Sie sind auf Seite 1von 504

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


G Walter de Gruyter Berlin New York 1999

Niels P. Kristensen (ed.)

Lepidoptera, Moths and Butterflies

Volume 1:
Evolution, Systematics,
and Biogeography


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

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

Library of Congress - Cataloging-in-Publication Data

Handbook of Zoology

agr 2 3 - 1 4 3 6

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

Die Deutsche Bibliothek - Cataloging-in-Publication Data lines of the ANSI to ensure permanence and durability.

Handbuch der Zoologie : eine Naturgeschichte der

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

27: 6 5 - 1 0 1 , 1996). But it was decided that the As an editor I am greatly indebted to the team
similarly innovative analysis of Gelechioid phy- of authors for their commitment, faith and pa-
logeny, which forms the basis of 19, should be tience. I similarly thank the numerous specialists
included, with the primary material printed as who provided reviews of individual chapters. Fi-
appendices. nally I thank D r Maximilian Fischer (former sole
While some contributions were completed just editor, Insecta section of the Handbook series)
before type-setting of the entire colume was initi- as well as P D Dr Mario Noyer-Weidner and Ms
ated, others had been completed considerably Christiane Bowinkelmann (Walter de Gruyter,
earlier. However, most authors were able to take Biosciences Division) and Ms Ingeborg Klak
the opportunity to make additions to their con- (Walter de Gruyter, Production Department) for
their collaboration on this enterprise.
tributions in 1997, so the volume as a whole will
appear reasonably updated by the time of its Copenhagen, April 1998
publication. Niels P. Kristensen

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.
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

P. R. Ackery Ronald W. Hodges

Department of Entomology Systematic Entomology Laboratory, USDA
The Natural History Museum Present address: 85253 Ridgetop Drive
Cromwell Road Eugene, OR 97405-9535, USA
London SW7 5BD, England email:
J. D. Holloway
D. J. Carter do Department of Entomology
Department of Entomology The National History Museum
The Natural History Museum Cromwell Road
Cromwell Road London SW7 5BD, England
London SW7 5BD, England email:
Marianne Horak
Donald R. Davis
Entomology Division, CSIRO
Department of Entomology
PO Box 1700, Canberra City
National Museum of Natural History
ACT 2601, Australia
Smithsonian Institution
Washington, DC 20560, USA
R. de Jong
John S. Dugdale Department of Entomology
do Landcare Research Naturalis/Nationaal Naturhistorisch Museum
Private Bag 6 Postbus 9517
Nelson, New Zealand NL-2300 RA Leiden, Netherlands
email: email:

E. D. Edwards Ian J. Kitching

Entomology Division, CSIRO Department of Entomology
PO Box 1700, Canberra City The National History Museum
ACT 2601, Australia Cromwell Road
email: London SW7 5BD, England
Marc E. Epstein
Department of Entomology Niels P. Kristensen
National Museum of Natural History Department of Entomology
Smithsonian Institution Zoological Museum
Washington, DC 20560, USA University of Copenhagen
email: epstein. DK-2100 Copenhagen 0 , Denmark
Brian Farrell
Museum of Comparative Zoology
Claude Lemaire
Harvard University
La Croix des Baux
Cambridge, MA 02138, USA
F-84220 Gordes, France

Patricia Gentili Jol Minet

Department of Entomology Musum National d'Histoire Naturelle
National Museum of Natural History Entomologie
Smithsonian Institution 45, Rue de Buffon
Washington, DC 20560, USA F-75005 Paris, France
email: MNHEN087@SIVM.SI.EDU email:

Henk Geertsema Charles Mitter

Department of Entomology and Nematology Dept of Entomology
University of Stellenbosch University of Maryland
Stellenbosch, 7600 South Africa College Park, Maryland 20742, USA
email: email:

Eugene Munroe Gaden S. Robinson

3093 Barlow Crescent Department of Entomology
R.R. 1 Dunrobin The Natural History Museum
Ontario, KOA ITO, Canada Cromwell Road
London SW7 5BD, England
Clas M. Naumann
Zoologisches Forschungsinstitut Malcolm J. Scoble
und Museum Alexander Koenig Department of Entomology
Adenauerallee 160 The Natural History Museum
D-53113 Bonn 1, Germany Cromwell Road
email: London SW7 5BD, England

Ebbe S. Nielsen Andrzej W. Skalski (deceased)

Entomology Division, CSIRO Pedagogical University in Czestochowa
PO Box 1700, Canberra City Poland
ACT 2601, Australia
M. Alma Solis
Systematic Entomology Laboratory, USDA
c/o National Museum of Natural History
Jerry Powell Washington, DC 20560, USA
Department of Entomological Sciences email:
201 Wellmann Hall
Gerhard M. Tarmann
University of California
Tiroler Landesmuseum
Berkeley, California 947200-0001, USA Museumstrasse 15
email: A-6020 Innsbruck, Austria
John E. Rawlins R. I. Vane- Wright
Department of Invertebrate Zoology The Natural History Museum
Carnegie Museum of Natural History Department of Entomology
4400 Forbes Avenue Cromwell Road
Pittsburgh, PA 15213, USA London SW7 5BD, England
email: email:
1. Historical Introduction
Niels P. Kristensen

While the references in the following chapters of Entomologica Systematica (179294) and Sys-
these volumes by necessity will be focussed on tema Glossatorum (1807, incomplete). P. A. Lat-
twentieth-century works, a brief outline is given reille similarly refined the system by dividing
below of principal pre-1900 contributions to the Linnean supraspecific taxa.
foundation of contemporary lepidopterology (as
covered in the present Handbook).
Early works on European Lepidoptera
While Fabricius' descriptive works all had a
global scope, Linnaeus himself had devoted a
Systematics and Bionomics Before 1860 special publication to the treatment of his local
(national) fauna: Fauna Svecica (1746, first no-
The Lepidoptera include some of the most con- menclatorically valid edition 1761), and this ex-
spicuous and attractive of all insects, and their ample was soon followed by workers in many
striking metamorphosis lends itself well to obser- European countries. Principal early 'faunas'
vation. Therefore, the book which demarcates from Central Europe, with many original species
the beginning of the current naming and classifi- descriptions, include J. A. Scopoli's Entomologia
cation of animals, C. Linnaeus' Systema Naturae Carniolica (1763), J. N. C. M. Denis & I. Schif-
(tenth edition, 1758), could already refer to a fermller Ankndigung eines systematischen Wer-
substantial body of published work on lepidop- kes von den Schmetterlingen der Wienergegend
teran natural history. Principal references include (1776), and F. von Schranck's Fauna Boica 13
T. Mouffet's Insectorum sive Minimorum Ani- (1798-1804); the lastmentioned work was the
malium Theatrum (published posthumously 1634 first to expand the Linnean classificatory hierar-
and including significant contributions by other chy for Lepidoptera (see Emmet 1991). Among
naturalists, notably the Swiss scientific pioneer British works M. Harris' The Aurelian or Natural
K. Gesner, 151665), J. Goedart's Metamorpho- History of English Insects, namely Moths and
sis Naturalis (1662-67), Maria S. Merian's il- Butterflies (1766) is renowned for its exquisite
lustrated accounts of European Lepidoptera as coloured illustrations, but truly comprehensive
well as her renowned Metamorphosis insectorum accounts of the British fauna only came with
Surinamensium (1705), J. Ray's Historia Insec- A. H. Haworth's Lepidoptera Britannica (1803
torum (1710), J. Petiver's Papilionum Britanniae 28), followed by two works of a more modern
icones (1717), . Wilkes' English Moths and But- scope, J. F. Stephen's Illustrations of British En-
terflies (1747-60), A. J. Rsel von Rosenhof's tomology: Haustellata (1827-35) and Curtis'
Die monatlich herausgegebene Insekten-Belusti- British Entomology (1823-40). Principal early
gung 1 - 4 (1746-61), A. R. Reaumur's M- works on Lepidoptera in the Mediterranean sub-
moires pour servir l'histoire des insectes 16 region are by J. P. Rambur, including Catalogue
(1734-42) and K. de Geer's similarly titled work systmatique des Lpidoptres de l'Andalusie
( 1 - 7 , 1752-78). (1858-66, 'Macrolepidoptera' only). Principal
pioneering contributions to the knowledge of the
lepidopteran fauna of Russia (including Siberia)
Linnaeus, Fabricius and Latreille
are the works of E. F. Eversmann, e. g. Fauna
Linnaeus' system profoundly influenced later Lepidopterologica Volgo-Uralensis (1844).
classifications. His basic groups in the Lepidop- Several important works on the European
tera Papilio (Eques, Danaus, Nymphalis, Plebe- fauna were of a broader geographical scope.
jus, Barbarus), Sphinx, and Phalaena (Bombyx, Early examples were E. J. C. Esper's Die Schmet-
Noctua, Geometra, Tortrix, Pyralis, Tinea, Alu- terlinge in Abbildungen nach der Natur 15
cita) persisted in some general zoology text- (177794 [second edition, with additions, by T.
books for more than a century, although in the von Charpentier 1829-39]) and M. B. Bork-
meantime Lepidoptera classification had un- hausen's Naturgeschichte der Europischen
dergone considerable refinement in specialists' Schmetterlinge 1 - 5 (1788-94). Later followed
writings. Linnaeus' pupil J. C. Fabricius de- the influential ten-volume treatise Die Schmetter-
scribed numerous lepidopteran species from linge von Europa (1807-34), initiated by F. Och-
many parts of the world and subdivided several senheimer and completed by his collaborator F.
Linnean groups into what were to become recog- Treitschke. Its title notwithstanding the Histoire
nized as valid genera; his latest and most com- Naturelle des Lpidoptres de France, initiated in
prehensive works dealing with Lepidoptera are 1820 by J. B. Godart and completed (in 11 vol-
2 Niels P. Kristensen

umes, with four supplements) in 1842 by P. A. J. Lepidoptera; so did D. Drury's Illustrations of

Duponchel also was Europe-wide in scope. Natural History 1 - 3 (1770-82). Hbner's
J. B. A. D. de Boisduval's Genera et Index Me- Sammlung exotischer Schmetterlinge (1806
thodicus Europaeorum Lepidopterorum (1840) [38]), completed by Geyer and supplemented by
catalogued the European 'Macrolepidoptera', Herrich-Schffer's Sammlung neuer oder wenig be-
with Guene's incomplete Europaeorum Micro- kannter auereuropischer Schmetterlinge (1850
lepidopterorum Index Methodicus (1845) as a 1869) time-wise overlapped the author's works
companion volume; these works were paralleled on the European fauna, and the scope of Her-
by Duponchel's Catalogue mthodique des Lpi- rich-Schaeffer's system was broadened by the in-
doptres d'Europe (1844-46). The artist J. clusion of exotic taxa. The Spcies Gnral des
Hbner's volumes of outstanding coloured Lpidoptres (eight volumes, 1836-75) by Bois-
plates, Sammlung europischer Schmetterlinge duval & A. Guene in 'Nouvelles Suites Buffon'
1796[1838] and Geschichte europischer Schmet- was planned as a complete treatise of the lepi-
terlinge [1793][1842], deserve particular men- dopteran species, but remained incomplete. E.
tion; he also published illustrations of extra- Doubleday & J. O. Westwood's The genera of di-
European Lepidoptera (cp. below) and intro- urnal Lepidoptera (1846-1852), illustrated by
duced a wealth of new taxon names. Hbner's W. C. Hewitson is the foundation for modern
work was continued and completed by G. A. W. butterfly classification, while F. Walker's List of
Herrich-Schffer and the painter C. Geyer, The Specimens of Lepidopterous Insects in the Collec-
former's six-volume supplement to Hbners tion of the British Museum (35 parts, 18541866)
Sammlung, entitled Systematische Bearbeitung is the last synopsis of the World's moths to have
der Schmetterlinge von Europa (1843 1856), is a been completed.
milestone in Lepidoptera systematics with a fam-
ily classification largely foreshadowing that
adopted today. Herrich-Schffer made extensive Systematics and Bionomics 18601900
use of characters provided by the wing venation
(preliminary studies on which had been made by C. Darwin's On the Origin of Species is a turning
some earlier workers), but also took characters point in biological history. The principal contri-
of e. g. vestiture, palpi, and legs into account. J. bution of Lepidoptera research to the early shap-
Lederer's Die Noctuiden Europas (1857) is one of ing of evolutionary biology came with the mim-
the first works in which characters in the male icry theory, first formulated in H. W. Bates' A
genital apparatus are used (an even earlier use Naturalist on the Amazonas (1861) and later elab-
was in Rambur's study of Andalusian skipper orated on by several lepidopterists including
butterflies). Of outstanding significance are P. C. A. R. Wallace and F. Mller. Towards the end of
Zeller 's numerous works on micro-moths; his the century K. Jordan's theoretical work {Novit.
1839 article (in Oken's Isis) Versuch einer natur- Zool. 3, 1896) rooted in Lepidoptera taxonomy
gemssen Eintheilung der Schaben (meaning Lin- laid an important foundation for the later preva-
nean 'Tinea', not cockroaches) was followed by lent species concept. Truly important progress
a suite of high-quality monographs of several towards an evolutionary higher classification of
micro-moth groups, and eventually his writings the order were made in the last decade of the
were extended to extra-European faunas. He fur- century, particularly through studies on imma-
ther contributed, as did H. Frey and J. W. Doug- ture stages by T. A. Chapman (e. g., Trans, ent.
las, to the unique and lavishly illustrated 13-vol- Soc. London 1893, 1896) and on wing structure
ume The Natural History of the Tineina (1855 by A. Spuler (Z. Wiss. Zool 53, 1892) as well as
73) published (with text in English, French, Ger- by J. Comstock and J. Needham (articles in Am.
man and Latin) by H. T. Stainton. Nat., also Comstock's Manual for the Study of
Insects, 1895).

Early works on the world fauna

Inventory of the World fauna
Expectedly, local/regional 'faunas' by resident
workers outside Europe were few in this period. Outstanding among the European 'fauna hand-
J. Abot & J. E. Smith's Natural History of Rarer books' of the period was Die Schmetterlinge
Lepidopterous Insects of Georgia 12 (1797) and Deutschlands und der Schweiz 1 - 2 (1863-77) by
E. Blanchard's treatment (1852) of the Lepidop- H. von Heinemann & M. F. Wocke. The Catalog
tera in C. Gay (ed.) Historia fisica y politica de der Lepidopteren Europas by O. Staudinger &
Chile are examples. M. F. Wocke (1861) and its later editions in 1871
Already in the earliest phase of descriptive and 1901 (the lastmentioned, covering the entire
taxonomy P. Cramer's De Uitlandsche Kapellen Palaearctic region, was published by H. Rebel)
in de drie warereld-deelen Asia, Africa en America have been invaluable tools. Major works on
(177582, with posthumous supplements by C. central- and eastern-Palaearctic taxa by H. T.
Stoll in 1782 and 1787-1790) provided substan- Christoph, S. N.-Alpheraky and O. Staudinger
tial pictorial documentation of extra-European appeared in the series Mmoires sur les Lpidop-
Historical Introduction 3

teres edited by . M. Romanoff from 1884 on- in A Handbook of British Lepidoptera, 1895,
wards. Principal comprehensive works on Lepi- takes the World fauna into account), G. F.
doptera from E. and SE. Asia include J. H. Hampson (author of monographs on pyraloid-
Leech's Butterflies from China, Japan and Corea grade moths, but best known for the monumen-
(1892-94), W. L. Distant's Rhopalocera Malay- tal Catalogue of the Lepidoptera Phalaenae in the
ana (188286), F. Moore's The Lepidoptera of British Museum [treating a major part of the
Ceylon 1 - 3 (1880-87) and G. F. Hampson's Noctuoidea] which started appearing in 1898),
four-volume treatment (1893-96) of the macro- and the remarkable team of W. Rothschild and
moths and pyraloids in the Fauna of British In- K. Jordan, whose first joint monograph (on
dies series. Early comprehensive works on Afro- Charaxes butterflies, Novit. Zool., 5 - 7 ) was pub-
tropical Lepidoptera were few. Suffice it to note lished 1898-1900.
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), Morphology and Physiology
completing ealier works by Boisduval and Ma-
bille. In the Nearctic J. G. Morris' Catalogue of Two of the most renowned early general works
the Described Lepidoptera of North America on insect anatomy deal with larger moths: M.
(1860) was followed by several monographs in- Malpighi's Dissertatio epistolica de Bombyce
cluding W. H. Edwards' Butterflies of North (1663) treating the silkworm moth Bombyx, and
America 1 - 3 (1868-97) and S. H. Scudder's P. Lyonet's remarkably accurate (with respect to
Butterflies of New England 1 - 3 (1888-89). A. S. description of structure, not to interpretation of
Packard's Monographs of the Geometrid Moths function) Trait anatomique de la chenille qui
or Phalaenidae of the United States (1876) and ronge le bois de saule (1762) treating a Cossus
Monograph of the Bombycine Moths of America larva. A comparative approach to insect mor-
North of Mexico ... I, 1 Notodontidae (1895) phology was only developed in the late 18th cen-
stand out for craftsmanship and attention to de- tury, mainly by French workers. Particularly
tail. J. Smith monographed several noctuid outstanding was J.-C. Savigny's account of the
groups. Principal N. American micro-moth au- mouth parts in Mmoires sur les Animaux sans
thors were B. Clemens, whose papers on moths Vertbres (1816).
of the lower grades were reprinted by Stainton
as Tineina of North America (1872), and V. T. Integumental structures
Chambers. The prodoxid/ Yucca symbiosis, one
of the most intriguing insect-plant relationships Lepidopteran vestiture was an attractive subject
known, was first discovered by C. V. Riley (Na- for early microscopists. 19th century studies on
ture 6, 1872). Outstanding in the New World lit- hair and scale structure (often focussing on scent
erature is the regional monograph Biologia organs) include papers by, i. a., R. Maddox, R.
Centrali-Americana edited by F. D. Godman & Schneider, C. Aurivillius, A. Spuler, V. Kellog,
O. Salvin. Lepidoptera parts include the editors' and particularly numerous contributions by F.
account of butterflies, 1 - 3 (1879-1901) and H. Mller and E. Haase. F. G. Hopkins (Phil.
Druce's of macro-moths, 1 - 3 (1881-1900) Trans. R. Soc., 186B, 1895) proved that excretory
(while T. de Grey Walsingham's treatment of the products are stored in wing pigments in pierid
'Tineina, Pterophorina, Orneodina, Pyralidina butterflies. There are numerous smaller 19th cen-
and Hepialina [part]' appeared later). tury accounts of defensive glands, urticating
Of global coverage were E. L. Ragonot's hairs and other integumental specializations of
Monographie des Phycitinae et des Galleriinae (in caterpillars. Dyar (Ann. New York Acad. Sci., 8,
Romanoff's 'Mmoires' 7, 8; 1893-1901) as well 1894 and later papers) made a pioneering study
as the accounts of butterfly genera in O. Stau- of the larval setal pattern, foreshadowing 20th
dinger's Exotische Tagfalter (188488) and the century taxonomic use of this system.
companion volumes Die Familien und Gattungen
der Tagfalter by E. Schatz & J. Rber (1885-92).
The head
W. F. Kirby's A synonymie catalogue of diurnal
Lepidoptera (1871), supplement 1877) was com- The coilable proboscis, a unique specialization
plete for its time, the same author's Catalogue of present in most adult Lepidoptera, had been
Lepidoptera Heterocera (1892) covers "Sphinges noted by early naturalists. Lepidopteran mouth-
and Bombyces". By the end of the 19th century parts were treated in some detail in Savigny's
Lepidoptera systematics of a worldwide scope aforementioned work. E. Burgess described the
was pursued by several workers whose most in- sucking pump in his succint, but influential, ac-
fluential works date from the 20th century. Par- count of the anatomy of the monarch butterfly
ticularly outstanding were E. Meyrick (all-time (Danaus) (Anniv. Mem. Boston Soc., 1880). The
most prolific descriptive systematist of micro- discovery that primitive biting mouthparts are
moths; his Lepidoptera classification presented 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).

Most early studies concern applications of wing
characters to systematics (see above). The tho- Worthwhile studies on lepidopteran embryogen-
racic trunk attracted little attention; the principal esis started with A. Kowalewsky (Mm. Acad.
early account of the musculature is by Amans Sci. St. Ptersb., (7) 16, 1871). Egg diapause was
(Annal. Sci. nat. (Zool.), (6) 19, 1885). The early discovered in the silkworm Bombyx, and its
tympanic organ of noctuoids was first described dependence upon temperature regimes was
by A. Swinton (Ent. month. Mag., 14, 1877). 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-
Abdomen gression of head capsule widths at moults (Psy-
che 5, 1890). M. Herold's Entwicklungsgeschichte
The early applications of genital characters to
der Schmetterlinge, anatomisch und physiologisch
Lepidoptera systematics were paralleled by in-
bearbeitet (1815) is a pioneer (albeit crude) ac-
quiries into their structure and function; often
count of metamorphic changes in the viscera of
the approaches were combined. The presence of
a Pieris butterfly; it also demonstrated the pre-
two female genital openings characteristic of
hatching determination of sex, as well as the de-
most female Lepidoptera had been recognized
velopment of male genital rudiments in a peri-
already by Malpighi, and the functional design
podial cavity. Principiai early studies on the mor-
of this ditrysian female apparatus was examined
phogenesis of wings, scales and hairs include
in more detail by C. T. von Siebold (Arch. Anat.
those of C. Semper (Z. wiss. Zool. 8, 1857) and
Physiol, wiss. Med., 1837) and H. Lacaze-Du-
H. Landois (Ibid. 21, 1871). E. Verson made sev-
thiers (Ann. Sci. Nat. (Zool.), (3), 19, 1853); the
eral important observations on the structure and
former also (Z. wiss. Zool. 3, 1850) studied the
metamorphosis of Bombyx, including the first
sphragis in Parnassius butterflies. Principal con-
discovery of insect dermal glands (e. g. Zool.
tributors to the discussion of the morphological
Anz. 13., 1890) and with E. Bisson he described
interpretation of the genital structures in the two
the morphogenesis of the genitalia of both sexes
sexes and to the development of the nomencla-
(Z. wiss. Zool., 61, 1896).
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 Information Sources
(Ibid. 2 (V), 1890) and A. Peytoreau, author of
Contributions l'tude de la morphologie de Principal sources for this outline are the chapters
l'armure gnitale des insectes (1895). J. Wood's by Tuxen, Lindroth, Richards and Wigglesworth
account of lepidopteran 'ovipositors', particu- in Smith et al. (eds.) 1973, as well as the litera-
larly the piercing types (Ent. month. Mag 27, ture compilations in Kirby (1897) and Kuznet-
1891) ist still useful. sov (1967).
Historical Introduction 5

References Richards, A.G. (1973): Anatomy and morphology.

Pp. 185-202 in Smith, R. F., Mittler, T. E. &
Kirby, W. F. (1897): A Hand-book to the Order Lepi Smith, C. N. (eds.) History of Entomology. Palo
doptera. IV. London, Allen & Co. Alto, Annual Reviews.
Kuznetsov, N. Y. (1967): Lepidoptera I. Introduction. Tuxen, S. L. (I 973): Entomology systematizes and de
Fauna of Russia and Adjacent Countries. Jerusa scribes: 1700-1815. Pp. 95-118 in Smith, R. F.,
lem, Israel Program for Scientific Translations. Mittler, T. E. & Smith, C. N. (eds.) History of Ento
Lindroth, C. H. (1973): Systematics specializes. Be mology. Palo Alto, Annual Reviews.
tween Fabricius and Darwin: 1800-1859. Pp. 119- Wigglesworth, V. B. (I 973): The history of insect phys
154 in Smith, R. F., Mittler, T. E. & Smith, C. N. iology. Pp. 203-228 in Smith, R. F., Mittler,
(eds.) History of Entomology. Palo Alto, Annual T. E. & Smith, C. N. (eds.) History of Entomology.
Reviews. Palo Alto, Annual Reviews.
2. Phylogeny and Palaeontology
Niels P. Kristensen & Andrzej W. Skalski*

The monophyly of the taxon Lepidoptera is nal scapus and pedicellus (3). Maxillary palp
firmly established by an impressive suite of syna- with points of flexion between segments 1/2 and
pomorphies of its constituent basal lineages. The 3/4; segment 4 longest; intrinsic palp musculature
position of the group within the insect hierarchy not comprising antagonistic pairs (4). Cranio-
is similarly well established: It has a sistergroup stipital muscle present, slender and running close
relationship to the Trichoptera (caddisflies), con- to craniocardinal muscle (5). Postlabium an
stituting with the latter the high-rank taxon Am- arched sclerite with long piliform scales (6). Ter-
phiesmenoptera. The Amphiesmenoptera, in minal segment of labial palp with group of sen-
turn, together with the Antliophora (Mecoptera silla in depresseion ('vom Rath's organ') (7). Sali-
+ Siphonaptera + Diptera) constitute the 'pan- varium devoid of dorsolongitudinal muscle (8)
orpoid' clade within the Endopterygota (see (note: this character needs additional scrutiny,
Kristensen 1991, and references therein, for high- since recent observations on Trichoptera and
level relationships). Mecoptera cast doubt on the primitive nature of
This chapter first treats the ground plan and a muscle of this kind in the panorpoid orders; M.
intrinsic phylogeny of the Lepidoptera, as de- Engvall & N. P. Kristensen unpublished). Labral
duced from the character distribution in extant nerve and frontal ganglion connective separating
taxa. Subsequently the palaeontological contri- immediately at their origin on the tritocerebrum
butions to the knowledge about Lepidoptera (9). Nervus recurrens running inside aorta until
reaching retrocerebral complex (10). Laterocer-
evolution are surveyed. No known fossil Lepi-
vical sclerite with proprioceptive 'hair plate'
doptera represent organisation types which
close to anterior apex (11). Prothoracic endoskel-
markedly differ from extant ones, and as in sev-
eton with prominent free arm arising from
eral other insect groups, fossils have so far con-
bridge between sternum and lower posterior cor-
tributed little, if anything, to the understanding
ner of pleuron (12). Mesothorax with 'tergopleu-
of phylogenetic interrelationships. But fossils, of
ral apdeme' issued from upper part of pleural
course, provide the only direct means for estab-
suture and accommodating insertion of a ter-
lishing the minimum ages of individual evolu-
gopleural muscle (13). Metathorax with 'prescu-
tionary lineages. tal arm' (14). Fore tibia with movable 'epiphysis'
Major ecological aspects of lepidopteran evo- on inner surface (15). Fore tibia with at most a
lution are dealt with in 120. single spur (15) (Shields 1993; note: not included
in the 1984-enumeration this regressive trait ap-
parently represents yet another lepidopteran au-
The Lepidopteran Ground Plan and tapomorphy, since 3 spurs can be ascribed to the
the Superorder Amphiesmenoptera trichopteran ground plan). Wings with dense
covering of broad scales (16). Metathoracic spi-
racle with single, anteriorly situated, external lip
Detailed accounts of the lepidopteran ground
(17). Tergum I extensively desclerotized, with
plan will be presented in the morphological
concomitant loss of external layer of 'short' dor-
chapters in volume 2 of the present work. Kris-
solongitudinal I/II muscles (18). Tergum I with
tensen (1984) identified 27 probable lepidopteran
lateral lobes extending posteroventral to articu-
autapomorphies, and another 20 groundplan
late with anterior corners of sternum II (19).
traits were identified as probable synapomor-
Male 'valve' (gonopod) primarily undivided (20).
phies with the Trichoptera, i. e., autapomorphies
Phallic protractor muscle originating inside valve
of the Amphiesmenoptera. The great majority of (21). Cerci lacking in both sexes (22). Abdominal
the characters in question are in the adult insect nerve cord with at most five ganglionic masses,
(Fig. 2.1). Subsequent work has so far entailed and unpaired connectives (23). Sperm of apyrene
few modifications to these conclusions. variety (24). Larva: Pleurostome elongated, cran-
The lepidopteran groundplan autapomorphies iocardinal articulation far behind mandibular
are briefly enumerated here; for more detailed base. Maxillary palp with fewer than 5 segments.
discussions see Kristensen (1984) and references
therein. Adult (numbers in brackets refer to Fig. Kristensen (1984), on the basis of Hessel's
2.1): Median ocellus lost (1). Corporotentorium (1969) information about neighbouring endop-
with posteromedian process (2). Intercalary scler- terygotes considered the dorsal curvature of the
ite present laterally in membrane between anten- adult's mesothoracic aorta to be another lepi-
dopteran groundplan autapomorphy. However,
* deceased 1996 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- cause they are "much more closely related to the
morphic at the panorpoid level. Trichoptera than to the Lepidoptera" (Hinton
In the light of this suite of synapopmorphies 1958) now have only historical interest. Two re-
of the basal lepidopteran lineages inclusive of the gressive traits in larval structure (loss of lacinia,
Micropterigoidea, earlier suggestions that the marked weakening of corporotentorium) are
latter should be assigned to a separate order, be- known to be shared by Trichoptera and non-
Phylogeny and Palaeontology 9

micropterigoid Lepidoptera, but on grounds of Lepidoptera Phylogeny above

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

NEOLEPIDOPTERA 1 Palaephatoidea
HETERONEURAL- Tischerioidea
















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 11

only one (not two) pairs of male accessory Basal Glossatan Phylogeny. The monophyly of
glands. The interpretation of the two lastmen- the Glossata is very strongly supported. Not only
tioned traits as apomorphies at the basic lepidop- do the glossa tan autapomorphies (listed in 15)
teran level is debatable, however; particularly the include some very remarkable neoformations/
occurrence of the coxal muscle in agathiphagoids modifications, but there is at present no conflict-
(and, as far as known, nowhere else among pan- ing evidence at all (i. e., evidence that subordi-
orpoid insects !) could well be an autapomorphic nate glossatan taxa should have their closest rel-
character reversal. The reason why the phylog- atives outside the Glossata).
eny here preferred has the Micropterigoidea Six basal clades are recognized within the
rather than the Agathiphagoidea as the first di- Glossata: the Eriocraniidae, Acanthopterocteti-
verged lepidopteran lineage, is that the suite of dae, Lophocoronidae, Neopseustidae, Exoporia
derived characters shared by non-micropteri- (Mnesarchaeoidea + Hepialoidea) and Hetero-
goids in addition to a number of regressive char- neura (all other Glossata). A phylogenetic analy-
acters includes some striking neoformations sis of these clades, based on 46 characters (Niel-
(metafurcal process, double-compartment sper- sen & Kristensen 1996) gave best support to the
mathecal duct, larval lateral head-capsule mus- arrangement illustrated in Fig. 2.2, viz., Eriocra-
cle, hypertrophied and angularly bent pupal niidae + (Acanthopteroctetidae + (Lophocoro-
mandible), while those shared by non-agathipha- nidae + (Neopseustidae + (Exoporia + Hetero-
goids all are regressions/simplifications which are neura)))). Apomorphies supporting each inter-
more likely to be homoplasious. This issue is not node are listed in 15. Names have been given
addressed in Shield's (1993) advocacy of the the- to three of the identified high-rank taxa: Coelo-
ory of the Agathiphagoidea being the sister lepida (the non-eriocraniid Glossata), Myoglos-
group of all other Lepidoptera. Ivanov (1994) sata (Neopseustidae + (Exoporia + Hetero-
has recently found support for the same theory neura)) and Neolepidoptera (Exopria + Hetero-
in characters in the wing base; these await eval- neura).
The next splitting event is here believed to Basal Heteroneuran Phylogeny. The monophyly
have been between the Agathiphagoidea and the of the Heteroneura is here accepted as a reason-
Heterobathmioidea + Glossata. Derived traits able working hypothesis supported by several
characterizing the latter entity include some very putative groundplan autapomorphies discussed
characteristic traits of the larval head (adfrontal in more detail by Nielsen & Kristensen (1996):
and hypostomal ridges, interrupted hypostomal Heteroneurous venation (hindwing Sc and R
bridge) in addition to the presence of sensilla au- fused beyond a short distance from wing base,
ricillica and loss of Sc-R crossvein in the adult, Rs unbranched). Male with composite frenulum.
and presence of coarse apical teeth on the mandi- Forewing jugum reduced, projecting little be-
ble in the pupa. Alternatively the dichotomy yond general wing contour. Dorsal tentorial
could be between the Agathiphagoidea + Heter- arms in adult strongly reduced. Prothoracic dor-
obathmioidea and the Glossata, but the derived sum with median sclerotization behind principal
traits of the former entity seem less impressive. paired 'warts', and precoxal bridge present. Mes-
Most notable are the Y-configuration of the an- osternum with suture-delimited 'mesoclidium',
terior tentorial arms and the posteriorly widened and fusion of the latter with the prospina. Two
postlabium in the adult, as well as the subapical discrete bundles, with different posterior inser-
ventral tooth on the mandible in the pupa; other tion sites, of mesofurco-metafurcal muscles.
similarities are in regressive traits (indistinct lat- Metaprescutal arm and sternum I obliterated. In
eral lobe from tergum I, loss of labral compres- embryogenesis 'fault-type' formation of amnion
sors, intrinsic maxillary muscles, and dorsal and serosa developed, germ band immersed in
postcerebral pharynx dilators in the endopha- yolk, and temporary epithelium-like membrane
gous larvae). The anastomosis pattern of the formed by peripheral yolk nuclei/vitellophages
mesepisternal para- and precoxal sutures is absent. This evidence arguably outweighs the dif-
somewhat different in the Agathiphagoidea and ficulties with heteroneuran monophyly empha-
Heterobathmoidea, and the suggested presence sized by Dugdale (1974) and Minet (1984), viz.,
(Kristensen & Nielsen 1979, 1983) in the Hetero- respectively, the somewhat similar female genital
bathmioidea of a phallotheca-type intromittent systems (with separate copulatory orifice and
organ, similar to that of the Agathiphagoidea, ovipore, and with ventral bursa copulatrix) in
has been proved wrong by a later reinvestigation. Exoporia and Heteroneura-Ditrysia (here con-
Needless to say, an Agathiphagoidea + Hetero- sidered parallelisms), and the absence of muscu-
bathmioidea monophyly model also necessitates lated, crochet-bearing prolegs in Heteroneura-
that the unique agathiphagoid plesiomorphies Nepticuloidea (here considered a secondary re-
listed above should have been lost three times gression, linked to endophagy).
independently (in the Micropterigoidea, Hetero- The basal splitting events within the Hetero-
bathmioidea and Glossata). neura remain problematical (see 16). A mono-
phyletic group comprising (Palaephatoidea +
12 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-
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-
Fig. 2.3. Davis' (1986) alternative proposals for the ment, in recent decades, of an increasing number
basal phylogeny of the Heteroneura. of superfamilies comprising a single or very few
families. In the classification adopted here 33 di-
trysian superfamilies are recognized. The scheme
Tischerioidea) + Ditrysia (Fig. 2.2) could be sup- of their hypothesized interrelationships shown in
ported by the presence of a medial ridge between Fig. 2.2 is mainly based on Minet's seminal 1991
the 'ovipositor' lobes in the two firstmentioned study, which should be consulted for further de-
taxa and a derived female frenulum type (few, tails.
closely approximated setae) in all (Nielsen 1985, A large assemblage of ditrysian superfamilies,
1989). Moreover, the eggs in Tischerioidea (pa- including the apparently first differentiated su-
laephatid egg ultrastructure remains unstudied) perfamilies, share some conspicuous (but plesio-
and Ditrysia share a unique trabecular chorionic morphic) traits with most non-ditrysians: They
layer (Fehrenbach 1995, 2 - 1 8 ) . Also, the mono- are generally small moths with a distinct CuP (at
phyly of all non-nepticuloid Heteroneura is now least in the hindwing; this vein, however, is re-
being supported by data from female 'calling' be- peatedly lost in subordinate clades, particularly
haviour (oviscapt extrusion) and 'long-chain' fe- those characterized by pronounced wing narrow-
male sex pheromenes (Lfstedt & Kozlov 1995, ing), and with larvae that are concealed feeders
2 - 1 3 ) . Alternatively, Davis (1986) advocated the and usually have the proleg crochets arranged in
phylogeny shown in Fig. 2.3, implying that all a circle/ellipse (which may be more or less in-
non-ditrysian Heteroneura constitute a mono- terrupted). Together with the non-ditrysians the
phyletic 'Monotrysia' the groundplan of which is members of this ditrysian grade are colloquially
characterized by the presence of a 'pectinifer' (a referred to as microlepidopterans or micro-
valve area bearing a comb of specialized stout moths ('micros').
setae) and a derived metafurcal configuration. A The monophyly of the non-tineid Ditrysia is
pectinifer, however, is absent in the Palaepha- only supported by a few regressive traits (Minet
toidea and Tischerioidea, and the ground plan 1991): shortening of the maxillary palp which is
of the 'monotrysian' metafurca is probably not at most 4-segmented (the only known exception
different from that of the Heteroneura (24). being some Alucitoidea-Alucitidae in the Obtec-
The monophyly of the Nepticuloidea + Tischeri- tomera; a character reversal?) and has lost the
oidea was supported only by the flattened head sharp bend between segments 3 and 4; absence
and reduction of larval legs (commonplace modi- of projecting lateral bristles on the labial palp.
fications in miners) as well as a narrowing of the The significance of the lastmentioned character
hind wings and loss of M2 in both wings. Recent is ambiguous; the bristles in question have tenta-
evidence from 18S rDNA data support the mo- tively been considered a tineoid autapomorphy
nophyly of Tischerioidea + Ditrysia as well as of by Robinson & Nielsen (1993, see also 1 - 7 ) , but
a clade comprising the remaining monotrysian these authors, like Minet, noted their occurrence
Heteroneura. However, the nepticulid examined also in a variety of non-ditrysian Heteroneura,
comes out as subordinate in the Incurvarioidea, and hence admitted their possibly plesiomorphic
which appears unacceptable on the basis of mor- nature at the ditrysian level. Robinson & Nielsen
phology, and it is emphasized, that the above- (1993) suggested that all Ditrysia apart from the
mentioned alternative phylogenies "are not Tineoidea and Gracillarioidea constitute a mo-
strongly contradicted by the 18S rDNA gene" nophylum, to whose ground plan they ascribed,
(Wiegmann 1994). as an autapomorphy, the presence of spines on
the abdominal terga in the adult moths.
Ditrysian Phylogeny. The monophyly of the Di- All Ditrysia, apart from the Tineoidea, Gracil-
trysia is corroborated by very notable ground- liarioidea, Yponomeutoidea and Gelechioidea
Phylogeny and Palaeontology 13

are tentatively united in the taxon Apoditrysia subsequently they were excluded from this taxon
(Minet 1983), whose monophyly is supported by when it was redefined (Minet 1991) by the addi-
an apomorphic sternum II structure. The apo- tion of another groundplan autapomorphy, viz.,
demes are shortened, with enlarged bases, not the differentiation of a dorsal lobe (sometimes
appearing as mere continuations of the 'venulae' merely appearing as a group of bristle-like pro-
(longitudinal costae on the sternal plate). Pro- cesses) on the adult's pretarsal pulvillus.
duced anterolateral corners are another charac- Eleven superfamilies of non-obtectomeran
teristic specialization of sternum II in most Apoditrysia are presently recognized; the Tortri-
Apoditrysia, but it is uncertain whether they can coidea are by far the largest of these, and the
be ascribed to the ground plan of the clade. Rob- whole assemblage may conveniently be referred
inson & Nielsen (1993) considered the loss of ter- to as the 'tortricoid grade'. Interrelationships
gal spines an additional apoditrysian autapo- above the superfamily level are largely unclari-
morphy (as a character reversal), but tergal fied within this grade. The Sesioidea (which as
spines are actually retained in some apoditrysian now delimited include the Brachodidae, Sesiidae
lineages, including Zygaenoidea, Epermenioidea and Castniidae) and Cossoidea may be each oth-
and Alucitoidea (1 11, 113). er's closest relatives (1 11), and it is just possible
The monophyly of the Gelechioidea + Apodi- that the two together are the sister group of the
trysia has been suggested on the basis of their Zygaenoidea. A possible synapomorphy accord-
shared reduction of the basal apdeme of the ing to Minet (1991) is the position of the sensil-
male valve and the dorsal displacement of the lum Pb on the larval head capsule, which in these
origin of the valve 'protractor' (Robinson & three superfamilies lies distinctly dorsad from,
Nielsen 1993; see also 24): Additional support rather than below, a line between the P2 setae;
for this clade has recently come from the discov- admittedly, however, the polarization of this
ery of putative synapomorphies in its members' character is difficult, because both states occur
proboscis structure: two (not one) rows of sen- within the Tineoidea. According to Ehnbom
silla styloconica per galea, and presence of (1948) the ocellus structure in Zygaenidae and
strongly sclerotized transverse ribbons (Ram- Sesiidae was unique among the Lepidoptera ex-
mert 1994); however, considerable homoplasy in amined by him because of the presence of an ex-
these traits render their significance questionable tra cell layer between the cornea and the retina
(Minet, pers. comm.). The close approximation (he had not correctly identified the vitreous body
of setae LI and 2 on larval segments IVIII may subsequently described from Eriocranioidea; see
be yet another groundplan autapomorphy of this 15, 212). A reinvestigation, including an ex-
clade; the wide spacing of these setae in many amination of ocellus structure in the Brachodi-
advanced Apoditrysia would, then, have be con- dae and Castniidae, is now an obvious priority.
sidered character reversals. In the light of the evi- The proposed monophyly of the Alucitoidea
dence for an alternative placing of the Gelechi- + Pterophoroidea (Minet 1991) is based on their
oidea (as the sister group of the Yponomeu- shared specialized configuration of the meso-pre-
toidea, see below) the question has here been coxal suture, and the resting posture with fore-
left open. wings markedly held out from the body.
The classification of the Apoditrysia is partic- The Alucitoidea have previously been in-
ularly problematical. Fig. 2.2 reflects Minet's cluded, together with the Epermenioidea, in the
view that a large assemblage of apoditrysian su- Copromorphoidea. Specializations shared by
perfamilies constitute a monophylum 'Obtec- these superfamilies include, in addition to the pu-
tomera'. When this taxon was first proposed pal type (see above), a process on the larval post-
(Minet 1986) its monophyly was supported only labium (Heppner 1987) and a bisetose protho-
by the immobility of pupal abdominal segments racic L group (the significance of the trisetose L
I - I V ; related to this immobility the pupal abdo- in Isonomeutis, hitherto placed in 'Copromorphi-
men has lost the dorsal transverse rows of spines dae' is ambiguous: see 113): Common (1970)
(which were acquired by ancestral Neolepidop- mentioned some further characteristics of the
tera) and are not protruded from the pupal shel- larval spiracle VIII of copromorphoids and aluc-
ter prior to eclosion. This character complex is itoids (larger size and more dorsal position than
subject to considerable homoplasy. In the non- the preceding spiracles), but these are actually of
apoditrysian grade it occurs also in the Ypono- widespread occurrence (Minet 1983) and ques-
meutoidea and Gelechioidea; for this reason the tionably apomorphic at the apoditrysian level.
monophyly of an entity comprising these two su- The arrangement in Fig. 2.2 reflects Minet's
perfamilies has been suggested by Minet (1991), rejection (1991) of the monophyly of the Copro-
who recently (1996) has found this monophyly morphoidea . lai., since only the Copromor-
supported by details in labial palp structure. phoidea s. str., but not the Epermenioidea and
Among the Apoditrysia the 'obtectomeran-type' Alucitoidea, possess the lobed pulvillus attrib-
pupa also occurs in the Epermenioidea and Alu- uted to the obtectomeran groundplan. But the
citoidea; hence these two superfamilies were ini- relatively immobile, non-spinose and non-pro-
tially included in the Obtectomera. However, truded pupa of Epermenioidea and Alucitoidea
14 Niels P. Kristensen & Andrzej W. Skalski

