogyris, 285
366, 367, 369, 370, 374, 376, 377, 378, 380, 381, 383, Oidaematophorus rogenhoferi, 221
384, 385, 386, 387, 390, 391, 392, 412, 417, 429, Oiketicinae, 107
431, 434 Oiketicoides, 107
Notodontinae, 363, 364, 387, 390 Oiketicus, 107; O. aristocosma, 105, 106; O. geyeri,
Notodontini, 363, 364 106; O. kirbyi, 96
Nyceryx, 348 Oinophila, 95, 99
Nyctelius, 271 Oinophilidae, 99
Nyctemera, 361, 367, 386; N. apicalis, 383 Olceclosera, 334
Nyctemerini, 383, 392, 394 Olera, 293
Nycteola, 361 Olethreutes, 211
Nycteolidae, 38 Olethreutinae, 199, 200, 201, 202, 203, 204, 2 1 0 - 2 1 3
Nymphalidae, 19, 20, 38, 39, 174, 265, 266, 268, 269, Olethreutini, 200, 202, 203, 210, 211, 212
274, 275, 276, 277, 278, 279, 281, 282, 284, 285-293, Oliera, 79; O. argentinana, 70
286, 291, 365, 366, 384, 413, 415, 417, 450, 451 Oligamatites, 355
Nymphalinae, 19, 20, 267, 287, 288-289, 291 Oligocentria, 364
Nymphalini, 288 Olindia, 209
Nymphalis, 1, 288 Omiodes, 250
Nymphicula, 235, 247 Ommatothelxis, 139
Nymphidium, 284 Omorphinini, 377
Nymphula, 247 Omphaliodes, 326
Nymphulinae, 234, 235, 236, 239, 246-247, 412 Omphax plantara, 311
Nymphulini, 247 Omphisa, 250
Nystaleinae, 357, 364, 365, 376, 387 Onceropyga, 173; O. anelia, 175
Nyx, 219 Oncocnemidini, 380-381
Oncopera, 61, 62
Oberthueriini, 334 Oneida luniferella, 241
Obtectomera, 10, 12, 13, 14, 28, 217, 219, 222, 224, Oospila, 316; O. albicoma, 314
225, 227, 406, 408, 412 Operophtera brumata, 312, 316
Ochlodes, 269, 274 Operophterini, 312, 316
Ochrognesiini, 315 Ophiderinae, 374
Ochromolopinae, 222 Opodiphlhera, 338, 341
Ochromolopis, 222 Opogona, 18, 92, 94, 99; O. nipponica, 93
Ochropacha, 305; O. duplaris, 303 Opostega, 67, 69, 71; O. salaciella, 74
Ochsenheimeria, 124; O. bubaleila, 123; O. vacculella, Opostegidae, 34, 41, 67, 68, 6 9 - 7 1 , 70, 74, 76, 78, 80,
124 82, 84, 85, 404, 407, 415, 436, 450
Ochsenheimeriidae, 405, 408, 409 Oposteginae, 71
Ochsenheimeriinae, 35, 119, 124, 450 Opostegoides, 69, 71; O. gephyraea, 67; O. scioterma,
Ochyrotica, 224; O. concursa, 221, 224 84
Ochyroticinae, 224 Opostegoidinae, 71
Ocinara, 334; O malagasy, 335 Opsiphanes, 290
Ocnerostoma, 122 Oreixenica, 290
Ocnogyna, 447 Orenaia, 249
Odiles, 143 Oreta, 305
Odonestis, 326 Oretini, 303, 304, 305
480 Index of Scientific Lepidoptera Names