may represent a genuine synapomorphy with the character reversals are invoked (Brock 1988, see
Obtectomera. If the Alucitoidea + Pteropho- also Robbins 1987).
roidea are indeed monophyletic, this would evi- As far as the butterflies are concerned Minet
dently require that the pupal mobility and spi- (1991) suggested that they constitute a subordi-
nosity in pterophorids are character reversals/ nate rather than a basal macrolepidopteran lin-
neoformations. In the case of the spinosity this eage, inasmuch as he proposed the monophyly
has already been claimed by Kuznetzov & Stekol- of a large entity comprising the Axioidea, Calli-
nikov (1979) and Scott (1986). Minet (1991) duloidea, butterflies (s. lat., including the Hedy-
noted that the Epermenioidea share with the loidea) and Drepanoidea + Geometroidea.
Alucitoidea and Pterophoroidea the apparently Probable synapomorphies of these taxa include:
apomorphic trait of having groups of lamellar concealment of pupal labial palps; reduction of
scales in the forewing fringe. But obviously the the metapostnotal 'fenestrae laterales' (see 24);
acceptance of a monophyletic Epermenioidea + internal laminae of secondary metafurcal arms
(Alucitoidea + Pterophoroidea) will require ad widened, their mesal edges adjacent and parallel,
hoc explanations for the specializations (in im- or posteriorly convergent. It was emphasized,
matures) shared by Epermenioidea and Aluci- however, that at least the first two of these traits
toidea but not Pterophoroidea. are pronouncedly homoplasious, occurring also
Witihin the Obtectomera there is an assem- in subordinate members of other macrolepidop-
blage of superfamilies, which have retained the teran lineages. No hypothesis on the precise posi-
'microlepidopteran' characteristics. The Pyra- tion of the butterflies s. lat. (or s. str. for that
loidea are by far the largest of these, and the matter) can at present be considered at all well
whole assemblage, including also the Copromor- founded in spite of considerable recent effort to
phoidea, Immoidea, Hyblaeoidea, Thyridoidea clarify this matter on the basis of morphological
and Whalleyanoidea, may conveniently be re- as well as molecular data; see 117 as well as
ferred to as the 'pyraloid grade'; indeed some of Weller & Pashley (1995), de Jong et al. (1996)
these taxa have previously been formally in- and Scoble (1996) for further details.
cluded in the Pyraloidea. However, as empha- It has been a central theme in recent discus-
sized by Minet (1983), there is no evidence that sions about ditrysian phylogeny whether a taxon
the superfamily, if more broadly delimited, is 'Macrolepidoptera' inclusive of the butterflies is
monophyletic. indeed a monophylum (Scott 1986, 1989; Scott &
The Simaethistoidea, with 'pyraloid' facies as Wright 1990; Brock 1988, 1990 a, b). In a recent
adults (immatures are unknown), are at present phylogeny reconstruction by Rammert (1994)
entirely unplaced within the Ditrysia. neither the Obtectomera nor the Macrolepidop-
The remaining Ditrysia, which include some tera came out as monophyla. Instead the basal
very speciose superfamilies, are mostly medium- dichotomy within the Apoditrysia was found to
sized to large, frequently rather broad-winged in- lie between to major lineages, one being Cos-
sects with predominantly exophagous larvae. soidea + (Zygaenoidea + (Hyblaeoidea + ((Eper-
Most members of this assemblage have long been menioidea + Thyridoidea) + (Sesioidea + 'but-
known to share a few structural specializations terfly assemblage')))), the other (Schreckensteini-
rarely encountered among 'microlepidopterans', oidea + Tortricoidea) + ((Copromorphoidea +
such as the complete loss of CuP, and the ar- (Alucitoidea + Pterophoroidea)) + Choreu-
rangement of larval proleg crochets in a 'mesose- toidea + (Pyraloidea + 'macromoths')). How-
ries'. In the tree in Fig. 2.2 these superfamilies ever, Rammert's results must be considered in-
constitute a monophylum Macrolepidoptera conclusive, since (1) the analysis was focussed on
supported by at least the apparently unique elon- characters in the abdominal base and omitted
gation of the 1st axillary sclerite in the forewing most other characters currently discussed in
base (Minet 1991). The Macrolepidoptera com- apoditrysian phylogeny, and (2) the scorings of
prise the mimallonoid/lasiocampoid/bombycoid many of the character states for individual super-
assemblage, the Axioidea, Calliduloidea, the but- families/superfamily groups were highly debat-
terflies (i. e., the hedyloid/hesperioid/papilionoid able (Minet, pers. comm.).
assemblage), the Drepanoidea, Geometroidea The abovementioned 'butterfly assemblage' in-
(including Uraniidae) and Noctuoidea. Scott cludes the Papilionoidea, Hesperioidea, Hedy-
(1986, Scott & Wright 1990; see also Scott 1989) loidea, Calliduloidea and Axioidea. The 'macro-
listed a suite of additional putative autapomor-
moths' include the Mimallonoidea, Lasiocam-
phies of an almost identically delimited taxon
poidea, Bombycoidea, Drepanoidea, Geome-
Macrolepidoptera (Mimallonidae excluded).
troidea and Noctuoidea.
These apomorphies, however, either occur else-
Any Apoditrysia cladogram based on current
where among apoditrysians, or the plesiomor-
morphological knowledge will contain so much
phic states are actually retained in scattered
homoplasy 'noise' that its value for making bio-
macrolepidopterans, so that the apomorphic
logical inferences is very limited. It remains to
state cannot be attributed to the macrolepidop-
be seen to what extent future morphological and
teran ground plan unless ad hoc hypotheses of
molecular investigations can remedy this situa-
Phylogeny and Palaeontology 15

tion. The field of comparative morphology still general accounts by Larsson (1978) and Poinar
contains vast, and largely untapped sources of (1992). General problems in the classification
information, e. g. several aspects of proboscis and taxonomy of fossil Lepidoptera were dis-
structure, myology, respiratory organs, internal cussed by Skalski (1976, 1977; at that time still
genitalia, larval mouthparts: also, a detailed re- from a non-cladistic viewpoint).
assessment of some already available morpho-
logical information (particularly the wealth of Fossil Sites. The Eocene/Oligocene (ca. 55
data on male genitalia musculature emerging 54 myr) Baltic amber, of which the main deposit
from the research programme of Kuznetznov, is located on the southeast shore of the Baltic
Stekolnikov and coworkers) from a cladistic Sea, is the richest source of fossil Lepidoptera.
viewpoint is challenging. The molecular ap- Another rich, and now intensively investigated,
proach is still in a very initial phase (Weller et al. source is the mostly younger Dominican amber
1994, Weller & Pashley 1995, review by Regier (ca. 4015 myr, Poinar 1992). Cretaceous amber
et al. 1995), but many more contributions can be from Lebanon, N. Siberia (Tajmyr) and N.
expected in the foreseeable future. It seems likely America have so far yielded only few Lepidop-
that 'total-evidence' studies integrating morpho- tera which, however, are of particular interest be-
logical and molecular data hold the greatest cause of their age.
promise for genuine progress in Lepidoptera The rocky sediments best known for lepidop-
phylogeny. 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
Fossil Lepidoptera continental deposits near Stavropol, N. Cauca-
sus in Russia (Danilewskij & Martynova 1962;
The fossil record of Lepidoptera is meagre com- Nekrutenko 1965; Kozlov 1988; Skalski 1988),
pared with that of the other major endopterygote calcareous marls of the gypsum quarries at Aix-
insect orders; this can be at least partly attrib- en-Provence in France (Theobald 1937 a, 1937 b;
uted to the delicateness of their bodies and Leestmans 1983), and argillaceous limestone of
wings. The record is also a short one, and unam- the Isle of Wight in England (Jarzembowski
biguous lepidopteran fossils of pre-Tertiary age 1980); all of these are similarly from the Tertiary.
were not recorded until 1970. It is estimated that Further important strata for fossil Lepidoptera
only some 600700 specimens of fossil Lepidop- are the Tertiary Gabbro in Italy (Rebel 1898),
tera are presently known, and of these about 500 Radoboj in Croatia (Scudder 1875), Rott (Scud-
are preserved in resins. Most of the remaining der 1875, Burgeff 1951) and Randecker Maars
are from fine-grained sediments: clay, sand, silt, (Zeuner 1942) in Germany, Mo-clay in Denmark
mud, shale, limestone, marl, asphalt and vol- (Larsson 1975), Creste (Nel & Descimon 1984)
canic ash/dust, and there is even an old record and Dauphin (Henrotay 1986) in France, as well
of a moth larva in a rock crystal (Kawall 1876). as some Mid-Late Jurassic/Early Cretaceous for-
Most lepidopteran fossils are adults, but also mations in Middle Asia (Skalski 1979 b, 1984;
fossil larvae, larval cases and pupae (Rebel 1934, Rasnitsyn 1983; Kozlov 1988, 1989).
Kusnezov 1941; MacKay 1969; Nel & Nel 1985), The main sites for fossil Lepidoptera were
and even a moth egg (Gall & Tiffney 1983) have mapped by Schepdeal (1974) and Skalski (1976,
been recorded. Mines in fossil leaves have with 1977, 1979 b, 1979 c, 1979 d). These localities are
more or less certainty been attributed to Lepi- all in the Northern Hemisphere. It is only quite
doptera (Opler 1973, 1974, 1982; Hickey & recently that Pre-Quarternary Lepidoptera have
Hodges 1975; Skalski 1979 a, 1990 a; Kuroko been reported from the Southern Hemisphere
1987; Kozlov 1988; Davis 1994; Labandeira et al. (Martins-Neto & Vulcano 1989; Rozefelds
1994). Lepidopteran scales have been identified 1988 a, b).
in fossil resins (Khne, Kubig & Schlter 1973;
Schlter 1974; Skalski 1976; Whalley 1978), and
together with cuticular remnants in alimentary Systematics of Fossil Lepidoptera
canal of fossil bats (Richter & Storch 1980). Numerous lepidopteran families (in the classifi-
A succinct account of fossil Lepidoptera de- cation adopted here) have been identified (Skal-
scribed up to 1983 is given by Carpenter (1992). ski 1990 b) in fossil resins, most of which in the
Other recent reviews include Skalski (1979 b, Baltic amber. The majority of the recorded fami-
1990 a), Whalley (1986) and Kozlov (1988), the lies belong to the microlepidopteran grade. Fos-
lastmentioned with a list of alleged Lepidoptera sil Lepidoptera recorded from rocky sediments
fossils which have subsequently been transferred include several families that have not been found
to other orders. Lepidoptera from fossil resins in resins (Skalski 1990 b).
have long attracted particular attention; they Lepidoptera inclusions in fossil resins are fre-
were monographed by Kusnezov (1941) and quently well preseved, and when lying in a fa-
Skalski (1976), and summarily reviewed in more vourable position they may be examined in some
16 Niels P. Kristensen & Andrzej W. Skalski

detail. In Lepidoptera fossils from rock beds dae etc.) do not exhibit any apomorphies to per-
usually only the wings lend themselves to closer mit their unambiguous assignment to the
examination; the venation and/or a wing pattern Trichoptera.
composed of light and dark elements are fre-
quently discernible (Figs. 2.7 8), but some early Evidence of Basal Splits. Interpretations of early
interpretations have been marred by artifacts. lepidopteran diversification are inextricably
An instructive example is the widely reproduced linked to interpretations of their host associa-
reconstruction (Rebel 1898) of the Tertiary papi- tions; see chapter 120 for a fuller discussion.
lionid butterfly Doritites bosniaskii, in which The bulk of the Mesozoic Lepidoptera so far de-
grooves on the rock surface near the wing mar- scribed are believed to belong to the non-glos-
gins appear as traces of a parnassiine-like wing satan grade, but it is admittedly only in excep-
pattern. tional cases (Parasabatinca) that head structures
A recurrent problem in the identification of are so well conserved that there is direct evidence
fossil Lepidoptera (as indeed of most fossil or- for this assumption. Two fossils, Protolepis cu-
ganisms) is that they so rarely display the auta- prealata (Fig. 2.5 A) and Karataunia lapidaria
pomorphies that are diagnostic of taxa recog- from the Upper Jurassic of Kazakhstan, have
nized in our classification. A very considerable been assigned to the Glossata-Ditrysia (Kozlov
proportion of the family assignments of fossil 1989), and while the evidence for this assignment
Lepidoptera proposed in previous literature are have proved spurious upon reexamination, the
based on superficial phenetic similarity and these presence of a coiled proboscis in Protolepis is still
interpretations can therefore be characterized at believed to be correct (M. Kozlov pers. comm.).
best as tentative, at worst as misleading. Several If the existence of the Glossata in the Jurassic
of these family assignments have now been can really be confirmed, then this would have
rightly replaced by transfers to the incertae sedis important consequences for the debate over
category by Carpenter (1992) and de Jong (in multiple origins of early lepidopteran angio-
press), but it is almost certain that several addi- sperm-feeding, since the earliest current records
tional transfers of this kind still remain to be of angiosperms are from the Lower Cretaceous
made for fossil moths. (Crane et al. 1995). In any case the attributing of
About one-third of the known lepidopteran mines (of gallery type, with no frass observable)
fossils have been described and named. Named in Upper Jurassic pteridospermophyte leaves
fossils have generally received more attention in from Queensland to the Heteroneura-Nepticuli-
the literature than those (sometimes more impor- dae (Rozefelds 1988 b) cannot easily be accepted
tant ones) which have been described, but not on the basis of present knowledge. The earliest
named. Mesozoic to Middle Tertiary Lepidop- convincing evidence for the existence of the Di-
tera fossils have usually been assigned to extant trysia are leaf mines from the N. American Da-
families but to extinct genera and species. Plio- kota Flora (97 myr BP) which can with great
cene and Pleistocene fossils are recognized as re- certainty be referred to the Gracillariidae-Phyl-
cent species. locnistinae (Labandeira et al. 1994); obviously,
therefore, the origin of the Glossata must have
been considerably earlier. It would be unsurpris-
Overview of fossil Lepidoptera
ing if future fossil discoveries reveal that several
The oldest known evidence of the Lepidoptera is splitting events even within the higher Ditrysia
from the Lower Jurassic (Whalley 1985). Triassic had taken place by the end of the Cretaceous.
fossil insects such as Eoses trissica (Tindale
1945), Eocorona iani (Tindale 1980), Mesoses op- Unplaced Homoneurous Moths. The oldest
tata and M. magna (Riek 1976), and Curvicubitus known fossil which can with great certainty be
triassicus (Hong 1984) have been referred to the referred to the Lepidoptera is the small scaly-
Lepidoptera on inadequate evidence; their sys- winged Archeolepis mane (Whalley 1985) from
tematic status is discussed by Hennig (1969), the Lower Jurassic (Lias) of Dorset, England. It
Skalski (1979 b), Sukatsheva (1982), Willmann was originally placed in a family of its own
(1984, 1989), Whalley (1986) and Kozlov (1988). (Archaeolepidae). Four moths from Upper Ju-
Also some younger Mesozoic fossils have been rassic to Lower Cretaceous aleuronitic tuftes in
included erroneously in the Lepidoptera; the Central Asia, described in the genera Eolepiop-
most widely cited are probably the Jurassic terix, Undopterix, Daiopterix and Palaeolepidop-
Palaeontinidae (Scudder 1875; Handlirsch terix (Skalski 1979 b, 1984; Rasnitsyn 1983; Koz-
19061908), which were eventually transferred lov 1989) were similarly assigned to a family, and
to be Hemiptera. On the other hand some of the even suborder, of their own (Eolepidopterigina-
Mesozoic fossils currently placed in the Trichop- Eolepidopterigidae), but there is no convincing
tera may well be stem-lineage Lepidoptera or support for the monophyly of the latter. These
stem-lineage Amphiesmenoptera; the majority of fossils are all female specimens in which extensi-
these fossils (Microptysmatidae, Cladochoristi- ble oviscapts and/or one or two pairs of long
dae, Dasyneuridae, Liassophilidae, Necrotaulii- oviscapt apophyses have been identified; hence
Phylogeny and Palaeontology 17

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- ae

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- Fig. 2.5. Mesozoic basal Lepidoptera. A. Protolepis cu-
port this placement. Two moths described by prealata. The appendage indicated by arrow has been
Martins-Neto & Vulcano (1989) from the same interpreted as the proboscis, but it cannot be ruled out
formation as Gracilepterix were assigned to Un- that it is a maxillary palp (which in extant basal Glos-
dopterix and Parasabatinca respectively (also sata is a much more conspicuous formation than the
these genera were both considered by the authors proboscis). B. Lophocoronid-like moth from Siberia,
to be micropterigids), but again there are no reli- forewing venation and male postabdomen. A after
Kozlov (1989).
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 from the Baltic and Saxonian amber (Jarzem-
immensipalpia, a Baltic amber moth with ex- bowski 1980; Skalski 1989, 1995, in press b).
traordinarily large maxillary palps, to the Eri-
ocraniidae (Kusnezov 1941) is also not sup- 'Eriocraniid-Grade' Families. Fossil evidence of
ported by convincing synapomorphies. It does Eriocraniidae comes in the form of Tertiary leaf-
have unpaired mesotibial spurs, but the mesotib- mines (Opler 1973; Jarzembowski 1980).
ial spur complement has been reduced indepen- An interesting moth (Fig. 2.5 B) in Upper Cre-
dently in several primitive moth lineages. Kozlov taceous amber from Siberia (Skalski 1979 a)
(1988) synonymized Electrocrania with Micro- shares two apomorphies with the Lophocoroni-
pterix (Micropterigidae), but this family assign- dae, viz., a postapical Rs4 and a bilobed valve
ment is contradicted by the presence of a meso- with a strong apical seta on the lower lobe (in
tibial spur. It is unknown whether Electrocrania extant lophocoronids this seta is, however, short
was indeed homoneurous (or glossatan, for that and blunt); the very long phallus has, beyond
sake). mid-length, a dorsal process which is without
counterpart in extant homoneurous moths. The
Micropterigoidea. Parasabatinca from lower Cre- moth is now being described in the Lophocoroni-
taceous amber of Lebanon (Whalley 1978) does dae (in a subfamily of its own, Skalski in press
exhibit some apomorphies (shortened labial c), but its assignment to this family, the extant
palps, pronouncedly moniliform antennae) range of which is restricted to Australia, must be
which may permit its placement in the socalled considered very tentative indeed; the support for
Sabatinca group of genera in the Micropterigidae this assignment is considered inadequate by Niel-
(Kristensen & Nielsen 1979). In the Tertiary the sen & Kristensen (1996).
family is represented by the extant genus Micro-
pterix, recorded from the Baltic amber and marls Exoporia. The surprise record of Mnesarchaei-
of the Isle of Wight, and Sabatinca-group taxa dae, presently restricted to New Zealand, in Up-
18 Niels P. Kristensen & Andrzej W. Skalski

per Cretaceous Siberian Amber (Zherihin & Su- scribed in the recent genus Adela. See also 'Un-
katsheva 1973) is inadequately substantiated; the placed Heteroneura'.
strong family autapomorphies are not in the
wing venation. Tineoidea. See 'Unplaced Heteroneura'. All de-
Hepialoids have been recorded from Palaeo- scribed tineoids are placed in fossil genera, ex-
gene Isle of Wight limestone (Robinson 1977, cept that species in the Dominican amber have
Jarzembowski 1980) and mid-Miocene continen- been referred to the extant tineid genera Choro-
tal deposits in China (Zhag 1989). pleca and Opogona (Skalski unpublished).
Psychidae are known mainly from cases
Unplaced Heteroneura. Stem-lineage Hetero- ('bags') in the Baltic amber; a few compression
neura were undoubtedly superficially similar to fossils of adults have also been reported (Kozlov
Incurvarioidea in many aspects, and records of 1988). Acrolophidae are recorded from Domini-
this superfamily from the Cretaceous (Whalley can amber (Skalski in press d). Bucculatricidae
1978, Skalski 1979) must be considered uncer- are recorded from the Late Cretaceous to Oligo-
tain, since the pertinent autapomorphies (16) cene on the base of leaf-mines (Opler 1973, 1982;
cannot be observed in these specimens. Strauss 1977; Crane & Jarzembowski 1980; Koz-
A larva from Canadian Cretaceous amber (the lov 1988).
first pre-Tertiary lepidopteran fossil to be discov-
ered, MacKay 1970) has been assigned to the Ti- Gracillarioidea. Leaf mines exhibiting detailed
neoidea, but while it certainly belongs in the similarities with those of extant Gracillariidae-
Glossata (it has a distinct spinneret), it cannot Phyllocnistinae are present in the mid-Creta-
be placed with any greater precision. About 30% ceous D a k o t a Flora (Labandeira et al. 1994),
and somewhat similar mines have been identified
of all lepidopteran inclusions in the Baltic amber
in (similarly N. American) Eocene deposits
have been referred to the Tineidae. However,
(Hickey & Hodges 1975). Adult Gracillaridae are
since the monophyly of this family remains de-
recorded from Baltic and Dominican amber.
batable, and that of the superfamily Tineoidea
is supported by characters which are difficult to
Yponomeutoidea. The families Yponomeutidae,
observe (17), many of these fossils may well
Plutellidae, Heliodinidae are recorded from Bal-
belong to basal grades in other ditrysian, or just
tic amber (see also 'Unplaced Heteroneura'). An
heteroneurous, lineages. Tineoid-grade moths
(undescribed) heliodinid species is represented by
are also numerous in Dominican amber, and
more than 70 specimens, and is a dominant
have been found in Mexican amber as well. A
taxon in this resin (Skalski 1976).
few have been found in Tertiary rock beds (Rebel
1934; Kusnezov 1941; Skalski 1974, 1977; Jar-
Gelechioidea. About 30% of the Baltic amber
zembowski 1980; Kozlov 1987). Prolyonetia from
Lepidoptera have been referred to the Oecopho-
the Baltic amber was assigned to a family of its ridae (almost as many as to the 'Tineidae'), but
own (Kusnezov 1941), close to the Lyonetiidae. since the Oecophoridae as previously delimited
Its venation is less modified than in extant Lyo- are paraphyletic, many of these fossils presuma-
netiidae, and it has no eye-cap. Its fully devel- bly belong in the stem-lineages of other gelechi-
oped maxillary palp precludes a systematic as- oid families. Oecophorid-like moths are also nu-
signment 'above' the tineoid grade. merous in the Dominican amber. Among 20 de-
scribed species 19 are placed in fossil genera and
Nepticuloidea. While the above-mentioned al- one is refered to the recent genus Schiffermuel-
leged presence of nepticulids in the Jurassic is leria.
questionable, several leaf mines in the mid-Creta- A few Gelechiidae and Elachistidae-Elachisti-
ceous Dakota Flora exhibit so detailed similari- nae are recorded from the Baltic amber, while
ties with those made by extant nepticulids in records of Cosmopterigidae have come from the
Stigmella and Ectoedemia that their assignment Mexican amber only (Rebel 1935/1936; Kus-
to these genera seems justified (Labandeira et al. nezov 1941; Skalski 1973 a, 1976, 1977, 1979 c;
1994). A minute undescribed moth from Cana- Kozlov 1988).
dian Upper Cretaceous amber (Mutuura & Kris-
tensen, unpublished) may also be nepticulid. The Zygaenoidea. Zygaenids are known from the Mi-
family is represented by two adults in the Baltic ocene of Germany (Reiss 1936; Burgeff 1951;
amber (one referred to Ectoedemia), and one in Naumann 1987) and Spain (Fernandez-Rubio
African copal (referred to the Recent genus Aca- et al. 1991). The systematic position of an Oligo-
lyptris (= Niepeltia)), and many leaf-mines from cene moth from Aix-en-provence, determined as
the Tertiary have been assigned to it (Skalski a Zygaena species (Leestmans 1983), remains un-
1976, 1979 a, 1990 a, 1992; Kozlov 1988, Roze- clear. Fossil zygaenids are discussed by Faille
felds 1988 a). (1994).

Incurvarioidea are recorded mainly from the Bal- Cossoidea. Cossidae are recorded from the Flo-
tic amber. Two adelid species have been de- rissant (Cockerell 1926); the alleged cossid Xy-
Phylogeny and Palaeontology 19

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 Fig. 2.6. An apparent hesperioid butterfly from the
from one specimen, well preserved in the Oligo- Upper Paleocene, Fur, Denmark; wings folded below
cene shales near Aix-en-Provence and described body. If correctly identified this is the oldest butterfly
in the extant genus Pterophorus (Bigot, Nel & fossil so far known.
Nel 1986).
tained in extant members of the family. About
Pyraloid-Grade Families. A copromorphid is re-
20 fossil species have been assigned to the
corded from English Palaeogene marls (Jarzem-
Nymphalidae, including 2 species tentatively re-
bowski 1980). Thyrididae are recorded from the ferred to the Libytheinae and 3 to the Satyrinae;
Baltic amber (Skalski 1992 a), but the Eocene in several cases an 'open cell' in the hindwing
Hexeritis from Colorado, described in this fam- suggests assignments to the Nymphalinae. All
ily almost certainly does not belong here (Whal- other butterfly families have very few known fos-
ley 1971). A few Pyraloidea are recorded from sil representatives: Lycaenidae s. lat. 1, Pieridae
the Baltic amber (Kusnezov 1941), and wing 3 (family assignments tentative, except in the
fragments in Palaeocene to Oligocene sediments case of a Pleistocene/post-Pleistocene African co-
from England and France are likewise referred pal specimen indistinguishable from an extant
to this superfamily (Jarzembowski 1980). Belenois species), Papilionidae 5 and Hesperiidae
1. Many fossil butterflies have the wing pattern
Bombycoidea. The German Pliocene Sphingid- well preserved, e. g. the nymphalids Neorinops
ites, known from remains of a larva with a poste- sepulta, Prodryas persephone (Fig. 2.7), Apanth-
rior horn, is presumably a sphingid (Kernbach esis leuce and 'Agais' karaganica. Most butter-
1967). flies from the Eocene to Miocene are placed in
extinct genera, but they are in some cases phenet-
Geometroidea. Geometridae are recorded from ically similar to, and probably closely related to,
the Florissant (Cockerell 1922). Wing fragments extant taxa. For instance, in the Papilionidae-
from Paleocene to Oligocene English sediments, Parnassiinae the Miocene Doritites from Italy is
and a larva from a German Pliocene deposit are probably the sister group of Bhutanitis + Luehd-
also referred to this family (Jarzembowski 1980, orfia, and the Oligocene Thaites from France is
Kernbach 1967). So are specimens in African co- close to the most recent common ancestor of
pal (Evers 1907, Skalski unpublished). Parnassius + Hypermnestra (de Jong in press).
On the other hand some previously reported
Hesperioidea + Papilionoidea. The fossil repre- cases of particularly close relationships between
sentatives of butterflies are unusually well under- fossil and recent taxa (e. g. between the Oligo-
stood, due to de Jong's (in press) critical review cene 'Vanessa' amerindica and the extant Vanessa
of previous accounts. Butterfly fossils are partic- indica, and between the Miocene 'Vanessa' kara-
ularly abundant in the Florissant formation ganica and the extant Agais urticae) were shown
(Scudder 1889; Brown 1976; Tindal 1985, Emmel by de Jong to be founded on inadequate evi-
el al. ); in other localities usually only single spec- dence. Reports of extant forms of younger (Plio-
imens have been found. The oldest named but- cene and Pleistocene) age are unsurprising
terflies are Praepapilio colorado, P. gracilis (Papi- (Zeuner 1942, 1943; Branscheid 1968, 1969; Fuji-
lionidae) and Riodinella nympha (Lycaenidae) yama 1968, 1983; Skalski 1976). While earlier
from the Eocene Green River Shale of Colorado claims of findings of butterflies in the Baltic am-
(Durden & Rose 1978), but an apparent hesperi- ber have not been confirmed, they have been
oid (de Jong 1. c.) butterfly (Fig. 2.6) has recently found in Dominican amber (Poinar 1992) and
been identified from the Upper Paleocene of Fur, African copal.
Denmark. Interestingly, the Praepapilio species
exhibit a plesiomorphic trait (trifid, not quadri- Noctuoidea. Fossil Notodontidae, Noctuidae and
fid hind margin of forewing discal cell) not re- Arctiidae are recorded from the Eocene to the
20 Niels P. Kristensen & A n d r z e j W. Skalski

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).

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
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-
Fig. 2.8. Stauropolia nekrulenkoi (Arctiidae?); a m o t h phasized that the monophyly of this genus group
fossil with unusually well preserved wing p a t t e r n . 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
Pleistocene (Schille 1914; Thobald 1937 a; Kern- Doritites amd Thaites were West Palaearctic,
bach 1967; Leestmans 1983; Kozlov 1988; Skal- their presumed extant sister taxa are East Pa-
ski 1988). The assignment of a Cretaceous lepi- laearctic.
dopteran egg to the Noctuidae is uncertain (see Phoretic/ectoparasitic relationships between
below). Most taxa have been identified on the Arachnida and Lepidoptera have (not unexpect-
basis of single wings, and family assignments are edly) existed since the Eocene, as evidenced by a
mostly weakly founded. Stauropolia nekrutenkoi, Baltic amber gelechiid m o t h carrying a phoretic
described as an arctiid, is notable for its well pre- pseudoscorpion (Skalski 1994). Poinar et al.
served wing pattern (Fig. 2.8). (1991) recorded parasitic erythraeid mites f r o m
Phylogeny and Palaeontology 21

a tineoid and a gracillarioid moth in Dominican Bryk, F. (1934): Baroniidae, Teinopalpidae, Parnassii-
amber. dae pars I. - Das Tierreich 64: 1 - 1 3 1 .
A lepidopteran egg in an Upper Cretaceous Burgeff, H. (1951): Die Meeralpengrenze der Zygaenen
deposit has aroused particular interest, because (Lep.), eine mit Hilfe der Populationsanalyse der
Arten der Gattung Zygaena (Lepidoptera) durch-
it was referred to the Noctuidae (Gall & Tiffney
gefhrte Untersuchung ber die Lokalisation und
1983). It has been commonplace to think of bat
die Bedeutung geographischer Rassen in ihrem Zu-
prdation as the chief selective agent promoting sammenhang mit der Eiszeit. - Biol. Zentralb. 70:
the evolution of tympanic organs in nocturnal 1-23.
Lepidoptera, and while tympanate moth lineages Carpenter, F. M. (1992): Arthropoda 4. Vol. 4: Super-
were therefore not expected to predate the bats class Hexapoda. In Moore, R. C. & Kaesler, R. L.
in the fossil record, the latter group are actually (eds.): Treatise on Invertebrate Paleontology. The
so far not known before the Eocene. Two caveats Geological Soc. of America & The University of
are in place. Firstly, the bat lineage may, of Kansas.
course, be much older than documented by its Cockerell, T. D. A. (1909): A fossil tortricid moth. -
fossil record. And secondly, no diagnostic auta- Canad. Entom. 39: 416.
pomorphies in the egg structure of the higher - (1916): Some American fossil insects. roc. U. S.
Nat. Mus. 51, 2146: 8 9 - 1 0 6 .
lepidopteran lineages, including the Noctuidae
- (1919): Two interesting insects in Burmese amber.
(or Noctuoidea), have actually been identified
- Entomologist 52: 193-195.
(Hinton 1981); the family assignment, based on - (1926): A new fossil moth from Florissant. Psy-
phenetic similarity, therefore must be considered che 33: 1 6 - 1 7 .
very tentative at best. Common, I. F. B. (1970): Lepidoptera. In C S I R O
As noted above, evidence for considerable an- (sponsor) The Insects of Australia, pp. 7 6 5 - 8 6 6 .
tiquity of some lepidopteran host relationships is Carlton, Melbourne University Press.
provided by leaf mines, closely resembling those Crane, P. R. & Jarzembowski, E. A. (1980): Insect leaf
of extant 'microlepidopteran' taxa, in leaves as- mines from the Palaeocene of southern England. -
signed to extant plant families (Cretaceous) or Journ. Nat. Hist. 14: 6 2 9 - 6 3 6 .
genera (Tertiary) (Hickey & Hodges 1975; Opler Danilewskij, A. S. & Martynova, . M. (1962): Lepi-
doptera. In Rohdendorf, . . (ed.). Osnovy paleon-
1973, 1974, 1982; Labandeira et al. 1994). See
tologa, pp. 303-307. Izd. Akad. Nauk SSSR,
Davis, D. R. (1986): A new family of monotrysian
moths from austral South America (Lepidoptera:
Palaephatidae), with a phylogenetic review of the
We are indebted to Dr. M. J. Scoble for critical Monotrysia. - Smiths. Contr. Zool. 434, 202 pp.
comments of a draft of this chapter, and to Drs. - (1989): Generic revision of the Opostegidae with a
synoptic catalog of the World's species (Lepidop-
M. V. Kozlov and J. Minet for valuable informa-
tera: Nepticuloidea). - Smiths. Contr. Zool. 478,
tion. Mr. G. Brovad and Mr. S. Langemark pro-
97 pp.
vided much appreciated assistance with illustra-
- (1994): New leaf-mining moths from Chile, with re-
tions. marks on the history and composition of Phylloc-
nistinae (Lepidoptera: Gracillariidae). Neotropical
Microlepidoptera XXV. - Tropical Lepidoptera 5:
References 65-75.
Bigot, ., Nel, . & Nel, J. (1986): Description de la de Jong, R. (inpress): The fossil butterflies in phyloge-
premire espce fossile connue de Ptrophore (Lepi- netic perspective. - Ent. scand.
doptera Pterophoridae). - Alexanor 14: 2 8 3 - 2 8 8 . - , Vane-Wright, R. I. & Ackery, P. (1996): The higher
Branscheid, F. (1968): Ein weiterer Schmetterlingsfl- classification of butterflies (Lepidoptera): problems
gel aus dem Pliozn von Willershausen. - Beih. and prospects. - Ent. scand. 27: 6 5 - 1 0 1 .
Ber. Naturhist. Ges. Hannover 6: 4 1 - 4 2 . Dugdale, J. S. (1974): Female genital configuration in
- (1969): Funde von Tagfaltern (Rhopalocera, Lepi- the classification of Lepidoptera. - N. Z. J. Zool.
dopt.) im Pliozn von Willershausen. - Beih. Ber. 1: 127-146.
Naturhist. Ges. Hannover 113: 101-106. Durden, Ch. J. & Rose, H. (1978): Butterflies from the
Brock, J. P. (1971): A contribution towards an under- Middle Eocene: the earliest occurrence of fossil
standing of the morphology and phylogeny of the Papilionoidea (Lepidoptera). - Texas Mem. Mus.,
Ditrysian Lepidoptera. - J. nat. Hist. 5: 2 9 - 1 0 2 . Pearce-Sellards Ser. 29: 1 - 2 5 .
- (1988): Reply to Scott's Criticism. - J. Res. Lepid. Ehnbom, K. (1948): Studies on the central and sympa-
26: 2 4 0 - 2 4 7 . thetic nervous system and some sense organs in the
- (1990 a): Pupal protrusion in some bombycoid head of neuropteroid insects. Opuse, ent. Suppl.
moths (Lepidoptera. Ent. Gaz. 41: 9 1 - 9 8 . 8, 162 pp.
- (1990 b): Origins and phylogeny of butterflies. In Emmel, T. C., Minno, M. C. & Drummond, B. A.
Kudrna, O. (ed.) Butterflies of Europe 2, pp. 2 0 9 - (1992): Florissant Butterflies. A Guide to the Fossil
233. Aula Verlag, Wiesbaden. and Present-Day Species of Central Colorado.
Brown, F. M. (1976): t Oligodonta florissantensis 118 pp. Stanford University Press.
gen.n., sp. nov. (Lepidoptera: Pieridae). - Bull. Al- Evers, J. (1907): Copal-Schmetterlinge. - Entom.
ly Mus. 37: 1 - 4 . Jahrb. 16: 129-132.
22 Niels P. Kristensen & Andrzej W. Skalski

Faille, L. (1994): Guide pour l'identification des es- Kernbach, K. (1967): ber die bisher im Pliozn von
pces franaises du genre Zygaena. 52 pp. Edition Willershausen gefundenen Schmetterlings- und
J.-M. Desse. Raupenreste. - Ber. naturh. Ges. Hannover 111 :
Fehrenbach, H. (1995): Egg shells of Lepidoptera - 103-108.
Fine structure and phylogenetic implications. - Knorre, D. von (1978): Fossile Insektenreste aus letzt-
Zool. Anz. 234: 1 9 - 4 1 . interglazialen Travertinen Thringens. - Quartr-
Fernandez-Rubio, F., Penlver, E. & Martinez- palontologie 3: 9 3 - 9 5 .
Delclos, X. (1991): Zygaena? turolensis, una nueva Koponen, M. & Nuorteva, M. (1973): ber subfossile
especie de Lepidoptera Zygaenidae del Mioceno de Waldinsekten aus dem Moor Piilonsuo in Sdfinn-
Rubielos de Mora (Teruel). Description y filogenia. land. Acta Entom. Fenn. 29: 384.
- Est. Mus. Cienc. Nat. de Alava 6: 7793. Koteja, J. (1986): Eopterum devonicum - should fossils
Freeman, T. . (1965): A lepidopterous leaf-mine from be given species names? - Przegl. Zool. 30, 3:
the Tertiary period. - Can. Ent. 97: 1069-1070. 257-267.
Fujiyama, I. (1968): A Pleistocene fossil Papilio from Kozhanchikov, I.V. (1957): A new representative of
Shiobara, Japan. - Bull. Nat. Sei. Mus. Tokyo 11: the family Cossidae from the Miocene deposits of
85-95. Caucasus (Lepidoptera, Insecta). Dokl. Acad.
- (1983): A Pleistocene nymphalid butterfly from Shi- Sci. USSR 113: 6 7 5 - 6 7 7 . [In Russian],
obara, Japan. - Bull. Nat. Sei. Mus. Tokyo, Ser. C. Kozlov, M. V. (1987): New tineoid-moths from Baltic
9: 121-128.
amber. - Paleont. Zhurn. 4: 5 9 - 6 8 . [In Russian].
Gall, L. F. & TifTney, B. H. (1983): A fossil noctuid
- (1988): Paleontology of the Lepidoptera and prob-
moth egg from the Late Cretaceous of Eastern
lems of phylogeny of the order of the Papilio-
North America. - Science 219: 507-509.
noidea: In Ponomarenko, A. G. (ed.) Cretaceous
Handlirsch, A. (1906-1908): Die fossilen Insekten und
biocenotic crisis and evolution of insects, pp. 1 6 - 6 9 .
die Phylogenie der rezenten Formen 13, 1430 pp.
Nauka, Moskva. [In Russian].
Engelmann, Leipzig.
- (1989): New Lepidoptera (Papilionida) from Upper
Hennig, W. (1953): Kritische Bemerkungen zum phylo-
genetischen System der Insekten. Beitr. Ent. 3 Jurassic and Lower Cretaceous. - Paleont. Zhurn.
(Sonderheft): 1 - 8 5 . 4: 3 7 - 4 2 . [In Russian],
- (1966): Phylogenetic Systematics. Illinois University Kristensen, N. P. (1984): Studies on the morphology
Press, 263 pp. Urbana. and systematics of primitive Lepidoptera (Insecta).
- (1981): Insect Phylogeny. (Pont., A. C. & Schlee, D., - Steenstrupia 10: 141-191.
eds.), 514 pp. J. Wiley & Sons, Chichester, New - (1991): Phylogeny of extant hexapods. In C S I R O
York, Brisbane, Toronto. (ed.) The Insects of Australia, 2nd edn, pp. 1 2 5 -
Henrotay, M. (1986): Dcouverte d'un nouveau gise- 140. Melbourne University Press.
ment de Lpidoptres, d'autres insectes et d'Araig- - (1997): Early evolution of the Lepidoptera +
nes fossiles a Dauphin (Hte Provence, France). Trichoptera lineage: phylogeny and the ecological
Linn. Belg. 10: 2 6 6 - 2 7 9 . scenario. Mem. Mus. natn. Hist. nat. 173: 253
Heppner, J. . (1987): Copromorphidae. Alucitidae. 271.
Carposinidae. Epermeniidae (Copromorphoidae); - & Nielsen, E. S. (1979): A new subfamily of micro-
Glyphipterigidae. Plutellidae (Yponomeutoidea). In pterigid moths from South America. A contribu-
F. W. Stehr (ed.) Immature Insects, pp. 3 9 9 - 4 0 5 tion to the morphology and phylogeny of the
Dubuque, Kendall/Hunt. Micropterigidae, with a generic catalogue of the
- (1991): Faunal regions and the diversity of Lepi- family (Lepidoptera: Zeugloptera). - Steenstrupia
doptera. Tropical Lepidoptera 2 (Suppl. 1): 1 - 8 5 . 5: 6 9 - 1 4 7 .
Hickey, L. J. & Hodges, R. W. (1975): Lepidopteran - (1983): The Heterobathmia life history elucidated:
leaf mine from the early Eocene Wind River For- Immature stages contradict assignment to suborder
mation of northwestern Wyoming. Science 189: Zeugloptera (Insecta, Lepidoptera). - Z. zool.
718-720. Syst. Evol.-Forsch. 21: 101-124.
Hinton, . E. (1958): The phylogeny of the panorpoid Khne, W. G., Kubig, L. & Schlter, T. (1973): Eine
orders. - A. Rev. Ent. 3: 181-206. Micropterigidae (Lepidoptera, Homoneura) aus
- (1981): The Biology of Insect Eggs 1 - 3 , 1125 pp. mittelcretazischen Harz Westfrankreichs. - Mitt.
Pergamon, Oxford. Dtsch. ent. Ges. 32: 6 1 - 6 5 .
Hiura, I. & Miyatake, Y. (1974): On the fossil Arthro-
Kuhn, O. (1951): Ein vermutlicher Schmetterling, Geis-
poda in Mizuami amber from Gifu Prefecture
feldia benkerti n. g. n. sp. aus dem Lias ... Nord-
(Pleistocene). Bull. Mizunami Fossil Mus. 1 :
frankens. Neues Jahrb. Geol. Palont. Monats-
heftel: 5 8 - 6 1 .
Hong, Y. (1984): Curvicubitidae fa. nov. (Lepidoptera?
Kuroko, H. (1987): A fossil leaf mine of Nepticulidae
Insecta) from Middle Triassic of Shaanxi. - Acta
(Lepidoptera) from Japan. - Bull Sugadaira Mon-
Palaeont. Sin. 2: 782-785.
tane Rs. Cent. 8: 119-121.
Ivanov, V. D. (1994): Comparative analysis of the wing
articulation in archaic Lepidoptera. - Ent. Obozr. Kusnezov, N. J. (1941): A revision of the amber Lepi-
73: 569-590. [In Russian], doptera. Izd. Akad. Nauk SSSR, M o s k v a - L e -
Jarzembowski, E. A. (1980): Fossil insects from the ningrad, 136 pp. [In Russian].
Bembridge Marls, Palaeogene of the Isle of Wight, Kuznetznov, V. I. & Stekolnikov, A. A. (1979): Classi-
Southern England. - Bull. Brit. Mus. (Nat. Hist.), fication and phylogenetic relationships of the fami-
Geology 33: 2 3 7 - 2 9 3 . lies and subfamilies of the Pyraloidea (Lepidoptera)
Kawall, J. H. (1876): Organische Einschlsse im of the Palaearctic fauna with regard of functional
Bergkrystall. Bull. Soc. Imper. Natur. Mose. 51 : morphology of male genitalia. Proc. Zool. Inst.
170-173. Acad. Sci. USSR 82: 4 3 - 7 3 . [In Russian].
Phylogeny and Palaeontology 23

Kyrki, J. (1983): Adult abdominal sternum II in the Nielsen, E. S. (1985): The monotrysiam heteroneuran
Ditrysian tineoid superfamilies - morphology and phylogeny puzzle: a possible solution. - Proc. 3rd.
phylogenetic significance (Lepidoptera). - Ann. Congr. Europ. Lep.: 138-143.
Ent. Fenn. 49: 8 9 - 9 4 . - (1989): Phylogeny of major lepidopteran groups. In
Labandeira, C. C., D. L. Dilcher, D. R. Davis, and Fernholm, ., Bremer, . & Jrnvall, . (eds.) The
D. L. Wagner (1994): Ninety-seven million years of Hierarchy of Life, pp. 281-294. Elsevier.
angiosperm-insect association: Paleobiological in- - & Kristensen, N. P. (1996): The Australian moth
sights into the meaning of coevolution. Proc. family Lophocoronidae and the basal phylogeny of
Natl. Acad. Sci. 91: 12278-12282. the Lepidoptera-Glossata. - Invertebrate Taxon-
Larsson, S. G. (1975): Paleobiology and mode of bur- omy 10: 1199-1302.
ial of the insects of the Lower Eocene Mo-clay of Opler, P. (1973): Fossil lepidopterous leaf mines de-
Denmark. - Bull. geol. Soc. Denmark 24: 193-209. monstrate the age of some insect-plant relation-
- (1978): The Baltic Amber. A Palaeobiological ships. - Science 179: 1321-1323.
Study. - Entomonograph 1: 1 - 1 9 2 . - (1974): Biology, ecology and host specificity of
Leestmans, R. (1983): Les lpidoptres fossiles trouvs Microlepidoptera associated with Quercus agrifolio
en France (Insecta Lepidoptera). - Linn. Belg. 9: (Fagaceae). - University Cai Pub. Ent. 75: 1 - 8 3 .
64-89. - (1982): Fossil leaf-mines of Bucculatrix (Lyonetii-
Lomnicki, A . M . (1894): Pleistocene insect from dae) on Zelkova (Ulmaceae) from Florissant, Colo-
Boryslaw. - Muz. im. Dzieduszyckich we Lwowie 4: rado. - J. Lepid. Soc. 36: 145-147.
6 - 1 1 6 . [In Polish], Philpott, A. (1926): The maxillae in the Lepidoptera.
Lfstedt, C. & Kozlov, M. V. (1996): A phylogenetic - Trans. . Z. Inst. 57: 721-746.
analysis of pheromone comminication in primitive Pierce, W. D. (1947): Fossil arthropods of California.
moths. In Card, R. T. & Minks, A. K.. (eds.) Phero- 14. A progress report on the McKittrick asphalt
mone research; New Directions, pp. 473-489. Chap- field. - Bull. South. Calif. Acad. Sci. 46: 138-143.
man & Hall, New York. Poinar, Jr., O. G. (1992): Life in Amber. Stanford Uni-
MacKay, M. R. (1969): Microlepidopterous larvae in versity Press, Stanford.
Baltic amber. - Can. Ent. 101: 1173-1180. - & Brown, J. W. (1993): A new fossil tortricid (Lepi-
- (1970): Lepidoptera in Cretaceous amber. - Sci- doptera: Tortricidae) from Dominican amber. -
ence 167: 379-380. Ent. scand. 24: 2 5 - 2 9 .
Martins-Neto, R. G. & Vulcano, . . (1989): Amphi- Rammert, U. (1994): Morphologische Untersuchungen
esmenoptera (Trichoptera + Lepidoptera) na zur Aufdeckung der stammesgeschichtlichen Verhlt-
Formacao Santana (Cretceo Inferior) Bacia do nisse der basalen Gruppen der ditrysen Lepidopteren
Araripe, Nordeste do Brasil. I - Lepidoptera (In- (Lepidoptera: Ditrysia). Dissertation, Universitt
secta). - An. Acad. bras. Ci., 61: 459-466. Bielefeld 193 pp., 25 pis. Flintbek.
Martynova, O. M. (1960): On the wing venation in Rasnitsyn, . P. (1983): First finding of the Lepidop-
Lepidoptera. - Ent. Obozr., 39: 296-299. [In Rus- tera of the Jurassic age. - Dokl. Akad. Nauk SSSR
sian]. 269: 467-471. [In Russian],
Minet, J. (1983): Etude morphologique et phylogn- - & Kozlov, M. V. (1990): A new group of fossil in-
tique des organes tympaniques des Pyraloidea. I sects: scorpion-fly with adaptation of cicadas and
gnralits et homologies. (Lep. Glossata). - Annls. moths. - Dokl. Akad. Nauk SSSR 310: 973-976.
Soc. ent. Fr. (N. S.) 19: 175-207. [In Russian].
Regier, J. C., Friedlander, T., Leclerc, R. F., Mitter,
- (1984): Contribution a l'analyse phylogntique des
C. & Wiegmann, . M. (1995): Lepidoptera phylog-
Nolpidopteres (Lepidoptera, Glossata). - Nouv.
eny and applications to comparative studies of de-
Rev. Ent. (N. S.) 2: 139-149.
velopment. In Goldsmith, M. R. & Wilkins, A. S.
- (1986): Ebauche d'une classification moderne de
(eds.) Molecular Model Systems in the Lepidoptera,
l'ordre des Lpidoptres. - Alexanor 14: 291-313.
pp. 107-139. Cambridge University Press, Cam-
- (1991): Tentative reconstruction of the ditrysian
phylogeny (Lepidoptera: Glossata). - Ent. scand.
Reiss, H. (1936): Zygaena miocenica, Ein Zygnenfund
22: 6 9 - 9 5 .
aus der Tertrzeit. - Entom. Rund. 53: 554556.
- (1996): Phylogeny of Ditrysia. Abstracts, XX Int.
Rebel, H. (1898): Fossile Lepidopteren aus der Mio-
Congr. Ent. Florence: 32.
cnformation von Gabbro. Sitzber. Akad. Wien
Naumann, C. M. (1987): On the phylogenetic signifi-
107: 731-745.
cance of two Miocene zygaenid moths (Insecta,
- (1934): Bernstein-Lepidopteren. Palaobiologica
Lepidoptera). - Palaeontol. Z. 61: 299-308.
6: 1 - 1 6 .
Nel, A. & Descimon, H. (1984): Une nouvelle espce - (1935/36): Bernstein-Lepidopteren. - Dtsch. En-
de Lpidoptre fossile du Stampien de Creste (04) tom. Ztschr. Iris 49: 162-186.
(Lepidoptera-Satyridae). - Gol. Mediterran 11: Richter, G. & Storch, G. (1980): Beitrge zur Ernh-
287-293. rungsbiologie eozner Fledermuse aus der "Grube
Nel, . & Nel, J.(1985): A propos de chenilles et de Messel". - Natur u. Mus. 110: 353-367.
chrysalides fossiles du stampien en France (Lepi- Riek (1976): A new collection of insects from the Up-
doptera). - Alexanor 14: 126-130. per Triassic of South Africa. - Ann. Natal. Mus.
- (1986): A propos de deux Lpidoptres fossiles du 22: 791-820.
Stampien d'Aix-en-Provence (Lepidoptera, Hespe- Rozefelds, A. C. (1985): The first record of fossil leaf
riidae et Nymphalidae Satyrinae). - Bull. Mus. mining from Australia. New Zealand Geol. Sur-
natn. Hist, nat., 4: 343-350. vey Record 9: 8 0 - 8 1 .
Nekrutenko, Y. P. (1965): Tertiary nymphalid butter- - (1988 a): Insect leaf mines from the Eocene Angle-
flies and some phylogenetic aspects of systematic sea locality, Victoria, Australia. - Alcheringa 12:
lepidopterology. - J. Res. Lep. 4: 149-158. 51-57.
24 Niels P. Kristensen & Andrzej W. Skalski

- (1988 b): Lepidoptera mines in Pachypteris leaves - (1975): Notes on present status of botanical and
(Corystospermaceae: Pteridospermophyta) from zoological studies of ambers. Studi a ricerche sulla
the Upper Jurassic/ Lower Cretaceous Battle Camp problematica dee'ambra. - ATTI della Cooperazi-
Formation, North Queensland. Proc. Roy. Soc. one Interdisciplinare Italo-Polacca, 1: 153-175.
Queensland 99: 7 7 - 8 1 . - (1976): Les lpidoptres fossiles de l'ambre. Etat ac-
Robbins, R. K. (1987): Logic and phylogeny: A cri- tuel de nos connaissances. - Linn. Belg. 6 - 7 : 1 5 4 -
tique of Scott's phylogenies to the butterflies and 169; 8: 195-208; 9: 221-233.
Macrolepidoptera. J. Lep. Soc. 41: 214216. - (1977): Studies on the Lepidoptera from fossil res-
Robinson, G. S. (1977): A taxonomic revision of the ins. Part I. General remarks and description of new
genus Callipielus Butler (Lepidoptera: Hepialidae). genera and species of the families Tineidae and Oe-
- Bull. Brit. Mus. nat. Hist. (Ent.) 35: 103-121. cophoridae from the Baltic amber. - Prace Muz.
- (1988): A phylogeny for the Tineoidea (Lepidop- Ziemi 26: 324.
tera). - Ent. scand. 19: 117-129. - (1979): Studies on the Lepidoptera from fossil res-
- & Nielsen, E. S. (1993): Tineid genera of Australia. ins. Part VII. Palaeodepressaria hannemanni gen. et
Monographs on Australian Lepidoptera 2: 1 - 3 4 4 . spec. nov. (Oecophoridae) from the Baltic amber.
Rohdendorf, . . & Zherihin, V. V. (1974): Paleontol- - Prace Muz. Ziemi 32: 101-107.
ogy and protection of nature. Priroda 5: 8291. - (1979 a): Records of oldest Lepidoptera. - Nota
[In Russian], lepid. 2: 6166.
Schepdael, J. van (1974): Macrolpidoptres fossiles - (1979 b): A new lower Cretaceous representative of
du domaine palarctique. La palontologie au ser- the family Micropterigidae (Lepidoptera) from
vice de l'cologie. - Natur. Belg. 55: 3 - 3 7 . Transbaikalia. - Paleont. Zhurn. 2: 9097. [In
Schille, F. (1914): Lepidoptera. In Wykopaliska starun- Russian],
skie. Muzeum im. Dzieduszuckich we Lwowie, 15, - (1979 c): Genesis of the lepidopterofauna of the
pp. 8 9 - 9 1 . Krakow. West Palearctic on the basis of fossil material. -
Schlter, T. (1974): Kritisches zum Nachweis von Verhandl. VII SIEEC, Leningrad: 279-283. [In
Schmetterlingsschuppen aus einem fossilen Harzen
der mittleren Kreide Nordwestfrankreichs. En- - (1984): A new lower Cretaceous Lepidoptera (Ho-
tern. Ztschr. 84: 253-256. moneura). Bull. Pol. Acad. Sci., Biol. Sci. 32:
Scoble, M. J. (1992): The Lepidoptera. Form, Function
- (1988): Stauropolia nekrutenkoi gen. et sp. n. (Lepi-
and Diversity, 404 pp. Oxford Univ. Press, Oxford.
doptera, Arctiidae) a new fossil tiger-moth from
- In search of butterfly origins: morphology and
Miocene of the Caucasus. - Vestn. zool. 4: 2 1 - 2 5 .
molecules. Trends. Ecol. Evol. 11: 274-275.
[In Russian].
Scott, J. A. (1985): The phylogeny of butterflies (Papi-
- (1989): Micropterigidae in fossil resins with special
lionoidea and Hesperioidea). J. Res. Lepid. 23:
emphasis on the past and present distribution of
this family. - Nota lepid. Suppl. 1: 44.
- (1986): On the monophyly of the Macrolepidop-
- (1990 a): Lepidoptera in fossil resins with emphasis
tera, including a reassessment of their relationship on new investigations. - Prace Muz. Ziemi 41:
to Cossoidea and Castnioidea, and a reassignment 163-164.
of Mimallonidae to Pyraloidea. J. Res. Lepid. 25: - (1990 b): An annotated review of fossil records of
30-38. lower Lepidoptera. Bull. Sugadeira Montane Res.
- (1989): Parallelism and Phylogenetic Trees. J. Center, Tsukuba Univ. 11: 125-128.
Res. Lepid. 27: 257-258. - (1992 a): The families Nepticulidae and Thyrididae
- & Wright, D. M: (1990): Butterfly phylogeny and in Baltic amber (Lepidoptera). Nota lepid. Suppl.
fossils. In Kudrna, O. (ed.) Butterflies of Europe 2, 4: 155-156.
pp. 152-208. Aula Verlag, Wiesbaden. - (1992 b): Studies on the Lepidoptera from fossil res-
Scudder, S. H. (1875): Fossil Butterflies. - Am. Ass. ins. Part III. Two new genera and species of Tortri-
Adv. Sci., Mem. 1: 1 - 9 9 . cidae from the Baltic amber. Roczn. Muz. Gr-
- (1889): The fossil butterflies of Florissant. - U. S. nosl. Bytom., Entom. 3: 137146.
Geol. Survey, Annu. Rep. 8: 433-474. - (1994): First fossil case of phoresy of pseudoscor-
Shields, O. (1976): Fossil butterflies and the evolution pion on moth. - Abstract XVI Congress of the Po-
of the Lepidoptera. - J. Res. Lepid. 15: 132-143. lish Zoologists.
- (1993): Is Micropterigidae or Agathiphagidae the - (1995): Study on the Lepidoptera from fossil resins.
most primitive lepidopteran family? - Tyo to Ga Part I. A new fossil genus Baltomartyria, and rede-
44: 152-156. scription of Baltomartyria proavitella (Rebel, 1936)
Skalski, A. W. (1973 a): Studies on the Lepidoptera comb. nov. (Lepidoptera, Micropterigidae) from
from fossil resins. Part II. Epiborkhausenites obscu- Baltic amber. - Amber & Fossils 1: 2627.
ro trimaculatus gen. et spec. nov. (Oecophoridae) - (in press, a): Study on the Lepidoptera from fossil
and tineid-moth discovered in the Baltic amber. resins. Part IX. A new remarkable micropterigid
Acta Palaeont. pol. 18: 153-160. moth, Saxibatinca meyi gen. et spec. nov. (Lepidop-
- (1973 b): Studies on the Lepidoptera from fossil res- tera, Micropterigidae) from the lower Miocene fos-
ins. Part VI. Tortricidrosis inclusa gen. et spec. nov. sil resin of Bitterfeld district in Saxony (Germany).
from the Baltic amber (Lep. Tortricidae). - Dtsch. - Nota lepid.
ent. Z., N. F, 20: 339-344. - (in press, b): Study on the Lepidoptera from fossil
- (1974): Zwei neue Gattungen und Arten der Familie resins. Part XII. A second sabatincoid-like micro-
Tineidae aus dem baltischen Bernstein. Studien an pterigid moth, and a new species of the genus
Lepidopteren aus fossilen Harzen. V. Beitr. Ent. Micropterix Hbner (Lepidoptera, Micropterigi-
24: 9 7 - 1 0 4 . dae) from the Eocene Baltic amber.
Phylogeny and Palaeontology 25

- (in press, c): Study on the Lepidoptera from fossil - (1978): New taxa of fossil and recent Micropterigi-
resins. Part VIII. A new fossil homoneurous-moth dae with a discussion of their evolution and a com-
from the Upper Cretaceous Siberien retinite (Lepi- ment on the evolution of Lepidoptera (Insecta). -
doptera, Homoneura). - Acta zool. cracov. Ann. Transvaal Mus. 31: 7 1 - 8 6 .
- (in press, d): Study on the Lepidoptera from fossil - (1985): The systematics and palaeogeography of the
resins. Part X. First fossil acrolophid-moth from Lower Jurassic insects of Dorset, England. - Bulle-
Dominican amber (Lepidoptera, Acrolophidae). - tin of the British Museum (Natural History), Geol-
Tropical Lepidoptera. ogy, 39: 107-189.
Strauss, A. (1977): Gallen, Minen und andere Frass- - (1986): A review of the current fossil evidence of
spuren im Pliokn von Willershausen am Harz. - Lepidoptera in the Mesozoic. - Biol. Journ. Linn.
Verh. Bot. Ver. Prov. Branderburg 113: 4 3 - 8 0 . Soc. 28: 253-271.
Sukatsheva, I. (1982): Historical development of the Wiegmann, . M. (1994): The Earliest Radiation of the
Trichoptera. - Trudy paleontol. Inst. USSR Acad. Lepidoptera: Evidence from 18S rDNA. PhD disser-
Sci. 197: 1 - 1 1 3 . [In Russian], tation, University of Maryland 230 pp.
Thobald, . (1937 a): Les insectes fossiles des terrains Wiggins, G. B. (1992): Comments on the phylogeny of
oligocnes de France. George Thomas, Nancy. pupation behaviour in Trichoptera: a response to
- (1937 b): Note complmentaire sur les insectes fos- Weaver. - Cladistics 8: 181-185.
siles oligocnes des gypses d'Aix-en-Provence. Willmann, R. (1984): Zur systematischen Stellung me-
Bull. Soc. Sci. Nancy (N. S.) 1937: 157-178. sozoischer und tertirer Mecopteren einschlielich
Tindale, . B. (1945): Triassic insects of Queensland. Eoses triassica Tindale (angeblich Lepidoptera) (In-
1. Eoses, a probable lepidopterous insect from the secta, Holometabola). - Palont. Z. 58: 231-246.
Triassic Beds of Mt. Crosby, Queensland. - Proc. - (1989): Evolution und phylogenetisches System
Roy. Soc. Queensland, 56: 3 7 - 4 6 . der Mecoptera (Insecta: Holometabola). - Abh.
- (1980): Origin of the Lepidoptera, with description senckenberg. naturforsch. Ges. 544: 1 - 1 5 3 .
of a new Mid-Triassic species and notes on the ori- Zeuner, F. E. (1931): Die Insektenfauna des Bttinger
gin of the butterfly stem. - J. Lep. Soc. 34: 2 6 5 - Marmors. - Forstschr. Geol. Palaeontol. 9: 274-
285. 406.
- (1942): Two new fossil butterflies of the familiy
- (1985): A butterfly-moth (Lepidoptera Castaniidae)
Pieridae. Ann. Mag. Nat. Hist. 11: 409-416.
from the Oligocene shales of Florissant, Colorado.
- (1943): On recent and fossil Pseudonaclia Butler
- J. Res. Lep. 24: 31 - 4 0 .
(Amatidae). Ann. Mag. Nat. Hist. 10: 140-144.
Verity, R. (1947): Le farfalle diurne d'Italia 3. Mar-
Zhag, J. (1989): Fossil insects from Shanwang, Shan-
zocco, S. ., Firenze.
dong, China. Shandong Science and Technology
Weaver III, J. S. (1992a): Remarks on the evolution of
Publishing House.
Trichoptera: A critique of Wiggins and Wichard's
Zherihin, V. V. & Sukatsheva, I. D. (1973): On Creta-
classification. - Cladistics 8: 171-180.
ceous "ambers" (retinites) with insects of North
- (1992b): Further remarks on the evolution of
Siberia. Doklady na XXIV ezhegodnom tshtenii
Trichoptera: a reply to Wiggins. - Cladistics 8: pamiatiN. A. Cholodkovskogo 1971: 3 - 4 8 . [In Rus-
187-190. sian].
Weiler, S. J. & Pashley, D. P. (1995): In search of but-
terfly origins. - Mol. Phyl. Evol. 4: 235-246.
- Martin, J. A. & Constable, J. L. (1994): Phylogeny Addendum
of noctuoid moths and the utility of combining in-
Ivanov (1996) claimed the elongated forewing 1st
dependent nuclear and mitochondrial genes. -
Syst. Biol. 43: 194-211.
axillary to be untenable as evidence for macro-
Whalley, P. (1971): The Thyrididae (Lepidoptera) of lepidopteran monophyly.
Africa and its Islands. A taxonomic and Zoogeo- Ivanov, V. D. (1996): Wing articulation of Lepidoptera
graphie study. - Bull. Brit. Mus. nat. Hist. (Ento- and the status of the Macrolepidoptera. - Ent.
mology). Suppl. 17: 1 - 1 9 8 . Obozr. 59: 516-529. [In Russian],
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 important (suprageneric) exceptions to

this principle are the categories at subfamily/
The systematics chapters in this Handbook are all family/superfamily level (for which the Interna-
written on the basis of 'Hennigian' phylogenetic tional Code of Zoological Nomenclature pre-
systematics or cladistics. Therefore, taxa which scribes specific suffixes). The system adopted re-
are identified as likely poly- or paraphyletic (i. e., flects the firm belief that the family category has
which do not include all descendants of the come to play such an important role in the lan-
members' last common ancestor) have been re- guage by which biologists communicate (thus it
jected, or they are accepted merely as prelimi- is pivotal in many catalogue projects) that its
nary groupings, pending further analysis. abandonment now, for whatever reason, would
Phylogenetic systematists are not, however, in be counterproductive. Hence every recent genus
agreement about how even a fully resolved phy- should be assigned to a family (a view with
logeny is best transformed into a written classifi- which 'pure systematizers' would not agree). A
cation. Indeed it has been argued that 'classifica- similar case may be made for superfamilies. Ar-
tion' is an inappropriate term in this context: guably the superfamily category has for some
phylogentic systematists are concerned not with time been so extensively used in Lepidoptera sys-
'classifying', but with 'systematizing', because tematics (the same may not be true in all other
monophyla/clades are individuals forming parts major insect groups) that it is desirable that every
of a 'system', rather than elements of definable family is assigned to a superfamily. For this
classes (Griffiths 1974). Adherents of this view reason new, and admittedly redundant, super-
claim that the Linnean categories have outlived families are here created for the three families
their usefulness (see e. g. Ax 1984/1987; Will-
which were left outside the superfamilies in pre-
mann 1987, 1989; de Queiroz & Gauthier 1992),
vious Lepidoptera systems. Needless to say, it is
and that the optimal written presentation of the
disturbing that no less than 27 of the 46 super-
phylogenetic subordination is an indented (or
families here recognized are redundant, compris-
numerically coded) list, in which all recognized
clades are named but not given categorial ranks. ing only a single family. Extensive amalgam-
The principal alternative approach has been ations of such superfamilies with others are
termed the 'annotated Linnean classification'. highly desirable, to the extent future research can
Here the Linnean categories are retained (even disclose convincing synapomorphies to support
an expanded array of them), but a suite of con- them. A subfamily classification has been
ventions, most importantly that of 'sequencing', adopted in 48 of the 121 lepidopteran families
may be adopted to permit an exact representa- here recognized; subfamilies have very variable
tion of the cladistic information while prolifera- status in the literature, but in some cases they
tion of taxon names as well as changes in previ- are entities of widespread and longstanding rec-
ous systems are minimized (for more detailed ac- ognition.
counts see e. g. Wiley 1981, Forey 1992). Table 1 presents a synoptic classification of the
lepidopteran superfamilies recognized in this
It seems immediately obvious that the two
kinds of approach each has its merits and prob- Handbook; phylogenetic systems of individual
lems, and the Lepidoptera system adopted here superfamilies have been worked out at very dif-
embodies elements of both. It is accepted that ferent levels of refinement, as discussed in chap-
the notion of categorial ranks, rooted in pre-evo- ters 14 through 1 - 1 9 . The table adopts the se-
lutionary systematics, may represent an unneces- quencing, plesion, incertae sedis and sedis muta-
sary impediment to efficient systematization. bilis conventions of the 'annotated Linnean clas-
Formal categories have therefore been largely sification' rather than a consistent subordina-
abandoned in the present system: no lepidop- tion. Names have been applied only to a selec-
teran 'suborders', 'infraorders' etc. are recog- tion of the more inclusive clades recognized; the
nized here. Use of the redundant names Zeug- choice at least partly reflects an assessment of
loptera, Aglossata and Heterobathmiina (for the the firmness of the support for the monophyly
superfamilies Micropterigoidea, Agathiphago- of the clades in question and/or the conspicuous-
idea and Heterobathmioidea respectively) is dis- ness of their autapomorphies. Naturally these
continued altogether, and named high-rank decisions are open to debate. Thus the hypothe-
monophyla (Glossata, Ditrysia, etc.) are simply sized clade comprising all non-micropterigoid
designated as 'clades'. 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)
Superfamilies 4 - 4 6 : clade GLOSSATA 28. ALUCITOIDEA (2)
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)
Superfamilies 8 - 4 6 : clade NEOLEPIDOPTERA 32. COPROMORPHOIDEA (2)
Superfamilies 8 - 9 : clade EXOPORIA 33. HYBLAEOIDEA (1)
Superfamilies 10-46: clade HETERONEURA Superfamilies 36-46: clade MACROLEPIDOP-
11. INCURVARIOIDEA (6) Superfamilies 3638, 39-45, 46: sedis mutabilis
Superfamilies 12, 13, 14-46: sedis mutabilis Superfamilies 36, 37, 38: sedis mutabilis
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)
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 tion, and the results should be checked against
that of, e. g., the Obtectomera. descriptions. This key will not work for brachyp-
It must be emphasized that the choice of clas- terous/wingless forms, for which reference
sificatory approach ultimately depends on judge- should be made to the reviews by Heppner
ments which are by necessity subjective. For ex- (1991) and, particularly, Sattler (1991).
ample, the systematizers' attitude to subordina-
tion/naming is dependent on the clearly sub- 1 Hindwing with Rs unbranched; fore-
jective belief that the (potentially very numerous) wing with jugal lobe not markedly pro-
new names needed for all recognized clades duced Heteroneura, 11
above the generic level do not "represent a com- Hindwing Rs with 3 or 4 branches; fore-
plication worth mentioning" (Willmann 1987: wing almost always with jugal lobe
228); the refusal of consistently naming such markedly produced 2
clades reflects the equally subjective view that 2 (1) Maxillary galeae forming a proboscis,
the opposite is true. Presumably, however, there usually spirally coiled in repose, some-
will be broad agreement that the principal con- times secondarily reduced or absent;
cern of phylogenetic systematists should be the mandibles often strongly reduced, artic-
cladograms, their support and the biological in- ulations with head capsule undeveloped
ferences that can be drawn from them, rather Glossata, 5
than the written classifications/systems per se. Maxillary galeae unmodified, not form-
ing proboscis; mandibles large; articula-
tions with head capsule well developed
Key to superfamilies, adults
Modified from Nielsen & Common 1991. All 3 (2) M4 present; ocelli absent; tibial spurs 1-
keys to higher taxa must be used with reserva- 4-4; larger, caddisfly-like moths; fore-
Classification and Keys to Higher Taxa 29

wing pattern mottled brownish, non-iri- iae with prominent spines

descent; Australia, SW Pacific Nepticuloidea ( 1 - 6 )
Agathiphagoidea (1 - 4 ) Antennal scape without eye-cap; hind
M4 absent; ocelli present; tibial spurs 0- tibiae without spines 13
0-4; smaller moths (wingspan at most 13(12) Proboscis scaled; labial palp with lateral
ca. 16 mm., usually much smaller; fore- bristles on segment 2; valvae with
wings often iridescent 4 prominent, blunt spines often forming
4 (3) Forewing Sc forked; pterostigma ab- comb (pectinifer); ovipositor piercing,
sent; hindwing without produced jugal without medial sensory ridge
lobe; all regions Incurvarioidea (16)
Micropterigoidea (14) Proboscis scaled or unsealed; labial palp
Forewing Sc simple; pterostigma pre- without lateral bristles; $ valvae with-
sent; hindwing with distinct jugal lobe out prominent, blunt spines; ovipositor
(Fig. 4.1)); temperate S. America . . . . non-piercing, with medial sensory ridge
Heterobathmioidea (14) 14
5(2) Mediumsized to very large, robust 14(13) Proboscis unsealed; broad-winged
moths; proboscis and maxillary palps micro-moths, wingspan up to 30 mm.
strongly reduced Southern S. America, Australia, S.
Hepialoidea (part, 15) Africa Palaephatoidea (16)
Mediumsized to small moths, wingspan Proboscis scaled at base; narrow-
not exceeding 27 mm, often much winged micro-moths, wingspan less
smaller; proboscis and maxillary palps than 12 mm; Holarctic, tropical
often well developed 6 Tischerioidea (16)
6(5) Forewing Rs4 postapical 7 15(11) Proboscis scaled 16
Forewing Rs4 apical or preapical . . . 9 Proboscis unsealed or absent 18
7(6) Forewing Rs3 postapical; larger, more 16(15) Labial palps recurved, apical segment
broad-winged moths (wingspan at least may exceed vertex, usually tapering;
ca. 12 mm, usually much larger) chaetosemata absent; sternum II of 'tin-
Hepialoidea (part, 15) eid type' (see couplet 40); tympanal or-
Forewing Rs3 preapical; small, narrow- gans absent Gelechioidea {1-9)
winged moths 8 Labial palps porrect, beak-like or as-
8 (7) Maxillary palps well developed, 5-seg- cending; chaetosemata present or ab-
mented, with marked bends between sent; sternum II of 'tortricoid type';
segments 1/2 and 3/4; Australia tympanal organs present at base of ab-
Lophocoronoidea (15) domen, or absent 17
Maxillary palps very small, at most 3- 17(16) Labial palps ascending; fore wing with
segmented, porrect; New Zealand . . . . CuP present near termen; tympanal or-
Mnesarchaeoidea (15) gans absent . . . Choreutoidea (1 13)
9 (6) Meso-tibiae with paired apical spurs; Labial palps porrect, beak-like or as-
larger, often markedly broad-winged cending; CuP usually absent; tympanal
moths (wingspan 1327 mm) SE. Asia, organs present at base of abdomen . . .
temperate S. America Pyraloidea (1 14)
Neopseustoidea (15) 18(15) Antennae gradually or abruptly
Mesotibiae with unpaired apical spurs; clubbed, tip some times hooked . . . 19
small (wingspan at most ca. 16 mm usu- Antennae filiform or tapered, often cili-
ally markedly smaller) narrow-winged ate, pectinate or plumose 21
moths 10 19(18) Hind wing with frenulum; ocelli pre-
10(9) Ocelli present, M l not stalked with Rs sent, large; chaetosemata absent
veins Eriocranioidea (15) Sesioidea (part, 111)
Ocelli absent, M1 stalked with Rs . . . Hind wing usually without frenulum;
Acanthopteroctetoidea (15) ocelli absent; chaetosemata present . 20
11(1) Sternum II with paired anterior apo- 20(19) Antennae widely separated at base; with
demes; wing membranes devoid of subapical thickening; fore wing with all
microtrichia; female genital system with veins from discal cell arising separately
copulatory orifice (on VIII) separate Hesperioidea (1 16)
from more posterior ovipore Antennae approximated at base, with
Ditrysia 15 apical thickening; fore wing usually
Sternum II without anterior apodemes; with at least some peripheral veins
wing membranes usually with more or stalked Papilionoidea (1 16)
less extensive microtrichiation; female 21 (18) Tympanal organs present in metathorax
with single genital orifice 12 or ventrally in base of abdomen . . . 22
12(11) Antennal scape with eye-cap; hind tib- Tympanal organs absent 26
30 David J. Carter & Niels P. Kristensen

22 (21) Tympanal organs in metathorax, coun- without apical lobe; 2 A in fore wing
ter-tympanal cavities usually at base of not sinuous 35
abdomen Noctuoidea (119) 35 (34) Robust medium-sized and relatively
Tympanal organs in abdomen . . . . 23 narrow-winged moths with aposematic
23 (22) Chaetosemata absent 24 colours . . . . Thyridoidea (part, 1 13)
Chaetosemata present Facies different 36
Geometroidea (1 17) 36(35) Pronouncedly narrow-winged (forewing
24 (23) Fore wing broad, often falcate w/1 ratio 1/4 or less) and delicate moths
Drepanoidea (1 17) 37
Fore wing narrow, not falcate . . . . 25 Wings broader, forewing w/1 ratio 1/3 or
25 (24) Fore wing reddish brown, with silvery more 38
marks Cossoidea (part, 1 - 1 1 ) 37 (36) Small moths, wingspan < ca. 15 mm;
Fore wing grey . Tineoidea (part, 1 7) wings entire; body and legs not unusu-
26 (21) Wings more or less deeply cleft into two ally long . Schreckeristeinioidea (113)
or more plumes 27 Mostly larger moths, wingspan often
Wings not cleft 28 considerably exceeding 15 mm (always
27 (26) Hind wing divided into 3 plumes . . . . if wings entire); wings often cleft; body
Pterophoroidea (part, 1 13) and legs unusually long
Hind wing divided into 6 - 7 plumes . . Pterophoroidea (part) (1 13)
Alucitoidea (part, 1 - 1 3 ) 38 (36) Abdominal tergum I with large post-
28 (26) CuP absent as a tubular vein in fore spiracular expansion laterad from 'ter-
wing 29 gal rim' (Fig. 13.3 H); medium-sized
CuP well developed, at least near mar- moths, often with cryptic 'netted' wing
gin, as a tubular vein in fore wing . 40 pattern . . . . Thyridoidea (part, 1 13)
29 (28) Abdominal segment VII with pocket- Abdominal tergum I without such ex-
like concavities associated with spira- pansions 39
cles; medium-sized moths with silvery 39 (38) Strong spines present on all tarsomeres;
markings on forewing (Figs. 15.2 A, B); facies as Fig. 13.41; Madagascar only .
sub-tropical W. Palaearctic Whalleyanoidea (113)
Axioidea (1 15) Strong spines present absent from api-
Abdominal segment VII without such cal foreleg tarsomere; facies variable;
concavities 30 Madagascar, Asia Calliduloidea (1 15)
30 (29) Broad-winged micro-moths (wingspan 40 (28) Sternum II of 'tineid' type: anterolateral
< 20 mm) with prominent ocelli; head corners not produced, apodemes long
vestiture or very narrow scales with and slender; usually small delicate spe-
deeply scalloped apices; Australasian . . cies, usually without raised scales on
Simaethistoidea (1 13) fore wing 41
Not as above 31 Sternum II of 'tortricoid' type: antero-
31 (30) Head vestiture of piliform scales, if lateral corners usually distinctly pro-
mixed with lamellar scales (Mimallo- duced, apodemes with broad bases, usu-
noidea, part) frenulum small or absent; ally short and not appearing as continu-
robust, medium-sized to very large ations of sternal ridges, ('venulae'); usu-
moths . 'Bombycoid assemblage' (1 18) ally robust, small to very large species;
Head vestiture of lamellar scales; frenu- sometimes with raised scales on fore
lum well developed; small to medium- wing 42
sized moths 32 41 (40) Abdominal segment VIII of $ with
32 (31) Hind wing CuA with pecten; fore wings prominent lobes; fore wing with Rs4
often with raised scales often to termen and M usually absent
Copromorphoidea (part, 1 13) from discal cell Yponomeutoidea ( 1 - 8 )
Hind wing CuA without pecten; fore Abdominal segment VIII of $ without
wing without raised scales 33 lobes; fore wing with Rs4 often to costa;
33 (32) Abdominal terga with anterior band of M usually present in discal cell
spines; sternum II with V-shaped sclero- Tineoidea (part, 17)
tization . . . . Alucitoidea (part, 1 - 1 3 ) 42 (40) Fore wing with raised scales on surface
Abdominal terga without spines; ster- or along dorsum 43
num II without V-shaped sclerotization Fore wing without raised scales . . . 44
34 43 (42) Fore wing with raised scales along dor-
34 (33) Labial palps beak-like; S hind coxa with sum; wing narrow, almost parallel-
apical process and tibia with hair-pen- sided Epermenioidea (13)
cil; fore wing with 2 A sinuous Raised scales on fore wing not restricted
Hyblaeoidea (1 13) to dorsum; wing relatively narrow, rect-
Labial palps not beak-like; hind tibia angular Copromorphoidea (part, 113)
Classification and Keys to Higher Taxa 31

44 (42) Chaetosemata present; ocelli absent; to larvae (Carter, 1984; Carter in Hollo way,
fore wing without chorda and M stem Bradley & Carter, 1987) to include as many fami-
in cell Immoidea ( 1 - 1 3 ) lies as possible. It is based largely on final instar
Chaetosemata present or absent; ocelli larvae alhough earlier instars are taken into ac-
present or absent; fore wing often with count where practically possible. It should be
both chorda and M in cell 45 borne in mind that larval instars may differ very
45 (44) Lower part of frons with ascending considerably. Many species with very hairy final
scales; ocelli and chaetosemata present; instar larvae have first instar larvae that bear pri-
ovipositor lobes leaf-like mary setae only.
Tortricoidea (1 12) The larvae of many Lepidoptera families are
Lower part of frons without ascending very poorly known and key characters for some
scales; ocelli and chaetosemata rarely are based on only a single species. Some families
both present; ovipositor lobes not leaf- have been omitted due to lack of sufficient infor-
like 46 mation to provide key characters for immature
46 (45) M tubular and forked in cell 47 stages.
M absent in cell, or M not present in In the preparation of this key, ease of use has
cell as a complete, forked tubular vein been considered of high importance. Stehr (1987)
48 provides a much more detailed and precise key
47 (46) Chaetosemata absent; ocelli rarely pre- to families of Nearctic Lepidoptera but omits
sent Cossoidea (1-11) many families that only occur in other parts of
Chaetosemata and ocelli present . . . . the World. In order to cover all families in the
Zygaenoidea (part, 1 10) present key without making it too long and diffi-
48 (46) Patagia large, extending ventrad be- cult to use, it has been simplified by keeping to
yond anteroventral corres of pronotum fairly short couplets and omitting reference to
Sesioidea (part, 1 11) exceptions within families, excepting major sub-
Patagia relatively smaller, not extending families and other important groups. The key
beyond anteroventral corres of prono- should therefore be used with caution and identi-
tum 49 fications checked against good family descrip-
49(48) Ocelli present tions.
Zygaenoidea (part, 1 - 1 0 ) Traditionally, the most important larval char-
Ocelli present 50 acteristics used for identification are relative se-
50 (49) Forewing CuP distinct throughout . . . tal positions. Unfortunately, study of larval
Zygaenoidea (part, 1 10) chaetotaxy has been hampered by a proliferation
Forewing CuP distinct only at margin of different systems of setal nomenclature. The
and (sometimes) at base. Some smaller system adopted here for body setae is that of
groups of apoditrysian moths with Hinton (1946), while nomenclature of cranial se-
poorly understood affinities will key out tae is based on that of Heinrich (1916) modified
here; they are currently placed in the su- by Gerasimov (1935) and Stehr (1987).
perfamilies . . . . Galacticoidea (1 13),
Not only are setae sometimes very difficult to
Urodoidea (1 13) and Zygaenoidea
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
Key to families: Larvae sides of the body.
Larval characters are not illustrated within the
Identification of lepidopterous larvae often key but the figures of general larval features (Fig.
proves to be difficult, largely due to the fact that 3.1,2) should act as a simple glossary of terms
the larvae of relatively few species have been de- used.
scribed in detail and comprehensive reference
collections of immature stages are scarce. This is 1 Case-bearing 2
because Lepidoptera classification is tradition- Not case-bearing 6
ally based on studies of imagines. However, 2(1) Pro thorax with L pinaculum fused
modern studies are increasingly taking into ac- with thoracic shield 3
count the immatures stages, which provide many Prothorax with L pinaculum separate
vital clues to systematic and phylogenetic rela- or not developed 4
tionships. Identification of immatures is also ex- 3 (2) Meso and metathorax with SV unise-
tremely important to the economic entomologist tose. Small larvae in flat, elongate-
as the larva stage is almost invariably the one oval cases of plant fragments on the
causing damage to crops and other commodities. ground Adelidae (16)
The present key has evolved by the refinement Meso and metathorax with SV bise-
and expansion of previous more restricted keys tose Psychidae (17)
32 David J. Carter & Niels P. Kristensen


Anterodorsal view of head-

setal nomenclature and general features


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

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


anal comb

chalaza scoli thoracic leg

uniserial jniserial uniserial multiserial

lili fff H
uniordinal biordinal triordinal uniordinal

!# l?
transverse circle mesopenellipse lateropenellipse mesoseries pseudocircle
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 David J. Carter & Niels P. Kristensen