Organodesma, 100 Pancalia, 147

Orgyia, 370, 386, 388; O. postica, 390; O. pseudotsu- Pandemis, 207
gata, 380, 388 Panemeria, 382
Orgyiini, 386 Panolis flammea, 384
Orgyinae, 386 Panosmia, 454
Oriodryas, 212 Pansepta, 139
Ormetica, 392 Panthea, 385; P. furcilla, 371, 390; P. pallescens, 390
Ormiscodes, 340; O. amphinome, 340 Pantheidae, 355, 356, 360, 361, 366, 367, 370, 371, 375,
Orna, 174; O. pecinicornis, 171 379, 381, 384, 385-386, 389, 390
Ornalivalva, 147 Panthuama, 372
Orneodes, 223 Pantoctaenia, 167
Ornichinae, 114 Pantographa acoetesalis, 248
Ornithoptera, 277, 454; O. alexandrae, 277; O. goli- Papaipema eryngii, 371
ath, 277 Papilio, 1, 277, 279, 454; P. dardanus, 277; P. demodo-
Orophia, 143 cus, 425; P. glaucus, 277, 455; P. machaon, 264, 286;
Orthenches, 119 P. memnon, 277; P. polytes, 277; P. ulysses, 282; P.
Orthocomotis, 206, 208 xuthus, 441
Orthogonioptilum, 341 Papilionidae, 19, 37, 39, 174, 264, 265, 269, 274, 275,
Orthotaelia, 121; O. sparganella, 123, 125 276, 276-279, 282, 286, 291, 292, 302,413,417,428,
Orthotelidae, 409 429, 450, 451,453, 455
Orthoteliinae, 119, 120, 125 Papilioninae, 265, 278-279, 291, 453, 454
Osminiini, 182, 184 Papilionini, 453, 454
Osrhoes, 58, 60; O. coronta, 56 Papilionoidea, 10, 14, 19, 28, 29, 257, 263, 267, 268,
Ostrinia, 250; O. nubilalis, 237 269, 274-293, 413
Othreis, 374 Parabarrovia, 380
Ourapterygini, 315 Parabraxas, 302; P. davidi, 303, 306
Oxycanus, 62 Parachma ochracealis, 245
Oxychirota, 222; O. paradoxa, 221 Paraclemensia, 81, 83; P. acerifoliella, 66, 85
Oxychirotidae, 233, 234, 450 Paracolax, 372
Oxycnemis, 380 Paracomotis, 206
Oxynetra, 272 Paracossus, 190
Oxysemaphora, 202 Paradaemonia, 340
Oxy stola, 143 Paradeuterogonia, 139
Oxytenidae, 321, 336, 450 Paradirphia, 340
Oxyteninae, 39, 337, 338, 340 Parafomoria, 69
Oxytenis, 340 Parahypopta, 192
Oxythres splendens, 378 Paralbara, 304
Paralethe, 287
Pachliopta, 277, 278, 279, 302, 454 Paraleucoptera, 128; P. coffeella, 128; P. meyricki, 128
Pachnistis, 146 Paralipsa, 242
Pachymeta, 329 Parallelia, 374
Pachymetana, 329 Paralopostega, 69, 71; P. callosa, 82
Pachynoa, 250 Parametriotes bipunctella, 140, 148
achy odes, 315 Parametriotinae, 157
Pachypasa, 329 Parametriotini, 139
Pachypodistes, 242 Paramorpha, 226
Paedes, 377 Paramyelois, 243
Pagara, 391 Paranthrene, 183
Palaeolepidopterix, 16, 28 Paranthreninae, 184
Palaeosabatinca, 17 Paranthrenini, 182, 184
Palaeoses, 60; P. scholastica, 56 Parantica, 291, 292, 452, 453; P. Corinna, 451; P. crow-
Palaeosetidae, 56, 58, 59, 60, 448, 450 leyi, 453; P. menadensis, 453; P. phyle, 453; P. pumila
Palaeotoma, 205 453; P. schenkii, 453; P. sita, 451, 453; P. toxopei,
Palaephatidae, 36, 67, 70, 74, 76, 78, 80, 82, 8 3 - 8 6 , 453; P. weiskei, 452
84, 404, 408, 449, 450 Parapoynx, 247; P. diminutalis, 247; P. seminealis, 248
Palaephatoidea, 10, 11, 12, 28, 29, 65, 8 3 - 8 6 , 408, Parapsestis, 305
413, 450 Pararge, 290
Palaephatus, 83, 86; P. falsus, 67, 70, 74, 76, 78, 80 arasa, 167
Palirisa, 330 Parasabatinca, 16, 17
Palla, 289 Paratalanta, 250
Pallini, 289 Parategeticula, 75, 77, 79; P. pollenifera, 66
Palpita, 250 Pardopsini, 288
Palumbina, 147 Pardopsis, 288
Palumbininae, 158 Parectopa, 114; P. robiniella, 113
Pammeces, 139 Parides, 278, 279, 302, 454
Pammene, 213 amara, 271
Panacela, 331 Parnassiinae, 19, 265, 278, 413, 437, 438, 453, 454, 455
Panacelinae, 331 Parnassiini, 454
Index of Scientific Lepidoptera Names 481

Parnassius, 4, 19, 257, 277, 278, 453, 454, 455, 456; P. Phiditia, 334, 387; P. maculosa, 332; P. minor, 347
apollo, 264, 278, 286; P. mnemosyne, 278 Phiditiinae, 331, 332, 333, 334
Parornix, 114 Phidotricha, 243
Parotis, 250 Phigalia titea, 314
Parthenos, 289 Philampelinae, 346
Pasiphila, 439, 447 Philampelini, 347
Passova, 273 Philobota, 143
Pathysa, 454 Philodoria, 114
Patissa, 246 Philoros, 392
Pazala, 454 Philotherma, 329
Pectinivalva, 67, 68, 69; P. commoni, 80 Phoberia atomaris, 383
Pectinivalvinae, 69, 407, 415 Phobetron pithecium, 175
Pectinophora, 149; P. gossypiella, 138, 142 Phocides, 269
Pedaliodes, 290 Phoebis, 281
Pediasia, 244 Phoenicladocera, 329
Peleopoda, 146 Phormoestes, 219
Peleopodidae, 135, 136, 138, 146, 157, 405 Phosphila, 384
Pelepodinae, 155, 157 Phosphilini, 384
Penestoglossa, 107 Phostria, 250
Penetes, 290 Phragmataecia, 191, 193
Penicillaria, 377 Phragmatobia, 392; P. flavata, 447; P. parvula, 447; P.
Pennisetiini, 183 thursbyi, 447
Penthophera, 388; P. morio, 386 Phricanthes, 205; P. asperona, 200
Pentobesa, 376 Phricanthini, 199, 200, 201, 203, 204, 205, 411, 412
Peoria, 243; P. roseotinctella, 245 Phryganidia californica, 376, 390
Peoriini, 238, 243 Phthorimaea, 147
Peragrarchis, 226, 228 Phthoropoeinae, 97
Pericopidae, 392 Phtonerodes, 139
Pericopini, 373, 392, 394 Phyciodes, 286, 288
Perdromo, 375 Phycitinae, 233, 234, 235, 238, 241, 242, 243, 406, 412
Periga, 337, 340 Phy codes, 182
Perigea, 370, 381 Phycomorpha, 226
Perigonia, 348 Phylledestes, 355
Perimede, 146 Phyllocnistinae, 16, 18, 114
Perinetoides, 242 Phyllocnistis, 114, 410; P. insignis, 108, 110, 111; P. po-
Periphoba, 340 puliella, 96; P. vitegenella, 113
Periploca, 146 Phyllodesma, 329
Perissomasticinae, 94, 99 Phyllonorycter, 113, 114; P. blancardella, 96, 111
Perissomastix, 99 Phylloxiphia, 346
Perittia, 139; P. eremonoma, 137, 148 Phylomictis, 136
Pero, 315 Phymatopus, 58, 61
erla, 167 Phyprosopus, 374
Persectania aversa, 371 Physcaneura panda, 291
Persicoptila, 147 Phy soptila, 138, 147; P. pinguivora, 148
Perthida, 66, 80, 81 Physoptilinae, 133, 135, 136, 147, 156, 158
Petrophila, 247; P. jaliscalis, 248 Pierella, 290
Pexicopia, 149 Pieridae, 19, 39, 174, 264, 265, 274, 275, 276, 279-281,
Pexicopiinae, 133, 135, 136, 147-149, 156, 158 282, 286, 291, 413, 450
Phaedropsis, 250 Pierinae, 265, 280, 281, 413
Phaegopterinae, 392 Pierini, 280
Phaegopterini, 373, 392, 394 Pieris, 4, 275, 280; P. brassicae, 264, 281, 286; P. ra-
Phaeoses, 99 pae, 281
Phalacra, 305 Pigritia, 144
Phalaena, 1, 233 Pigritiinae, 157
Phalaenoides glycinae, 379 Piletocera, 250
Phalanta, 288 Pilocrocis, 250
Phalera, 363, 366 Placodoma, 107
Phalerinae, 357, 358, 363, 364, 365, 387 Placodominae, 107
Phalonidia, 206 Placosaris, 250
Phaos, A4! Plagiomimicus, 382
Pharmacis, 61, 62 Planotortrix, 207
Pharmacophagus, 453, 454 Platagarista tetraplaura, 379
Phasis, 285 Plathypena, 376, 380
Phassus, 62 Platychasma, 363, 366; P. virgo, 364
Phauda, 174; P. cantonensis, 171 Platychasmatinae, 358, 363, 364
Phaudinae, 160, 163, 170, 171, 172, 173, 174 Platymota, 208
Phenacodes, 235, 244 Platyptilia, 224; P. carduidactyla, 224
Phiala, 331; P. fuscodorsata, 325 Platyptilinae, 223
482 Index of Scientific Lepidoptera Names

Platysenta, 370, 381 Procometis, 146

Plebejus, 1, 285 Procridinae, 170, 171, 172, 173, 174
Plesiophatus, 85, 86 Procris statices, 444
Pleurota, 143 Proditrix tetragma, 123
Pleurotinae, 157 Prodoxidae, 34, 35, 66, 67, 70, 71, 74, 7 5 - 7 9 , 76, 80,
Plodia interpunctella, 243 84, 86, 404, 407, 408, 450
Plotheia, 389 Prodoxinae, 7 7 - 7 9 , 408
Plumana, 107 Prodoxoides, 75, 77, 408
Plusia, 377 Prodoxus, 76, 77, 79; P. quinquepunctellus, 66, 80, 84;
Plusiinae, 360, 367, 368, 369, 370, 375, 3 7 7 - 3 7 9 , 381, P. y-inversus, 66, 81
382, 387, 388, 390 Prodryas persephone, 19, 20
Plusiini, 3 7 7 - 3 7 8 Proeulia, 208
Plusiniini, 305 Prolyonetia, 18
Plutella, 121, 124, 125; P. xylostella, 123 Pronophila, 290
Plutellidae, 18, 35, 36, 119, 120, 121, 123, 124, 217, Prophyllocnistis, 113
409, 410 Prorifrons, 329
Pococera, 243 Proselena, 205
Podalia, 164 Prosetomorpha, 99
Podosesia, 183 Protaephagus, 81
Poecilasthena, 439, 441 Protambulyx, 346
Poecilmitis, 285 Proterospastis, 99
Poecilocampa, 274, 326, 327, 329; P. alpina, 328 Prothoe, 289
Poecilocampinae, 326, 327, 328, 329 Prothoini, 289
Polia, 384 Protobactra, 211
Poltes, IIA Protographium, 453, 454
Pollanisus, 172, 173, 174 Protolechia, 147
Polydactylini, 305 Protolepis cuprealata, 16, 17
Polygonia, 288 Protoploea, 451, 452
Polygrammodes, 250 Protopterna, 209
Polyhymno, 147; P. luteostrigella, 145 Protolheora, 58, 60; P. petrosema, 56
Polylopha, 209; P. epidesma, 200, 202, 203 Prototheoridae, 56, 58, 59, 60, 448, 450
Polyommatinae, 285 Pryeria, 174; P. sinica, 171
Polyommatini, 285 Psacaphorinae, 157
Polyommatus, 285; P. icarus, 270 saltica, 141
Polyortha, 209 Psaphida electilis, 357; P. thaxteriana, 371
Polyorthini, 199, 200, 202, 203, 204, 209 Psaphidinae, 357, 360, 369, 370, 371, 381, 382, 387
Polyploca, 305 Psaphidini, 371, 381
Polyplocini, 305 Psara, 250
Polypogon, 372; P. inconspicualis, 387 Psephea, 375
Polyptychus, 346 Pseitdacraea, 286, 289
Polytela, 382 Pseudaegeria, 143
Polytelodes, 382 Pseudaletia, 384, 432, 433
Polythysana, 340 Pseudanapaea trigona, 175
Polythysanini, 338, 340 Pseudantheraea, 338
Polyura, 289 Pseudaphelia, 331
Poncelia, 363 Pseudapheliini, 341
Porela, 329 Pseudarbela, 107; P. semperi, 96
Poritia, 283, 284; P. hewitsoni, 270 Pseudargyrotoza, 207, 208
Poritiinae, 266, 283, 284 Pseudatteria, 209
Poritiini, 284 Pseudautomeris, 340
Praepapilio colorado, 19; P. gracilis, 19 Pseudergolis, 289
Praetaxila, 284 Pseuderotis, 146
Praezygaena, 174 Pseudethmia, 138
Praydinae, 120, 122 Pseudobankesia, 107
Prays, 121, 122; P. citri, 122; P. fraxinella, 123; P. Pseudobunaea, 341
oleae, 122 Pseudocomotis, 210
Prepona, 289 Pseudocossinae, 191
Preponini, 289 Pseudocossus, 182, 191
Preptos, 331 Pseudodirphia, 340
Princeps, 454 Pseudodoxia, 141
Prionapteryx, 244 Pseudojana, 330
Prionoxystus, 192 Pseudometa, 329
Prismosticta, 332, 333, 334 Pseudoplusia, 435; P. includens, 378
Prismostictinae, 332, 333, 334 Pseudopontia, 280; P. paradoxa, 264, 280, 281
Prismostictini, 334 Pseudopontiinae, 265, 280281
Prochola, 139 Pseudopostega, 69, 71; P. bistrigulella, 70, 76, 80, 82
Prochoreutis, 219 Pseudoschinia elautalis, 248
Proclesis, 143 Pseudosphex, 394
Index of Scientific Lepidoptera Names 483