- Antennae much shorter than head. 18 (17) Larvae slug-like, often with scoli.
Strongly tapered prolegs not present Head strongly retracted into thorax
on abdominal segments 1 - 8 7 Limacodidae (1 10)
7 (6) Head without adfrontal sutures or ec- Larvae subcylindrical, without sec-
dysial lines. Thoracic legs and prolegs ondary setae. Head not retracted into
absent. Body setae minute and diffi- thorax 19
cult to detect. Larvae feeding within 19(18) Small larvae, less than 3 cm long,
seeds of kauri pine (Agathis) without patches of spicules on dorsal
Agalhiphagidae (14) surface 20
Head with adfrontal sutures and ecdy- - Large larvae, more than 5 cm long
sial lines 8 with prominent dorsal patches of spic-
8 (7) Thoracic legs absent or very weakly ules Castniidae (part, 111)
developed. Prolegs with or without 20 (19) Meso and metathorax and abdominal
crochets 9 segments 18 each with a pair of dor-
- Thoracic legs present 17 sal calli. Head with spinneret not de-
9 (8) Numerous secondary setae present . . veloped . . . Heterobathmiidae (14)
Cecidosidae ( 1 - 6 ) - Dorsal paired calli absent from thorax
Secondary setae absent 10 and abdomen. Head with spinneret de-
10 (9) Head with adfrontal suture passing veloped 21
behind antenna . Eriocraniidae (15) 21 (20) Abdominal segments 1 - 8 with LI di-
Head with adfrontal suture in front of rectly caudad to spiracle
Acanthopteroctetidae ( 1 - 5 )
antenna 11
- Abdominal segments 1 - 8 with LI not
11 (10) Head with frons triangular, upper
directly caudad to spiracle 22
edge not or hardly included in forma-
22 (21) Pro thorax with two L setae and with
tion of hind edge of head capsule . 12
spiracle on sclerotised plate
Head with frons parallel-sided or
Glyphipterigidae (18)
shaped as an inverted trapezium . 14
- Prothorax with three L setae. Spiracle
12(11) Anal shield with a dorsal, hooked
not on sclerotised plate 23
prong Douglasiidae
23 (22) A few rudimentary crochets present
(Tinagma, overwintering form) (17)
on abdominal prolegs; abdominal se-
Anal shield without a dorsal pro-
tae short . . . Gelechiidae (part, 1 - 9 )
jection 13
- Abdominal crochets absent; abdomi-
13 (12) Dorsal plates or calli present on tho- nal setae long . . Douglasiidae (17)
racic and/or abdominal segments . . . 24(17) Crochet-bearing prolegs present on
Heliozelidae (16) abdominal segments 2 - 7 . Slug-like
Dorsal plates or calli absent, larva larvae with deciduous, jelly-like tuber-
usually pink or red when preserved . . cles Dalceridae (1 10)
Prodoxidae (16) - Abdominal segments 2 and 7 without
14 (11) Prolegs with crochets, at least on anal crochet-bearing prolegs. Body without
prolegs Tischeriidae (16) jelly-like tubercles 25
Prolegs without crochets 15 25 (24) Crochets of abdominal prolegs ar-
15(14) Head with frons rectangular. Weakly ranged in medial longitudinal bands
developed, stump-like, unsegmented or curves (mesoseries), sometimes with
legs present on meso- and metathorax rudimentary lateroseries, forming a
Nepticulidae (16) pseudocircle or broken pseudocircle
Head with frons shaped as an inverted 82
trapezium; thoracic legs absent . . 16 - Crochets not in a mesoseries . . . . 26
16(15) Head with frontal bridge connecting 26(25) Abdominal segment 10 with crochets
hind margins of adfrontal ridges . . . in a complete or almost complete
Gracillariidae (part, 1 - 7 ) ellipse, with large biordinal crochets
Head without frontal bridge connect- anteriorly and smaller uniordinal or
ing hind margins of adfrontal ridges biordinal crochets posteriorly
(Mandible sometimes with large dorsal Mimallonidae (1 18)
conical projection) Long, very slender - Abdominal segment 10 with crochets
larvae Opostegidae (1 6) not as above 27
17 (8) Larvae with thoracic legs but without 27 (26) Body with secondary setae, sometimes
crochet-bearing prolegs or with only a only in SV region 28
few crochets (less than 5 on each pro- - Body without secondary setae . . . 33
leg) 18 28 (27) Head large and rounded with many
Larvae with thoracic legs and prolegs, secondary setae, sometimes on chala-
the latter usually with crochets . . 24 zae. Prothorax narrowed to form dis-
Classification and Keys to Higher Taxa 35

tinct neck. Anal comb sometimes pre- 39 (35) Prothorax with L group trisetose . . 40
sent Hesperiidae (1 16) Prothorax with L group bisetose . . 70
Head not large and rounded; cranial 40 (39) Prothorax with L pinaculum fused
setae not on chalazae. Prothorax not with thoracic shield 41
narrowed to form a "neck" 29 - Prothorax with L pinaculum separate
29 (28) Prolegs with crochets in a mesopenel- or not developed 43
lipse that may be mistaken for a meso- 41 (40) Prolegs with crochets in a uniserial cir-
series Pterophoridae {1-13) cle, small larvae 42

Crochets otherwise arranged . . . . 30 Prolegs with crochets in a biserial or
30 (29) Secondary setae on Verrucae; setae multiserial circle. Larvae medium to
with minute lateral branches large Hepialidae (15)
Lecithoceridae {1-9) 42(41) Abdominal segments 1 - 6 with LI and
Secondary setae not on Verrucae; setae L2 on a common pinaculum
without lateral branches 31 Galacticidae (1-13)
31 (30) Abdominal prolegs with crochets in - Abdominal segments with LI and L2
uniordinal circle. Tarsi of thoracic legs remote, not on a common pinaculum
with flattened setae Tineidae (part, 17)
. . ElachistidaeAgonoxeninae (19) 43 (40) Abdominal prolegs with crochets in
Abdominal prolegs with crochets not uniserial transverse bands 44
uniordinal. Tarsi of thoracic legs with- Abdominal prolegs with crochets in a
out flattened setae 32 circle or penellipse 48
32(31) Abdominal prolegs with crochets 44(43) Abdominal prolegs usually with one
biordinal. Secondary setae not con- transverse band of crochets, occasion-
fined to SV group. Thoracic segments ally with two bands. Anal shield with
with SV trisetose six setae. Small larvae in fruit and
. . . XyloryctidaeScythridinae (19) buds
Abdominal prolegs with crochets tri- . . . ProdoxidaeLamproniinae (16)
ordinal to multiordinal. Secondary se- (early instar, leaf-mining larvae of In-
tae confined to SV group or to anal curvariidae may also key out here)
prolegs only. Meso- and metathorax - Abdominal prolegs each with two
with SV unisetose bands of crochets. Anal shield with
. . . XylorytidaeXyloryctinae (19) eight setae 45
33 (27) Abdominal seg 6 without prolegs . . . 45 (44) Abdominal segments with LI and L2
Gracillariidae (part, 17) always remote from each other . . . .
Abdominal seg 6 with prolegs . . . 34 Bucculatricidae (17)
34 (33) Body globular. Head reduced to small Abdominal segments with LI and L2
triangular capsule. Stemmata grouped close together 46
on a raised tubercle. Larva parasitic 46 (45) Anal prolegs with crochets divided
on Hemiptera . . Epipyropidae (1 - 1 0 ) into two groups
Body not globular. Head normal. Gelechiidae (part, 19)
Stemmata not on tubercle 35 - Anal crochets in undivided bands . 47
35 (34) Abdominal spiracles on pinacula or 47 (46) Dorsal surface of body with raised
small sclerotized conical protuber- patches of spicules. Large larvae (c
ances 36 7 cm) Castniidae (part, 1 - 1 1 )
Abdominal spiracles not on pinacula - Dorsal surface of body without raised
or protuberances 39 patches of spicules. Small to medium
36 (35) Spiracles on raised or conical protu- sized larvae Sesiidae (111)
berances 37 48 (43) Small larvae mining in grasses
Spiracles not raised 38 . . . ElachistidaeElachistinae (19)
37 (36) Prothorax with L group trisetose; - Not mining in grasses 49
some dorsal setae spatulate or bifur- 49 (48) Abdominal segment 3 with LI and L2
cate . . . . Schreckensteiniidae (1 13) remote, or in very small larvae L2
Prothorax with L group bisetose; dor- missing 50
sal setae normally developed Abdominal segment 3 with LI and L2
Epermeniidae (1 13) close, often on same pinaculum . . 54
38 (36) Pinacula surrounding spiracles ex- 50 (49) Crochets in a uniserial circle or penel-
tended dorsocranially. Prolegs with lipse 51
few crochets or without crochets . . . - Crochets in a multiserial circle or a
Ypsolophidae Ochsenheimeriinae (1 8) penellipse surrounded by a circle of
Pinacula surounding spiracles not ex- small crochets 53
tended dorsocranially. Prolegs with 51 (50) Prolegs long, almost as long as tho-
crochets in a circle racic legs Plutellidae &
Acrolepiidae (18) Ypsolophidae Ypsolophinae (18)
36 David J. Carter & Niels P. Kristensen

Prolegs considerably shorter than tho- 64 (54) Head with LI further from A3 than
racic legs 52 A3 is from A2
52 (51) Abdominal segments with SD2 absent Gelechiidae!Oecophoridae (part, 19)
. . . . YponomeutidaeArgyresthiinae Head with LI nearer to A3 than A3 is
(part, 1 - 8 ) to A2 or equidistant 65
Abdominal segments with SD2 pre- 65 (64) Metathorax with distance between
sent Tineidae (part, 17) coxae twice as great as their breadth
53 (50) Abdominal prolegs with crochets in a Cosmopterigidae (19)
multiserial circle Metathorax with distance between
Yponomeutidae- Yponomeutinae ( 1 - 8 ) coxae less than twice their breadth 66
Abdominal prolegs with crochets in a 66 (65) Abdominal segments with D1 and D2
penellipse surrounded by a circle of close together . . . Heliodinidae { 1 - 8 )
small crochets Plutellidae (part, 18) Abdominal segments with D1 and D2
54 (49) Abdominal segment 9 with setae D2 remote 67
closer together than setae D1 of abdo- 67 (66) Head with frons reaching only half
minai segment 8, often on a common way to vertical triangle 68
pinaculum 55 - Head with frons reaching at least 2/3
Abdominal segment 9 with setae D2 at distance to vertical triangle 69
least same distance apart as setae D1 68 (67) Abdominal segment 9 with LI and L2
of abdominal segment 8 64 on separate pinacula
55 (54) Abdominal prolegs slender and pencil- Cossidae Cossinae (1 11)
like, almost as long as thoracic legs. . _ Abdominal segment 9 with LI and L2
56 on a common pinaculum
Abdominal prolegs short, distinctly . . Elachistidae-Stenomatinae (1-9)
shorter than thoracic legs 57 59 Prolegs with crochets usually biserial.
56 (55) Abdominal prolegs each with a sclero- Anal segment, excluding anal shield,
tised collar . . Gelechiidae (part, 19) with more than nine setae on each
Abdominal prolegs without sclerotised side . . . . Oecophoridae {part, 1 - 9 )
collars Choreutidae {1-13) _ Crochets uniserial. Anal segment, ex-
57 (55) Prothorax with spiracle dorsad of L cluding anal shield, with not more
pinaculum than nine setae on each side . Ypono-
. ColeophoridaeBlastobasinae (19) meutidaeArgyresthiinae (part, 1 - 8 )
Prothorax with spiracle posterior to L Abdominal segments with LI and L2
70 (39)
pinaculum 58 remote (LI distinctly posterior to spi-
58 (57) Metathoracic legs swollen and club- racle) . . . Tineidae-Scardiinae (17)
shaped Chimabachidae (19) Abdominal segments with LI and L2
Metathoracic legs not swollen . . . 59 close together, often on a common pi-
59 (58) Head rugulose. Each abdominal seg- naculum 71
ment with all four D setae on a large
71 (70) Crochets of abdominal prolegs in two
sclerotised plate
transverse bands 72
. . . . CossidaeMetarbelinae (1 11)
Head smooth, D setae not on large Crochetsof abdominal prolegs in a cir-
plates 60 cle or penellipse 73
60 (59) Head wedge-shaped. Large larvae. . . 72 (71) Head with frons reaching 3 or more
CossidaeZeuzerinae (1 11) to vertical triangle Coleo-
Head not wedge-shaped 61 phoridae- Momphinae (part, 19)
61 (60) Prolegs of abdominal segments 36 ~~ Head with frons reaching less than Vi
with crochets in a multiserial penel- way to vertical triangle
lipse Palaephatidae (16) Brachodidae (1 11)
Prolegs with crochets otherwise ar- 73 (71) Meso- and metathorax with SV group
ranged 62 bisetose 74
62(61) Prolegs each with a surrounding collar ~ Meso- and metathorax with SV group
and/or with a sclerotised, central foot unisetose 75
patch Gelechiidae (part, 19) 74(73) Meso- and metathorax with L2 absent
Prolegs without sclerotised collar or Thyrididae (1 13)
foot patches 63 - Meso- and metathorax with L2 pre-
63 (62) Abdominal segment 8 with SD usu- sent Pyraloidea (part, 1 14)
ally distinctly anterior to spiracle; anal 75 (73) SV group above prolegs with four
comb sometimes present macroscopic setae 76
Tortricidae (1 12) SV group, on or above prolegs, with
Abdominal segment 8 with SD ante- three macroscopic setae 78
rodorsad of spiracle. Anal comb ab- 76 (75) Abdominal segments 1 6 with LI and
sent . . . . Oecophoridae (part, 19) L2 on a common pinaculum . . . . 77
Classification and Keys to Higher Taxa 37

Abdominal segments 16 with LI and Abdominal prolegs with crochets in

L2 more widely separated, not on a simple mesoseries 88
common pinaculum 86 (85) Anal prolegs reduced to small lobes
ColeophoridaeMomphinae without crochets. Anal shield extended
(part, 1 - 9 ) to a single point or process. Secondary
77 (76) Anal prolegs with 2 rows of crochets; setae in SV region only
abdominal segment 9 with D1 absent Drepanidae (part, 1 17)
or minute. . Copromorphidae {113) Anal prolegs normally developed.
Anal prolegs with 1 row of crochets; Anal shield not extended to a single
abdominal segment 9 with D1 nor- point or process 87
mally developed. Carposinidae (113) 87 (86) Head with dorsal protuberances. Anal
78 (75) Prothorax with SD absent (ie protho- shield bifurcate extending to form two
racic shield with only 10 setae); lateral long processes. Anal comb present . .
leaf-like gland present on abdominal Hedylidae( 1 - 1 6 )
segment 1 . . . . Callidulidae {1-15) - Head without protuberances. Anal
Pro thorax with SD present (pro tho- shield not modifidied. Anal comb ab-
racic shield with 12 setae); no lateral sent. Dorsal eversible gland present on
gland on abdominal segment 1 . . 79 prothorax . Papilionidae (part, 1 - 1 6 )
79 (78) SV setae situated low-down on abdo- 88 (85) Larvae without numerous secondary
minai prolegs. Metathorax with SD setae, except in some cases in the SV
approximately equal in length to SD2. region and on the prolegs 89
Abdominal prolegs with crochets tri- Larvae with many secondary setae (in
ordinal Hyblaeidae (1 13) some cases, secondary setae are micro-
SV setae situated at base of abdominal scopic) 98
prolegs. Metathorax with SD con- 89 (88) Prothorax with L group trisetose; pro-
siderably longer than SD2 80 legs long and slender 90
80 (79) Crochets of prolegs biordinal or trior- - Prothorax with L group bisetose . 91
dinal (exceptionally uniordinal) . . . . 90 (89) Abdominal prolegs medially con-
Pyraloidea (part, 1 14) stricted Urodidae (1 13)
Crochets of prolegs uniordinal. . . 81 - Abdominal prolegs not medially con-
81 (80) Abdominal prolegs long and slender; stricted Immidae (1 13)
crochets in a mesal penellipse 91 (89) Abdominal prolegs with secondary se-
Tineodidae (1 13) tae 92
Abdominal prolegs short; crochets in Abdominal prolegs without secondary
a circle Alucitidae (1 - 1 3 ) setae 94
82 (25) Abdominal prolegs long, often as long 92 (91) Setae on raised pinacula. Head with
as thoracic legs; with crochets in epicranial index more than 0.8; stipital
strongly curved mesoseries, closely re- lobe absent
sembling a penellipse NoctuidaeDilobinae (1 19)
Uraniidae (1 17) Setae not on raised pinacula. Head
Abdominal prolegs not long, or if so, with epicranial index less than 0.6;
crochets no arranged in a strongly stipital lobe present 93
curved mesoseries resembling a penel- 93 (92) Abdominal segment 1 with only 1 MD
lipse 83 seta; thorax swollen
83 (82) Thoracic legs very short, with adhesive Doidae-Doa (1-19)
pads on tarsi; body with broad lateral - Abdominal segment 1 with 2 MD se-
flanges. Small head usually retracted tae or thorax not swollen
into thorax. . . Cyclotornidae {1 10) Notodontidae (part, 1 19)
Thoracic legs normally developed . 8 4 94 (91) Thorax and abdomen with crown-like
84 (83) Well developed prolegs present on ab- circlets of spines scattered over the
dominal segments 6 and 10 only; dorsal and lateral surfaces
sometimes with small or rudimentary Heterogynidae (1 10)
prolegs on abdominal segments 4 and - Thorax and abdomen without circlets
5; anal prolegs strongly developed . . of spines 95
Geometridae (1-17) 95(94) Abdominal segments 1 - 8 with two
Well developed prolegs present, at extra L setae, one caudad of spiracle,
least on abdominal segments 5 and 6 one distinctly ventrocaudad of spira-
85 cle . DrepanidaeThyatirinae ( 1 17)
85 (84) Abdominal prolegs with an additional Abdominal segments without extra L
lateroseries of crochets, forming a setae caudad of spiracle 96
pseudocircle or broken pseudocircle . 96 (95) Meso- and metathorax with SV group
86 bisetose 97
38 David J. Carter & Niels P. Kristensen

- Meso- and metathorax with SV group long and thin, with enlarged plantae
unisetose Noctuidae (part, 107
(including Aganainae; 1 - 1 9 ) Spiracles large and usually elliptical.
97 (96) Body covered with minute spinules. Prolegs short 108
D2 and LI arising from elongate tu- 107 (106) All setae on distinct Verrucae; protho-
bercles or conical projections espe- racic L group with more than 3 setae
cially on meso- and metathorax. NycteolidaeNolinae (1 19)
Crochets homoideous Only some setae on Verrucae, others
NycteolidaeEarias (1 19) scattered; prothorax with L group tri-
- D1 and D2 not arising from elongate setose Pterophoridae (1 13)
tubercles. Prolegs with ends of plantae 108 (106) Prolegs with crochets biserial and with
drawn out. Crochets usually heteroi- 3 or less SV setae
deous Arctiidae (part, 1 - 1 9 ) Oenosandridae (1 19)
98 (88) Additional rudimentary prolegs, with- Prolegs with crochets uniserial and
out crochets, on abdominal segments with more than 3 SV setae . . . . 109
2 and 7 99 109 (108) Some setae arranged on Verrucae or
- Abdominal segments 2 and 7 without flat, verruca-like plates. Head with
prolegs 100 stipi tal lobes absent 110
99 (98) Prothorax and abdominal segments Setae not grouped on Verrucae or sim-
18 with peg-like, glandular protu- ilar plates. Head with stipital lobes
berance adjacent to spiracle present . . Notodontidae {part, 119)
Megalopygidae (1 10) 110(109) Abdominal segment 8 with spiracle at
- Prothorax and abdominal segments least 1.5 the height of that in abdo-
without peg-like protuberances adja- minal segment 7. Head with mandibles
cent to spiracle lacking secondary setae
Somabrachyidae (1 10) Noctuidae (part, 1 - 1 9 )
100(98) Abdominal segments 6 and 7 with Abdominal segment 8 with spiracle
mid-dorsal, eversible glands, or, if ab- less than 1.5 (usually about 1.25 x)
dominal segment 6 is without a gland, the height of that on abdominal seg-
then abdominal segment 8 with a ment 7. Head with mandibles bearing
large, mid-dorsal hair pencil. Body secondary setae. Larvae usually living
usually with series of dorsal hair tufts communally
or pencils . . . . Lymantriidae ( 1 - 1 9 ) . . . Notodontidae Thaumetopoeinae
- Abdominal segments 6 and 7 without (1-19)
mid-dorsal glands 101 111 (104) Secondary setae very variable in
101 (100) Prolegs with crochets heteroideous, or length (some at least 5 length of oth-
ends of plantae drawn out beyond ex- ers), not arising from raised Verrucae
tent of crochets 102 or scoli 112
- Prolegs with crochets homoideous; Secondary setae fairly equal in length
ends of plantae not drawn out beyond or larvae with distinct warts or spines
the extent of the crochets 103 114
102(101) Mesothorax with only one verruca 112(111) Pointed anal lobe present between
above the spiracle anal prolegs; body sometimes with lat-
. . . . ArctiidaeCtenuchinae { 119) eral or subventral lappets
- Mesothorax with more than one ver- Lasiocampidae (1 18)
ruca above the spiracle Anal lobe absent; lappets not present
Arctiidae (part, 1 19) 113
103 (101) Abdominal prolegs with a row of soft 113 (112) Mid-dorsal tapered hair tufts or pen-
basal plates below the crochets; head cils present
retracted into thorax . . BombycidaeApatelodinae (1 18)
Zygaenidae (110) Mid-dorsal hair tufts or pencils absent
- Abdominal prolegs without soft basal Lemoniiidae (1 18)
plates below crochets 104 114(111) Secondary setae minute, many flat-
104 (103) Prolegs with crochets uniordinal 105 tened and scale-like. Dorsal paired fil-
Prolegs with crochets biordinal or tri- aments on meso and metathorax and
ordinal Ill abdominal segment 9 and mid-dorsal
105 (104) Anal shield produced to form a pair of filament on abdominal segment 8 are
anal projections. Head with stemma 3 lost in the final instar
enlarged Brahmaeidae (118)
. . . . NymphalidaeSatyrinae (1 16) Secondary setae if minute not scale-
- Anal shield not forked 106 like. Body without deciduous fila-
106(105) Spiracles small and circular. Prolegs ments 115
Classification and Keys to Higher Taxa 39

115 (114) Abdominal segment 8 with a mid-dor- Prothorax without a dorsal eversible
sal horn, tubercle or spine . . . . 116 gland 125
- Abdominal segment 8 without a mid- 125 (124) Head and body without protuber-
dorsal horn or spine 122 ances 126
116(115) Body with spines or large protuber- Head and/or body with spines, scoli or
ances or expansions on thorax and other conspicuous protuberances 129
elsewhere 117 126(125) Primary setae reduced to small stout
- Body without spines or projections on cones; abdominal spiracles surrounded
thorax and rarely elsewhere, except on by eyespots
abdominal segment 8 120 . . . . Carthaeidae (final instar, 1 18)
117 (116) Head angular or covered with spines. Primary setae not reduced to cones;
Abdominal prolegs with crochets usu- circumspiracular eyespots absent 127
ally triordinal 127 (126) Some setae grouped on Verrucae; setae
Nymphalidae (part, 1 16) sometimes plumose
- Head rounded and smooth. Abdomi- . . . . AntheiidaelEupterotidae (1 18)
nal prolegs with crochets biordinal . . Verrucae absent; setae short, not plu-
118 mose 128
118(117) Dorsal and subdorsal primary setae 128 (127) Head and body densely covered with
clubbed or reduced to stout cones; short hairs, often on small warts; abd
eyespots surround spiracles seg 8 without mid-dorsal patch; anal
Carthaeidae (penultimate instar, 1 18) comb sometimes present
- Primary setae not so modified or lat- Pieridae (1-16)
eral circumspiracular eyespots absent Body with minute secondary setae, ap-
119 pearing completely smooth. Abdomi-
119(118) Metathorax expended laterally, with nal segment 8 with a sclerotised dorsal
dorsal eyespots; Horn on abdominal patch . . . . Sphingidae (part, 1 - 1 8 )
segment 8 with bifid tip 129(125) Head with protuberances
. . . SaturniidaeOxyteninae (1 18) Nymphalidae (part, 1 16)
- Metathorax not expanded laterally or Head without protuberances
without dorsal eyespots; Uorn/tuber- Saturniidae (part, 118)
cle on abdominal segment 8 without
bifid tip . . . Saturniidae (part, 1 18)
120 (116) Abdominal segments each divided into Acknowledgments
six to eight annuii. Prolegs not widely NPK thanks Dr. N. M. Andersen and Mr. O.
separated . . Sphingidae (part, 1 - 1 8 ) Karsholt for critical perusal of the introductory
Abdominal segments indistinctly di- section. The abandonment of sub- and infraor-
vided into two or three annuii. Prolegs ders in Lepidoptera systematics was laid before
often widely separated 121 of a number of leading Lepidoptera systematists,
121 (120) Abdominal prolegs very widely sepa- several, but not all of whom sympathized with
rated and splayed. Thoracic segments the step; all expressions of views were informa-
often enlarged tive and are appreciated.
Bombycidae (part, 1 18)
- Abdominal prolegs not widely sepa-
rated or splayed. Thoracic segments References
taper towards head
Ax, P. (1984): Das Phylogenetische System. Stuttgart,
Endromidae (1 18)
New York G. Fischer [English edition 1987 Thephy-
122 (115) Abdominal prolegs with crochets in a logentic system. Chichester, New York J. Wiley]
mesoseries divided by a central gap, or Carter, D . J . (1984): Pest Lepidoptera of Europe with
reduced centrally, with a pad-like, special references to the British Isles. - Series Ento-
central protuberance. Head usually re- mologica 31. W. Junk, The Hague,
tracted into thorax de Queiroz, K. & Gauthier, J. 1992. Phylogenetic tax-
Lycaenidae (1 16) onomy. - A. Rev. Ecol. Syst. 23: 4 4 9 - 4 8 0 .
123 (122) Metathorax dorsally expanded into Forey, P. L. (1992): Formal classification. Pp. 1 6 0 - 1 6 9
in Forey P. L., Humphries, C. J., Kitching, I. J.,
forward-pointing conical peak; anal
Scotland, R. W., Siebert, D. J. & Williams, D. M.
shield produced into a cone-like back-
Cladistics. A practical Course on Systematics. The
ward-pointing structure Systematics Association Publication N o 10. Ox-
. Saturniidae Cercophaninae (1 18) ford, Clarendon Press.
- Metathorax and anal shield not modi- Gerasimov, A. M. (1935): Zur Frage der Homodyna-
fied in this way 124 mie der Borsten von Schmetterlingsraupen. Zool.
124(123) Prothorax with a dorsal eversible Anz. 112: 1 1 7 - 1 9 4 .
gland situated in a groove or pit . . . Griffiths, G. C. D. (1974): On the foundations of bio-
Papilionidae (part, 1 16) logical systematics. Acta Biotheor. 23: 8 5 - 1 3 1 .
40 David J. Carter & Niels P. Kristensen

Heinrich, C. (1916): On the taxonomic value of some Sattler, . (1991): A review of wing reduction in Lepi-
larval characters in the Lepidoptera. - Proc. ent. doptera. - Bulletin of the British Museum (Natural
Soc. Wash. 18: 154-164. History) (Entomology) 60: 243-288.
Heppner, J. B. (1991): Brachyptery and aptery in Lepi- Stehr, F. (1987): Order Lepidoptera. Pp. 288-340 in
doptera. Tropical Lepidoptera 2: 1140. Stehr, F. (ed.) Immature Insects. Kendall/Hunt, Du-
Hinton, H. E. (1946): On the homology and nomencla- buque.
ture of the setae of lepidopterous larvae, with some Wiley, E. O. (1981): Phylogenetics. The Theory and
Practice of Phylogenetic Systematics. New York
notes on the phylogeny of the Lepidoptera. -
etc., J. Wiley.
Trans. R. ent. Soc. Lond. 97: 1 - 3 7 .
Willmann, R. (1987): Phylogenetic systematics, classi-
Holloway, J. D Bradley, J. D. & Carter, D. J. (1987): fication and the plesion concept. - Verh. naturwiss.
Lepidoptera. CIE Guides to Insects of Importance Ver. Hamburg (NF) 29: 221-233.
to Man 1. CAB International Institute of Ento- (1989): Palaeontology and the systematization of
mology & British Museum Natural History, natural taxa. - Abh. naturwiss. Ver. Hamburg (NF)
London. 28: 267-291.
4. The Non-Glossatan Moths
Niels P. Kristensen

Three lepidopteran clades are primitively devoid MICROPTERIGOIDEA

of the coilable proboscis and other apomorphic Micropterigidae. Probable autapomorphies: An-
traits which characterize the huge monophylum tennae with 'ascoid' sensilla (branched basiconic
Glossata. These clades are the Micropterigoidea sensilla, Fig. 4.1 E; paralleled in Nepticuloidea-
Micropterigidae, Agathiphagoidea - Agathipha- Opostegidae). Labrum extensively desclerotized.
gidae and Heterobathmoidea-Heterobathmii- Strong mandibles asymmetrical, incisor cusps on
dae; their phylogeny is discussed in 12 and left mandible only present apically. Labial palps
their formal ranking in 13. Inferences about greatly shortened. Spurs 0-0-4 (absence of meso-
the ground plan of lepidopteran structure and tibial spurs paralleled in the Heterobathmi-
function are by necessity based largely on condi- oidea). Prothorax unusually small, with anterior
tions in the non-glossatan clades, and their mor- notopleural articulation and with unsclerotized
phology is therefore treated in some detail in vol- areas in notum. Spiracle VIII nonfunctional.
ume 2. Male with sternal sclerotization on VIII strongly
reduced, represented by small paired plates or
The adult moths in the non-glossatan grade
altogether absent. Phallus with close-set radial
share a suite of plesiomorphic traits. Heads with
folds around gonopore. Female postabdominal
'rough' vestiture of hair-scales. Dorsal tentorial
segments without apophyses. 6 malpighian tu-
arms variably developed, but never large. La-
bules grouped into two bundles (of 3), each dis-
brum large, movable, with frontal retractors.
charging through a common duct (paralleled in
Mandibles have well developed articulations the Neolepidoptera). Brain usually with distinct
with the head capsule and strong muscles. Maxil- 'deutocerebral loop' (paralleled in Glossata-
lae generalized, with long, folded 5-segmented Acanthopteroctetoidea and Lophocoronoidea).
palps, distinct, sclerotized laciniae and non-elon- Larval head with antennae unusually long (seg-
gate galae. Labial palps with vom Rath's organ ment 2 particularly elongate). Anterior tentorial
well developed. Hypopharynx broad, with varia- pits close to antennal base. Larval trunk approx-
bly armed infrabuccal cavity. Laterocervicale imately hexagonal in cross section. Trunk integu-
with 'hair plate' close to the apex. Pronotal plate ment with cuticle specializations unique among
not subdivided, with at least one pair of seta- arthropods: exocuticle high, with liquid-filled
bearing 'warts'. First thoracic spiracle of 'primi- chambers (each corresponding to one epidermal
tive type' (see 2 - 8 ) . Wing-'coupling' jugo-fre- cell) in a honeycomb-like pattern, exo-and endo-
nate (but there is presumably no functional cuticle separated by fluid-filled space, con-
coupling, judging from the photographic obser- tinuous with 'honeycomb' chambers via pores in
vations on the overall similar Glossata-Eriocran- lower exocuticular layer; cuticle overlaid by
iidae by Grodnitsky & Kozlov 1985). Wings held sticky pellicle to which foreign bodies often ad-
tectiform in repose, set with microtrichia ('acu- here. Thoracic legs short, with at most 4 seg-
lei'); wing surface scales (not marginal 'fringes') ments, subterminal one presumably a tibiotarsus.
'primitive type' exclusively, with 'herringbone'
pattern (see 2 - 2 ) . Sternum I a distinct, discrete It is surprising, that a Micropterix species has
plate with produced corners. In male genitalia been found to have eupyrene sperm exclusively,
valve consistently devoid of movable medial while an Epimartyria like other Lepidoptera in-
(morphologically distal) appendage. Female with vestigated has both eupyrene and apyrene sperm.
single genital opening, common oviduct commu- However, the monophyly of the Micropterigidae
nicating with genital chamber ventrad from is strongly supported by the numerous synapo-
bursa copulatrix. Cephalic CNS with free trito- morphies of Micropterix and other micropterigid
cerebral commissure. All three suborders in genera, including Epimartyria.
ground plan with well-developed sternum V Small moths (wingspan up to ca. 15 mm, usu-
gland (at least in male) and with hindwing Rs ally ca. 10 mm or less). Head with headcapsule
four-branched; secondary reductions in these only moderately extending above upper margin
traits occur in Micropterigidae. The larvae in this of compound eyes. Ocelli usually prominent,
grade are more or less distinctly prognathous, very rarely lost. Antennae moniliform to fili-
without a spinneret on the apex of the prelabio- form, scaling variable, without typical sensilla
hypopharyngeal lobe; larval structure otherwise auricillica (probably a plesiomorphy); antennae
very diverse. The decticous, exarate pupae have and palps with a diversity of little understood
the cephalic setae very prominent; clypeus dis- sensory (and glandular?) organs (Le Cerf 1926,
tinctly demarcated, setose. Faucheux 1992). In cibarium epipharynx usually
with complex asymmetrical armature of sclerites
42 Niels P. Kristensen

and microtrichial brushes (which serve to clean male accessory glands. Each ovary with 4 or 5
the mandibles, Hannemann 1956) and hypophar- ovarioles.
ynx with 'triturating basket', i. e. infrabuccal Chorion in the Micropterix egg highly special-
cavity set with strong, transversely aligned spines ized, with clubbed protuberances formed within
(similar cibarial specializations present in Heter- one hour after oviposition (see 218, 219); egg
obathmiidae, parallelism or symplesiomorphy?). in Neomicropteryx covered with hygroscopic, ge-
Labial palps usually 2-segmented (3 segments re- latinous mass. Larvae (Fig. 4.2 A) strikingly dif-
tained in a few 'Sabatinca', only a single present ferent from other lepidopteran caterpillars. Head
in Neomicropteryx). Metathoracic furcal stem in repose completely retractible into thorax (an-
without anterior process (apparently a uniquely tennal length surely related to this extraordinary
primitive condition in the Lepidoptera, absence retractability). Antennae 3-segmented. A median
of this formation elsewhere in the order interpre- seta on frontoclypeus between antennae proba-
ted as secondary loss). Protibial epiphysis lost in bly homologous with a campaniform sensillum
a few genera. Wings narrow, but outline variable, in caddisfly larvae, but without obvious homo-
apex sometimes pointed, sometimes obtusely logues in other Lepidoptera. 5 or 6 stemmata,
rounded. Forewing pattern often iridescent, very contiguous or at least close-set. Headcapsule
variable (Figs. 4.3 AC), in some genera promi- without adfrontal ridges and with short hyposto-
nent transverse fasciae are commonplace. Fore- mal bridge (plesiomorphies shared with Aglos-
wing (Fig. 4.1 A) with Sc forked (except in an sata), tentorial arms and corporotentorium
undescribed dwarfish New Caledonian taxon), stout, labrum with both medial and lateral re-
crossvein Sc-R usually present, R forked or sim- tractors retained, maxilla with both galea and
ple, position of Rs4 relative to apex variable, A lacinia distinct (unique plesiomorphies within
veins sometimes in typical 'double-Y' configura- Lepidoptera). Trunk setae simple (some Saba-
tion, or A 2 and A 3 strongly reduced; hindwing tinca) or thickened, truncate or davate; Hinton's
with configuration of Sc-R region variable, posi- (1958) claim of the distribution of primary setae
tion of Rs4 variable as in fore wing, in Squami- being "entirely different" in zeuglopteran and
cornia only three Rs branches present (Fig. other lepidopteran larvae has not been con-
4.1 B). Sternum V gland orifice in some genera firmed (Davis 1987), but the ground plan of the
on sculptured process which may, or may not, be former still remains to be worked out. Abdomen
set with hair scales, in other genera simple, or usually without appendages, but Micropterix and
gland lost. Segment IX usually a completely scle- a species group within 'Sabatinca' with pointed,
rotized ring, but dorsalmost part sometimes non-musculated and non-crochet-bearing pro-
membranous, its posterodorsal margin in Micro- legs on IVIII. Abdominal end in Micropterix
pterix with a variable complement of processes with a threelobed 'sucker', with which the larva
(Fig. 4.1 C). Phallus not of phallotheca-type; in may attach itself to the substrate. First thoracic
Epimartyria (Fig. 4.1 D) with accessory ventral and all eight abdominal spiracles functional in
branch. Segment X variable, presumably primi- some genera; Hinton's report of a functional spi-
tively with paired tergal lobes. Female abdomen racle on metathorax may refer to a topographi-
(Fig. 4.1 F) with ten distinct segments and termi- cally forwards shifted spiracle I (Davis 1987).
nal cloaca; segment IX sclerotization variable, a Pupa (Fig. 4.2 B) with mandibles not hypertro-
complete ring or reduced/absent on dorsum; seg- phied.
ment X sclerotization a pair of lateral setose
plates; segment XI probably represented only by Adults usually diurnal, but nocturnal activity
circumcloacal membrane. Terminal abdominal is reported for some species. They are mostly
segments in repose telescoped into segment VIII, pollen eaters. Micropterix species feed in the
extensible through blood pressure. Bursa copu- flowers of a great variety of dicot and monocot
latrix in some genera with highly characteristic plants, often in trees. In contrast, the 'Sabatinca-
'signa' sclerotizations (Fig. 4.1 F). Ductus sper- group' genera have almost exclusively been ob-
mathecae, as far as known, without double-com- served on sedges; particularly interesting excep-
partment structure (a unique plesiomorphy tions are the feeding of some New Caledonian
within the Lepidoptera). Nature of formation taxa on pollen of Winteraceae and fern spores,
termed 'circumcloacal chamber' (Dugdale 1974) the latter is also a food source of Malagasy taxa.
awaiting reinvestigation. Gut without stomo- The larvae may be broadly characterized as soil
daeal crop and with long midgut commencing animals with high moisture demands. Most
in thorax, few rectal papillae (3 in Micropterix). known larvae in the 'Sabatinca-group' feed on
Nerve cord with 4 or fewer (2 in some genera) folise liverworts, but the taxon with distinct
abdominal ganglia; abdominal nerve sheath dor- prolegs has been found in rotten logs (feeding
sally with enlarged cells. Tracheal system in on fungus hyphae?), and Micropterix larvae in
Micropterix without transverse commissures in addition to detritus/fungus hyphae eat green an-
abdominal segments, but this condition is not giosperm leaves. Pupation takes place in an oval
general in the family. Testes follicles minute, tes- cocoon (Fig. 4.2 B), which may be a pure-silk
tes of the two sides not united; a single pair of formation (Sabatinca, Epimartyria, Neomicro-
pteryx) or may have soil particles incorporated
The Non-Glossatan Moths 43

into the outer wall (Micropterix); at emergence it Pirngruber (1944), Lorenz (1961), Carter & Dug-
is cut open in the anterior end, which at least dale (1982), Tuskes & Smith (1984) and Pellmyr
in Micropterix is less dense than other parts. In et al. (1990).
temperate zones adult micropterigids are pre-
dominantly spring/early summer insects.
Clade Agathiphagoidea + Heterobathmioidea +
121 described species, but about half again as
many are recognized and still unnamed. The
family has been provisionally classified into the Probable autapomorphies of the ground plan of
'Micropterix group' (Micropterix only, 67 species, this clade (Kristensen 1984 f) include: Loss of
Palaearctic, most speciose in Mediterranean sub- paraglossae. Presence of anterior process (ac-
region; Fig. 4.3 C) and the 'Sabatinca group' commodating origin of sternal trochanter de-
comprising all other genera; the principal puta- pressors) on metafurcal stem. Ductus sperma-
tive autapomorphy of the latter is the complete thecae with thickwalled and thinwalled compart-
absence of discrete male venter VIII sclerotiza- ment. Larval head with tentorium delicate, me-
tions, but its monophyly remains questionable. dian labral retractor lost, only single maxillary
One apparently monophyletic subgroup com- endite present, cranial flexor of dististipes lost,
prises 4 genera with an amphipacific N. Hemi- unique dorsoventral muscle laterally spanning
sphere distribution: Taiwan/China/Japan (where foramen magnum. Pupa (Fig. 4.2 D) with mandi-
Neomicropteryx has been studied in some detail) ble hypertrophied, angularly bent (this special-
- N.America (Epimartyria only; Fig. 4.3B). ization lost again in higher Glossata).
Otherwise the group is mainly SW Pacific: Sa- In the currently preferred phylogeny of this
batinca s. lat. occurring in E. Australia, New clade the Heterobathmioidea and the Glossata
Zealand and particularly New Caledonia, which constitute a monophylum; derived characters
has a 'swarm' of about 50 species, of which only shared by the Agathiphagoidea and Heterobath-
three are so far named. There is one small genus mioidea are therefore considered homoplasious.
(Agrionympha) in S. Africa, one monotypic in
austral S. America, and a recently discovered,
still un-named taxon in Madagascar; apart from AGATHIPHAGOIDEA
the monotypic Squamicornia from montane Agathiphagidae. Probable autapomorphies: Ocelli
Equador the absence of records of micropterigids absent. Antennae with bifurcate sensilla. Ster-
from the continental tropics is conspicuous. num V gland in male opening on long, smooth
process; gland structure uniquely specialized, se-
Principal references on taxonomy/biogeogra-
cretory section a long and twined tube leading
phy include Issiki (1931; with extensive discus-
into large thinwalled reservoir; gland absent in
sion of previous works on systematics and integ-
female. Male segment IX ring greatly indented in
umental structure), Whalley (1978), Kristensen &
dorsal and ventral midline. Genital chamber in
Nielsen (1979, 1982), Kaltenbach & Speidel
roof with distinctive complement of sclerotiza-
(1982), Minet (1985), Gibbs (1983, 1989), Kozlov
tions, including median 'spiny plate'. Larva with
(1988-90). The morphological literature on
but two stemmata (without corneal lenses) pre-
adults is relatively extensive; because of the long-
sent on each side. Antenna reduced, one-seg-
recognized phylogenetic key position of the
mented. Endophagous larva with trunk unpig-
micropterigids they (usually at least a Micro-
mented and with greatly reduced setae, thoracic
pterix species) are considered in many compara-
legs and abdominal appendages absent.
tive accounts of lepidopteran structures, see
Additional agathiphagid apomorphies are
vol. 2. Studies devoted to this family deal with
shared with the Heterobathmioidea, but must be
the adult head (Le Cerf 1926, Tillyard 1923 a,
homoplasious autapomorphies of both taxa, if
Hannemann 1956, Chauvin & Faucheux 1981,
the Heterobathmioidea are indeed the sister
Faucheux 1992), wings (Tillyard 1919, Philpott
group of the Glossata (12): Anterior tentorial
1923), pregenital abdomen (Kristensen 1984 c),
arms fused into Y-shaped formation. Postlabium
genitalia (Philpott 1934, 1924, Hannemann 1957,
proximally widenend. Mesepisternal pre- and
Kristensen 1984 a), brain (Buxton 1917), tracheal
paracoxal sutures anastomosing below anapleu-
system (Kristensen 1984 b), spermatogenesis
ral cleft. Tergum I with lateral process indistinct
(Friedlnder 1983, Sonnenschein & Huser 1990,
and hind corners fused with tergum II. In larval
Hamon & Chauvin 1992), ovary, egg structure
head labral compressors, intrinsic maxillary
and embryogenesis (Chauvin & Chauvin 1980,
muscles and dorsal postcerebral pharynx dilators
Ando & Kobayashi 1978, Kobayashi 1994, Ko-
lost. Pupal mandible with strong subapical tooth
bayashi & Ando 1981, 1982). The extraordinary
on lower surface.
larval structure has so far been treated only in a
Medium-sized micro-moths, wingspan up to
preliminary way (Tillyard 1923 b, Mutuura 1956,
ca. 27 mm. Facies singularly caddisfly-like, with
Hinton 1958, Lorenz 1961, Yasuda 1962, Davis
the antennae stretched forward in repose (other
1987, Kozlov 1991, Kristensen 1990). Principal
primitive moths have them upward, in a V-like
works on natural history and behaviour include
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 45

antenna claws
I labial palp

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).

(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
Fig. 4.3. Micropterigidae: - C ; Agathiphagidae: D; area possibly pertaining to XI; phallus of phallo-
Heterobathmiidae: E. A, Sabatinca lucilia-, B, Epimar- theca-type, with well-developed eversible aedea-
tyria pardella; C, Micropterix aureatella\ D, Agathi- gus. Each testis comprising four large saccular
phaga vitiensis; E, Heterobathmiapseuderiocrania. Uni- follicles with narrow apex (unique condition in
form magnification, scale (milimetres) in D. Lepidoptera, most probably a plesiomorphy);
seminal vesicle close to testis, questionably ho-
mologous with vesicles in other Lepidoptera; two
and cryptic, brownish mottled (Fig. 4.3 D). pairs of long (one particularly long, twined), tu-
Headcapsule extending far above compound bular accessory glands (similarly unusual lepi-
eyes, with high internal crest medially above an- dopteran condition, interpretation questionable).
tennae. Antennae filiform. Dorsal tentorial arms Female segment VIII (Fig. 4.11) posteriorly nar-
short. Mandibles large, but without incisivus rowing, tergal and ventral plate separate, each
cusps, post-ecdysial function unknown; adductor with pair of long apophyses on anterior margin;
muscle huge, origin covering large parts of dorsal segments I X - X I partly retractible into VIII,
headcapsule, including median crest. Basal forming narrow, weekly sclerotized probe with
maxillary piece with costa between cardo and ba- Y-shaped sclerotization in dorsal wall, arms of
sistipes; galea a large membranous pad with nu- ' continuing into long internal apophyses; clo-
merous sensilla basiconica. Cibarium with simple acal opening subterminal. Bursa copulatrix
epipharynx; armature in infrabuccal pouch slen- small, without signa. Colleterial glands very
der, scattered microtrichia (not strong, aligned large sacs. Very numerous (40 +) ovarioles per
spines as in 'triturating baskets' of Micropteri- ovary (isolated case among homoneurous Lepi-
goidea and Heterobathmioidea; another family doptera, probably a plesiomorphy). Gut with
autapomorphy?). Legs with spur formula 1-4-4 large, thinwalled stomodaeal crop extending into
46 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.

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.
Fig. 4.4. Agathiphaga vitiensis, larva (length ca. 6 mm.) Spurs 0-0-4 (loss of mesotibial spurs paralleled
in Agathis seed.
in Micropterigoidea). Forewing Sc and R simple.
Forewing scales on obverse surface with smooth
rounded plates on inter-ridge areas. Male seg-
abdominal base; midgut short; six malpighian tu- ment VIII desclerotized in middle (parallelism
bules each discharging separately into gut; nine with Micropterigoidea); segment IX (Fig. 4.1 K)
rectal papillae. Nerve cord with five abdominal with short, stout (sometimes markedly bifid)
ganglia (discrete ganglia I I - V , plus composite ventromedian process; two pairs of variably
terminal ganglion); sheath thickened dorsally, shaped processes arising f r o m posterolateral cor-
but thickened mass apparently mainly cellular, ners of dorsal plate; base of phallus with posteri-
rather t h a n composed of extracellular material. orly directed apdeme; postgenital complex with
variable armature of spines.
Egg undescribed. Larva (Fig. 4.3 F) endopha-
gous (seed miner). Head capsule weekly mela- If the phylogenetic hypothesis preferred here
nized; adfrontal ridges and distinct ecdysial lines (see above, and chapter 2) is correct, suites of
lacking; complete hypostomal bridge present; specializations shared with the Micropterigoidea
short tentorial dorsal arms present. Maxilla (shortened labial palps, lack of mesotibial spurs,
without discrete cardo. Labial glands very long, medial desclerotization of male venter VIII, and
thick tubes. perhaps cibarial armature) and with the Agathi-
Adults are nocturnal and have been encoun- phagoidea (listed under the latter above) are ho-
tered in nature on but very few occasions; it is moplasious autapomorphies of the Heterobath-
u n k n o w n whether they feed at all. The eggs are mioidea as well as the other taxa.
believed to be inserted below the cone scales of Small m o t h s (Fig. 4.3 E), wingspan usually ca.
the kauri pines (Agathis, Araucariaceae) in the 10 mm. Headcapsule extending markedly above
seeds of which the larvae subsequently feed. The c o m p o u n d eyes. Ocelli present. Antennae with
excrements are incorporated into the lining of sensilla auricillica (probable s y n a p o m o r p h y with
the larval cell. U p o n maturity the larva coats the Glossata). Mandibles with strong incisivus cusps.
cell wall with a (labial gland?) secretion, produc- Wings narrow (forewing index < 0.3) but not
ing a smooth, hard layer. Larvae probably al- acutely pointed. Forewings iridescent purple,
ways enter a diapause, which seems to be termi- with silvery-white scales forming more or less ex-
nated only under high moisture regimes, and tensive suffusion and often distinct spots at least
which under laboratory conditions is very diffi- in tornai area; superficial similarity with Glos-
cult to terminate anyway. Before eclosion the sata-Eriocraniidae sometimes striking. Forewing
pharate adult grinds an emergence whole by Sc and R unforked, crossvein Sc-R absent (but
means of the hypertrophied pupal mandibles. In- weak basal crossvein S c - R + R S sometimes pre-
festation of pine seeds may locally attain pest sent); base of 1A sometimes appearing forked;
levels. pterostigma distinct in forewing, small in hind-
The single genus Agathiphaga comprises two wing. Hindwing base (Fig. 4.1 J) with distinct,
closely related species, occurring in Australia rounded jugal lobe (apparently a unique plesio-
(Queensland) and Fiji/Solomon Islands/New m o r p h y in the Lepidoptera). Sternum V gland
Hebrides/New Caledonia respectively. present in b o t h sexes, orifice on sculptured pro-
The original description of Agathiphaga by cess. Male segment IX (Fig. 4.1 K) usually com-
Dumbleton (1952) contained much information pletely divided into dorsal and ventral plate, but
on biology and integumental structure; subse- in Hetorobathmia valvifer an anteriorly complete
quent biological observations are reviewed by sclerotized ring; phallus not of phallotheca-type
Robinson & Tuck (1976) and C o m m o n (1990). (contrary to statement in original description),
Detailed structural accounts deal with the anten- ejaculatory duct partly melanized. In female
The Non-Glossatan Moths 47

postabdomen (Fig. 4.1 L) segment VIII with sim- mesothorax to VIII, venter I excepted. Terminal
ple tergum and functional spiracle; large subgeni- segment a trilobed formation, with subanal lobes
tal plate on venter VIII covering base of mem- forming paired 'anal feet' (without crochets).
branous ventral side of terminal segments. Seg- Pupa with apical teeth on hypertrophied mandi-
ment IX bilaterally flattened, high tergal plate bles markedly asymmetrical; right mandible with
with variably developed process arising above prominent tooth on lower edge, one-third from
anterolateral corner; in the best known species, apex.
Heterobathmia pseuderiocrania, this process is All known heterobathmiids are associated
very short indeed, in H. valvifer it is a genuine, with deciduous Nothofagus. The adults are very
albeit rather short, apophysis. Segment X proba- early spring insects and only fly actively in sun-
bly represented by small paired, unmelanized shine. They visit Nothofagus flowers and are be-
and setose lobes which are immovably fused with lieved to be pollen feeders. Eggs are deposited on
tergum IX; genital chamber and rectum opening the lower surface of the host leaf. The full-depth
separately in shallow median groove of membra- mine starts as a narrow gallery, but after the first
nous ventral surface of terminal segments. Duc- moult a blotch mine is formed, often obliterating
tus bursae opening on prominent, thickwalled the gallery. Frass is initially deposited at the egg
papilla in genital chamber; intima in ductus and site, which thereby becomes very conspicuous.
proximal corpus bursae with delicate spines and As leaf miners heterobathmiid larvae are unusual
ridges; no true signa. in being able to leave one leaf and start mining in
The species H. valvifer is unique in the Lepi- another. Feeding is rapidly completed. Pupation
doptera in having a pair of long, flattened ap- takes place in the earth, in a silken cocoon cov-
pendages arising from the ventral VIII/IX ered with soil particles.
boundary region, their bases being laterally very The single genus Heterobathmia comprises at
close to the anterolateral corners of tergum IX. least 9 species, of which 3 have been named so
The only musculature associated with these ap- far. They occur with deciduous Nothofagus in
pendages is the ventralmost fibre bundle of a austral. S. America.
broad muscle extending from the subgenital Principal references: Kristensen & Nielsen
plate to the membranous body wall adjacent to (1979, 1983, in press), Nielsen (1985).
the anterolateral margin of tergum IX; this bun-
dle inserts immediately in front of the appendage
base. The morphological interpretation of these Acknowledgements
appendages is highly problematical; if they are in Communication of unpublished information on
any way derivable from an element in the gener- Malagasy Micropterigidae from D. C. Lees in
alized insect ovipositor, they can only be consid- gratefully acknowledged.
ered an autapomorphic character reversal. The
female postabdomen of H. valvifer is also notable
for the presence of ribbonshaped longitudinal References
sclerites flanking the median groove on the mem-
Ando, H. & Kobayashi, Y (1978): The formation of
branous ventral surface of the terminal segments. germ rudiment in the primitive moth, Neomicro-
Gut without stomodaeal crop; long midgut pteryx nipponensis Issiki (Micropterygidae, Zeug-
commencing in thorax; 6 malpighian tubules loptera, Lepidoptera) and its phylogenetic signifi-
opening separately into gut. Ventral diaphragm cance. - roc. Jap. Soc. syst. Zool. 15: 4 6 - 5 0 .
with well developed segmental alae. Other as- Buxton, P. A. (1917): On the protocerebrum of Micro-
pects of visceral anatomy still remain to be pteryx. - Trans, ent. Soc. Lond. 1917: 1 1 2 - 1 5 3 .
worked out. Carter, D. J. & Dugdale, J. S. (1982): Notes on collect-
ing and rearing Micropterix (Lepidoptera: Micro-
Egg undescribed, covered by a jellylike sub-
pterigidae) larvae in England. - Ent. Gaz. 33:
stance. Leafmining larva (Fig. 4.2 C) progna- 43-47.
thous. Headcapsule with prominent adfrontal Chapman, T. A. (1917): Micropteryx entitled to ordi-
and hypostomal ridges, hypostomal lobes medi- nal rank; order Zeugloptera. Trans, ent. Soc.
ally separated by distinct membranous area Lond. 1917: 3 1 0 - 3 1 4 .
(noteworthy probable synapomorphies with Chauvin, G. & Faucheux, M. (1981): Les pices buc-
Glossata). Antennae short, 3-segmented. 7 stem- cales et leurs rcepteurs sensoriels chez l'imago de
mata (this high number a unique plesiomorphy Micropterix calthella L. (Lepidoptera: Micropterigi-
within the Lepidoptera) present on each side. La- dae). - Int. J. Insect Morph. Embryol. 10: 4 2 5 - 4 3 9 .
Chauvin, J. T. & Chauvin, G. (1980): Formation des
brum with even epipharyngeal surface sclero-
reliefs externes de l'oeuf de Micropteryx calthella L.
tized. Mandible with medial tuft of barbed hairs.
(Lepidoptera: Micropterigidae). - Can. J. Zool. 58:
Prothorax with well developed notai and sternal 761-766.
sclerites, trunk otherwise membranous and un- Common, I. F. B. (1990): Moths of Australia, vi +
pigmented. Thoracic legs well developed, with 535 pp., 32 pis. Leiden.
large, subdivided trochanters (unique plesiomor- Davis, D. R. (1987): Micropterigidae. Pp. 3 4 1 - 3 4 3 in
phy within the Lepidoptera). Eversible calli pre- Stehr, F. (ed.) Immature insects. Kendall/Hunt, Du-
sent on dorsum and venter of all segments from buque.
48 Niels P. Kristensen

Dugdale, J. S. (1974): Female genital configuration in - (1984 b): Respiratory system of the primitive moth
the classification of Lepidoptera. - N. Z. J. Zool. Micropterix calthella (Linnaeus) (Lepidoptera:
1: 127-146. Micropterigidae). - Int. J. Insect. Morph. Embryol.
Dumpleton, L. F. (1952): A new genus of seed-infest- 13: 137-156.
ing micropterygid moths. - Pacific. Sci. 6: 1 7 - 2 9 . - (1984 c): The pregenital abdomen of the Zeuglo-
Faucheux, M. J. (1990): Antennal sensilla in adult ptera (Lepidoptera). Steenstrupia 10: 113-136.
Agathiphaga vitiens Dumbl. and A. queenslandensis - (1984 d): The larval head of Agathiphaga (Lepidop-
Dumbl. (Lepiodoptera: Agathiphagidae). - Int. J. tera Agathiphagidae) and the lepidopteran ground
Insect Morphol. & Embryol. 19: 2 5 7 - 2 6 8 . plan. - Syst. ent. 9: 6 3 - 8 1 .
- (1992): Structures sensorielles et glandulaires parti- - (1984e): The male genitalia of Agathiphaga (Lepi-
culires de l'antenne de Micropterix calthella L. doptera, Agathiphagidae) and the lepidopteran
(Lepidoptera: Micropterigidae). - Bull. Soc. Se. ground plan. - Ent. scand. 15: 151 178.
Nat. Ouest de la France 14: 114-118. - (1984 f): Studies on the morphology and systemat-
Friedlnder, M. (1983): Phylogenetic branching of ics of primitive Lepidoptera (Insecta). Steen-
Trichoptera and Lepidoptera: an ultrastructural strupia 10: 141-191.
analysis on comparative spermatology. - J. - (1990): The trunk integument of zeuglopteran lar-
Ultrastruct. Res., 83: 141-147. vae: one of the most aberrant arthropod cuticles
Gibbs, G . W. (1983): Evolution of Micropterigidae known. - Bull. Sugadaira montane Res. Cen. 11:
(Lepidoptera) in the S. W. Pacific. - GeoJournal 101-102.
7.6: 5 0 5 - 5 1 0 . - & Nielsen, E. S. (1979): A new subfamily of micro-
- (1989): Local or global? Biogeography of some pterigid moths from South America. A contribu-
primitive Lepidoptera in New Zealand. . . J. tion to the morphology and phylogeny of the
Zool. 16: 6 8 9 - 6 9 8 . Micropterigidae, with a generic catalogue of the
Hamon, C. & Chauvin, G. (1992): Ultrastructural family (Lepidoptera: Zeugloptera). - Steenstrupia
analysis of spermatozoa of Korscheltellus lupulinus 5: 6 9 - 1 4 7 .
L. (Lepidoptera: Hepialidae) and Micropterix cal- - (1980): The ventral diaphragm of primitive (nondi-
thella L. (Lepidoptera: Micropterigidae). Int. J. trysian) Lepidoptera. A morphological and phylo-
Insect Morphol. Embryol. 21 (2): 149-160. genetic study. - Z. zool. Syst. Evol. Forsch. 18:
Hannemann, H . J . (1956): Die Kopfmuskulatur von 123-146.
Micropteryx calthella (L.) (Lep.). Morphologie und - (1981c): Abdominal nerve cord configuration in
Funktion. - Zool. Jb. (Anat.) 75: 177-206. adult non-ditrysian Lepidoptera. Int. J. Insect
- (1957): Die mnnlichen Terminaba von Micro- Morph. Embryol. 10: 8 9 - 9 1 .
pteryx calthella L. (Lep. Micropterygidae). - Dt. - (1982): South American micropterigid moths: two
ent. Z. (N. F.) 4: 2 0 9 - 2 2 2 . new genera of the Sabatinca group. Ent. scand.
Hinton, H. E., 1958. The phylogeny of the panorpoid 13: 5 1 3 - 5 2 9 .
orders. - A. Rev. Ent. 3: 181-206. - (1983): The Heterobathmia life history elucidated:
Issiki, S. T. (1931): On the morphology and systemat- Immature stages contradict assignment to suborder
ics of Micropterygidae (Lepidoptera: Homoneura) Zeugloptera (Insecta, Lepidoptera). - Z. zool.
of Japan and Formosa, with some consideration on Syst. Evol. Forsch. 21: 101-124.
the Australian, European and North American - (1998): Heterobathmia valvifer n. sp.: a moth with
Forms. - Proc. zool. Soc. Lond. 1931: 999-1039. long apparent ovipositor valves. - Steenstrupia
Kaltenbach, T. & W. Speidel (1982): Eine neue Ur- (in press).
motte aus China (Micropterigidae). - Nota lepid. Le Cerf, F. (1926): Contribution a l'etude des organes
5(1): 3 1 - 3 6 . sensoriels des Lpidoptres. Pp. 133158 in Le
Kobayashi, Y. (1982): The early embryonic develop- Cerf, F. (ed.) Lepidoptera I, 3. Paris, P. Lechevalier.
ment of the primitive moth, Neomicropteryx nippo- Lorenz, R. E. (1961): Biologie und Morphologie von
nensis Issiki (Lepidoptera, Micropterygidae). - J. Micropterix calthella (L.). Dt. ent. Z. (N. F.) 8:
Morph. 172: 2 5 9 - 2 6 9 . 1-23.
- (1994): Ovarian structure of a zeuglopteran moth, Minet, J. (1985): Description de nouveaux Sabatinca
Neomicropteryx nipponensis issiki (Lepidopteran, no-caldoniens et redfinition du genre en fonction
Micropterigidae). Jap. J. Entomol. 62, 1: 93 100. de considrations phylogntiques (Lepidoptera,
Kobayashi, Y. & Ando, H. (1981): The embryonic de- Micropterigidae). - Nouv. Revue. Ent. (N. S.) 2:
velopment of the primitive moth, Neomicropteryx 285-297.
nipponensis Issiki (Lepidoptera, Micropterygidae): Mortimer, J. F. (1965): The alimentary canals of some
Morphogenesis of the embryo by external observa- adult Lepidoptera and Trichoptera. Trans. R.
tion. - J. Morph. 169: 4 9 - 5 9 . ent. Soc. Lond. 117: 6 7 - 9 4 .
Kozlov, M. V. (1988-90): A brief review and a key to Mutuura, A. (1956): On the homology of the body
Palearctic species of the genus Micropterix (Lepi- areas in the thorax and abdomen and new system
doptera, Micropterigidae) 14. - Vestn. Zool. 1988 of the setae on the lepidopteran larvae. Bull.
(4): 8 - 1 4 ; 1989 (6): 2 6 - 3 1 ; 1990 (2): 2 1 - 2 6 ; 1990 Univ. Osaka Prefect. (B) 6: 9 3 - 1 2 2 .
(3): 2 8 - 3 3 . Nielsen, E. S. (1985): Primitive (non-ditrysian) Lepi-
- (1991): Die Grundplanmerkmale und Autapomor- doptera of the Andes: diversity, distribution, bio-
phien im Bau der Micropterigiden-Raupen (Lepi- logy and phylogenetic relationships. - J. Res.
doptera). - XII Int. Symp. ber Entomofaunistik Lepid., Suppl. 1, 1985: 1 - 1 6 .
Mitteleuropa Verhandlungen, Kiev. Pp. 326330. Pellmyr, O., Thien, L. B. Bergstrm, G . & Groth, I
Kristensen, . P. (1984a): Skeletomusculatur anatomy (1990): Pollination of New Caledonian Wintera-
of the male genitalia of Epimartyria (Lepidoptera: ceae: opportunistic shifts or parallel radiation with
Micropterigidae). - Ent. scand. 15: 9 7 - 1 1 2 . their pollinators? - PI. Syst. Evol. 173: 143-157.
The Non-Glossatan Moths 49

Philpott, A. (1921): A study of the venation of the New and a comment on the evolution of the Lepidoptera
Zealand species of Micropterygidae. - Trans. Proc. (Insecta). - Ann. Transv. Mus. 31: 7286.
. Z. Inst. 54: 155-161. Yasuda, T. (1962): On the larva and pupa of Neomicro-
- (1923): The genitalia in Sabatinca and allied genera pteryx nipponensis Issiki, with its biological notes
(Lepidoptera Homoneura), with some observations (Lepidoptera, Micropterygidae). - Kontyu 30:
on the same structures in the Mecoptera. Trans, 130-136.
ent. Soc. Lond. 1923: 3 4 7 - 3 6 6 .
- (1927): Notes on the female genitalia in the Micro-
pterygoidea. - Trans, ent. Soc. Lond. 1927: 319 Addendum
Robinson, G. S. & Tuck, K. R. (1976): The kauri moth A SEM account of the antennal sensilla of Mi-
Agathiphaga. A preliminary report, 8 pp. British cop ter ix calthella was given by Faucheux (1997)
Museum (Natural History), London. and the larval integument of the same species
Sonnenschein, M. & Huser, C. L. (1990): Presence of
was described by Hamon & Chauvin (1995). A
only eupyrene spermatozoa in adult males of the
genus Micropterix Hbner and its phylogenetic sig-
S. African micropterigid taxon phenetically sim-
nificance (Lepidoptera: Zeugloptera, Micropterigi- ilar to generalized SW Pacific Sabatinca has been
dae). Int. J. Insect Morphol. & Embryo!. 19: discovered (Kristensen unpublished).
269-276. Upton (1997) reported on Agathiphaga larvae
Tillyard, R. J. (1919): On the morphology and system- staying alive after 12 years in diapause.
atic position of the family Micropterygidae (Sens.
Lat.). - Proc. Linn. Soc. N. S. W. 44: 9 5 - 1 3 6 . Faucheux, M. J. (1997): Sensory organs on the anten-
- (1923 a): On the Mouth-parts of the Microptery- nae of Micropterix calthella L. (Lepidoptera,
goidea (Order Lepidoptera). - Trans ent. Soc. Micropterigidae). - Acta zool. 78: 1 - 8 .
Lond. 1923: 181-206. Hamon, C. & Chauvin, G. (1995): Larval integument
- (1923 b): On the larva and pupa of the genus Saba- and its differentiation in Micropterix calthella L.
tinca (Order Lepidoptera, family Micropterygidae). Lepidoptera: Micropterigidae): anatomy and ultra-
- Trans, ent. Soc. Lond. 1923: 4 3 7 - 4 5 3 . structure. - International Journal of Insect Mor-
Tuskes, P. M. & Smith, N. J. (1984): The life history phology and Embryology 24: 213-222.
and behaviour of Epimartyria pardella (Micropteri- Upton, M. S. (1997): a twelve-year larval diapause in
gidae). - J. Lepid. Soc. 38 (1): 4 0 - 4 6 . the Queensland kauri moth, Agathiphaga queenslan-
Whalley P. (1978): New Taxa of fossil and recent densis Dumbleton (Lepidoptera: Agathiphagidae).
Micropterigidae with a discussion of their evolution - The Entomologist 116: 142-143.
5. The Homoneurous Glossata
Niels P. Kristensen

As outlined in 1 - 2 the monophyly of the clade 4.1 C, see also 24): the sclerotized abdominal
Glossata is very strongly corroborated, while the apex is flattened, pointed and has serrate edges,
monophyly of the clade Heteroneura, compris- forming an 'oviscapt saw'; a subterminal mem-
ing the vast majority of the glossatan superfami- branous area below the saw bears a pair of elon-
lies, and characterized i. a. by the 'heteroneu- gate, near-tubular sclerotizations ('ventral rods'
rous' wings, is a reasonably supported working in 2 - 4 , 'tertiary apophyses' of Davis 1978), at
hypothesis. Within the 'homoneurous' grade in the posterior apex of which the cloacal orifice is
the Glossata five extant lineages are currently located. The oviscapt saw is in repose completely
recognized, four of them with but one included retracted into the preceding 'oviscapt cone', a
family. In the presently preferred cladogram more or less extensively synscleritous ring which
(12, Fig. 2.2) the six extant basal glossatan lin- tapers posteriorly. The oviscapt saw and oviscapt
eages are seen to have arisen in a single sequence cone both have long paired apophyses invagi-
of five splitting events. 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
Primarily characterized by notable specializa- do not form a monophylum, it is arguably a glos-
tions in the adults' mouthparts: mandibles non- satan groundplan trait.
functional after pupal/adult ecdysis, without The name Dacnonypha (introduced by Hin-
clearly defined articulations with head capsule ton, 1946) has been used for a high-rank taxon
(and often strongly reduced); maxilla without centred on the Eriocraniidae. However since the
distinct, sclerotized lacinia, galeae elongate, Eriocraniidae do not appear to form a monophy-
forming a proboscis, which is coiled in repose lum together with any other subset of the Glos-
and primitively has the walls of the food canal sata (Nielsen & Kristensen 1996), the category
lined with discrete cuticular plates; in maxillary Dacnonypha is redundant and here abandoned.
palp muscle between segments 2/3 lost; prelabio-
hypopharyngeal lobe narrowed, hypopharynx
without infrabuccal pouch; on head capsule epi- ERIOCRANIOIDEA
stomal suture reduced, not accompanied by cu- Eriocraniidae. Probable autapomorphies: tibial
ticular thickening (regression surely related to re- spur formula 0-1-4 (unpaired mesotibial spur
duction of mandibular function). Dorsal tentor- paralleled in the Acanthopteroctetidae). Valvae
ial arms considerably larger than in non-glos- greatly shortened and phallus with accessory
satans, but apparently reduced again in ground ventral branch (Fig. 5.1 E, F). Female with com-
plan of the Heteroneura. Trend towards complex plex 'vaginal sclerite' (sensu Davis 1978) in poste-
fusion of adult brain and suboesophageal gan- rior part (behind spermathecal orifice) of genital
glion recognizable already in ground plan chamber encircling egg passage (Fig. 5.1 D).
through incorporation of tritocerebral commis- Ventral diaphragm musculature exceedingly re-
sure in suboesophageal mass. Larva with spin- duced, a single fibre pair present per segment (re-
neret on apex of prelabio-hypopharyngeal lobe. duction parallelled in Acanthopteroctetidae).
Pupal clypeus non-setose, not distinctly demar- Male accessory gland with swollen and thin-
cated. walled (non-secretory?) apical portion, adhering
All lineages in the homoneurous grade have to counterpart on other side. Larval head Fig.
(in their ground plans) retained microtrichiated 5.2 B) with dorsal ecdysial line running laterad
('aculeate') wing surfaces and distinct jugal lobes from antennal base, and only a single pair of
on the forewings. Members of the first differenti- stemmata. Legs and prolegs absent.
ated lineages (Eriocraniidae, Acanthopterocteti- Small moths, wingspan usually ca. 10 mm, al-
dae, Lophocoronidae) are all small moths (wing- ways below 15. Head capsule (Fig. 5.1 A) extend-
span nearly always less than 15 mm). Like the ing far above compound eyes. Ocelli present. La-
non glossatans they have a rough cephalic vesti- brum large, movable, with frontal retractors.
ture of hair-scales, long (5-segmented) and folded Epiphysis present or (in most taxa) absent. Fore-
maxillary palps, and they are relatively narrow- wing (Fig. 5.1 B) often with distinct pterostigma;
winged. The antennae are simple and filiform. forking of Sc and R retained in some taxa; pres-
The proboscis is devoid of an intrinsic muscula- ence of only one Rs 1 + 2 branch (also in hind-
ture. The females in these families have a charac- wing) characteristic of the dominant Palaearctic
teristic 'Eriocrania-type' piercing oviscapt (Fig. 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 53

ration. Hindwing with several distinct 'frenu- 24 species are named. The family is restricted
lum'-bristles, wing-coupling apparently non- to the Holarctic. 6 genera are currently recog-
functional (Grodnitsky & Kozlov 1985). Wing- nized, but the family is structurally and biolo-
surface scales of 'primitive-type', without 'her- gically overall homogeneous, and it is debatable
ringbone crests'. Central hindwing scales of vari- whether this degree of generic splitting is justi-
able shape, sometimes hairlike. Forewings irides- fied. Dyseriocrania, Eriocraniella and Eriocrania
cent. Orifice of sternum V gland on sculptured (Fig. 5.3 A) are the principal genera.
process. Sternum IV in females nearly always Davis (1978; principal reference), Wood (1890,
with unpigmented and un-microtrichiated 'win- 1891), Kristensen (1968), Birket-Smith & Kris-
dow' on areas to which gland reservoir adhere; tensen (1974), Kobayashi & Ando 1987; a review
similar 'windows' present in males of some taxa; of Old World eriocraniids by Karsholt, Kris-
sternum V glands altogether lost in some male tensen & Kozlov is in preparation. Like the
Eriocrania. Male segment IX (Fig. 5.1 F) a com- Micropterigidae the Eriocraniidae are treated in
plete synscleritous ring, with paired lateroventral many comparative accounts of lepidopteran
apophyses from its anterior margin accomodat- structure; see 24. A report by Koricheva (1994)
ing the origins of phallic retractors; segment X provides an entry into the ecological literature
with paired tergal lobes. Gut with short stomo- on eriocraniids, and Kozlov et al. (1996) give ref-
daeum widened posteriorly, but much narrower erences to works on their pheromone biology.
than midgut; midgut large, extending from meso-
thorax well into abdomen; 6 malpighian tubules
discharging separately into gut; 3 - 6 rectal papil- Clade COELOLEPIDA
lae. 3 - 4 abdominal ganglia. Testes small, not The clade Coelolepida (Nielsen & Kristensen
united. 4 ovarioles per ovary; colleterial gland 1996), comprising all non-eriocraniid Glossata,
relatively small, lobed or simple. is characterized primarily by the presence of
Egg oval, with thin, smooth chorion. Larva 'normal-type scales (22) on the wing surfaces,
(Fig. 5.2A) permanently leafmining, progna- and of the 'normal type' (28) of first thoracic
thous, head partially retracted within prothorax. spiracle in the adult. Ocelli are absent in all ho-
Mandible with or without bundle of barbed bris- moneurous Coelolepida as well as in nearly all
tles on inner base. Melanized plates present on non-ditrysian Heteroneura. Ocelli probably dis-
prothoracic dorsum and venter, anal shield usu- appeared in ancestral Coelolepida, and their re-
ally poorly defined; trunk with variable comple- appearance in heteroneuran lineages may be seen
ment of dorsal and ventral eversible calli. Pupa as apomorphic character reversals.
with hypertrophied, angularly bent mandibles. Three basal lineages are recognized within the
Coelolepida: The Acanthopteroctetoidea, the
Adults are usually diurnal, although nocturnal
Lophocoronoidea and the high-rank clade My-
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
Acanthopteroctetidae. Probable autapomorphies:
been proved (Kozlov et al. 1996). Eggs are in-
apices of dorsal tentorial arms fused with head
serted in the host leaves, in pockets cut by the capsule. Epiphysis absent. Tibial spurs 0-1-4 (see
female's oviscapt saw. The host range is almost Eriocraniidae). Labial palp greatly shortened.
restricted to the Fagales (Quercus and Betula be- Wings (Fig. 5.1 G) unusually narrow. M l
ing the principal host genera), but a few taxa are 'stalked' with Rs in both wing pairs; forewing
known to utilize hosts belonging to Salicales with only two A veins, both extending to wing
(120) and Rosales (Holodiscus, Sorbus). Mines margin; hindwing M-Cu crossvein vestigial,
are fulldepth blotch mines, which may or may hence 'cell open'. Sternum II with anterior rim
not start as a distinct gallery. Frass deposition separate from main plate. Sternum V gland ab-
usually highly characteristic, in thread-like pieces sent. Valve with membranous bulge on inner sur-
(sometimes very long and twined, Fig. 5.2 C). face; prominent, posteriorly produced and
Larval growth is rapidly completed; full-grown ventrally serrate transtilla present (Fig. 5.11);
larvae drop to the ground and bury into the soil, phallus (Fig. 5.1 H) with apparently eversible
where the cocoon (covered by soil particles on membranous vesica set with cornuti (a similar
the outer surface) is constructed. Most eriocrani- phallic configuration is commonplace in Hetero-
ids are early spring insects, the adult activity neura; this may be a parallelism).
period coinciding with the leafing of the host,
Moths (Fig. 5.3 B) small, a single species with
and the larvae completing their feeding stage
ca. 15 mm wingspan, others smaller. Head ex-
while the leaves are still young. A few eriocraniid
tending far above compound eyes. Labrum nar-
species occasionally occur in such large numbers
row, movable, with frontal retractors. Labial
that they attain pest levels, particularly at high palp 3-, 2- or 1-segmented. Forewing pattern
latitudes. 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 first seven abdominal segments with paired dor-
completely reduced in Acanthopteroctetes unifas- sal and ventral eversible calli (in thorax arising
cia and Catapterix; the former with two (stout) on coxal membranes). Mine a full-depth blotch
frenulum bristles, the latter, remarkably, with mine in the leaf; larva feeding in summer/autumn
only a single large bristle recalling that of typical and again in spring, leaving mine before pupa-
male Heteroneura. Male segment IX (Fig. 5.11) tion, which takes place in a cocoon on the
a synscleritous ring, anteroventral margin more ground. Pupa with hypertrophied, angularly
or less strongly concave; paired tergum X lobes bent mandibles.
small and narrow. Female genital chamber in Two genera are currently recognized, Acan-
some taxa with prominent sclerotization (extend- thopteroctetes with 4 species in western N. Amer-
ing into wall of bursa copulatrix) in front of ica and the monotypic Catapterix from Crimea.
spermathecal orifice (hence apparently nonho- Sinev & Zagulajev (1988) described the latter in
mologous with eriocraniid 'vaginal sclerite'). a family of its own, but clearly recognized its
Few data on soft anatomy available, all pertain- closest affinities to be with Acanthopteroctetes;
ing to Acanthopteroctetes unfascia: brain with indeed it is likely, that it is actually cladistically
'deutocerebral loop', and optic lobes pressed subordinate within the latter. There is an unde-
against suboesophageal ganglion (as in Lopho- scribed taxon from montane Peru.
coronidae); 3 abdominal ganglia; ventral dia- Davis (1978; principal reference), Davis
phragm very reduced; dorsal nerve-cord sheath (1987), Sinev & Zagulajev (1988), Sinev (1988),
not thickened. Gut without stomodaeal crop, Zagulajev (1990), Nielsen & Kristensen (1996).
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, In the currently best supported hypothesis of
inserted in upper epidermis. Larva (Fig. 5.2 D) coelolepidan phylogeny (Nielsen & Kristensen
prognathous, head partially concealed within 1996) the Lophocoronoidea are the sister group
prothorax; 6 stemmata, one of which with re- of the Myoglossata. Derived characters shared
duced lens; spinneret very prominent. Prothorax by the two include: frontal labrum retractors
with melanized tergal and sternal plates. Tho- lost. Pronotum with two sclerotizations, the an-
racic legs present, with indistinct coxae. Prolegs terior articulating with propleuron (parallelled in
present only on X, no crochets. Thoracic and Micropterigoidea). Vein R always unforked.
The Homoneurous Glossata 55

Mesosternum markedly produced anteromedi- Clade MYOGLOSSATA

ally. Thickened dorsal sheath of ventral nerve
cord with copious connective tissue. 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-
Lophocoronidae. Probable autapomorphies: cra-
tapomorphies include the somewhat enlarged
nium with weakly developed dorsal condyles at
median mesonotal wing process, as well as the
antennal base (in addition to commonplace
upwards bend of acrotergite I and concomitant
ventral ones, apparent parallelism with Exopo-
insertion of the indirect hind wing depressor on
ria). Mandibles completely devoid or muscula-
its anterior-facing surface (Nielsen & Kristensen
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 NEOPSEUSTOIDEA
basal setae; phallus in apical half with promi- Neopseustidae. Probable autapomorphies: facial
nent, near-tubular infolding from outer wall, and scales restricted to paired lateral, usually promi-
tergum X lobes small, set with characteristic con- nently swollen, patches. Antennae submonili-
ical sensilla. Stomodaeum with expanded crop form/subserrate, usually (always?) longer than
followed by narrow, tubular section. Only four forewing; scape with distinct condyle fitting pedi-
malpighian tubules present, each opening into cellar base; flagellar scales longitudinally aligned.
gut. Brain with 'deutocerebral loop', and optic Proboscis scaled, with a unique double-tube
lobes pressed against suboesophageal ganglion structure, the inner wall of each galea being
(similarities with Acanthopteroctetidae, presum- strongly concave and forming by itself a func-
ably parallelisms). Abdominal ganglion II tionally closed tube. Well-developed labial palp
discrete, so 5 ganglia are discernible (autapomor- without distinctly concave vom Rath's organ.
phic character reversal?). Prominent apodemal plate invaginated from up-
Small moths, wingspan 10-15 mm. Head cap- per base of propecoxal bridge. Mesofurcal arms
sule not markedly extending above compound fused into transverse bridge. Male sternum VII
eyes. (Fronto)clypeolabral boundary indistinct. with medial spinose process; 'socius' setae with
Mandibles medium-sized, simple lobes. Spurs 0- prominently swollen base.
2-4. Forewing pattern quite diverse in the few Intriguing similarities with the Heteroneura in
known species: unicolorourous fuscous, cryptic thoracic structure include presence of distinct
('bird-dropping' type, Fig. 5) or with bold black- proprecoxal bridge, transverse suture delimiting
and-white fasciae. Forewing (Fig. 5.1 J) Sc 'mesoclidium', and fusion of the latter with the
forked; A veins in 'double-Y' pattern; 'coupling' prospinasternum.
jugofrenate. Sternum V gland present in both Medium-sized moths, wingspan ca. 15
sexes, opening on sculptured process. Male seg- 27 mm. Head capsule not markedly extending
ment IX (Fig. 5.1 K) a synscleritous ring. Female above compound eyes. Labrum large and dis-
genital chamber and bursa without sclerotiza- tinctly demarcated. Mandibular lobes with well
tions. Large, delicate stomodaeal crop extending developed musculature. Maxillary palp similarly
into abdominal base; malpighian tubules each primitive, long and 5-segmented. Epiphysis
opening separately into gut. Ventral diaphragm much reduced in S. American taxa. Tibial spurs
with well developed segmental alae. 0-2-4. Wings in Nematocentropus and Synempora
Early stages unknown. Since the female has a with rather commonplace proportions and dense
piercing oviscapt type, the larva is supposed to scaling, in Neopseustis (Fig. 5.1 L) and, in partic-
be an internal feeder, and since the adult moth is ular, Apoplania wings unusually (for homo-
completely devoid of mandibular muscles (there neurous moths) broad and thinly scaled (scales
are also no apodemes for such muscles), the narrow and widely spaced). Rs (of both wing
pupa must be adecticous. The moths are noctur- pairs) 4-branched only in Nematocentropus, in
nal and mainly autumn fliers. Nearly all lopho- other genera Rsl + 2 represented by single vein.
coronid sites are in eucalypt forest, with growth Rs4 to apex. Forewing, except in Nematocentro-
of Melaleuca and usually Atriplex. Females have pus, with 'anal pocket', formed by wing mem-
been encountered much more rarely than males. brane behind curved base of 1 A covering base
The single genus Lophocorona (6 species, Fig. of Cu. Forewing A veins forming 'single Y \
5.3 C) occurs in the southern parts of West and Frenulum bristles undifferentiated. Female usu-
South Australia, with outliers in the ACT and ally with hyaline 'fenestrae' on IV, but sternum
N. S. Wales. V glands have not been identified. Genital seg-
Common (1973: benchmark paper with origi- ments in both sexes highly modified and inade-
nal description), Nielsen & Kristensen (1996 quately understood. Male (Fig. 5.1 N) segment
comprehensive account). IX ring with variably developed lateroventral
56 Niels P. Kristensen