Psidopalini, 305 Rachiplusia ou, 378

Psilocorsis, 146; P. cirrhoplera, 138 Ragada, 290, 291
Psycharium, 164, 165 Ragadiini, 290
Psyche, 107 Ramesa, 363
Psychidae, 18, 31, 91, 92, 93, 94, 95, 96, 98, 101, 103, Ramphis, 147
104-107, 170, 405,408,414 Raphia, 372; R frater, 387; R. pallula, 368
Psychinae, 107, 409 Raphiinae, 360, 368, 369, 370, 372, 387
Psychocampa, 324 Ratarda, 189; R. excellens, 191, 193
Psychoides, 100 Ratardidae, 188
Psychonoctua, 193 Ratardinae, 189, 191, 193, 450
Psychostrophia, 302; P. nymphidiaria, 306 Rebelia, 107
Pterocerota, 331 Rectiostoma fernaldella, 136; R. xanthobasis, 136
Pterodecta, 259; P. felderi, 260 Recurvara, 147
Pterogyne, 107 Reissita, 174
Pterolocera, 324, 326 Renia discoloralis, 368
Pterolonche, 138, 144 Rhabdotenes, 205
Pterolonchidae, 405 Rhagastis, 348
Pterolonchinae, 135, 144, 450 156, 157 Rhamphura, 140
Pteronycta, 372 Rhesala, 369
Pteronymia, 293 Rhigognostis, 125
Pterophoridae, 35, 38, 218, 221, 223-224, 233, 406, Rhinoprora, 447
435 Rhipidarctia danieli, 390
Pterophorinae, 223, 224 Rhodobates, 97
Pterophoroidea, 10, 13, 14, 19, 28, 30, 222, 223-224, Rhodobatinae, 97
233 Rhodometra, 316
Pterophorus, 19, 218, 224 Rhodometrini, 316
Pterostoma, 363 Rhodoneura, 229
Pterothysaninae, 258, 259, 331 Rhodoneurini, 229
Pterothysanus, 258, 259; P. laticilia, 260 Rhodostrophini, 316
Pterourus, 453 Rhomboceros, 207
Ptilodon kuwayamae, 364 Rhomboristini, 315
Ptilophorinae, 363 Rhynchopacha, 147
Ptilopsaltis, 103 Ridiaschinidae, 80
Ptiloscola, 340 Riodina, 284; R, lysippus, 270
Ptisanora, 81 Riodinella nympha, 19
Ptochoryctis, 146 Riodinidae, 274, 281
Ptychandra, 290 Riodininae, 265, 279, 282, 283, 284, 286, 287, 291
Ptycholoma, 207 Risobinae, 360, 373, 389, 390
Ptyssoptera, 67, 86 Risobini, 370
Pycnarmon, 250 Rivula, 376; R. propinqualis, 390
Pycnostola, 147 Rivulinae, 376
Pydnella, 363 Roelmana, 323
Pygaera, 363 Roelofa, 323
Pygaerinae, 358, 363, 364, 387 Roelmana maloba, 328
Pygoctenucha, 391 Roeslerstammia, 110, 409; R. erxlebella, 96, 108, 110
Pyralidae, 233, 234, 235, 238, 240-243, 244, 245, 310, Roeslerstammiidae, 96, 107, 108, 109-110, 405, 409
410, 412,413, 417, 435 Roeselia strigula, 369
Pyraliformes, 234 Rolepa, 334, 387
Pyralinae, 234, 235, 236, 239, 242-243, 406, 412 Rosema, 363, 364; R. deolis, 363
Pyralis, 1, 233, 243; P. farinalis, 245; P. manihotalis, Rothschildia, 340, 341
243 Rupela, 246; R. tinctella, 245
Pyraloidea, 9, 10, 14, 19, 28, 29, 36, 37, 131, 219, 228, Rusicada, 424, 426
230, 233-250, 257, 321, 406, 412, 414 Ruttellerona lithina, 441
Pyramidobela, 138
Pyrausta, 250; P. tyralis, 248; P. unifascialis, 248 Sabalia, 342; S. tippelskirchi, 328, 342
Pyraustinae, 217, 234, 235, 236, 240, 2 4 9 - 2 5 0 Sabaliinae, 342
Pyraustini, 250 Sabatinca, 17, 20, 43, 44, 45, 405, 445, 448, 449; S.
Pyrginae, 264, 272, 273 lucilia, 45
Pyrgus, 269, 273; P. malvae, 271 Sagalassa, 182, 183
Pyroderces, 147 Salagena, 193
Pyromorpha, 174; P. cuchumatana, 173 Salassa, 341; S. mesosa, 335
Pyrrhia umbra, 387 Salassinae, 338, 340, 341
Pyrrhopyge, 269, 272; P. araxes, 273; P. phidias, 271 Salatura, 452
Pyrrhopyginae, 263, 2 7 2 - 2 7 3 Salbia, 250; S. haemorrhoidalis, 250
Samanta, 279
Quadricalcarifera, 361 Samea multiplicalis, 250
Quentalia, 334 Samia, 341
Quercusia, 285 Sangala, 312
484 Index of Scientific Lepidoptera Names