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 (Data on soft anatomy available only for Sy-
usually (always?) fused with IX ring. 'Anellus' nempora).
with armature of processes. Phallus simple and The immature stages are unknown, but since
tubular in Nematocentropus, highly modified in mandibular muscles are strong in the adult, the
other genera; paired processes ('parameres') in pupa is believed to be decticous (and surely exa-
Neopseustis are probably phallic derivatives. rate). Adult activity has been recorded by day as
Postgenital complex with paired tergum X lobes; well as by night; feeding has not been observed.
variably shaped 'tegumenal lobes' usually pre- While Synempora show the tectiform resting pos-
sent between IX ring and tergum X, sometimes ture which is commonplace among primitive
apparently synscleritous with the former; anus moths, the broad-winged Apoplania hold the
flanked by mostly conspicuous patches of setae wings flat over the abdomen (one overlapping
with raised sockets. Testes not united in midline. the other), and in complete rest the antennae are
Female (this sex poorly known in Nematocentro- hidden below the wings (Fig. 5.2 E).
pus) postabdomen (Fig. 5.1 M) with deep pocket This small and little-known family has a pecu-
invaginated behind sternum VII. Tergum VIII liarly disjunct distribution. Nematocentropus
with short, blunt 'apophyses anteriores' from an- (= Archepiolus, 2 species), the sister group of the
terolateral corners. 'Apophyses posteriores' aris- remaining taxa, occurs in Assam, Burma and
ing from minute terminal sclerotization, which Szechuan, Neopseustis (5 species, Fig. 5.3 E)
usually bears a pair of distinct lateral teeth and ranges from Assam to Taiwan, while Synempora
which may have a rasping/piercing function. The (monobasic) and Apoplania (3 species) occur in
presence of an apparently distinct segment IX, austral. S. America.
with more or less divided tergal plate, between Davis (1975; principal reference), Davis &
the apophysis-bearing VIII and the terminal unit Nielsen 1980, 1985), Kristensen & Nielsen
on which the posterior apophyses are born, is (1981), Nielsen & Kristensen (1996), Kristensen
morphologically intriguing (see 2-4). Alimen- (in preparation).
tary canal without stomodaeal crop and with
large midgut; 6 malpighian tubules, each opening
separately into gut. Ventral nerve cord with Clade NEOLEPIDOPTERA
paired connectives between suboesophageal and
All Myoglossata apart from the Neopseustidae
prothoracic ganglia and between the 4 abdomi-
probably constitute a monophylum, Neolepidop-
nal ganglia; autapomorphic neotenic trait?
tera, characterized by adecticous, obtect pupae.
Nerve cord sheath not thickened dorsally;
In the lower neolepidopteran grades the pupae
ventral diaphragm fibres completely absent.
have backwards-pointing spines on the dorsum
The Homoneurous Glossata 57

of several abdominal segments; at eclosion this thoracic L-pinaculum fused with notai shield in
spinosity aids the movement of the pharate adult second and later instars in known hepialoids and
out of the (usually silk-lined) pupal shelter. some, but not all mnesarchaeoids (Nielsen &
Correlated with their loss of function, even in Kristensen 1989). Apomorphies paralleled in Lo-
the pharate stage, the adult mandibles (which in phocoronoidea include the presence of (some-
several neolepidopterans are retained as quite times strongly developed) dorsal cranial condyles
distinct lobes) have at most a vestigial muscula- at the antennal base, and the distinctly postapi-
ture. The complement of crochet-bearing prolegs cal Rs4.
on abdominal segments IIIVI and X in the Homologies of exoporian postabdominal
larva is another notable apomorphy ascribed to structures (in both sexes) with those of other
the ground plan of the Neolepidoptera. Addi- Lepidoptera have been difficult to establish. In
tional probable neolepidopteran autapomorphies males the prominent dorsal sclerotization pre-
include the lateral displacement of the free pro- viously identified as the tegumen most probably
furcal arm, the concealment of the upper part pertains to segment X (Nielsen & Kristensen
of the mesopleural suture below a mesepimeral 1989), and it is now referred to as the 'pseudoteg-
bulge, the disassociation of the aorta from the umen' (or 'pseudoteguminal plates', since it is
metathoracic dorsum, the general absence of usually a distinctly paired formation); conspicu-
sternum V gland, the arrangement (non-aliform) ous processes from the pseudotegumen and the
of ventral diaphragm fibres in II, the presence of tergal area just above it occur in several exopo-
a spacious stomodaeal crop; the arrangement (in rian lineages. In females the main morphological
2 groups of 3) of the malpighian tubules, the problem concerns the segmental assignment of
coiling of the spermathecal duct. Some of these the sclerite, which forms the lower lip of the cop-
traits are paralleled in one or more non-neolepi- ulatory orifice.
dopteran clades; caveats concerning their phylo- Female pheromone activity has recently been
genetic interpretation are discussed by Nielsen & recorded from the hindwing of a hepialid
Kristensen (1996). (Kuenen et al. 1994) and also from mnesarchaeid
All homoneurous Neolepidoptera belong to wings (Kozlov, pers. comm.). The nature and ex-
the clade Exoporia. act source of exoporian female pheromones re-
main to be identified.
Larvae of ancestral exoporians were probably
primarily 'soil animals' with generalized feeding
The monophyly of this clade is now well substan- habits similar to those of extant mnesarchaeids.
tiated (summary in Nielsen & Kristensen 1996), A checklist of the World Exoporia (by E. S.
primarily by the unique configuration of the fe- Nielsen, G. S. Robinson & D. L. Wagner) is
male genital apparatus (detailed account in forthcoming.
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 MNESARCHAEOIDEA
is (in the exoporian ground plan) no internal Mnesarchaeidae. Mnesarchaeids are by far the
communication between the two parts of the smallest exoporian moths, and the only ones
genital system. Sperm deposited in the bursa with functional mouthparts in the adults. They
must be transported to the spermatheca via the were long grouped with the 'eriocranioid-grade'
ovipore, the body wall below the latter being families, their close relations to the hepialoids
typically grooved one way or another. Accessory only being revealed in the 1970s. Probable auta-
(colleterial) glands lost. No apodemal apophyses pomorphies: wings narrow, lanceolate. Rsl + 2
present. Male genitalia also distinctive: phallus in both wing pairs represented by one vein only
membranous and devoid of protractor and re- (Fig. 5.4 A). Wing coupling achieved by band of
tractor muscles; segment X with transverse (su- long piliform scales on lower side of forewing
pra- and subrectal) muscles very strongly devel- dorsum entangling with correspond band of pili-
oped. Maxillary palp always small, hidden in the form scales on hindwing costa. In male genitalia
vestiture and at most 3-segmented. Wing scales anterodorsal corner of valve produced and fused
mostly with strong 'secondary ridges' (2-1). with the lateral part of the pseudoteguminal
High number (5 or 6) of abdominal ganglia in plate (which in turn is indistinctly separated
adult probably an autapomorphic character re- from the counterpart of the hepialoid trulleum).
versal. Egg shell structure and embryogenesis Elongate and apically bifid 'dorsal plate' on fe-
distinctive (2-18, 2-19); eggs turn black after male postabdominal apex comprising fused terga
being deposited. The 'epicranial notch' of the of VIII and following segments; anterolateral
larval head is unusually small, and there are corner of dorsal plate more or less extensively
several potential autapomorphies in larval chae- fused with elongate subgenital plate (the latter
totaxy, including e. g., the absence of the thoracic forms the lower lip of the copulatory orifice and
MXD1 and the approximation of the meso-/ presumably at least partly pertains to segment
metathoracic MD1 to the two MSD setae; pro- 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 have occasionally been observed, but not as a
tracts). Pupa (Fig. 5.2 F) on mesothorax and ab- stable condition in any species.
dominal segments I - V I I I with mid-dorsal Egg relatively very large, with smooth
spines only. chorion. Larval facies characteristic, with very
Antennae filiform, fully scaled, in one species long spinneret, anteriorly tapering thorax, and
group with 3-5-branched trichoid sensilla. La- slender prolegs with biserial crochets. Pupa with
brum very small (in anterior view practically hid- cephalic cocoon-cutter.
den by bulging frontoclypeus). Maxillary palp 3- Adults mainly diurnal, but have occasionally
segmented, or second and third segments more been attracted to light. Males fly frequently
or less completely fused; proboscis (Fig. 5.4 B) around low vegetation, while females are rarely
well developed, coilable. Labial palp 3-seg- encountered by day; adults not flower-frequent-
mented; apical segment with sensilla cluster at ing. Eggs are attached singly to moss or liverwort
most slightly depressed. Protibial epiphysis vari- leaves. Larvae live in silken sheets and galleries
ably developed, occasionally absent. Spurs 0-2-4. spun in liverwort/moss periphyton on moist soil,
Forewing in one species group with Ml stalked logs etc., feeding on leaves and rhizoids, fern
with stem of Rs3 + 4; only one A vein present. sporangia, fungal hyphae and other detritus. Pu-
Sc branching and presence of inter-M crossvein pation in an oval, debris-covered cocoon.
The Homoneurous Glossata 59

14 mnesarchaeid species are known (half of ruled out that it pertains to VIII (as the mnes-
them still unnamed), all included in the genus archaeid subgenital plate apparently does) or is
Mnesarchaea (Fig. 5.3 D). The family is endemic a composite formation. Occasionally there is a
to New Zealand. distinct sclerotization between sternum VII and
Gibbs (1979); comprehensive treatment by the antevaginal lamella. Primitively narrow and
Gibbs & Kristensen in progress. paired 'subanal plates' (pertaining to X or IX?)
flank the grooved (or otherwise modified) 'inter-
genital field' between ostium bursae and ovipore;
HEPIALOIDEA. their anterior/lateral corners may be closely ap-
The Hepialoidea are the first differentiated lepi- proximated to, or fused with, the anterolateral
dopteran lineage in which the number of extant corners of the dorsal plate.
species runs into several hundreds; they are also Male scent organs on wings and hindlegs have
the first differentiated lineage which includes evolved independently on a number of occasions
truly large and robust taxa (indeed some of the in hepialoids. It is noteworthy that male lekking,
largest moths belong here). A long-recognized an unusual behavioral trait in the Lepidoptera,
hepialoid autapomorphy is the strong regression may also be evolved on more than one occasion
of the proboscis, which is at most as long as the within the superfamily: in Palaeosetidae-Ogj-
head capsule (usually much shorter, or com- gioses (Davis et al. 1995) and at least once within
pletely absent), never truly coilable and probably the Hepialidae s. str. (Wagner & Rosovsky 1991).
always non-functional; the food canal walls are Or will the lekking 'palaeosetids' actually
without discrete plates. Other autapomorphies prove to be cladistcally subordinate within the
include: intercalary sclerite between scapus and latter?
pedicellus elongate, partly lowered into pocket in Larvae of most of the basal hepialoid lineages
intersegmental membrane (primitive, i. e., small are unknown, so diagnostic traits of the super-
and superficial, sclerite in some palaeosetids now family remain unidentified. Known pupae (Fig.
interpreted as autapomorphic character rever- 5.2 H) with dorsal abdominal spine rows basi-
sal). Posterior Rs fork backwards displaced, Rs3 cally similar to those of the lower Heteroneura;
reaching termen, not costa (Figs. 5.4 D, I). Meta- this presumably represents the neolepidopteran
furca without anterior process, sterno-trochan- ground plan condition.
teral muscle originating directly from (anteriorly The Hepialoidea are currently classified into
tilted and convex) furcal stem. Male genitalia five families, of which three are monogeneric and
with characteristically hinged 'juxta + trulleum' the remaining two without unambiguous auta-
sclerites (at least partly homologous with 'me- pomorphies. A total of 12 terminal taxa (species
dian plate' in lepidopteran ground plan, see and assumedly monophyletic supraspecific as-
24) below phallocrypt entrance (Fig. 5.4 F). semblages) can at present be recognized within
Antennae structurally diverse; a marked short- the superfamily, and a re-analysis of their inter-
ening is commonplace, but can at most be an relationships, based on an extensive character
'underlying synapomorphy' of the members of set, is much needed. While Scoble's (1992) inclu-
the superfamily. The same is probably true of the sion of all 12 in a single family may have much
sensillum-bearing 'nipples' on the vertex and to recommend it, the time-honoured family cate-
pronotum found in a variety of hepialoids. While gories are retained here, pending results from an
the maxillary palp in the Prototheoridae (Fig. analysis of the said kind.
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- Family key
duced (with indistinct segmentation) or com-
pletely lost in other hepialoids. Hairs on hind- 1 Inter-M crossvein (between M3 and M2 or
wing front margin soft, with no structural differ- stem of M1 + M2) present, at least in fore-
entiation between 'frenular' and 'costal' hairs wing (Fig. 5.4 D) 2
(sensu Braun 1924, see 2-4). Pro tibial epiphysis Inter-M crossvein absent (Fig. 5.41) . . . .
and tibial spurs variably developed. Abdominal Palaeosetidae
venter I usually (but according to Brner 1938 2(1) Metatibia almost always without spurs,
not always) membranous. Female postabdomen very rarely 1 or 2 p r e s e n t . . . . Hepialidae
(Figs. 5.4 H, J) with tergum VIII separate from Metatibia with 4 spurs 3
'dorsal plate', the latter posteriorly more or less 3 (2) Entire wing surfaces with microtrichia
deeply indented (in Neotheoridae apparently ('aculei'). All wing scales of 'normal type'
completely divided, but confirmation of this is (hollow, with internal trabeculae and per-
desirable). Distinct sternum VII sometimes pre- forated upper lamella). Forewing with anal
sent, but ventral surface often extensively unscle- loop. S. African Prototheoridae
rotized behind VI. The sclerotized 'antevaginal Wings microtrichiated at base only. Wings
lamella' anteriorly bordering the ostium bursae with lower layer of 'primitive type' wing
probably belongs to IX; however, it cannot be scales (devoid of perforations and trabecu-
60 Niels P. Kristensen

lated lumen). Forewing without anal loop each galea densely spined. Labial palp long, por-
4 rect and densely scaled. Wings entirely micro-
4 (3) Labial palps long and snout-like, pro- trichiated; Rs3 + 4 'stalked' in both wings; fore-
jecting far beyond antennal pedicellus; api- wing anal loop present (Fig. 5.4 D). Protibial
cal segment longer than two basal seg- epiphysis well developed. Spurs 0-2-4. Hind tibia
ments together. Protibial epiphysis present. unusually long, more than twice femoral length.
Neotropical Neotheoridae Eggs and larvae undescribed; a pupa has been
Labial palps shorter, not projecting beyond incidentally found "in moss"; it had no kind of
antennal pedicellus; middle segment equal- silken enclosure. The moths are nocturnal. Fe-
ling basal and distal segments together. male specimens are rarely observed, their flight
Protibial epiphysis absent. Australian . . . activity is likely to differ markedly from that of
Anomosetidae males.
The nine described species are now all in-
Palaeosetidae. It is debatable whether the diag- cluded in the genus Prototheora (Fig. 5.31), re-
nostic lack of an inter-M crossvein (Fig. 5.41) is stricted to Southernmost Africa (Cape Floristic
apomorphic or not. Moths unusually small (wing Region and Afromontane forests, northwards
span no more than ca. 12.5 mm in some Ogy- to Natal).
gioses), slender-bodied and broad-winged for Davis (1996; exhaustive account of available
hepialoids, Palaeoses itself least pronouncedly knowledge.)
so. Maxillary appendages completely reduced (as
in most Hepialidae).
Neotheoridae. Monobasic. Neotheora chiloides
Protibial epiphysis absent. Spurs 0-1-2 (Fig. 5.3 J) is known from a single, incomplete
(Osrhoes), 0-1-1 (Genustes), otherwise 0-0-0. female specimen. It is unique among exoporians
Male metatibia in Osrhoes, Genustes and (Tai-
in having several (9) round and crested signa in
wanese) Ogygioses thickened, in two latter gen-
the bursa copulatrix, and the antevaginal lamella
era with hair-pencil (surely a scent apparatus) in
preceded by an asymmetrical sclerite with two
a groove. Male Ogygioses with hair pencil in
posterior processes.
folded expansion of hindwing anal area. 'Primi-
Maxillary palp short, with indistinct segmen-
tive type' wing scales absent from the genera so
far examined for this trait (Osrhoes, Ogygioses). tation; proboscis vestige distinct, medial galea
surface more distinctly concave than in other
Osrhoes is notable for its unusually reduced
hepialoids examined, somewhat spinose. Labial
dorsal tentorial arms and, in particular, for its
palp long, porrect and densely scaled. Protibial
unique female genital apparatus (Fig. 5.4J): a
epiphysis present; metatibia just above twice
long, narrow 'ductus seminalis' (presumably a
femoral length, with 4 spurs. Wings devoid of
functional analogue to the similarly termed duct
microtrichia (forewing base excepted), with ter-
in Ditrysia) extends backwards from the ductus
men slightly convex, apices produced; vestiture
bursae to an opening adjacent to the ovipore.
with lower layer of 'primitive type' scales:
Information on biology and immatures avail-
Rs3 + 4 'sessile'; forewing without anal loop. Ap-
able only for Ogygioses: egg with prominent tu-
parent tergum VIII narrow, anterolateral corners
bercles on chorion. 1st instar larva (only instar
produced. 'Dorsal plate' probably completely
known) with only 3 stemmata. Slender prolegs
with crochets in a uniordinal circle. Adult moths
diurnal, males exhibiting lekking behaviour. Brazil (Matto Grosso).
Four genera with a total of 8 species are in- Kristensen (1978 b.)
cluded in the Palaeosetidae: Osrhoes (Colombia,
monobasic; Fig. 5.3 G), Palaeoses (Queensland, Anomosetidae. Monobasic. Anomoses hylecoetes
2 species, one named; Fig. 5.3 F), Genustes (As- (Fig. 5.3 H) is distinctive by the enormously de-
sam, monobasic) and Ogygioses (Taiwan, Thai- veloped male trulleum and a 'chambered' roof
land, 4 species). of the female genital sinus hidden by the large
Issiki & Stringer (1932), Common (1990), antevaginal lamella.
Kristensen & Nielsen (1994), Davis et al. (1995), Maxillary palp short, with indistinct segmen-
Heppner et al. (1995). tation; proboscis vestige distinct, medial galea
surface densely spinose. Labial palp not conspic-
Prototheoridae. Probable autapomorphies: geni- uously long. Protibial epiphysis absent; hind
talia with unique apparent copulation locking tibia only about 1.5 times femoral length; spurs
device, comprising a median 'conjugal process' 0-2-4. Wings devoid of microtrichia (bases ex-
on the female antevaginal lamella and a corre- cepted); vestiture with lower layer of 'primitive
sponding male 'conjugal pouch', the roof of type' scales; Rs3 + 4 variable, 'sessile or stalked;
which is formed by the arched ventral trulleum forewing without anal loop; male hind wing with
wall. 'Primitive type' wing scales absent. hair-pencil concealed in folded posterior margin.
Maxilla (Fig. 5.4 E) with 3-segmented palp Immature stages unknown. The adult moth is
and distinct proboscis vestige; medial surface of 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 Fig. 5.5. Hepialidae: A, Hepialus humuli, male (the
and apparently of taxonomic importance. Fe- 'ghost moth' par excellence, the name refers to the ag-
male 'intergenital lobes' (strengthened by sub- gregations of pendulating white moths seen at dusk);
anal plates) flanking groove between ovipore B, Sthenopis argenleomaculatus; C, Zelotypia stacyi.
and ostium bursae, in some taxa fused medially, Scale 50 mm.
thereby forming an internal tubular sperm pas-
sage; this variability and its phylogenetic implica-
tions deserving further scrutiny. and the only hepialids 5. lat. with sizable probos-
Larvae with development of melanized pina- cis vestiges; known larvae grass-feeders. Afro-
cula variable between taxa, sometimes also theora and Antihepialus (8 and 4 named species,
markedly variable between instars of the same respectively) are Afrotropical; their immatures
species. Proleg crochets uni-, bi- or multiserial, undescribed. Gazoryctra (Holarctic, 13 species)
the latter arrangement (with crochet size increas- have meso- and metatibial spurs; like the Hepia-
ing towards centre) probably representing the lidae s. str., but unlike Fraus their pupae have
ground plan condition in mature larve. two dorsal spine rows (instead of a single) on
Adults crepuscular/nocturnal; mass emer- abdominal segments IIIVII.
gences stimulated by rainfall being of regular oc- Remaining hepialid genera, 'Hepialidae s. str. '
currence in some taxa. Mating behaviour diverse probably constituting a monophylum, as indi-
(see below). Egg numbers may be remarkably cated by specialized male genitalia (trulleum
high, > 30000 in larger species; in some taxa fused with pseudoteguminal plates), reduction of
they are broadcast by the females in flight. Many pseudempodial seta, and presence of a ventral
larvae tunnel in soil, or construct silken galleries spine row on pupal segment VII. Medium-sized
among litter, and feed on roots (or, at night, on to very large, robust moths. Pretarsal arolium
leaves adjacent to retreat) of pteridophytes, gym- and pulvilli sometimes greatly reduced. Ca. 500
nosperms and angiosperms; others are root/stem/ named species, currently in 50 + genera; cosmo-
branch borers, some feeding exclusively on callus politan. Previous subfamily classifications con-
tissue. Fungivory is probably widespread, the sidered untenable (Nielsen & Robinson 1983).
transition from fungivory to phytophagy during An assemblage characterized by androconial
larval ontogeny being recorded in a number of scales on the male metatibia may prove mono-
cases. phyletic or at least have a monophyletic core;
Four genera of medium-sized hepialids, re- well-known genera include Oncopera, Hepialus
ferred to by convenience term 'primitive Hepiali- (Fig. 5.5 A), Zenophassus, Sthenopis (Fig. 5.5 B,
dae', are devoid of specializations shared by Nearctic, 4 species), Phymatopus (with metatibia
other members of the family. Fraus (Australian, club-like, and metatarsi lost, Fig. 5.5), and the
25 structurally diverse species) include some spe- probably closely related Aenetus (with many
cies with almost conventional 'micro-moth fa- large and colourful species) and Endoclita whose
late-instar larvae are wood-borers, sometimes
cies' (slender bodies, relatively long antennae)
62 Niels P. Kristensen

pests. While female pheromone-calling m a y still new superfamily (Lepidoptera: Neopseustoidea). -

occur in s o m e Oncopera, calling is 'reversed' else- Smithson. Contr. Zool. 210: i - i i i , 1 - 4 5 .
where, with male territoriality and 'pendulating' - (1978): A revision of the North American moths of
lek flights being d o c u m e n t e d in s o m e taxa. the superfamily Eriocranioidea with the proposal of
a new family Acanthopteroctetidae (Lepidoptera).
A m o n g the largest and/or m o s t striking hepialids
- Smithson. Contr. Zool. 251: i - i i i , 1 - 1 3 1 .
(indeed m o t h s ) are the Australian Zelotypia
- (1987): Eriocraniidae, Acanthopteroctetidae. Pp.
stacyi (Fig. 5.5 C; w i n g s p a n u p t o 25 cm), the S.
3 4 4 - 3 4 7 in Stehr, F. (ed.) Immature Insects, Ken-
African Leto venus, and the N e o t r o p i c a l Tricho- dall/Hunt, Dubuque - (1996): A revision of the
phassus giganteus, all with w o o d - b o r i n g larvae. southern African family Prototheoridae (Lepidop-
While the t w o first-mentioned taxa probably are tera: Hepialoidea). - Eni. scand. 27: 3 9 3 - 4 3 9 .
closely related to the EndoclitalAenetus lineage , Karsholt, O., Kristensen, N. P. & Nielsen, E. S.
the latter appears isolated. Important pasture (1995): Revision of the genus Ogygioses (Palaeoseti-
pests occur in, e. g., Gorgopis, Dalaca, Oncopera, dae). - Invertebr. Taxon. 9: 1231-1263.
Oxycanus and Wiseana. Other m a j o r and/or - & Nielsen, E. S. (1980): Description of a new genus
wellstudied genera include, e. g., Bipectilus, Eu- and two new species of Neopseustidae from South
dalaca, Callipielus, Phassus, Aepytus, Aoraia, Tri- America, with discussion of phylogeny and biologi-
cal observations (Lepidoptera: Neopseustoidea). -
odia, Korscheltellus, Pharmacis, Thitarodes, Tri-
Steenstrupia 6: 2 5 3 - 2 8 9 .
ctena and Abantiades.
- (1985): The South American neopseustid genus
Hepialids are treated in m a n y regional hand- Apoplania Davis: a new species, distribution records
b o o k s o n 'macro-moths', including v o l s 2, 6, 10, and notes on adult behaviour (Lepidoptera: Neo-
14 o f Die Groschmetterlinge der ErdelThe pseustina). - Ent. scand. 15: 4 9 7 - 5 0 9 .
Macrolepidoptera of the World/Les Macrolpi- Dugdale, J. S. (1994): Hepialidae (Insecta: Lepidop-
doptres du Globe (A. Seitz ed.). Recent w o r k s tera). - Fauna of New Zealand 30, 163 pp.
providing references t o the v o l u m i n o u s previous Gibbs, G. W. (1979): Some notes on the biology and
literature o n the m o r p h o l o g y , b i o l o g y and taxon- status of the Mnesarchaeidae (Lepidoptera). -
o m y o f the family include R o b i n s o n & Nielsen N. Z. Ent. 7: 2 - 9 .
(1983), G r e h a n (1989), Mallet (1984), Wagner - (1989): Local or global? Biogeography of some
(1987, 1988), Wagner et al. (1989), Nielsen & primitive Lepidoptera in New Zealand. N. Z. J.
Zool. 16: 6 8 9 - 6 9 8 .
Scoble (1986), Nielsen (1988), Nielsen & Kris-
Grehan, J. R. (1989): Larval feeding habits of the Hep-
tensen (1989: principal account o f a n a t o m y ) ,
ialidae (Lepidoptera). - J. Nat. Hist. 23: 8 0 3 - 8 2 4 .
C o m m o n (1990), Wagner & R o s o v s k y (1991),
Grodnitsky, D. L. & Kozlov, M. V. (1985): Functional
Leonard et al. (1992) and D u g d a l e (1994: exten- morphology of the wing apparatus and flight char-
sive treatment centred o n rich N e w Zealand acteristics of primitive moths (Lepidoptera: Micro-
fauna). pterigidae, Eriocraniidae). - Zool. Zh. 64: 1 6 6 1 -
1671 [In Russian].
Heppner, J. B., Balczar-L., . . & Wang, H. Y.
Acknowledgments (1995): Larval morphology of Ogygioses caliginosa
from Taiwan. Tropical Lepidoptera 6: 149154.
I a m indebted to Drs. D . R . D a v i s , G . W. Gibbs,
Hinton, H. E. (1946a): On the homology and nomen-
M . V. K o z l o v and E. S. Nielsen for i n f o r m a t i o n
clature of the setae of lepidopterous larvae, with
and illustrations and t o Drs. G . S. R o b i n s o n and
some notes on the phylogeny of the Lepidoptera.
M . J. Scoble for c o m m e n t s o n parts o f the m a n u - - Trans. R. Ent. Soc. Lond. 97: 1 - 3 7 .
script. Issiki, S. & Stringer, H. (1932): On new oriental genera
and species of the Hepialoidea (Lepidoptera Homo-
neura) I - I I . - Sty lops 1: 7 1 - 7 2 , 7 3 - 8 0 .
References Kobayashi, Y. & Ando, H. (1987): Early embryonic
Birket-Smith, S. J. R. & Kristensen, N. P. (1974): The development and external features of developing
skeletomuscular anatomy of the genital segments of embryos in the primitive moth, Eriocrania sp. (Lep-
male Eriocrania (Insecta, Lepidoptera). Z. idoptera, Eriocraniidae). Arthropod. Embryol.
Morph. Tiere 77: 157-174. Soc. Jpn. 1987: 159-180.
Braun, A. F. (1924): The frenulum and its retinaculum - & Gibbs, G. W. (1995): The early embryonic devel-
in the Lepidoptera. - Ann. ent. Soc. Am. 17: opment of the mnesarchaeid moth Mnesarchaea fu-
234-256. silella Walker (Lepidoptera: Mnesarchaeidae) and
Cerf, F. le (1926): Contribution a l'etude des organes its phylogenetic significance. - Aust. J. Zool 43:
sensoriels des Lpidoptres. Pp. 1 3 3 - 1 5 8 in Leche- 479-488.
valier, P. (ed.) - Lepidoptera I, 3. Paris. Koricheva, J. (1994): Air pollution and Eriocrania min-
Common, I. F. B. (1973): A new family of Dacnony- ers: Observed interactions and possible mecha-
pha (Lepidoptera) based on three new species from nisms. - Rep. Dept. Biol., University of Turku 44.
Southern Australia, with notes on the Agathiphagi- Kozlov, M. V., Zhu, J., Philipp, P., Francke, W., Zver-
dae. - J. Aust. ent. Soc. 12: 1 1 - 2 3 . eva, E. L., Hansson, B. S. & Lfstedt, C. (1996):
- (1990): Moths of Australia, vi + 535 pp., 32 pis. Pheromone specificity in Eriocrania semipurpurella
E. J. Brill, Leiden. (Stephens) and E. sangii (Wood) (Lepidoptera: Eri-
Davis, D. R. (1975): Systematics and zoogeography of ocraniidae) based on chirality of semiochemicals. -
the family Neopseustidae with the proposal of a J Chem. Ecol. 22: 4 3 1 - 4 5 4 .
The Homoneurous Glossata 63

Kristensen, . P. (1968): The anatomy of the head and Nielsen, E. S. (1988): The peculiar Asian ghost moth
the alimentary canal of adult Eriocraniidae. - Ent. genus Bipectilus Chu & Wang: taxonomy and sys-
Meddr 36: 2 3 9 - 3 1 5 . tematic position (Lepidoptera: Hepialidae s. str.). -
- (1978): Ridge dimorphism and second-order ridges Syst. Ent. 13: 171-195.
on wing scales in Lepidoptera: Exoporia. - Int. J. - & Kristensen, N. P. (1989): Primitive Ghost Moths.
Insect. Morph. Embryol. 7: 297-299. Monographs on Australian Lepidoptera. 1, 206 pp.
- (1978 a): Observations on Anomoses hylecoetes (An- - (1996): The Australian moth family Lophocoronide
omosetidae), with a key to the hepialoid families and the basal phylogeny of the Lepidoptera -
(Insecta, Lepidoptera). - Steenstrupia 5: 1 - 1 9 . Glossata. - Invertebr. Taxon. 10: 1199-1302.
- (1978 b): A new family of Hepialoidea from South - & Robinson, G. S. (1983): Gost Moths of southern
America, with remarks on the phylogeny of the sub-
South America (Lepidoptera: Hepialidae). - Ento-
order Exoporia (Lepidoptera). - Ent. Germ. 4:
monograph 4, 192 pp.
- & Scoble, M. J. (1986): Afrotheora, a new genus of
- (in preparation). Rediscovery of Nematocentropus
primitive Hepialidae from Africa (Lepidoptera:
omeiensis, with a reassessement of the ground plan
Hepialoidea). - Ent. scand. 17: 2954.
of the Neopseustidae (Lepidoptera).
- & Nielsen, E. S. (1980): The ventral diaphragm of Sattler, . (1991): A review of wing reduction in Lepi-
primitive (nonditrysian) Lepidoptera. A morpho- doptera. Bull. Br. Mus. nat. Hist. (Ent.) 60:
logical and phylogenetic study. Z. zool. Syst. 243-288.
EvolForsch. 18: 123-146. Scoble, M. J. (1992): The Lepidoptera: Form, Function
- (1981a): Abdominal nerve cord configuration in and Diversity. 352 pp. Oxford University Press/
adult non-ditrysian Lepidoptera. Int. J. Insect Natural History Museum, Oxford.
Morph. Embryol. 10: 8 9 - 9 1 . Sinev, S. Y. (1988): Systematic Position of the Catap-
- (1981 b): Intrinsic proboscis musculature in nondi- terigidae (Lepidoptera) and the Problem of the Nat-
trysian Lepidoptera-Glossata: Structure and phylo- uralness of the G r o u p Heteroneura. - Ent. Obozr.
genetic significance. In Cederholm, L. (ed.): Ad- 67: 6 0 2 - 6 1 4 . [In Russian; English translation Ent.
vances in insect systematics and phylogeny. Ent. Rev. 69: 1 - 1 4 , 1990]
scand. Suppl. 15: 2 9 9 - 3 0 4 . Wagner, D. L. (1987): Hepialidae (Hepialoidea).
- ( 1 9 8 1 c ) : Double-tube proboscis configuration in Pp. 3 4 7 - 3 4 9 in Stehr, F. (ed.) Immature Insects,
neopseustid moths (Lepidoptera: Neopseustidae). Kendall/Hunt, Dubuque. & Rosovsky, J. 1991.
- Int. J. Insect Morph. Embryol. 10: 4 8 3 - 4 8 6 . Mating systems in primitive Lepidoptera, with em-
- (1994): Osrhoes coronta, the New World palaeosetid phasis on the reproductive behaviour of Korschel-
moth: A reappraisal, with description of a new type tellus gracilis (Hepialidae). Zool. J. Linn. Soc.
of female genital apparatus (Lepidoptera, Exopo- 102: 2 7 7 - 3 0 3 .
ria). - Ent. scand. 24: 3 9 1 - 4 0 6 .
Wood, J. H. (1890): Micropteryx larvae. - Ent. month.
Kuenen, L. P. S Wagner, D. L., Wallner, W. E. & Mag. Second series 1: 1 - 6 .
Card, R. T. (1994): female sex pheromone in
- (1891): On oviposition, and the ovipositor, in cer-
Korscheltellus gracilis (Grote) (Lepidoptera: Hepia-
tain Lepidoptera. Ent. month. Mag. Second series
lidae). - Can. Ent. 126: 3 1 - 4 1 .
2: 175-185, 2 5 3 - 2 5 8 .
Leonard, J. G Grehan, J. R. & Parker, B. L. (1992):
Zagulajev, A. K. (1992): New and little known Micro-
First instar description of Korscheltellus gracilis
(Grote) and Sthenopis auratus (Grote) (Lepidop- lepidoptera (Lepidoptera: Incurvariidae, Tineidae,
tera: Hepialidae) with a consideration of cladistic Psychidae, Alucitidae) of the fauna of the USSR.
relationships between setae. - J. New York Ento- V. Ent. Obozr. 71: 1 0 5 - 1 2 0 [In Russian],
mol. Soc. 100: 594-614. - & Sinev (1988): Catapterigidae fam. n. - a new
Mallet, J. (1984): Sex roles in the ghost moth Hepialus family of lower Lepidoptera (Lepidoptera, Dacno-
humuli (L.) and a review of mating in the Hepiali- nypha). - Ent. Obozr. 67: 593-601. [In Russian;
dae (Lepidoptera). Zool. J. Linn. Soc. 79: 6 7 - 8 2 . English translation - Ent. Rev. 68: 3 5 - 4 3 , 1989],
6. The Monotrysian Heteroneura
Donald R. Davis

Including more than 99% of the extant species of a feature of the glossatan ground plan (Nielsen
Lepidoptera (Nielsen 1985, Heppner 1991), the 1985 b, Davis 1986), supports that Incurvari-
infraorder Heteroneura overwhelmingly consti- oidea be considered the most basal of the mono-
tutes the most successful clade within the order. trysian superfamilies. Preliminary studies involv-
The monotrysian Heteroneura, a questionably ing the 18S rDNA gene tentatively support this
monophyletic assemblage of four distantly re- view with the tendancy to position prodoxine
lated superfamilies, are believed to represent the genera basal to all Heteroneura (Wiegmann
basal group of the infraorder, as indicated by the 1994). In contrast, the basis for considering the
retention of a primitive monotrysian repro- vestigial jugum in Nepticulidae as indication of
ductive system (Figs. 6.8 G, LN, Dugdale this taxon's basal position (Hennig 1953, Minet
1974) and more generalized distribution of wing 1984) has been overemphasized (Davis 1989). Al-
microtrichia. Understanding the phylogeny of though sequence data from 18S rDNA suggest
this archaic group is important because one of the Incurvarioidea to be paraphyletic, our cur-
its components probably is the sister group of rent knowledge based on morphology strongly
most of the remaining Lepidoptera (Nielsen supports its monophyly (Nielsen and Davis
1985 b). Major synapomorphies shared with the 1985). Resolution of phylogenetic relationships
ditrysian Heteroneura include the reduction of at this level utilizing the above, very conservative
the hindwing Rs system (i. e., heteroneurous ve- gene was often shown to be inconclusive. Data
nation), the frenulum-retinaculum wing cou- from genes with higher levels of sequence varia-
pling, and loss of the first abdominal sternum tion are needed to test hypotheses on heteroneu-
(Fig. 6.7, Kristensen 1984). Monophyly of the ran relationships (Wiegmann 1994). Hypothe-
monotrysian Heteroneura (or Monotrysia) is in- sized morphological relationships between Nep-
adequately supported, with only two possible sy- ticuloidea and Tischerioidea have been estab-
napomorphies proposed (Davis 1986): (1) pres- lished only on the basis of reduction characters
ence of spine combs (pectinifers, Figs. 6.8 B, D) in the adult (loss of M2) and larva (depressed
on the male valva (absent in Tischerioidea and body, stemmatal reduction) - the latter often as-
Palaephatoidea), and (2) metafurcasternum with sociated with a leaf-mining habit (Davis 1986,
two pairs of dorsal tendons and without second- 1989). Palaephatoidea and the highly apomor-
ary arm lamellae (Figs. 6.6 I -U). Unfortunately, phic Tischerioidea are suprisingly linked by two
the metafurcasterna of most ditrysian genera re- morphological synapomorphies the presence
main too poorly known for adequate evaluation, of a median ovipositor lobe (Figs. 6.9 GH) and
and it cannot be ruled out that the monotrysian enlarged, closely set female frenular setae (Davis
configuration represents the groundplan state in 1989, Nielsen 1989). The tineid-like Palaepha-
the Heteroneura. The subdivision of sternum 2 toidea had been suggested as a possible sister
into a smaller anterior sclerite (S2a) and a much group to the ditrysian Heteroneura (Davis 1986),
larger caudal region (S2b, see Fig. 6.7) may con- but Wiegmann's (1994) molecular studies more
stitute another synapomorphy of the monotry- strongly supported the monophyly of Tischeri-
sian Heteroneura, although considerable varia- oidea + Ditrysia, as proposed by Nielsen
tion in this condition has developed within the (1985 b). See also 1 - 2 .
non-incurvarioid Monotrysia (Figs. 6.7 N - R ,
Kristensen and Nielsen 1980, Davis 1986, 1989).
The four superfamilies comprising the mono- Heteroneura - Monotrysia
trysian Heteroneura have been partitioned into
two subgroups (Davis 1986): (1) Incurvariina Key to the Superfamilies and Families
represented only by Incurvarioidea, charac-
terized by an extensible, piercing ovipositor with 1 Female with elongate piercing ovipositor
its associated tendons for inserting eggs into (Figs. 6.8 G - K ) . Male genitalia usually with
plant tissue (Figs. 6.8 G - K ) , and an elongate elongate, sagittate, or deeply furcate juxta
cloaca terminating on or near the apex of the (Figs. 6.8 C, F, Incurvarioidea) 2
tenth abdominal segment (AIO, Fig. 6.8 G); and Ovipositor short, non-piercing. Juxta absent
(2) Nepticulina - including Nepticuloidea, or variably shaped, never sagittate nor
Tischerioidea, and Palaephatoidea, all with deeply furcate 7
greatly shortened, nonpiercing ovipositors and a 2 Hindwing with discal cell open (M-CuA
separate ovipore opening on A8A9 (Figs. 6.8 crossvein absent, (Figs. 6.5 A B)
L - N ) . Acceptance of the piercing ovipositor as 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; of first (Fig. 6.3 M, Tischerioidea)
A36 with biserial crochets Crinopterygidae Tischeriidae
Male pectinifer multispined or absent. Late - Hindwings broader, width 0.33 the length or
instar larvae usually exophytic; A 3 - 6 with more; third antennal segment less than
uniserial crochets Incurvariidae length of first (Palaephatoidea)
6 Adult with haustellum usually reduced, less Palaephatidae
than first labial palpal segment in length, or
absent; maxillary palpy short, 3-segmented
to absent. Frenulum-retinaculum absent. NEPTICULOIDEA.
Larva cecidogenic Cecidosidae Includes the smallest Lepidoptera; length of
Adult with haustellum developed; maxillary forewing 1.5-8.3 mm. Vertex of head rough,
palpi elongate, 4-5-segmented. Frenulum-ret- with erect piliform scales. Ocelli and chaetosem-
inaculum present. Larva usually boring in ata absent. Antennae with scape typically ex-
fruits, leaves or stems, rarely cecidogenic . . panded to form large eyecap (Figs. 6.3 J - K ) ; fla-
Prodoxidae gellum, with specialized, often branched sensilla
7 Discal cell open in forewing; antennal scape (Figs. 6.9 DE). Maxillary palpi 5-segmented,
usually greatly broadened, covering eye folded. Haustellum naked, short, 0.75 1.5 the
(Nepticuloidea, Figs. 6.3 J - K ) 8 length of labial palpi. Prosternum reduced
Discal cell closed in forewing; scape unmodi- (Davis 1986). Wings with reduced venation; dis-
fied, never forming eyecap 9 cal cell usually absent in both wings or reduced
8 Forewing with branched or stalked, often ar- in forewing; hindwing with (Nepticulidae) or
cuate veins (Fig. 6.5 I) Nepticulidae without (Opostegidae) frenulum; a dense row
Forewing with all veins separate and rela- of elongate subcostal pseudofrenular setae (Fig.
tively straight (Fig. 6.5 J) . . . . Opostegidae 6.9 C) usually present in hindwing of both sexes;
9 Hindwings slender; width 0.23 the length or microtrichia scattered over all wing surfaces in
less; third antennal segment 2 . 5 - 3 X length Nepticulidae, or reduced in Opostegidae. Legs
68 Donald R. Davis

with tibial spur pattern of 0-2-4; epiphysis ab- sometimes completely fused to tegumen or ab-
sent; hindtibia with prominent spinose scales. sent. Tegumen a narrow to broad, dorsal band.
Abdomen with S2a separate in Nepticulidae (Fig. Vinculum usually well developed, elongate,
6.7 N), fused to lateral ridges (venulae) in Opos- either U-shaped or like an inverted U (with lat-
tegidae (Fig. 6.7 P); tergosternal connection usu- eral arms), less commonly V-shaped or truncate.
ally absent, preserved in some Nepticulidae Gnathos a well developed U-to -shaped bridge
(Nieukerken 1986). Ovipositor extremely short, with or without a median caudal process, rarely
non-piercing, largely enclosed ventrally by en- absent. Anellus membranous. Juxta sometimes
larged S7 (Fig. 6.8 L). Larva apodal leaf or stem present as a lightly sclerotized, broad plate. Val-
miners, with a single pair of stemmata. Pupa vae relatively large and broad, often tapering to
with all coxae exposed and tergal spines on a narrow, distal process or apex, occasionally
A 2 - 8 ; pupa partially extruded from oval, lentic- with slender lobes arising from costal or mesal
ular cocoon (Fig. 6.11 E) prior to eclosion. surfaces; an elongate, sessile pectinifer present in
many species of Pectinivalva (Fig. 6.8 D).
Nepticulidae (Fig. 6.2 BD). Very small moths, Transtilla either present or absent, usually well
with forewings 1.54.6 mm in length. Head (Fig. developed. Aedoeagus a relatively broad, elon-
6.3 K): Vertex rough; lower portion of frons and gate tube usually heavily armed with numerous
clypeus naked; occipital scales either relatively spinose internally as well as various spinose scle-
broad or piliform. Ocelli absent. Eyes relatively rotizations externally in the surrounding dia-
small, interocular index 0.671.0; cornea naked. phragma; base of vesica almost always enclosed
Antennae 0.40.6 the length of forewing, rarely by a thickened collar, or cathrema (Scoble 1983,
up to 1.0; scape usually greatly enlarged, cover- Nieukierken 1986, Johansson et al. 1990). Female
ing dorsum of eye, reduced in some genera; pec- genitalia: Ovipositor short, with two pairs of
ten present or absent; flagellum filiform, possess- apophyses; posterior pair usually longer and
ing paired, variably branched sensilla vesicu- more slender, terminating on T9, sometimes in
loclada (Fig. 6.9 E, Nieurkerken and Dop 1987) paired, setose pads (papillae anales); anterior
per segment and a single row of slender, biden- apophyses usually shorter and stouter, some-
tate scales encircling each segment. Pilifers and times indistinct, arising from T8 which continues
mandibles absent. Haustellum short, usually less as a short band across tergum. Oviporus located
than, rarely up to 3 X, the length of labial palpi terminally, between S8 and 9. Vagina short, usu-
(Scoble 1983). Maxillary palpi long, 5-seg- ally enlarging anteriorly into thick walled vesti-
mented. Labial palpi 3-, rarely 2-segmented, bulum; latter with or without spicules. Sperma-
drooping; lateral bristles present or absent. Tho- theca with a lateral lagena. Corpus bursae usu-
rax: Moderately depressed (Davis 1989). Meta- ally gradually enlarged, rarely reduced, with or
furca (Fig. 6.6 Q) with primary apophyses free, without accessory (appendix) bursae; signa pre-
erect, elongate. Forewings (Fig. 6.5 I) slender to sent or absent, variable, usually with a large pair
lanceolate; index 0.22-0.25; venation reduced, of oval, reticulated signa in Trifurculini (Johans-
usually faint, without true crossveins; forewing son et al. 1990), or with numerous spicules scat-
with Sc extremely short; M anastromosing with tered over inner walls.
R and sometimes basally with Cu; R with 1 - 4
Egg: Largely undescribed due to cuplike cov-
branches terminating on costa; M with 12
ering secreted by over egg (Johansson et al.
branches; Cu reduced, unbranched; 1A + 2A
1990); egg very flat, laid singly and cemented to
usually fused, rarely with anal fork; $ retinacu-
epidermis of host. Larva (Fig. 6.10 LM):
lum consisting of several stiff curled scales from
Small, usually less than 5 mm, sometimes up to
costal fold as well as subdorsal scales in some
9 mm long; body cylindrical to slightly de-
species. Hindwing lanceolate with 3 6 major
pressed, apodal, with translucent cuticle ranging
veins and no closed cells; index 0,160.31; cilia
in color from yellow to green or whitish. Epicra-
24X wing width; sex scaling frequently present
nial notch deep, extending ca. 0.5 the length of
in S frenulum a single stout seta, sometimes
head; lobes of vertex well developed, extending
accompanied by a series of stout, subcostal pseu-
deep into T l ; frontoclypeus triangular to rectan-
dofrenular setae; frenulum composed only of
gular, generally outlined by strong internal
subcostal pseudofrenular setae. Pretarsi with un-
ridges. One pair of stemmata. Antenna usually
guitractor plate composed of rows of divided
reduced to one segment, 3-segmented in Pectini-
scutes. Abdomen: S2 (Fig. 6.7 N) divided into a
valva (Johansson et al. 1990). T 2 - 3 with paired
short, broad, anterior fragment and a much
ventral ambulatory calli, sometimes with dorsal
larger posterior sclerite; a pair of small tubercu-
calli. Chaetotaxy reduced; D2 absent on A l - 9 .
late plates present in pleura; wart- like processes
Paired ventral calli usually present on Al8,
absent. S4 of $ with a pair of small, circular fe-
sometimes lost on A8; dorsal calli present in
nestra; $ sometimes with sex scaling mid-dor-
some Ectoedemia\ AIO with a slender pair of
sally on A68, or with paired lateral tufts on
sclerotized anal rods. Larvae are miners through
A48; corythrogyne absent. Male genitalia. Un-
all instars, usually in leaves, less commonly in
cus usually distinct and either acute or bilobed,
bark of woody twigs, in fruit, or in galls. Leaf
The Monotrysian Heteroneura 69

mines are typically slender and serpentine, with tellum short, 0 . 7 5 - 1 . 5 the length of labial palpi.
a relatively broad median frass trail (Figs. Maxillary palpi long, geniculate, 5-segmented.
6.12 - C ) , occasionally widening to form oval Labial palpi 3-segmented, moderately short,
blotch mines in some species. More than 30 plant slightly drooping; lateral bristles absent. Thorax:
families reported as hosts, with Betulaceae, Fa- Depressed, relatively broad. Metafurca (Fig. 6.6
gaceae, Rhamnaceae, Rosaceae, and Salicaceae RS) variable, usually with furcal apophyses
particularly favored in the Holartic region free; apophyses connected to secondary arms in
(Nieukerken 1986). Larvae usually with 4 instars, Pseudopostega. Forewings (Fig. 6.5 J) lanceolate,
less commonly 5 - 8 (Johansson et al. 1990). The index 0.20.23; venation extremely reduced, typ-
number of generations can vary according to ically only 4 major veins present, without sec-
species, with some obligatorily univoltine and ondary branches or crossveins; microtrichia gen-
others undergoing as many as 3 generations ac- erally restricted to base of ventral forewing sur-
cording to climatic conditions. Pupa: Vertex face. Hindwing index 0.140.17; a uniform
smoothly rounded. Antennal scape usually series of 418 subcostal pseudofrenular setae
greatly enlarged. Wings extending to A9. A28 (Fig. 6.9 C) arising from base of hindwing in
with one or more rows or small, tergal spines. both sexes to couple with a subdorsal retinacu-
Al8 apparently moveable in both sexes lum consisting of a row of stiff scales on un-
(Mosner 1916). Cremaster usually consisting of derside of Cu in forewing. Hindtibiae with rough
a small pair of dorsal and/or ventral spines. Pu- spinose scales; pretarsus with unguitractor plate
pation rarely inside mine, usually occurring out- composed of several rows of undivided scutes.
side, either attached to some plant surface or in Abdomen. S2 (Fig. 6.7 O) large, weakly sclero-
ground detritus beneath host, within an oval, tized, usually with a large, central, partially or
lenticular cocoon of densely woven brown to yel- completely divided hyaline area. Coremata and
lowish silk. corethrogyne absent. Male genitalia: Uncus re-
Currently 11 genera and nearly 800 species duced, typically a slender bridge between a pair
have been described from every major region ex- of widely separated setose lobes (socii?), rarely a
cept Antarctica (Braun 1917, Borkowski 1969, medium connate lobe (in Eosopostega). Tegumen
1972, Wilkinson and Scoble 1979, Newton and reduced to a narrow dorsal ring. Vinculum usu-
Wilkinson 1982, Puplesis 1984 a, b, 1985, 1994, ally a narrow, rounded ventral ring, anterior
Kemperman and Wilkinson 1985, Nieukerken margin deeply concave in Opostegoides and Eo-
1985, 1990, Donner and Wilkinson 1989, Hepp- sopostega. Gnathos fused, variable, usually a
ner 1991, Hoare et al. 1997). Serpentine mines, well developed arch. Transtilla typically absent,
allegedly caused by Nepticulidae (but see 1 - 2 ) , rarely present as a slender, medium bridge. Juxta
have been reported on fossilized leaves of one of either absent, or, if present, a broad menbranous
the most primitive seed plants, Pachypteris (Pte- plate. Valvae variable in outline, with a distinct
ridospermophyta) from the upper Jurassic/lower costal apophysis extending inward from base but
Cretaceous of north Queensland, Australia articulated in Pseudopostega, apex of valva (cu-
(Rozefields 1988). Two subfamilies are recog- cullus) an ovoid, pectinated lobe joined to basal
nized (Scoble 1983). half of valva by a slender pedicel. Aedoeagus
Pectinivalvinae. With pectinifer on S valva. usually absent, present in Notiopostega, Opos-
One genus, Pectinivalva, restricted to Australia. tega, Paralopostega, and Eosopostega. Female
Nepticulinae. With pectinifer lost, subdorsal genitalia: Ovipositor (Fig. 6.8 L) non-piercing,
retinaculum present on forewing. Acalyptris, Are- short, barely extending beyond A7. Dorsal,
ticulata, Bohemannia, Ectoedemia, Enteucha, paired, anal papillae usually present and setiger-
Parafomoria, Simplimorpha, Stigmella, and Tri- ous, sometimes single, absent in Opostegoides.
furcula (often treated as a subgenus of Ectoede- Anterior apophyses usually absent; posterior
mia), and Varius. apophyses always present, either short or elon-
gate. Oviporus located terminally due to extreme
Opostegidae (Figs. 6.2 EG). Extremely to mod- reduction of A8, 9 + 1 0 . Ductus usually mem-
erately small moths, with forewings 1.8 8.3 mm branous. Corpus bursae usually a large membra-
in length. Head (Fig. 6.3 J): Vestiture variable; nous sac, without a well defined signum but
vertex typically rough with erect piliform scales often with inner walls variously covered with
possessing minutely bidentate apices; most of minute spicules.
frons and cranium caudad to vertex covered with Egg: Cylindrical, elongate-oval, up to 1.1 mm
broad, lamellar scales with 35 dentate apices; long. Chorion smooth and transparent, whitish
lower frons naked. Ocelli absent. Eyes small to to yellow. Laid singly on epidermal surface of
large; interocular index 0 . 5 - 1 . 2 ; cornea naked. plant host. Larva (Figs. 6.10 N - O ) : Body whit-
Antennae 0.70.9 the length of forewing; scape ish in color, extremely slender, 8 - 2 5 mm long,
usually greatly enlarged; pecten absent; flagellum and apodal, with paired ventral callosities on
filiform, typically with 3 ascoid sensilla per seg- T23. Head (Fig. 6.10 O) depressed, triangular,
ment (Fig. 6.9 D) and one row of slender biden- with shallow epicranial notch and poorly defined
tate scales. Pilifers and mandibles absent. Haus- 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 71

Simular to that of Nepticulidae. Cuticle rela- gyne absent; without coremata. Male genitalia
tively smooth except for concentrations of min- typically with 1 - 3 rows of comb-like spines
ute tergal spines on A28. Antennal scape usu- (pectinifers) on valva (Figs. 6.8 AB); pectinifers
ally extremely broad. Wings, antennae, and fore- absent in Incurvariidae, Adelinae, or reduced in
legs usually extending to or slightly surpassing Crinopterygidae and some Prodoxidae; juxta
abdomen. All coxae flat and exposed. Pupation usually elongate, either sagittate (Fig. 6.8 C) or
occurring in a tough, silken, lenticular cocoon deeply furcate (Fig. 6.8 F). Female with elongate,
(Fig. 6.11 F) outside of mine, usually in leaf lit- extensible, ovipositor specialized for piercing
ter. Family recently revised (Davis 1989) wherein plant tissue (Figs. 6.8 GK); 2 or more pairs of
6 genera were recognized. Two subfamilies. threadlike retractor apodemes attached to vari-
Opostegoidinae. Aedoeagus present. Notiopos- ous sites along the ovipositor and associated in-
tega (1 sp.), Eosopostega (1 sp.), Opostegoides ternal organs (Fig. 6.8 G); cloaca well developed,
(7 sp.), Paralopostega (6 sp., all in Hawaii). elongate; cloacal opening terminating near apex
Oposteginae. Aedoeagus absent. Widely dis- of ovipositor (AIO). Larva with crochets con-
tributed in all major regions except Antarctica. sisting of relatively simple spines typically ar-
Opostega (58 sp.), and Pseudopostega (29 sp.). 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
INCURVARIOIDEA. spines; partially extruded from larval/pupal case
Small to very small moths; length of forewings prior to eclosion.
1.716 mm. Head usually with rough, piliform Monophyly of the Incurvarioidea has been de-
scales, smooth and broadly scaled in most Helio- termined by the presence of (1) an extensible,
zelidae. Chaetosemata and ocelli absent. Anten- piercing ovipositor, similarly developed in all
nae filiform, rarely pectinate in some <3 Incurv- members of the clade; (2) two or more pairs of
arla, scape without eyecap; pecten usually pre- retractor apodemes; (3) the enclosure of the $
sent. Haustellum usually moderate in length, eighth abdominal segment largely inside greatly
typically elongate in Adelidae, to greatly reduced enlarged seventh sternum; and (4) sagittate juxta
or absent in most Cecidosidae; usually unsealed, in <5. Cladistic analysis of the six incurvarioid
basally scaled in Adelidae and a few Prodoxidae. families + a hypothetical outgroup was per-
Maxillary palpi variable, usually 5-segmented, formed using the computer program Hennig86
less commonly reduced or absent. Labial palpi (Farris 1988). A morphological data set of 15
usually 3-segmented, with apical sensory organ characters produced two parsimonius clado-
of vom Rath on segment III and erect bristles grams 36 steps long with consistency and reten-
on II; palpi rarely 2-segmented or absent. Wings tion indices of 69 and 54 respectively. One tree
usually with complete heteroneurous venation was identical to a manually derived cladogram
and distinct discal cell in forewing; venation previously proposed (Nielsen and Davis 1985).
more reduced in Heliozelidae and Dicranoses Successive weighting selected the second tree that
(Cecidosidae); microtrichia generally scattered differed only in the position of Prodoxidae +
over all wing surfaces, reduced in some genera; Cecidosidae being inverted basal to Crinoptery-
male frenulum usually consisting of a single large gidae + Incurvariidae.
bristle that inserts into a triangular, subcostal
lobe (retinaculum) under base of Sc in forewing; Heliozelidae (Figs. 6.1 A - C ) . Generally very
frenulum absent in Cecidosidae; $ frenulum usu- small moths with forewings 1.77.0 mm. Head
ally indistinct, consisting of 2 or more small, cos- (Fig. 6.3 A): Vertex typically smooth with broad,
tal bristles along base of hindwing. Legs usually laminate, iridescent scales directed down over
with tibial spur pattern of 0-2-4, epiphysis usu- similarly smooth frons; head rough with pilform
ally present, absent in at least one genus in al- scales in one undescribed South American genus
most every family. Abdomen with second ster- (Nielsen 1985). Eyes small, interocular index
num (Figs. 6.7 A, C, G, H, J, L) divided trans- 0.6-0.7; cornea with sparse, scattered inter-
versely into a mostly membranous anterior third facetal microsetae. Antennae 0.40.6 the length
(S2a) and a more sclerotized, posterior two- of forewing; scape smooth, entirely covered with
thirds with slender, anterolateral processes and a usually iridescent scales; pecten absent; flagellum
bilateral pair of minute wart-like protuberances filiform, fully scaled to apex but more sparsely
(Fig. 6.7 A); usually with two rows of tuberculate scaled ventrally; a single ring of scales per seg-
plates present in pleura, one near lateral margin ment. Pilifers present or absent. Mandibles ves-
of sterna and one in a posterodorsal position to tigial. Haustellum moderate to long, 1.5-2.0 the
spiracle (Nielsen 1980); anterolateral processes of length of labial palpi, unsealed, occasionally with
S2b articulating with first sternum by means of base concealed by elongate, lamellar scales from
slender tergosternal connection posterior (Fig. labrum. Maxillary palpi reduced in length, not
6.7 A) to Al spiracle (Kyrki 1983). ? with A8 geniculate, usually 3 - 5 short segments, reduced
reduced and largely enclosed by enlarged S7; T7 to 1 minute segment in Coptodisca. Labial palpi
(Fig. 6.7 E) less than half the size of S7; corethro- moderately short, 3-segmented, drooping. Tho-
72 Donald R. Davis

Fig. 6.4. 9 Tegeticula yuccasella (complex) collecting pollen from anther of Yucca elata.

rax: Lateral angles of laterocervical sclerites lobed. Vinculum well developed with an elon-
broad, triangular (Fig. 6.6 A). Metafurca (Fig. gate, broad, V-shaped saccus. Valvae with a sin-
6.6 I) with dorsal apophyses moderate to well de- gle pair of pectinifers, each bearing about 520
veloped, arising perpendicular from mesal la- spines. Juxta slender, sagittate. Aedoeagus an
mella free from secondary arms of metafurca- elongate tube usually equalling length of saccus;
sternum. Forewings (Fig. 6.5 AB) broadly lan- cornuti present. Female genitalia: Ovipositor
ceolate, index 3.4-3.6; microtrichia mostly re- with depressed, moderately broad apex terminat-
stricted to subhumeral and subanal areas; scales ing in 45 minute cusps. Spermatheca with lat-
often lustrous; radius with 3 or 4 branches; Rs4 eral lagena (Razowski and Wojtusiak 1978).
stalked to Ml and terminating on costa near Ductus bursae sometimes with small spines; cor-
apex; discal cell either open or closed; accessory pus bursae entirely membranous, without signa.
cell absent; M with 1 or 2 branches; CuA with Egg\ Poorly known; inserted singly into host
12 branches; 1A + 2A without basal fork; <? tissue. Larva (Figs. 6.10 AB): Body white to
retinaculum a triangular lobe from costa; $ reti- pale yellow or green, usually with brownish to
naculum consisting of 34 stout, curved spines black pronotum and anal plate, sometimes with
from a small swelling at base of Sc. Hindwings a series of nearly contiguous, mid-dorsal dark
more slender, lanceolate, index 0.20.28; M- plates down length of body; 46 mm in length.
CuA crossvein absent, discal cell open; M with 1 Head mostly prognathous, strongly depressed
or 2 branches; 1A + 2A usually fused, rarely di- with 2 - 5 pairs of stemmata. Legs usually absent,
vided near margin; frenulum a single large bristle well developed, 5-segmented in some Heliozela
in male with 46 smaller costal setae and with (Davis 1987). Coxal plates, if present, separated.
usually 2 small bristles in . Legs with tibial spur Paired ventral and dorsal ambulatory calli some-
pattern of 0-2-4; epiphysis usually present, ab- times present on T1 and 2. Prolegs either absent
sent in Coptodisca. Abdomen. Anterior third of or nearly so; crochets usually absent, present on
S2 with broad U-shaped caudal rim (Fig. 6.7 A); A3 6 in some Heliozela and consisting of
S7 of ? with caudal margin rounded (Fig. 6.7 B). multiserial rows of simple, enlarged spines re-
Male genitalia: Uncus indistinct, shallowly bi- sembling those of Adelidae except planta some-
The Monotrysian Heteroneura 73

times not separated mesally (Fig. 6.9 K); sent in Cauchas; flagellum filiform, usually fully
crochets always absent on AIO. Some Antispila scaled to apex with two annuii of appressed
and Coptodisca with fused, ventral ambulatory scales per segment, more thinly scaled ventrally;
calli on A3 6; paired dorsal and ventral calli basal 0.20.6 of antenna roughly scaled in some
also present on A8 in Coptodisca, but only a sin- Adela; $ of some Adela and all Nemophora with
gle mid-dorsal callus on A8 in Antispila. Most variable number of mediodorsal spines (Fig.
first instar larvae begin a short, serpentine mine 6.9 A) along base of flagellum (Nielsen 1980).
that is abruptly enlarged into a small, full depth Pilifers present. Mandible vestigial. Haustellum
blotch with the frass retained inside the mine. All usually elongate, 1.52.5x the length of labial
instars mine except the last, which constructs a palpus; reduced, about equal to labial palpus in
flat, oval case by cutting sections from the upper some African Ceromitia; basal '-'/ scaled.
and lower epidermis of the mine and joining Maxillary palpi 2-5-segmented. Labial palpi 3-
them with silk. The lenticular shape of the case segmented, variable in length, usually upturned;
provides the origin of this family's common segments II and III usually rough and clothed
name, 'shield bearers'. In most instances the lar- with long piliform scales. Thorax: Laterocervical
vae release a silk strand to lower themselves usu- sclerites with slender, elongate, lateral processes
ally to the ground to pupate amongst leaf litter. (Fig. 6.6 B). Metafurca (Fig. 6.6 J) with dorsal
Sometimes the cases are attached to the bark of apophyses well developed, arising perpendicular
the host plant. Abandoned mines with small oval from mesal lamella free from secondary arms of
holes are characteristic of the leaf damage caused metafurcasternum. Forewings (Fig. 6.5 C) slen-
by these insects (Fig. 6.11 A). Some Heliozela der, index 0.30.37; microtrichia usually pre-
mine twigs, petioles, leaf ribs, or initiate galls sent, reduced in some genera; radius with 5
(Needham et al. 1928, McGufin and Neunzig branches, Rs2 and 3 either separate, connate or
1985). At least 13 families of mostly woody trees, stalked; Rs4 usually to costa in most genera, ter-
shrubs, and vines have been reported as hosts minating on termen below apex in Nematopogon\
with the Betulaceae, Cornaceae, Fagaceae, and accessory cell present; base of M faint, rarely
Vitaceae particularly favored. Pupa: Very small, forked within cell; + 2A with short basal
length 2 - 3 mm. Cuticle thin, transparent. Head fork; S retinaculum a long costal fold extending
smoothly round; antenna short, less than half the under base of Sc. Hindwings similar to forewings
length of wing; scape cylindrical. Abdomen with in width, index usually 0.35; Ml and 2 either sep-
dense patches of minute, scattered dorsal spines; arate or stalked; + 2A with short basal fork;
segments A 2 - 8 movable in A 2 - 7 in $ i frenulum a single stout bristle, usually accom-
(Mosher 1916). Spiracles either raised or flush. panied by several smaller setae along costal mar-
Pupation inside larval case with pupa partially gin; $ frenulum consisting of 3 - 4 smaller bristles
extruded at eclosion. Most species are probably in a row along base of costa. Legs with tibial
univoltine, less commonly bivoltine. spur pattern of 0-2-4; epiphysis present. Abdo-
Because of their minute size and diurnal habit, men: Anterior third of S2 with a slender, U-
Heliozelidae are seldom collected. The neotropi- shaped caudal rim (Fig. 6.7 C); S7 of $ triangular
cal fauna is especially poorly known. Approxi- to subrectangular (Fig. 6.7 C), caudal margin
mately 12 genera and over 100 species (Heppner rounded to slightly emarginate. Male genitalia:
1991) have been reported from nearly every ma- Uncus reduced, often bilobed; tegumen a moder-
jor region except Antarctica and New Zealand. ate to narrow, dorsal band. Vinculum well devel-
Most of the species are included within the gen- oped with an elongate, broad to slender, V- to Y-
era Heliozela, Antispila, and Coptodisca. The shaped saccus. Valvae with 0 - 3 pairs of either
principle autapomorphy for the family is the loss sessile or pedunculate pectinifers; pectinifers ab-
of the M-CuA crossvein in the hindwing. sent in Adela, Cauchas, Nemophora, and some
subgenera of Ceromitia. Juxta slender, sagittate.
Adelidae (Figs. 6.1 D - ) . Small moths with Aedoeagus an elongate tube; cornuti usually pre-
forewing length 3.5-12 mm. Head (Figs. 6.3 sent, apex sometimes also armed with large, ex-
BC): Vertex rough, densely covered with pili- ogenous spines. Female genitalia (Fig. 6.8 G):
form scales; frons usually rough, occasionally Ovipositor with a compressed, acute apex usu-
covered by smooth, broad scales. Eyes small to ally minutely serrated along ventral keel. Sper-
large, interocular index 0.5-1.7; interfacetal matheca without lateral lagena. Ductus and cor-
microsetae scattered; eyes dimorphic in most pus bursae membranous, without spines or
Adela and Nemophora with that of enlarged, signa.
more approximate at vertex, and usually with Egg: Poorly known; inserted singly into plant
facets in upper 2 of eye enlarged. Antennae usu- tissue. Larva: Body slightly depressed, white to
ally longer than forewing in both sexes, up to green with darkly pigmented plates and head;
3X forewing and arising nearly contiguous in 712 mm in length. Head mostly prognathous
antennae shorter in Cauchas, 0.51.2 the length with 6 pairs of stemmata; AF2 absent; adfrontal
of forewing in both sexes; scape sometimes sclerite elongate, extending to deeply incised epi-
slightly swollen in 3; pecten usually present, ab- 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 (1894), during late spring and early summer the
prespiracular sclerite. Coxal plates fused, at least eggs of Nematopogon metaxella are inserted into
on Tl. Legs well developed; tarsi without en- any convenient herbaceous plant. Upon hatch-
larged, squamiform seta. Prolegs greatly reduced ing, the larva immediately drops to the ground
on A 3 - 6 , absent on AIO; crochets arranged in where it constructs a flattened, oval case from
several more or less definite transverse rows that soil particles and eventually dead leaves (Chrt-
gradually decrease in size from center. Largely ien 1894). The larva feeds on both living and
because of the secretive and sometimes omnivo- dead plants and does not complete its develop-
rous feeding habits of the larva, surprisingly little ment until the following spring. Kuroko (1961)
is known about the biology of most species in reports a somewhat different life history for
this relatively well known family of moths. Eggs Nemophora raddei that may more accurately re-
are inserted into plant tissue in which the larva flect the univoltine norm for the family. In this
may or may not feed (Heath and Pelham-Clinton species the eggs are inserted into the ovaries of
1976, Nielsen 1985). According to Chrtien Salix sieboldiana in spring. The first instar larva
The Monotrysian Heteroneura 75

feeds on the ovules as well as the ovary wall. Prodoxidae (Figs. 6.1 I - O ) . Small moths, with
After moulting, it constructs a small, oval case forewings 4 to 16 mm in length. Head (Fig.
and descends on a silken thread to the ground 6.3 E): Vestiture usually rough, with dense pili-
where it prefers to feed on dead leaves of the form scales; frons and vertex covered with ap-
host Salix and Castanea crenata. The mature pressed, lamellar scales in Prodoxoides, with
larva (6th instar) pupates near the end of Octo- erect tufts of slender, bidentate scales arising
ber, with the adult emerging the following more caudad on occiput. Eyes small to large; in-
spring. The eggs of most Adela are inserted into terocular index 0.71.5; cornea either naked or
the flower ovary of their host wherein the first with sparse, scattered interfacetal microsetae.
instar larvae feed on the developing seeds. From Antennae 0.330.6 the length of forewing; scape
the second instar on, the larvae become caseb- usually smooth, with pecten present or absent;
earers and feed on the lower or fallen leaves of flagellum filiform, mostly naked except for dense
their host (Heath & Pelham-Clinton 1976). First sensilla chaetica overall and a single row of nar-
instar larvae of some Adelidae may mine leaves row, dorsal scales per segment over basal seg-
(Common 1990). Over 20 families of angio- ments which gradually disappear out to basal
sperms and one gymnosperm (Pinaceae) have third. Pilifers usually well developed, reduced in
been reported as hosts (Kppers 1980). Pupa: Tetragma. Mandibles vestigial but relatively
Length 5 - 7 mm. Vertex smoothly rounded. An- prominent. Haustellum usually unsealed, basally
tennae usually surpassing abdomen, then loosely scaled in Tridentaforma and Greya pectinifera,
encircling caudal end of abdomen 1 - 4 times 12X length of labial palpi. Maxillary palpi
(Figs. 6.10 C - D ) . Abdomen with 1 - 2 rows of elongate, 0.5 1.2 the length of haustellum, 3 to
dorsal spines on most segments; segments 3 - 7 5-segmented; an elongate, unsegmented, coiled
movable (Common 1990). Pupation inside larval tentacle arising from intersegmental membrane
case with the pupal exuvium partially extruded. of basal segment of most Tegeticula and Para-
tegeticula (Fig. 6.3 D; Davis 1967). Labial palpi
Adelidae is a widely distributed family of
relatively short, either porrect or upturned, usu-
nearly 300 species that occur on all continents
ally 3-segmented, 2-segmented in Parategeticula;
except Antartica and are absent from New
lateral bristles present. Thorax: Lateral angles of
Zealand (Heppner 1991, Janse 1945, Meyrick
lateral cervical sclerites triangular, not produced
1912 a, b). Autapomorphies for the family in-
(Fig. 6.6 C - ) . Metafurca with apophyses either
clude the elongate antenna (particularly in the
free (Fig. 6.6 M) or joined as a bridge to second-
<$), slender lateral processes of the lateral cervical
ary arms (Fig. 6.6 N). Forewings (Fig. 6.5 D)
sclerites, and possibly the high number of ovari-
slender; index 0.24-0.32; microtrichia usually
oles per ovary (10-20, Nielsen 1980, Nielsen and
scattered over all surfaces, absent from dorsal
Davis 1985). Two subfamilies have been pro-
base of forewing in Tetragma, Rs4 terminating
posed (Kppers 1980):
on costa; Rs3 and 4 stalked in Tetragma, Acces-
Adelinae (maxillary palpi 2-3-segmented; $ sory and intercalary cells present; 1A + 2A with
valvae without pectinifers), including Adela, basal fork; S retinaculum a triangular lobe in
Cauchas, and Nemophora. costal membrane anterior to and tightly curling
Nematopogoninae (maxillary palpi usually 4- under base of Sc. Hindwings ca. same width as
5-segmented; $ valvae usually with pectinifers), forewings, index 0.33-0.44; intercalary cell pre-
including Ceromitia and Nematopogon. Recent sent; $ frenulum a single stout seta; $ usually
mtDNA sequence data also links the problem- with 23 sometimes indistinct frenular setae;
atic North American genus Tridentaforma to this frenulum lacking in both sexes of Parategeticula,
family (Brown et al. 1994). The lateral cervical sex scales typically absent from wings, a slender
sclerites of Tridentaforma (Fig. 6.6 C), however, hair pencil contained in hindwing anal fold of
differ from the typical adelid form (Fig. 6.6 B). Tetragma. Legs with tibial spur pattern of 0-2-4;
In contrast to most Nematopogoninae which epiphysis usually present, absent in Parategeti-
tend to be drab in color and crepuscular or noc- cula. Abdomen: Caudal rim of S2a U to W-
turnal, the adults of Adelinae are often metallic shaped (Fig. 6.7 G); S7 of $ with caudal margin
and predominantly diurnal. The males of many rounded. Male genitalia (Figs. 6.8 AC): Uncus
species of Adela and Nemophora swarm, usually variously fused to tegumen, usually with 2,
near the species' host plant or oviposition site. sometimes 1 terminal lobes. Tegumen a narrow
The enlarged compound eyes in these males are dorsal band. Vinculum usually well developed,
an adaptation for swarming (McAlpine and either V- or Y-shaped; saccus often well devel-
Munroe 1968, Downes 1969). The development oped, total length 1 - 2 . 5 X the length of valva.
of specialized, spinose setae and scales near the Anellus usually membranous. Valvae often with
base of the antennae of swarming males may be a pectinifer (Fig. 6.8 B), variably developed,
further adaptations associated with this courship sometimes reduced to a few short, contiguous
behavior (Nielsen 1980), possibly for use in spines, or absent. Juxta a well defined, sagittate
sound production or visual signaling (Bland sclerite. Aedoeagus an elongated tube, usually
1977). 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 77

elongate; apex (Figs. 6.8 JK) usually com- Prodoxoides and Tridentaforma). A typical life
pressed with dorsal and ventral margins often cycle for a member of this subfamily normally
raised and serrated, or smooth. Spermatheca involves two feeding phases interrupted by a lar-
without lateral lagena. Corpus bursae usually val diapause. Eggs are inserted singly into young
with paired, stellate signa which are occasionally fruit in late spring to early summer, with the
reduced or lost. early instar larvae first feeding on the developing
Egg: Poorly known; white, shape highly vari- seeds. In mid to late summer, the larva (probably
able, pedicellate in Tegeticula, ca. 2 mm long and third instar) leaves the fruit and spins a haberna-
0.1 mm in diameter (Riley 1892) to oval, 0.3 culum in some crevice lower on the host or in
0.5 mm long and 0.20.3 in diameter (Davis the soil where it overwinters. Early next spring
et al., 1992). Chorion smooth with reduced the larva moves back up the stem and bores into
micropylar reticulation. Larva (Fig. 6.10 K): developing buds or shoots, often entering and
Body white, green, or reddish in color, cylindri- damaging several. Pupation normally occurs in
cal to fusiform, 6 - 2 2 mm in length. Head light the shoot with the adult emerging soon af-
to dark brown, with frontoclypeus extending terwards in late spring. Lampronia (28 sp.),
0.5-0.66 the distance to epicranial notch. Usu- Tetragma (1 sp.), Prodoxoides (1 sp.), and Triden-
ally 6 pairs of stemmata, reduced to 3 pairs in taforma (1 sp.). Relationships of Tridentaforma
apodal forms (Davis 1987). Legs usually present, remain problematical, both from the standpoint
absent in Agavenema and Prodoxus; pretarsus of morphological and molecular data. Morpho-
often with a large squamiform seta from lateral logically, the genus possesses characters shared
base of claw. L series either bi- or trisetose on with both Adelidae (basally scaled haustellum)
T l . Prolegs reduced or absent; crochets often ab- and Incurvariidae valva with rows of flattened
sent, at least in early instars, with a single spines). Molecular data reported to date present
transverse row on A 3 - 6 in Lampronia and Mes- divergent results. Work with mtDNA (Brown
epiola or as biserial circle in Greya; crochets al- et al. 1994) associates the genus with Adela;
ways absent on AIO. Larva endophagous, boring whereas, analyses involving 18S rDNA places
inside plant shoots, leaves, buds, flower recepta- Tridentaform either at the base of the Hetero-
cles, fruits, or seeds. Typically there is a single neura or near Cecidosidae (Wiegmann 1994).
generation per year, with the larva overwinter- Prodoxinae. Adults with metafurcal apophyses
ing. In synchrony with the irregular flowering joined to secondary furcal arms (Fig. 6.6 N).
habit of their host, last instar larvae of some Pro- Larvae either with legs and crochets or apodal.
doxinae are capable of prolonged diapause, All genera with the exception of Greya (17 sp.)
which in Prodoxus y-inversus may exceed 20 are restricted to the plant family Agavaceae. The
years (Powell 1992). Pupa: Vertex usually with larval biology of those Greya studied appears to
a prominent frontal beak, but smooth in Greya. resemble that of most Lamproniinae, with the
Wings extending to A 5 - 7 . A 2 - 7 movable in early instars boring in the seeds, fruits, or pedun-
both sexes (Mosher 1916); A 2 - 8 usually with a cles of Saxifragaceae and Umbelliferae and then
single anterior row of tergal spines. Cremaster exiting to the ground where they eventually over-
usually consisting of a pair of dorsal spines or winter. Larvae complete their feeding the
tubercules on AIO, sometimes accompanied by following spring on the flower buds or foliage of
paired ventral spines. Pupation usually occuring their host (Davis et al. 1992). Among the Agava-
in silken cocoon either externally and attached ceae feeders, an unusual pollination mutualism
to host, or within larval gallery or underground, has evolved between two genera and their plant
with the adults of most species emerging in hosts. Most females of Tegeticula (3 sp.) and Par-
spring to early summer. ategeticula (1 sp.) purposefully pollinate their
Prodoxidae are almost entirely restricted to host Yucca while visiting the inflorescenses for
the Holarctic region and are most diversified in oviposition without nectaring. This insures the
the Nearctic Region, with a single monotypic ge- development of fruit upon which their larvae will
nus present in southern South America (Nielsen feed (Riley 1892; Powell and Mackie 1966; Davis
and Davis 1985). Dugdale (1988) has reported 1967; Powell 1984, Pellmyr and Thompson
the discovery of an unidentified, questionable 1992). Among all insects, only the females of
prodoxid larva from cortical mines on Weinman- these two genera have developed specialized
nia sp. (Cunoniaceae) in New Zealand. The maxillary tentacles (Fig. 6.3 D) which are used
paired, typically stellate signa are a major auta- to collect pollen from the anthers of their hosts
pomorphy for the family. Two subfamilies are (Fig. 6.4). The females carry the pollen in a com-
recognized, consisting of a total of 12 genera and pact mass under the head and force portions of
nearly 80 species. their load down the stigmatic tube of the yucca
Lamproniinae. Adults with metafurcal apoph- during visits to the flower. Tegeticula oviposits in
yses free. Larvae with thoracic legs and crochets; the ovary, their larvae later consuming only a
borers in seeds, fruits, or stems of Rosaceae, Sax- portion of the seed crop. Parategeticula oviposits
ifragaceae, and possibly Myrtaceae, or in stem in sterile tissue, usually in the flower scape or
galls of Betulaceae (larval biology unknown for 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 79

fruit to feed in a cyst within the fruit (Powell genera, absent in Dicranoses. Abdomen: S2a with
1984). The interrelationships between Tegeticula broad, U-shaped caudal rim (Fig. 6.7 H); ter-
and their host yuccas are complex, involving sev- gosternal connection absent. S7 of with caudal
eral sibling species including sympatric nonpolli- margin rounded (Fig. 6.7 I). Male genitalia. Un-
nators, or "cheaters", with vestigial maxillary cus usually shallowly bilobed, deeply bilobed in
tentacles (Davis 1967, in press; Miles 1983, Addi- Dicranoses. Socii large. Valva slender, long, with
cott et al. 1990; Powell 1992). Females of the bo- an elongated, sessil pectinifer along ventral mar-
gus yucca moths, Prodoxus (10 sp.), also do not gin at about half length of valva or with a promi-
transport pollen, but their larvae often depend nent swollen setose lobe along ventral margin.
upon the development of inflorescenses and, Vinculum Y-shaped. Aedoeagus simple, slender
hence, the efforts of Tegeticula and Parategeti- and tubular; cornuti absent. Juxta elongate, slen-
cula pollinators. Larvae of Prodoxus bore in the der, usually with caudal 0.2-0.3 deeply divided
flower stalks, sterile fruit tissue, or leaves of and partially encircling aedoeagus. Female geni-
Yucca (Wagner and Powell 1988), and those of talia. Apex of ovipositor compressed, often sagit-
Agavenema (2 sp.) in the flower stalks of Agave. tate, ventral ridge with or without minute serra-
Larvae of Mesepiola (1 sp.) bore in the seeds of tions. Apophyses short, less than A7 in length.
Nolina and Dasylirion (Frack 1982). All prodox- Spermatheca without lateral lagena; caudal part
ine larvae feeding on Agavaceae complete their of ductus spermatheca not coiled. Vestibulum
feeding in one season and are strictly endopha- without sclerotized structures; ductus and corpus
gous. The subfamily is largely restricted to the bursae membranous, signa absent.
Nearctic Region, with three species of Greya oc- Egg: Oval, up to 293 in length; chorion
curring in eastern Russia and Japan (Kozlov relatively hard, up to 23 thick, brownish-yel-
1996). low to dark brown; micropyle subapical, finely
reticulate, with reticulations enlarging toward
Cecidosidae (Figs. 6.1 PQ). Small moths, with periphery and gradually dissipating. (Known
forewings 3.513 mm in length. Head: (Figs. 6.3 only for Cecidoses, Wille 1926). Larva: Body of
F - G ) Vestiture moderately smooth, with elon- varying shades of greenish-yellow; length up to
gate, slender, lamellar scales mostly curved for- 11 mm. Head prognathous, with most structures
ward to smooth frons; all body scales piliform in reduced, lightly sclerotized, light brown; stem-
Dicranoses. Eyes large, interocular index 0.7 mata absent. Thoracic legs reduced to circular,
1.1. Antennae short, 0.5-0.6 length of forewing; minute, unsegmented tubercules. Prolegs and
scape smooth except for dense pecten; flagellum crochets absent. Body setae often reduced in
filiform, apparently 2 scale rows per segment but length, sometimes with numerous secondary se-
actually with slender scales scattered only over tae. The larvae of all species are univoltine and
dorsal half; ventral half with elongate sensilla ca. cecidogenic, developing within galls on Rhus and
1.5x the length of flagellomere. Labrum greatly Schinus (Anacardiaceae). Galls (pyxidiocecidea)
reduced. Pilifere absent. All mouthparts absent of the South American Cecidoses and Eucec-
in Dicranoses (Fig. 6.3 G). Mandible reduced to idoses develop as hard, thick-walled, sessile
minute, sclerotized stubs or absent. Galea (haus- spheres (Figs. 6.11 GH) up to 18 mm in dia-
tellum) usually greatly reduced to minute setose meter on the woody, terminal branches of Schi-
lobes or absent (Figs. 6.3 G, 6.9 B), but moder- nus (Brthes 1916, Curtis 1835, Wille 1926, Mani
ately well developed in Scyrotis. Maxillary palpi 1964). Pyxidial galls are characterized by the
0-3-segmented, reduced to a minute, unseg- presence of a cap-like covering, or operculum
mented lobe from vestigial galea (Fig. 6.9 B) in the formation of which is sometimes unclear. In
Cecidoses; labial palpi 3-segmented or absent, Cecidoses, operculum formation is largely inde-
without apical pit or lateral bristles. Thorax: An- pendent of larval development. Past a certain
terior arms of laterocervical sclerites long and developmental stage, the operculum forms even
slender (Fig. 6.6 F). Metafurca with slender, after the young larva dies or is surgically removed
elongate dorsal apophyses free from secondary (Wille 1926). Prior to emergence, the pharate
arms (Figs. 6.6 KL). Forewings (Figs. 6.5 adult easily forces open the operculum and exits.
EF) broad to slender, index 0.300.32; micro- Oliera and Dicranoses form colonies of much
trichia usually present, reduced in some genera; smaller, elliptical to spindle-shaped galls en-
radius usually with 5 free branches, 4 in Scyrotis; closed within swollen stems of Schinus (Brthes
M 3-branched; accessory cell present; wing vena- 1916, Keiffer & Jrgensen 1910). Stems contain-
tion extremely reduced in Dicranoses (Fig. 6.5 F), ing Dicranoses eventually rupture, exposing the
with most major veins indistinct; retinaculum ab- galls inside (Fig. 6.11 I). The larvae of the South
sent. Hindwing nearly equal to forewing in African Scyrotis form similar galls on Rhus, ex-
width, more lanceolate in Dicranoses; frenular cept their galls eventually dehisce and fall to the
bristles absent in both sexes; wing coupling con- ground. While on the ground and probably as a
sisting of long frenular scales arising in two to means of avoiding excessive heat from the sun,
five irregular rows near base of costa. Legs with the very active pupa is capable of propelling the
spurs 0-2-4 or 0-2-2; epiphysis present in most 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 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
" 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 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.
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). 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
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- a small f a m i l y (7 species) of exclusively gall
idoses. Pupa: M a x i m u m l e n g t h 11 m m . H e a d f o r m i n g m o t h s restricted t o the southern H e m i -
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 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-
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 noses (= Ridiaschinia) 2 sp., Eucecidoses
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 - 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
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 f r o m southern South America and two genera
The Monotrysian Heteroneura 81