Saptha, 219; S. smaragditis, 221 Selepa, 358, 370, 375, 388, 389
Sarbia, 273 Semanophorae, 346
Saridoscelinae, 120, 122 Sematura, 307; S. lunus, 306
Saridoscelis, 122, 123 Sematuridae, 301, 303, 3 0 5 - 3 0 7
Sarmalia, 331 Sematurinae, 307
Sarrothripinae, 388 Semioptila, 163; S. hilaris, 161
Sarrothripini, 389 Semioscopis, 138
Sarsina, 386 Semioscopidinae, 156
Sasakia charonda, 289 Semiothisa gambaria, 311
Sataspes, 346 Semiothisini, 312, 315
Satole, 242 Sena, 329
Saturnia, 337, 338, 341; S. pavonia, 340 Sericinus, 278, 454
Saturniidae, 39, 321, 322, 323, 335, 336-341, 347, Sericostola, 126
417, 450 Sesamia, 384
Saturniinae, 337, 338, 340, 341 Sesia, 184; S. apiformis, 182
Saturniini, 341 Sesiidae, 13, 35, 181, 182-184, 406
Satyrinae, 19, 38, 267, 272, 279, 286, 287, 290-291, Sesiinae, 182, 183-184, 346
292, 384, 413 Sesiini, 184
Satyrini, 290 Sesioidea, 10, 13, 19, 28, 29, 31, 181-188, 226
Satyrium, 285 Sesommata, 83, 85, 86; S. leuroptera, 67
Satyrus, 290; S. actaea, 268 Sesquiluna, 334
Scaeosophidae, 158 Setina aurita, 373
Scalidomia, 99 Setomorpha, 92, 94, 99
Scardia, 94, 99; S. anatomella, 98, 101 Setomorphinae, 99, 409
Scardiinae, 36, 94, 97, 99, 409 Sibine, 167
Sceliodes, 250 Siculodes triumphans, 228
Schacontia, 246 Siculodinae, 229
Schausiella, 337, 340 Siculodini, 229
Schiffermuelleria, 18 Siga, 235, 250
Schistomitra, 302; S. funeralis, 302 Silos ca, 100
Schistonoea, 141; S. fulvidella, 137 Siloscinae, 100
Schistonoeidae, 134, 135, 137, 141, 157 Simacauda, 83
Schistonoeinae, 134, 135, 141, 156, 157 Simaethis tricolor, 221
Schistophleps, 391; S. bipuncta, 390 Simaethistidae, 217-219, 221
Schizovalva, 147 Simaethislis, 217
Schizura, 364, 365; S. ipomoeae, 387; S. unicornis, 356 Simaethistoidea, 10, 14, 28, 30, 2 1 7 - 2 1 9
Schoenobiinae, 234, 235, 236, 239, 244, 245, 412 Similipepsini, 183
Schoenotenes, 205, 206 Simplicia, 372
Schoenotenini, 199, 200, 201, 205-206, 411 Simplimorpha, 69
Schreckensteinia, 127, 218, 220; S. festaliella, 220, 221 Simyra, 370, 372
Schreckensteiniidae, 35, 218, 220, 221, 406 Sinoprinceps, 454
Schreckensteinioidea, 10, 14, 28, 30, 220 Siskiwitia falcata, 131
Scieropepla, 139 Sisurcana, 208
Scirpophaga, 246 Sisyroxena, 226
Scodes, 147 Sitotroga, 149; 5. cerealella, 148
Scolesa, 337 Smerinthinae, 345, 346
Scolioplecta, 205 Smerinthus, 346
Scoliopteryx, 375 Snellenia, 143
Scoparia, 235, 236, 244 Sobanga rutilalis, 248
Scopariinae, 234, 235, 236, 239, 244, 412 Solenobia, 105, 107
Scopelodes, 167 Somabrachyidae, 38, 159, 160, 161, 1 6 4 - 1 6 5
Scopula, 316; S. aniara, 311 Somabrachys, 160, 163, 164, 165; S. aegrota, 161
Scopulini, 316 Sorhagenia, 146
Scoriodyta, 105, 107 Sorocaba, 333, 334, 387
Scoriodytinae, 107 Sorolopha cyclotoma, 200, 203
Scotinocerides, 168 Spalgis, 285
Scrancia, 363 Sparganopseustis, 208
Scranciini, 363 Sparganothidini, 412
Scrobipalpa, 147 Sparganothini, 199, 200, 2 0 7 - 2 0 8
Scrobipalpula, 147 Sparganothis, 208; S. pilleriana, 200
Scyrotis, 79, 81 Sparganothoides, 208
Scythrididae, 405, 410, 411 Speyeria, 288
Scythridinae, 35, 131, 132, 133, 134, 136, 139-141, Sphenograptis, 110
156, 157, 411, 450 Sphingidae, 39, 321, 322, 339, 341, 343, 344-348, 429,
Scythris, 139, 140; S. basilaris, 145; S. mixaula, 137 431, 434, 435, 438
Scythropia, 122; S. crataegella, 123 Sphingidites, 19
Scythropiinae, 120, 122 Sphinginae, 346, 347
Selania, 213 Sphingoidea, 321
Index of Scientific Lepidoptera Names 485

Sphingomorpha chlorea, 368 Styphlolepis, 246

Sphinx, 1, 347; S. ligustri, 343 Styx, 284; S. infernalis, 270, 284
Spialia, 273 Sumatratarda, 193
Spica, 303, 304, 305 Surattha, 244
Spiladarcha, 220 Surcossus, 192; S. per lar is, 191
Spilarctia, 392 Susica, 167
Spilomelinae, 234 Swammerdammia, 122
Spilomelini, 250 Syllectra erycata, 368
Spilonota, 212 Syllepte, 250
Spilosoma, 392; S. dubia, 390 Symmachia, 284
Spindasis, 285 Symmerista, 365
Spiramiopsis, 342, 344; S. comma, 343, 344 Symmoca, 146; S. oenophila, 140
Spodoptera, 369, 382, 384, 424, 435; S. exempta, 436; Symmocidae, 405
5. mauritia, 390 Symmocinae, 134, 135, 146, 156, 157
Spoladea, 250 Symphytophleps, 309
Spragueia marmorea, 387 Synallagma, 144
Spuleria, 139 Synanthedon, 183; S. coccidivora, 183; S. exitiosa, 184;
Squamicornia, 42, 43, 44 S. pictipes, 184; S. spheciformis, 182; S. tipuli-
Squamura, 193 formis, 184
Staetherinia, 362 Synanthedonini, 184
Stagmatophora, 147 Synchlorini, 315
Stalachtis, 284 Synclera, 235
Stamnodini, 316 Synegia, 439
Staphylinochrous, 163, 164 Synemon, 186, 188; S. collecta, 187; S. magnifica, 186,
Stathmopoda, 143; S. lychnacma, 137; S. pedella, 142, 187; S. nupta, 186; S. plana, 186, 187
145 Synemonini, 186, 188
Stathmopodidae, 405 Synempora, 52, 56
Stathmopodinae, 133, 134, 143, 156, 157, 410 Syngamia florella, 248
Stathmopolitinae, 100 Syngrapha, 366, 378
Stathmopolitis, 100 Syntominae, 355, 358, 373, 383, 390, 3 9 1 - 3 9 2
Stauropolia nekrutenkoi, 20 Syntomini, 373, 391, 392
Stauropus, 364 Syntomis , 383
Stegasta, 147 Syringopainae, 134, 135, 143-144, 156, 157
Stegommata, 128 Syringopais temperatella, 137, 145
Stemorrhages, 250 Syssphinx, 340
Stenoglene, 331
Stenoma, 136 Taeniarchis, 207, 209
Stenomatidae, 405, 410, 411 Tagiades, 273
Stenomatinae, 36, 131, 133, 134, 136, 156,411 Talaeporia, 107
Stenoptilia, 224 Taleporia, 107
Stephensia, 139 Taleporiinae, 409
Slericta, 243 Taleporinae, 107
Sterrila, 310 Talis, 244
Sterrhinae, 311, 313, 314, 316 Tancoa calitas, 237
Sterrhini, 316 Tanyenema, 222; T. anomala, 221
Sthenopis, 61; S. argenteomaculatus, 61 Tanyzancla, 143
Stibochiona, 287, 289 Taraka, 285
Stictoptera, 377; S. griveaudi, 368 Tarchon, 333, 334
Stictopterinae, 360, 368, 369, 370, 375, 376-377, 417, Targalla, 377
431 Tarsolepis, 363; T. taiwana, 364
Stigmella, 18, 69, 87; S. juglandiella, 82; S. malella, 67 Tascina, 187; T. metallica, 185; T. nicevillei, 185
Stilbosis, 146 Tascininae, 187
Stiphrometasia, 246 Tatochila, 281
Stiria, 382 Tebenna, 219
Stiriinae, 360, 370, 371, 381, 382, 387, 390 Tecmerium, 144
Stiriini, 371 Tegeticula, 75, 77, 79; T. yuccasella, 66, 70, 72
Stoeberhinus, 146; S. testaceus, 138 Tegostoma, 247
Stomopteryx, 147 Teichobiinae, 100
Stomopteryginae, 158 Teinopalpini, 454
Stomphastis, 114 Teinopalpus, 278, 279, 453, 454
Streblote, 329 Telchin licus, 186
Strepsicrates, 212; S. semicanella, 202 Teldenia, 305
Strepsimanes, 374 Telecrates, 139
Strepsimaninae, 360, 368, 370, 3 7 2 - 3 7 4 Telethera, 110
Striglina, 225, 229 Telicota, 274
Striglininae, 229 Telipna, 284
Striphnopteryx, 331 Tellervinae, 287
Strymon, 285 Tellervini, 292, 451, 452
486 Index of Scientific Lepidoptera Names

Tellervo, 292 Tinea, 1, 94, 95, 99, 233; T. pellionella, 93, 98, 106
Telphusa, 147 Tineidae, 18, 33, 35, 36, 81, 91, 92-100, 101, 103, 106,
Temnora, 348 405, 409, 435
Templemania, 208 Tineinae, 94, 95, 99, 409
Tephroniopsis, 222 Tineodes, 222; T, pterophoralis, 221
Tepilia, 334, 387 Tineodidae, 37, 221, 222, 223, 233, 234, 450
Teragra, 193 Tineoidea, 10, 12, 13, 18, 28, 30, 91-107, 217, 321,
Terastia, 250 403, 408, 413, 414
Terpne, 315 Tincla, 95, 99; T. bisselliella, 93
Terpnini, 315 Tinissa, 94, 99
Tetanocentria, 137, 139 Tinissinae, 99
Tethea, 305; T. ocularis, 306 Tinthia myrmosaeformis, 182
Tetheini, 305 Tinthiinae, 183
Tetragma, 15, 11 Tinthiini, 183
Tetragonus, 259; T. catamitus, 260; T. lycaenoides, 258 Tiquadra, 92, 94, 99; T. aeneonivella, 93
TTiaites, 19, 20 Tiracela, 358
Thalaina clara, 311 Tiradelphe, 452, 453
Thalamarchellidae, 156 Tirathaba, 242
Thalassodini, 315 Tirathabini, 242
Thalatha, 370 Tirumala, 292, 452, 453
Thalathoides, 370 Tischeria, 87; T. citrinipennella, 80; T. malifoliella, 76,
Thambotricha, 218, 221, 222 78, 82, 84; T. quercitella, 74; T. zelleriella, 67, 70
Thaumatographa, 202, 210; T. excellens, 200 Tischeriidae, 34, 67, 70, 74, 76, 78, 80, 82, 84, 8 6 - 8 7 ,
Thaumetopoea, 274 87, 404, 408, 415, 450
Thaumetopoeinae, 38, 357, 358, 362, 363, 364, 387 Tischerioidea, 10, 12, 28, 29, 30, 65, 8 6 - 8 7 , 408, 413,
Thaumatopsis, 244 450
Theages, 392 Tisis, 143; T. chalybaeella, 137
Thecla, 270, 285 Tissanga, 331
Theclinae, 285 Tissanginae, 331
Theclinesthes, 285 Titaea timur, 335
Theclini, 283, 285 Titanio, 247
Thecobathra, 121 Tithorea, 293
Theope, 284 Titya, 329
Theretra, 348 Tollophorinae, 157
Thereutis, 110 Tolype, 329
Thermochrous, 164; T. stenocraspis, 161 Tolypida, 323; T. amaryllis, 328
Thiacidas, 372 Tonica, 138
Thiotricha, 149 Torodora, 143
Thitarodes, 62 Torodorinae, 143, 157
Thliptoceras, 250 Torone, 227
Thosea, 167 Tor tilia, 143
Thudaca, 139 Tortricidae, 36, 81,199-213, 406, 413, 414, 435
Thyatira, 305 Tortricinae, 199, 200, 201, 202, 203, 204-209, 210,
Thyatiridae, 302 411, 412, 447
Thyatirinae, 37, 303, 305, 381 Tortricini, 199, 200, 201, 204, 205, 411
Thyatirini, 305 Tortricoidea, 10, 13, 14, 19, 28, 31, 199-213, 404,
Thymaridae, 163 411, 413
Thyretini, 373, 391-392 Tortrix, 1, 205
Thyrididae, 19, 36, 225, 227, 228, 228-229, 230, 233, Toxidia, 273
406, 450 Trabala, 329
Thyridinae, 229, 304 Tracholena, 202, 205, 206
Thyridoidea, 10, 14, 28, 219, 228-229, 230, 257 Trachydora, 147
Thyridopteryx, 107; T. ephemeraeformis, 96, 98, 103, Trachysmia, 206
106 Trapezites, 273, 291; T. symmomus, 271
Thyris, 225, 228 Trapezitinae, 265, 272, 273, 291, 451
Thyrocopa apatela, 139 Triaxomera, 99
Thysania agrippina, 374 Trichaea, 250
Thysanoplusia orichalcea, 378 Trichiura, 329
Timaea, 94 Trichophaga, 99; T. swinhoei, 95 T. tapetzella, 93
Timandrini, 316 Trichophassus, 62
Timandromorphini, 315 Trichoplusia ni, 378
Timocratica, 136; T. melanocosta, 13 Trichopterygini, 316, 439, 447, 450
Timyridae, 157 Trichosea, 385
Timyra pastas, 142, 145 Triclonella, 147 T. xuthocelis, 138
Tinacrucis, 208 Trictena, 62
Tinaegeriidae, 157 Tridentaforma, 15, 77; T. fuscoleuca, 76
Tinagma, 34, 110; T. balteolellum, 110; T. obscurofas- Tridrepana, 304, 305
ciella, 108, 110; T. perdicillum, 96 Trifurcula, 69, 87
Index of Scientific Lepidoptera Names 487

Trifurculini, 68 Whittleia, 107

Triocnemidini, 380, 381 Wingia, 143
Triodia, 62 Wiseana, 62
Tripteridia, 441 Wockia, 220
Tripudia, 369 Wurthia, 249
Triscaedecia, 223; T. dactyloplera, 221 Wurthiinae, 234, 236, 239, 2 4 7 - 2 4 9
Trissodoris, 147
Trisuloides, 361, 385, 386 Xanthodes, 379; X. transversa, 371, 385, 387
Trogonoptera, 454 Xanthopan, 347
Trogoptera, 324; T. erosa, 328; T. notata, 325 Xanthopastis, 382; X. limais, 387
Troides, 277, 219, 453, 454; T. helena, 264 Xanthorhoini, 312, 316
Troidini, 279, 413, 439, 453, 454, 455 Xenosphingia, 345, 346; X. jansei, 343
Trosia, 164 Xestia, 438; X. triangulum, 371
Trosiinae, 164 Xubida, 244
Trotosema, 372 Xylenini, 369, 384
Trymalitis, 200, 201, 210; T. climacias, 199, 202 Xyleutes, 193, 195; X. encalypti, 191; X. strix, 190
Tympanistes, 370; T. fusimargo, 390 Xyleutites miocenicus, 18
Tympanota, 441 Xylophones, 347, 348
Typhoniidae, 107 Xylorycta, 139; X. thrasycosma, 137
Tyria, 391; T. jacobaeae, 394 Xyloryctidae, 35, 134, 136, 137, 115, 139-141, 157,
405, 410, 411
Udea, 235, 250 Xyloryctinae, 35, 133, 134, 139, 156, 157
Ufeinae, 360, 367, 369, 370, 371, 372, 384, 385, 390 Xyrosaris, 122
Ufeus, 384; U. carnea, 384; U. satyricus, 371, 385, 390
Ulmara, 323; U. rotunda, 328 Yponomeuta, 121, 122, 416, 418; Y. padella, 123
Ukamenia, 211 Yponomeutidae, 36, 119, 120, 124, 127, 405, 409, 410,
416
Undopterix, 16, 17, 28
Yponomeutinae, 36, 120122, 123
Undulambia, 246
Yponomeutoidea, 10, 12, 13, 18, 28, 30, 107, 119-128,
Urania, 274, 278, 282, 307, 308, 310; U. boisduvalii, 306
Uraniidae, 14, 37, 301, 303, 305, 306, 307-310, 417, 131, 132, 169, 222, 405, 408, 409
450 Ypsolopha, 121, 124; Y. dentella, 123
Ypsolophidae, 35, 119, 120, 121, 123, 124, 410
Uraniinae, 308, 309
Ypsolophinae, 35, 124
Uranioidea, 321, 429, 434
Ypthima, 290
Uranothauma, 285
Yramea, 288
Urapteritra, 309; U. piperita, 303, 306
Urapteroides, 309
Zacorisca, 203
Urbanus proteus, 271
Zale, 374
Uresiphita reversalis, 248
Zanola, 334
Urodela, 144
Zapyrasta, 144
Urodidae, 37, 217, 220, 221, 406 Zaretini, 289
Urodoidea, 10, 28, 31, 220 Zaretis, 289
Urodus, 220; U. venatella, 221 Zatrephes, 392
Utetheisa, 391, 392, 394, 435; U. pulchelloides, 436 Zebeeba, 376
Utetheisini, 373, 392, 394 Zeiraphera, 212
Uzucha, 139 Zelleria, 122, 123
Uzuchidae, 157 Zelotypia, 62; Z. stacyi, 61
Zenophassus, 61
Vanessa, 288, 432, 433; V. amerindica, 19; V, atalanta, Zerynthia, 278, 454; Z. polyxena, 444; Z. rumina, 444
432, 433; V. buana, 433; V. dejeanii, 433; V. indica, Zerynthiini, 445, 454, 456
19, 432, 433; V. mounseyi, 433; V. polychloros, 266; V. Zethera, 290
samani, 433; V. tameamea, 432, 433; V. vulcania, 433 Zeugloptera, 9, 27, 405, 413
Vanessini, 288 Zeuxidia, 286, 290
Vanna, 99 Zeuzera, 193
Vespina, 81; quercivora, 83, 85 Zeuzerinae, 36, 188, 189, 190, 191, 192, 1 9 2 - 1 9 3
Viminia, 372 Zizeeria, 285
Vindula, 288 Zizina, 285
Virbia, 358, 392; V. rosenbergi, 383 Zizula, 285
Vitessa, 242; V. splendida, 245 Zonosomini, 316
Zurobata vacillans, 371
Walshia, 146; W. miscecolorella, 138 Zygaena, 18, 173, 174-177, 274; Z. carniolica, 173; Z.
Walshiidae, 157 trifolii, 171, 175, 176
Watsonalla binaria, 303 Zygaenidae, 38, 159, 160, 163, 169, 170-177, 193, 229,
Wegneria, 100 406, 450
Westermannia anchorita, 373 Zygaeniformia, 159
Westermanniinae, 360, 373, 389 Zygaeninae, 170, 171, 172, 173, 174-177
Whalleyana, 229; IV vroni, 228 Zygaenoidea, 10, 13, 18, 28, 31, 159-180, 181, 188,
Whalleyanidae, 228, 2 2 9 - 2 3 0 429, 434
Whalleyanoidea, 10, 14, 28, 30, 2 2 9 - 2 3 0 Zygaenoprocris, 174
 Instructions for the Authors
Handbook of Zoology
Volume IV Arthropoda: Insecta
The Handbook of Zoology, Vol. IV, Arthropoda:
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Handbook of Zoology, Volume IV Arthropoda: Insecta

Volume IV Arthropoda: Insecta will appear in parts with consecutive numeration in connection with the previous numera-
tion of the instalments. Beginning with 28 each part will be delivered bound.

Already published

Author Title Contribution Instalment

Alfred Kaltenbach Embiodea (Spinnfer) 8 1

Fritz Zumpt Human- und veterinrmedizinische Entomologie 9
Erwin Schimitschek Insekten als Nahrung, in Brauchtum, Kult und Kultur 10 2
Douglas St. Quentin
& Max Beier Odonata (Libellen) 6 3
Max Beier Mantodea (Fangheuschrecken) 12 4
Hermann Priesner Thysanoptera (Physopoda, Blasenfer) 19 5
Max Beier Phasmida (Stab- oder Gespenstheuschrecken) 10 6
Joachim lilies Ephemeroptera (Eintagsfliegen) 5 7
Maximilian Fischer Die Verwandlung der Insekten 16 8
Max Beier Diagnose 1
Max Beier Geschichte der Entomologie 2
Boris B. Rohdendorf Palontologie 3 9
Boris B. Rohdendorf Phylogenie 4
Max Beier Klassifikation 5
Stefan von Kler f Mallophaga (Federlinge u. Haarlinge) 17 10
Herbert Franz & Max Beier Die geographische Verbreitung der Insekten 6 11
Friedrich Schaller Collembola (Springschwnze) 1 12
Herbert Weidner Isoptera (Termiten) 14
Herbert Weidner Zoraptera (Bodenluse) 15 13
Heinz Janetschek Protura (Beintastler) 3 14
Ragnar Kinzelbach Strepsiptera (Fcherflgler) 24
Horst Aspck
& Ulrike Aspck Raphidioptera (Kamelhalsfliegen) 25 15
Karl H. C. Jordan Heteroptera (Wanzen) 20 16
Max Beier Saltatoria (Heuschrecken u. Grillen) 9 17
Herbert Weidner Copeognatha (Staubluse) 16 18
Erwin Schimitschek Pflanzen-, Vorrats- und Materialschdlinge 8 19
Willi Hennig Diptera (Zweiflgler) 31 20
Hans Malicky Trichoptera (Kcherfliegen) 29 21
Max Beier Blattariae (Schaben) 13 22
Klaus Gnther
& Konrad Herter Dermaptera (Ohrwrmer) 11 23
Felix Mainx Zytologie (Karyologie) 12 24
Alfred Kaltenbach Mecoptera (Panorpata, Schnabelhafte) 28 25
Peter Zwick Plecoptera (Perlaria, Uferfliegen) 7 26
Herbert Weidner Morphologie, Anatomie und Histologie 11 27
Hans Striimpel Homoptera (Pflanzensauger) Part 28
Werner Nachtigall Bewegungsphysiologie (Laufen Schwimmen Fliegen) Part 29
Timothy R. New Planipennia (Netzflgler) Part 30
Wolfgang Schedi Hymenoptera: Unterordnung Symphta (Pflanzenwespen) Part 31
Feliks Piotrowski Anoplura (echte Luse) Part 32
Timothy R. New
& Gnther Theischinger Megaloptera (Alderflies, Dobsonflies) Part 33
Lajos Zombori
& Henrik Steinmann Dictionary of Insect Morphology Part 34
Niels P. Kristensen Lepidoptera, Moths and Butterflies
Vol. 1: Evolution, Systematics, and Biogeography Part 35