(Ptisanora - 1 sp., Scyrothis - 1 sp.) f r o m South culum well developed, 1 - 2 X the length of val-
Africa. Except for two gall-producing species of vae, U - to V-shaped. Valvae without a distinct
Prodoxidae (Lampronia fuscatella and possibly pectinifer but with flattened spines often ar-
Prodoxus y-inversus), the Cecidosidae are the ranged in 1 - 3 groups. Juxta (Fig. 6.8 F) with
only Incurvarioidea with this habit. In addition 1 - 2 pairs of usually elongate caudal arms par-
to their cecidogenic biology, other m a j o r autapo- tially enclosing aedoeagus. Aedoeagus tubular,
morphies of this family include: (1) host restric- usually with 1 or more large cornuti. Female gen-
tion to Anacardiaceae, (2) loss of the retinacu- italia: Ovipositor with a depressed, acute apex
lum-frenulum wing coupling in male, and (3) loss usually bearing minute, lateral serrations (Fig.
of the tergal-sternal connection between T1 and 68 I). Spermatheca without lateral lagena. Duc-
S2 of the abdomen (Fig. 6.7 H). Together with tus bursae often with internal sclerotizations;
the Prodoxinae, Cecidosidae are the only incurv- signa absent.
arioids to have adapted to arid and semiarid en- Egg: Poorly known; inserted singly into plant
vironments. Early reports associated Cecidosi- tissue. Larva: Body slightly depressed, variable
dae with either Tortricidae or Tineidae (Curtis in color f r o m white, pale yellow to gray;
1835, Brthes 1916). Biezanko (1961, actually 5 - 9 m m in length. Head mostly prognathous
Forbes in litt. ) and later Becker ( 1977) were the with 6 pairs of stemmata; A F 2 absent; adfrontal
first to note the family's true affinities with the sclerite elongate, extending to deeply incised epi-
Incurvarioidea. cranial notch. Thorax with prespiracular sclerite
bearing spiracle usually separate f r o m prono-
Incurvariidae (Figs. 6.1 R - T ) . Small m o t h s with tum. Sternal plates reduced, coxae separate. Legs
forewing length 3 . 5 - 9 m m . Head (Fig. 6.3 H): well developed; tarsi sometimes with an enlarged,
Usually entirely rough with piliform scales; irons squamiform seta adjacent to claw (Davis 1987).
smooth in Paraclemensia, with broader, laminate Prolegs greatly reduced; crochets present on
scales. Eyes moderately reduced; interocular in- A36 and arranged in a uniserial, transverse
dex 0.61.0; cornea without interfacetal micro- band; AIO usually without crochets but present
setae. Antennae 0.40.8 the length of forewing; in bi- to triserial rows in Excurvaria, Paraclemen-
scape smoothly scaled; pecten present; flagellum sia, and Vespina. Eggs are frequently inserted
usually filiform, bipectinnate in $ Incurvarla, an- into the underside of the host leaf where the first
nuii usually indistinct with scales scattered over instar develops an irregular blotch mine (Dziur-
flagellomere. Pilifers moderate to well developed. zynski 1958). O n completion of the mining stage,
Mandibles vestigial. Haustellum unsealed, short, most larvae cut through the upper and lower epi-
ca. Vi the length of labial palpus. Maxillary palpi dermal layers of the mine and construct an oval,
5-segmented, geniculate. Labial palpi 3-seg- lenticular case (Figs. 6.11 B E) of the two sec-
mented, relatively short, drooping, with erect tions by sewing the edges together. At this stage,
bristles f r o m apex of second segment. Thorax'. leaf damage becomes most obvious as nearly cir-
Ventral half of lateral cervical sclerites broad, cular holes begin to appear on the host, some-
with lateral angles only slightly produced (Fig. times accompanied by circular patches of skele-
6.6 G). Metafurca (Fig. 6.6 O) similar to Adeli- tonized leaf (Figs. 6.11 B, D, Jensen 1932). Drag-
dae, with well developed dorsal apophyses aris- ging the case, the larva continues to feed exter-
ing perpendicular f r o m anteromedial process, nally on its host (Alloclemensia, Basileura, Para-
free f r o m secondary arms. Forewings (Fig. clemensia, Vespina, Davis 1972, Nielsen 1981,
6.5 G) slender, index 0 . 2 4 - 3 . 3 ; microtrichia usu- Parra & Ibarra 1994 a, Ross 1958) or descends
ally present on all wing surfaces. Radius with often to the ground to feed on dead or partially
45 branches, Rs4 to costa above apex; acces- withered leaves (Incurvarla). As the larva devel-
sory cell present; M with 23 branches; base of ops, it enlarges the case by cutting oval sections
M faint or absent within cell; 1A + 2A with f r o m leaves and adding these to the top and bot-
short basal fork; $ retinaculum either a long cos- tom of the smaller, older case (Fig. 6.11 C). A
tal fold or a short flap extending under base of few genera (e. g., Perthida, Protaephagus) con-
Sc. Hindwings nearly as b r o a d as forewings; M tinue to mine throughout their larval develop-
with 3 branches; base of M usually preserved and ment and do not abandon the mine to construct
forked; 1A + 2A with short basal fork; frenulum a case until ready to p u p a t e ( C o m m o n 1969,
a single stout bristle in $ accompanied by several Scoble 1980). Hosts include members of the
smaller, costal setae; female with usually an un- Aceraceae, Betulaceae, Caprifoliaceae, Corna-
differentiated, basal series of much smaller costal ceae, Ericaceae, Fagaceae, Myrtaceae, Pro-
setae. Legs with tibial spur pattern of 0-2-4; teaceae, and Rosaceae ( C o m m o n 1990, H e a t h
epiphyses either present or absent. Abdomen. and Pelham-Clinton 1976, Scoble 1980) Pupa:
Caudal rim of S2a broadly U-shaped (Fig. Length 410 mm. Vertex smooth. Haustellum
6.7 L); caudal margin of S7 truncate (Fig. short, usually extending to end of labial palpi.
6.7 M). Male genitalia. Uncus usually indistinct Wings extending to A 5 - 7 movable in $, A 2 - 6
and shallowly bilobed, consisting of an elongate in ? ( C o m m o n 1990). A 2 - 8 with scattered tergal
rodlike, caudal appendage in Alloclemensia. Vin- 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 83

Scoble 1980), South America (Basileura, Sima- arranged in two uniserial rows; crochets absent
cauda Nielsen and Davis 1981, Parra and Ibarra on AIO. According to Nolcken (1882), the larva
1994 a), and North America (endemic Vespina constructs an elongate (3.54.0 mm long, 1.5
quercivora, Davis 1972) and one species each of 2.0 mm wide), approximately cylindrical case
Alloclemensia and Paraclemensia (Davis 1983, (Fig. 6.10 E - G ) from the leaf epidermis of its
Nielsen 1981, 1982). One autapomorphy of the host, Cistus (Cistaceae). Prior to feeding, the
family is the presence of large, noncontiguous larva attaches its case to the underside of a leaf
spines on the $ valvae, in lieu of a pectinifer. The and then enters through a small opening to mine
depressed apex of the ovipositor and trunctate the leaf in a manner similar to that of Coleo-
caudal margin of A7 (?), synapomorphies shared phoridae. Pupa: Maximum length ca. 3.5 mm
with its nearest relative, Crinopterygidae, consti- A28 with a large patch of minute, scattered,
tute other diagnostic features. tergal spines (Petersen 1978).
Crinopterygidae is represented by a single
Crinopterygidae (Fig. 6.2 A). Small moths with species, Crinopteryx familiella, from southern
forewing length 3 - 3 . 5 mm. Head (Fig. 6.3 I): France and Sicily. It has only been reared from
Entirely rough, with piliform scales. Eyes moder- Cistus, a genus of evergreen shrubs found most
ately reduced; interocular index ca. 1.0. Antenna abundant in the Mediterranean region. On the
0.6 the length of forewing, smoothly scaled; pec- basis of the depressed ovipositor, Crinopteryx
ten present; flagellum filiform; annuii indistinct, shows greatest affinity to Incurvariidae. How-
with probably scattered slender scales. Labrum ever, significant autapomorphies (e. g., coleo-
and pilifers reduced. Mandibles minute, ca. phorid-like larval biology, elongate-cylindrical
the length of maxillary palpus I. Haustellum un- larval case, angulate juxta, and a single large
sealed, short, ca. 0.5 the width of head. Maxillary spine on the $ valva), exclude the genus from all
palpus 4-5-segmented, geniculate; apical segment other incurvarioid families (Nielsen and Davis
minute, sometimes lost. Labial palpi 3-seg- 1985).
mented, with erect bristles from second segment.
Thorax. Lateral projections of lateral cervical
sclerites broadly triangular (Fig. 6.6 H). Meta- PALAEPHATOIDEA
furca (Fig. 6.6 P) with anterior margin of ante- Palaephatidae (Figs. 6.2 IL). Small moths with
romedial process (APM) broad, flared; dorsal forewings 3.8-16 mm in length. Head (Fig.
apophyses moderately short, free of secondary 6.3 L): Vestiture rough to semi-rough, consisting
arms. Forewings (Fig. 6.5 H) slender, index 0.27. of long piliform scales with acute to bidentate
Radius with 5 separate branches; M with 3 sepa- apices. Ocelli usually absent, present in Sesom-
rate branches; base of M simple within cell of mata. Eyes small to large, interocular index 0.6
both wings; accessory cell and chorda present; 1.2; cornea naked. Antennae 0.50.9 the length
+ 2A completely fused, without basal fork; of forewing; pecten usually present, absent in
S retinaculum a relatively elongate, triangular, subgenus Palaephatus; flagellum filiform, each
costal fold extending under base of Sc. Hind- segment with two rows of slender, bidentate
wings nearly as broad as forewings, index 0.25; scales dorsally and a narrow strip ventrally
M with 3 separate branches; S frenulum a single which is bare except for dense sensilla chaetica.
large bristle accompanied by several smaller cos- Pilifers usually well developed, reduced in Palae-
tal setae; $ with a basal series of much smaller, phatus. Mandibles vestigial, varying in size from
undifferentiated costal setae. Legs with tibial minute to moderately large. Haustellum short to
spur pattern of 0-2-4; epiphysis present. Abdo- moderate in length, seldom exceeding length of
men: Caudal rim of S2a W-shaped (Fig. 6.7 J); maxillary palpi. Maxillary palpi usually long, 5-
caudal margin of S7 truncate (Fig. 6.7 K). segmented, reduced to 4 short segments in Apo-
Male genitalia: Uncus indistinct, with a broad, phatus. Labial palpi 3-segmented, curved slightly
shallow, bilobed caudal margin. Vinculum upwards; second segment with bristles usually
broadly V-shaped. Valvae with a single large sub- arising apically. Thorax: Metafurca (Fig. 6.6 T)
apical spine. Juxta broadly angulate (Fig. 6.8 E). with apophyses free, usually projecting dorso-
caudally between secondary arms. Forewings
Aedoeagus short and relatively stout, with nu-
(Fig. 6.5 K) slender to moderately broad, index
merous short cornuti (Petersen 1978). Female
0.27-0.36; Radius 5-branched, Rs4 terminating
genitalia: Ovipositor similar to some Incurvarii-
on costa; accessory and intercalary cells present;
dae, with a depressed, laterally lobed apex (Fig.
1A + 2A with basal fork; microtrichia distrib-
6.8 H). Bursa copulatrix completely membra-
uted over all surfaces of both wings, restricted
nous, signa absent.
on dorsal surfaces of forewing of Sesommata and
Egg: Poorly known; inserted singly into plant Metaphatus to apical fifth; retinaculum arising
tissue. Larva: Morphology poorly known. Body from costal membrane, usually forming a trian-
cylindrical, ca. 3.44.0 mm in length. Head gular lobe curling under base of Se, or an elon-
mostly prognathous with 6 pairs of stemmata. gate fold in Sesommata. Hindwings usually as
Thoracic legs well developed, coxae separate. broad as forewings, index 0.330.49; S frenulum
Prolegs reduced; crochets present on A 3 - 6 and
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 vae variable in form, from elongate and slender
without a smaller, anterior sclerite (Fig. 6.7 P, to broadly rounded, often with lobes or pro-
S2a); tergosternal connection well developed cesses from distal third. Aedoeagus highly vari-
(Fig. 6.7 Q). Coremata usually absent, sometimes able, usually as long as genital capsule, with vari-
with pleural sex scales on pleuron of A23; cory- ably developed exogenous spines or lobes and
throgyne absent. Male genitalia: Uncus usually cornuti. Female genitalia (Fig. 6.8 M): Ovipositor
fused to tegumen, with rounded to bilobed apex. short and barely extending beyond A8, nearly
Tegumen a narrow dorsal band. Vinculum usu- truncate and non-piercing, with a slightly raised
ally reduced, broadly triangular. Gnathos usu- median sensory ridge (Fig. 6.9 H). Two pairs of
ally present, absent in Metaphatus and Plesiopha- apophyses with anterior pair from A8 and more
tus. Juxta usually absent, reduced in Sesommata slender. S8 short and broad; caudal margin
and Plesiophatus. Anellus usually membranous, either divided or entire and often with paired
sclerotized in Sesommata and Metaphatus. Val- spine patches. Oviporus between S8 and 9 + 10.
86 Donald R. Davis

Vagina relatively broad. Ductus bursae often terocular index ca. 2.2; cornea naked; dorsal rim
with thickened, internally spined walls, gradually slightly concave beneath antenna. Tentorial pits
enlarging to elongate corpus bursae; internal situated high on frons and touching rim of eye.
walls of corpus bursae usually with dense spic- Antennae 0.6-0.7 the length of forewing; scape
ules; signum present or absent. mostly smooth, with a prominent tuft of slender
Egg\ Unknown. Larva (Fig. 6.10 P): Small, ca. pecten projecting over eye; flagellum filiform,
712 mm in length. Body cylindrical, pale green with 2 or more rows of slender scales per seg-
to white, with dark brown head, pronotal and ment over dorsal half; venter naked except for
anal plates; setal pinacula extremely small, dense sensilla chaetica which are 23X longer in
slightly pigmented. Frontoclypeus extending ca. $ than 9; sensilla trichodea with strongly re-
0.50.6 the distance to epicranial notch; 6 stem- curved bases (Fig. 6.9 F, Davis 1986); flagellom-
mata present in a closely set anterodorsal ellipse. ere 1 (third antennal segment) 2.5 3 X longer
Lateral series on T1 trisetose. Legs well devel- than 2. Pilifers well developed. Mandibles re-
oped; pretarsus with large squamiform seta from duced to a pair of minute lobes. Haustellum
lateral base of claw (Fig. 6.9 I); coxae widely sep- moderately long, over 2.5X the length of labial
arated. Prolegs well developed on A36 and palpus; basal half scaled; distal half naked, scle-
AIO; crochets on A 3 - 6 biserial with outer series rotized, covered with overlapping plates. Maxil-
interrupted along mesal side by an elongate, lary palpi reduced to either 1 or 3 short seg-
crescentic brownish plate (Fig. 6.9 J). Anal plate ments. Labial palpi moderately short, 3-seg-
with 3 pairs elongate setae; crochets of AIO with mented, drooping, smooth scaled. Thorax: Meta-
ca. 10 hooks in a curved row. Larvae feed on furca (Fig. 6.6 U) with apophyses connected as a
various species of Proteaceae (e. g., Embothrium, bridge to secondary arms. Forewings (Fig. 6.5 L)
Lomatia in South America and Persoonia in Aus- lanceolate; index 0.17-0.28; microtrichia re-
tralia), at first mining in leaves and later living stricted to basal area of both wings; forewings
in shelters formed by webbing together adjacent usually with all 5 branches of R terminating on
leaves (Nielsen and Common 1991, Parra and costa; Rs2 sometimes absent; M 2-branched,
Ibarra - Vidal 1994 b). Pupa: Head with vertex base of M present, usually faint within cell; CuA
smooth. Wings extending almost to A5. Abdo- unbranched; CuP faint. 1A + 2A usually with-
men with a single, anterior row of small spines out basal fork; retinaculum a broad lobe aris-
on A 2 - 9 ; A 2 - 8 movable in . Cremaster con- ing in costal membrane immediately anterior to
sisting of a pair of large, dorsal spines on AIO. and tightly coiling under base of Sc. Hindwings
Pupation inside a loose shelter of plant material usually narrower than forewings, sometimes
spun together. Pupa partially extruded from broader in $ than in ; index 0.15-0.23; M and
shelter prior to emergence. Cu usually unbranched; discal cell open; $ frenu-
lum a single stout seta; with 2 closely set frenu-
The family currently includes ca. 60 species lar setae. Legs with tibial spur pattern of 0-2-
equally divided between southern South America 4; epiphysis present. Abdomen: 2S divided into
(Davis 1986) and Australia (Nielsen 1987; Com- 3 sclerites with caudal most sternite possessing
mon 1990). Five genera (Sesommata, 6 sp.; Meta- elongate lateral arms extending anteriorly (Fig.
phatus, 6 sp.; Plesiophatus, 1 sp.; Palaephatus, 6.7 R); postspiracular tergosternal connection
13 sp.; and Apophatus, 2 sp.) have been described present but weak. 2 pairs of tuberculate plates
from South America and two (Azaleodes, 4 sp. present, one on lateral margin of 2S, other near
and Ptyssoptera, 25 sp.) from Australia. Autapo- 2T; plates evident on most of caudal segments;
morphies for the Palaephatidae include the ter- wart-like protuberances apparently absent. Core-
minal, median ridge covered with campaniform mata and corethrogyne absent. Male genitalia:
sensilla of the ovipositor (Fig. 6.9 H) and possi- Uncus strongly bifurcate, often fused to short or
bly the mesal plate on the abdominal prolegs of moderately long tegumen. Vinculum variable, a
the larva (Fig. 6.9 J). The squamiform seta lat- very slender to broad ventral ring, U-V- or Y-
erad to the pretarsal claw on the larva (Fig. 6.9 I) shaped, saccus present or absent. Gnathos ab-
is an unusual feature also shared with some In- sent. Juxta sometimes present. Anellus weakly
curvariidae and Prodoxidae. Unlike most mono- sclerotized. Transtilla present, sometimes sepa-
trysian Heteroneura, adult male Palaephatidae rated. Valvae usually simple, broad to slender.
have also developed a diverse assortment of sex Aedoeagus usually extremely slender, with a
scale pockets, most of them located in various strongly bifurcate apex with gonopore at or near
regions of the wings (Fig. 6.5 K). 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-
TISCHERIOIDEA ing short, stout "peg setae" and a median, termi-
Tischeriidae (Fig. 6.2 H). Small moths with fore- nal, smooth ridge without campaniform sensilla
wings 2.75 mm length. Head (Fig. 6.3 M): Ves- (Fig. 6.9 G). T9 represented by a bilateral pair of
titure of vertex generally rough with slender to subdorsal, usually smaller, flatter and finely se-
moderately broad scales directed mostly for- tose lobes which articulate with a moderately
ward; frons smooth. Ocelli absent. Eyes large; in-
The Monotrysian Heteroneura 87

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- larval stages and actively spin silk in all instars.
row sclerotized ring which is constricted laterally Feeding may proceed in one direction (trumpet
where T8 is weakly joined to the greatly attenu- mine) or in all directions (irregular blotch mine).
ated anterior internal arms (i. e., apophyses) of Some species construct a firm, silk-lined nidus
S8; in some species the median portion of S8 is within the mine into which the larva often re-
extended slightly anteriorly (the patibulum of treats when alarmed or not feeding (Fig. 6.12 D).
Braun 1972) before dividing to form the apophy- As many as 4 broods may occur. Hosts include
ses. S9 extremely reduced to a minute ventral the Asteraceae, Combretaceae, Ericaceae, Faga-
band from which arise a usually short, slender, ceae, Malvaceae, Rhamnaceae, Rosaceae, Ster-
lateral and more ventral pair of apophyses as culiaceae, and Tiliaceae (Braun, 1972). Pupa:
well as a broader and more dorsal, internal pair. Head with vertex usually smooth, rarely with
Both pairs of apophyses were collectively re- frontal process. Wings to anterior margin of A4.
ferred to as the prela by Braun (1972). Oviporus A 3 - 7 movable in , A 3 - 6 in $ (Mosher 1916).
situated in membrane of vestigial S9. Vestibulum Concentrations of irregular, tergal spines on A2
often enlarged with thickened walls. Bursa copu- or 37; D1 and 2 approximate on the movable
latrix membranous, without signum but often segments. Cremaster consisting of 2 dorsal hooks
with minute, spinulose sclerotizations. on AIO. Pupation occurs within mine, usually in
Egg. Poorly known. Elliptic to broadly oval in a flimsy cocoon.
shape, laid singly and cemented to leaf surface. The Tischeriidae are a small, primarily holarc-
Larva (Figs. 6.1 H - J ) : Small, not exceeding tic family of ca. 80 species, all included in one
6 mm. Body slightly flattened, pale green to yel- genus, Tischeria. The greatest concentration of
low, with brownish head, pronotal and anal species occurs in the United States (47 sp.) with
plates. Head flattened; frontoclypeus and ecdy- only a few described from the Neotropical, Ethi-
sial lines converge at epicranial notch; 6 stem- opian, and Indo-Malayan regions, and none
mata, mostly horizontal. Thoracic legs greatly from the Australian region or Oceania.
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 Acknowledgments
rudimentary with crochets in multiserial bands I am indebted to Vichai Malikul and George
or incomplete ellipses. D1 and 2 of A36 closely Venable of the Department of Entomology,
approximate (Fig. 6.1 I), widely separated else- Smithsonian Institution, Washington, D. C.,
where. Larvae are leaf-miners throughout the USA and Biruta Akerbergs Hansen of Liverpool,
88 Donald R. Davis

Pennsylvania for the drawings used in this chap- - (1972): Careospina quercivora, a new genus and spe-
ter. Similarly, I wish to thank Victor Kranz and cies of moth infesting live oaks in California (Lepi-
Carl Hansen of the Smithsonian Photographic doptera: Incurvariidae). Proc. Ent. Soc. Washing-
Laboratory and Susann Braden and Walter ton, 74 (1): 121-128.
- (1986): A new family of monotrysian moths from
Brown of the Smithsonian Scanning Electon
austral South America (Lepidoptera: Palaephati-
Microscope Laboratory for their photographic
dae), with a phylogenetic review of the Monotrysia.
assistance. Ebbe S. Nielsen of CSIRO, Canberra,
- Smithson. Cont. Zool., Washington, 434:
Australia graciously provided some illustrations i-iv, 1-202.
and notes. - (1987): Micropterigidae, Eriocraniidae, Acanthop-
teroctetidae, Nepticulidae, Opostegidae, Tischerii-
dae, Heliozelidae, Adelidae, Incurvariidae, Prodox-
idae, Tineidae, Psychidae, Ochsenheimeriidae, Lyo-
Addicott, J. F., J. Bronstein & Kjellberg, F. (1990): netiidae, Gracillariidae, Epipyropidae. p. 341378,
Evolution of mutualistic life-cycles: yucca moths 456, 459, 460. In F. W. Stehr, ed., Immature Insects.
and fig wasps, 143-161. - In F. S. Gilbert (ed.), - Kendall/Hunt Pubi. Co., Dubuque, Iowa.
Insect life cycles: genetics evolution and coordina- - (1989): Generic revision of the Opostegidae, with a
tion. Springer-Verlag, London, Berlin, New York. synoptic catalog of the world's species Lepidoptera:
Becker, V. O. (1977): The taxonomic position of the Nepticuloidea). Smithson. Contr. Zool., Washing-
Cecidosidae. Brthes (Lepidoptera). - Polskie ton, 478: i - i i i , 1 - 9 7 .
Pismo Ent., 47: 7 9 - 8 6 .
- (in press): Biology of yucca moths (Lepidoptera:
Biezanko, C. M. (1961): Olethreutidae, Tortricidae,
Prodoxidae). In T. Hirowatari (ed.), Biology of
Phaloniidae, Aegeriidae, Glyphipterygidae, Ypono-
Lepidoptera [in Japanese], Osaka.
meutidae, Gelechiidae, Oecophoridae, Xylorictidae,
- , O. Pellmyr & Thompson, J. N. (1992): Biology and
Lithocoletidae, Cecidosidae, Ridiaschinidae, Acro-
Systematics of Greya Busck and Tetragma, new ge-
lophidae, Tineidae et Psychidae da Zona Suesta do
nus (Lepidoptera: Prodoxidae). Smiths. Cont.
Rio Grande do Sul. Arquivos Ent., (A) 13: 1 16.
Bland, K. P. (1977): Notes on the biology of the genus Zool., Washington, 524: 1 - 8 8 .
Adela (Lepid.: Incurvariidae). - roc. Brit. Ent. Donner, H. & Wilkinson, C. (1989): Nepticulidae In-
Nat. Hist. Soc. 10: 8 8 - 9 1 . serta: Lepidoptera). Fauna of New Zealand, Wel-
Borkowski, A. (1969): Studien an Stigmelliden (Lepi- lington, no. 16, p. 1 - 8 8 .
doptera). Teil 1. Zur Verbreitung, Biologie und Downes, J. A. (1969): The swarming and mating flight
kologie der Stigmelliden in den polnischen Sude- of Diptera. - Ann. Rev. Ent. 14: 2 7 1 - 2 9 8 .
ten. - Polskie Pismo ent., 34: 9 5 - 1 2 2 . Dugale, J. S. (1988): Lepidoptera - annotated cata-
logue, and keys to family-group taxa. - Fauna of
- (1972): Studien an Nepticuliden (Lepidoptera). Teil New Zealand, 14: 1 - 2 6 2 .
V. Die europischen Arten der Gattung Nepticula Dziurzynski, A. (1958): The mines of Lepidoptera in
Heyden von Eichen. Polski Pismo ent., 45: Cornus sanguinea L. and Vaccinium myrtillus L.
leaves with the particular consideration of mines
Braun, A. F. (1917): Nepticulidae of North America. and development of Incurvarla oehlmanniella Tr.
Trans. Amer. Ent. Soc., Philadelphia, 48: 155-209. Acta Zool. Cracov., Polska Akademia Nauk, 3(6):
- (1972): Tischeriidae of America north of Mexico 199-245.
(Microlepidoptera). - Mem. Amer. Ent. Soc., Phila-
Emmet, A. M. (1974): The early stages of Lampronia
delphia, 28: 148.
praelatella (Denis & Schiffermller) (Lepidoptera:
Brthes, J. (1916): Estudio fito-zoolgico sobre al-
Incurvariidae). - Ent. Ree., 86: 180-182.
gunos Lepidpteros Argentinos productores de
- (1976): Tischeriidae, p. 2 7 2 - 2 7 6 . In J. Heath (Ed.),
agallas. - An. Soc. Cient. Arg., 82: 113-140.
The moths and butterflies of Great Britain and Ire-
Brown, J. ., O. Pellmyr, J. . Thompson & Har-
land, vol. 1. - The Curwen Press, London.
rison, R. G. (1994): Phylogeny of Greya (Lepidop-
tera: Prodoxidae), based on nucleotide sequence Farris, J. S. (1988): Hennig86, version 1.5, Computer
variation in mitochondrial cytochrome oxidase I program distributed by its author.
and II: congruence with morphological data. - Frack, D. C., Jr. (1982): A systematic study of Prodox-
Mol. Biol. Evol., Chicago, 11(1): 128-141. ine moths (Adelidae: Prodoxinae) and their hosts
(Agavaceae), with descriptions of the subfamilies of
- (1994 b): Mitochondrial D N A phylogeny of the
Adelidae (s. lat.). Masters thesis, Calif. State
Prodoxidae (Lepidoptera: Incurvarioidea) indicates
Polytechnic Univ., Pomona, vii+209 p.
rapid ecological diversification of yucca moths.
Ann. Ent. Soc. Amer., Lanham, 87(6): 7 9 5 - 8 0 2 . Gerasimov, A. M. (1952): Lepidoptera, Part 1 Caterpil-
Common, I. F. B. (1969): A new genus Perthida for lars. Zool. Inst. Avad. Nauk USSR. New Ser.
the western Australian jarrah leaf miner P. glyphopa No. 56, Fauna USSR. Insects-Lepidoptera. 2(1):
sp. n. and Tinea phoenicopa Meyrick (Lepidoptera: 1 - 3 3 8 . [In Russian],
Incurvariidae). - J. Australian Ent. Soc., 8: 1 2 6 - Heath, J. & Pelham-Clinton, E. C. (1976): Incurvarii-
130. dae, p. 2 7 7 - 3 0 0 . In J. Heath (Ed.), The moths and
- (1990): Moths of Australia. Melbourne University butterflies of Great Britain and Ireland, vol. 1. - The
Press, Carlton, Victoria, vi+ 535 p. Curwen Press, London.
Curtis, J. (1835): On a species of moth found inhabit- Heppner, J. B. (1991): Faunal regions and the diversity
ing the galls of a plant near to Montevideo. of Lepidoptera. Trop. Lepid. 2 (suppl. 2): 185.
Trans. London Zool. Soc., 1: 312313. Hoare, R. J. B., R. Johansson, E. J. van Nieuker-
Davis, D. R. (1967): A revision of the moths of the ken, & E. S. Nielsen. 1997: Australian Nepticulidae
family Prodoxinae (Lepidoptera: Incurvariidae). (Lepidoptera): redescription of the named species.
U. S. Nat. Mus. Bull., Washington, 255: 1 - 1 7 0 . - Ent. scand., 28: 126.
The Monotrysian Heteroneura 89

Janse, A. J. T. (1945): Adelidae. - The Moths of South - (1912 b): Lepidoptera Heterocera (Tineae), Fam.
Africa, 4(2): 7 9 - 1 4 8 , pis. L X I I I - L X X X I . Adelidae. Genera insectorum 133, p. 1 - 1 2 ,
Jensen, A. D. (1932): Studier over Incurvarla koerner- 1 col. pi.
iella Zell. (Lepidoptera, Incurvariidae). - Kg I. Dan- - (1917): A jumping cocoon. - Ent. Mon. Mag., liii:
ske Vidensk. Selskab, Biol. Medd., 10: 1 - 4 9 . 62.
Johansson, R., E. S. Nielsen, E. J. van Nieukerken & Miles, N. J. (1983): Variation and host specificity in the
Gustafsson, B. (1990): The Nepticulidae and Opos- yucca moth Tegeticula yuccasella (Incurvariidae): a
tegidae (Lepidoptera) of west Europe. - Fauna Ent. morphometric approach. - J. Lep. Soc., 37(3):
scand., Leiden, New York, Copenhagen, 23(1-2): 207-216.
1-739. Mosher, E. (1916): A classification of the Lepidoptera
Kemperman, T. C. M. & Wilkinson, C. (1985): Japan- based on characters of the pupa. - Bull. Illinois
ese species of the genus Stigmella (Nepticulidae: State Lab. Nat. Hist., Urbana, 12(2): 1 - 1 6 5 .
Lepidoptera). - Ins. Matsum. (n. s.) 32: 1 107. Needham, J. G., S. W. Frost & Tothill, . (1928): Leaf-
KiefFer, J. J. & Jrgensen, P. (1910): Gallen and Gal- mining Insects. Williams & Williams Co., Balti-
lentiere aus Argentinien. - Centralbibl. Bakter., more.
Parasitent, 27: 363-444. Newton, P.J. & Wilkinson, C. (1982): A taxonomic
Kozlov, M. V. (1987): Moths of the subfamilies Incur- revision of the North American species of Stigmella
variinae and Prodoxinae (Lepidoptera, Adelidae) of (Lepidoptera: Nepticulidae). - Syst. Ent., 7(4):
the Far East, p. 14-24. In P. A. Ler, V. A. Kirpich- 367-463.
nikova & Kononenko, V. S. (eds.), Lepidoptera of Nielsen, E. S. (1980): A cladistic analysis of the holarc-
the Far East of U. S. S. R - Biological and Soil tic genera of adelid moths (Lepidoptera: Incurvari-
Institute, Vladivostok [in Russian]. oidea). - Ent. scand. 11: 161-178.
- (1996): Incurvariidae and Prodoxidae (Lepidop- - (1981): A taxonomic revision of the species of Allo-
tera) from Siberia and the Russian Far East, with clemensia n. gen. (Lepidoptera: Incurvariidae). -
descriptions of two new species. - Ent. Fennica, 1: Ent. scand. 12: 271-294.
55-62. - (1982): The maple leaf-cutter and its allies: a revi-
Kuprijanov, . V. (1994): Towards a natural system of sion of Paraclemensia (Incurvariidae s. str.). Syst.
the Incurvarioidea, Part 2: Procacitas gen. nov. for Ent. 7: 217-238.
Alloclemensia orientella Kozlov, 1987, with new - (1985 a): A taxonomic review of the adelid genus
data on the distribution (Lepidoptera, Incurvarii- Nematopogon Zeller (Lepidoptera: Incurvarioidea).
dae s. str.). - Atalanta, Mnnerstadt 23(3): 6 3 7 - Ent. scand. suppl. 25: 1 66.
642. - (1985 b): Primitive (non-ditrysian) Lepidoptera of
- (1992): Towards a natural system of the Incurvari- the Andes: diversity, distribution, biology, and phy-
oidea. Part 3. Excurvaria gen. nov. for Incurvarla logenetic relationships. - J. Res. Lepid., Suppl. 1:
praelatella ([Denis & Schiffermller], 1775) (Lepi- 1-16.
doptera, Incurvariidae). - Atalanta, Mnnerstadt, - & Davis, D. R. (1981): A revision of the Neotropi-
25: 391-398. cal Incurvariidae s. str., with the description of two
Kppers, P. V. (1977): Zur taxonomischen Neuorien- new genera and two new species (Lepidoptera: In-
tierung der Adelidae Deutschlands. Adeliden-Stu- curvarioidea). Steenstrupia, 7(3): 2 5 - 5 7 .
dien I. - Beitr. naturk. Forsch. SiidwDtl. 36: 2 3 9 - - (1985): the first southern hemisphere prodoxid and
246. the phylogeny of the Incurvarioidea (Lepidoptera).
- (1980): Untersuchungen zur Taxonomie und Phylo- - Syst. Ent., 10: 307-322.
genie der Westpalaktischen Adelinae (Lepido- Nielsen, E. S. & Johansson, R. (1980): Cauchas bre-
ptera: Adelidae). - Wissenschaft. Beitr. Karlsruhe, viantennella n. sp. from N W Europe and C. brun-
7: 1 - 4 9 7 . nella n. sp. from Uzbekistan, with a check-list of
Kuroko, H. (1961): The life history of Nemophora rad- the Palearctic Cauchas species (Lepidoptera: Adeli-
dei Rebel (Lepidoptera, Adelidae). Sci. Bull. Fac. dae). - Ent. scand. 11: 145-153.
Agri. Kyushu Univ. 18(4): 323-334, pis. 37, 38. Nieukerken, E. J. van (1985): A taxonomic revision of
Kyrki, J. (1983): Adult abdominal sternum II in ditry- the western palaearctic species of the subgenera
sian superfamilies - morphology and phylogenetic Zimmermannia Hering and Ectoedemia Busck s. str.
significance (Lepidoptera). - Ann. Ent. Fenn., 49: (Lepidoptera, Nepticulidae) with notes on their
89-94. phylogeny. Tijds. v. Ent., 128: 1 - 1 6 4 .
McAlpin, J. F. & Munroe, D. D. (1968): Swarming of - (1986): Systematics and phylogeny of holarctic gen-
Longchaeid flies and other insects, with descrip- era of Nepticulidae (Lepidoptera, Heteroneura:
tions of four new species of Longchaeidae (Dip- Monotrysia). - Zool. Verhand., Leiden, no. 236,
tera). - Can. Ent. 100: 1154-1178. p. 1 - 9 3 .
McGiffen, K. C. & Neunzig, H. H. (1985). A guide to Parra, L. E. & Ibarra-Vidal, H. (1994 a): Nuevo
the identification and biology of insects feeding on esqueletizador del "coihue" (Nothofagus dombeyi)
muscadine and bunch grapes in North Carolina. en Chile, Basileura osornoensis sp. n., (Lepidoptera:
N. C. Agr. Res. Service. Bull. 470, 93 p. Incurvariidae). - Trop. Lep., 5(1): 5764.
Mani M . S . (1964): Ecology of plant galls. - Mono- - (1994 b): Description de los estados immaduros y
graphiae Biologicae, Vol XII, xii + 434 p., Dr. W. notas biolgicas sobre Metaphatus ochraceus (Lepi-
Junk Publishers, The Hague. doptera: Palaephatidae), defoliator del notro (Em-
Meyrick, E. (1909): New South African Micro-lepidop- bothrium coccineum). - Rev. Chilena Ent., 21: 7 7 -
tera. - Ann. South Afr. Mus., 5: 349-379. 84.
- (1912 a): Adelidae, Micropterygidae, Gracilariadae. Petersen, G. (1978): Zur systematischen Stellung der
- Lepidopterorum Catalogus, pars 6, p. 2 5 - 6 8 . Gattung Crinopteryx Peyerimhoff, 1871. (Lepido-
90 Donald R. Davis

ptera: Incurvariidae). Beitr., Eni., Berlin, 28 (2): Ross, D. A. (1958): The maple leaf cutter, Paraclemen-
217-220. sia acerifoliella (Fitch) (Lepidoptera: Incurvarii-
Powell, J. A. (1984): Biological interrelationships of dae), descriptions of stages. Canadian Ent., 90:
moths and Yucca schottii. - Univ. Calif. Pubi. Ent., 541-555.
100: 1 - 9 3 . Rozefelds, A . C . (1988): Lepidoptera mines in
- (1992): Interrelationships of yuccas and yucca Pachypteris leaves (Corystopermaceae: Pterido-
moths. - TREE, Cambridge, 7(1): 10-15. spermophyta) from the Upper Jurassic/Lower Cre-
- and R. A. Mackie (1966): Biological interrelation- taceous Battle Camp formation, North Queens-
ships of moths and Yucca whipplei. - Univ. Calif. land. - Proc. R. Soc. Qd. 99: 7 7 - 8 1 .
Pubi. Eni., 42: 1 - 5 9 . Scoble, M . J . (1980): a New incurvariine leaf-miner
Puplesis, R . K . (1984a): sisteme molej-maljutok from South Africa, with comments on structure,
(Lepidoptera, Nepticulidae) palearkticeskoj fauny. life-history, phylogeny, and the binominal system
- Ent. Obozr., 63: 582-597. of nomenclature (Lepidoptera: Incurvariidae). - J.
- (1984 b): Obzor vidov roda Microcalyptris (Lepi- Ent. sth. Afr., 43 (1): 7 7 - 8 8 .
doptera, Nepticulidae) iz pystyn Mongolii i SSSR - (1983): A revised cladistic classification of the Nep-
(A review of the nepticulid moths of the genus ticulidae (Lepidoptera) with descriptions of new
Microcalyptris (Lepidoptera, Nepticulidae) from taxa mainly from South Africa. - Trans. Mus.
the deserts of Mongolia and the USSSR). - Nasek- Monog. no. 2, Pretoria, xi + 105 p.
omye Mongolii, 9: 484-507. Wagner, D. L. & Powell, J. A. (1988): A new Prodoxus
- (1985): Contributions to the classification of the from Yucca boccata: first report of a leaf-mining
palaearctic Nepticulidae (Lepidoptera). - Ent. Prodoxine (Lepidoptera: Prodoxidae). - Ann. Ent.
Rev., Washington, 63: 149-164. [English transla- Soc. Amer., 81: 547-553.
tion of Puplesis 1984 a]. Wiegmann, . M. (1994): The earliest radiation of the
- (1994): The Nepticulidae of eastern Europe and Asia: Lepidoptera: Evidence from 18S rDNA. - Ph. D.
western, central and eastern parts. 299 p. Leiden. thesis, Univ. Maryland, College Park, ix + 230 p.
Razowski, J. & Wojtusiak, J. (1978 a): Family-group Wilkinson, C. & Scoble, M . J . (1979): The Nepticuli-
taxa of the Adeloidea (Lepidoptera). - Polskie dae (Lepidoptera) of Canada. - Ent. Soc. Can., Ot-
Pismo Ent., 48: 3 - 1 8 . tawa, Mem. 107, p. 1 - 1 2 9 p.
- (1978 b): Polish genera of the Adelidae (Lepidop- Wille, J. (1926): Cecidoses eremita Curt, und ihre Galle
tera). - Polskie Pismo Ent., 48: 19-33. an Schinus dependens Ortega. Z. Morph. Okol.
Riley, C. V. (1892): The yucca moth and Yucca pollen- Tiere, 7: 1 - 1 0 1 .
ation. Missouri Bot. Gard., Third Ann. Rept., Zimmerman, E. C. (1978): Microlepidoptera. - Insects
p. 9 9 - 1 5 8 . 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. 1983); a minute pair of warts (Fig. 7.4 B) usually

The Tineoidea are generally recognized to repre- near caudal end of rods; two pairs of small tu-
sent the most plesiomorphic group of ditrysian berculate plates present in pleura; ventral pair
Lepidoptera; until 1988 the superfamily included typically near or on margin of S2; dorsal pair
the groups now assigned to the Gracillarioidea. near T2; plates often repeated but progressively
The principal autapomorphy of the Tineoidea is smaller on posterior segments. Male genitalia:
the presence of a slender pair of ventral pseuda- Valvae with elongate, internal apodemes; 7 pairs
pophyses within AIO of most females (Robinson of genital muscles present (Kuznetsov and Stek-
1988; Davis 1988); the telescopic 'ovipositor' ol'nikov 1976, 1977). Female genitalia'. Oviposi-
itself is unusually elongated. Other characteris- tor often greatly extensible (Fig. 7.4 M), usually
tics useful in distinguishing Tineoidea from with 2 elongate pairs of apophyses, sometimes
Gracillarioidea and most ditrysian Lepidoptera reduced to 1 short posterior pair (Acrolophidae,
are the presence of erect scales on the frons, la- Fig. 7.4 L, some Psychidae); a ventral pair of
bial palpi with lateral bristles, and haustellum slender pseudapophyses usually present within
with short, disassociated galeae (Dugdale 1988). AIO (Bradley 1951); $ of Eriocottidae with an
Head: Generally rough, with mostly erect pili- additional pair (rarely 2 pairs) of dorsal pseuda-
form scales (Fig. 7.3 B). Ocelli usually absent, pophyses within A8 (Fig. 7.4 M, N); ostium
present in a few Eriocottidae and Psychidae. An- opening on S8.
tennae usually less than length of forewing; scape Egg: Oval, flat type, with micropylar axis par-
with pecten present or absent; flagellum usually allel or subparallel to oviposition substrate;
filiform, occasionally pectinate in $ Eriocottidae, chorion relatively smooth to finely reticulated,
Acrolophidae, Psychidae, and $ Arrhenophani- micropylar end finely reticulated. Eggs usually
dae; usually 1 row of slender scales per segment, deposited externally and singly, sometimes in
rarely 2, either encircling segment or restricted mass. Larva: Often living within tube or portable
to dorsal half. Chaetosemata absent. Pilifers and case (Fig. 7.6 A - N ) ; all instars with fully devel-
mandibles either present or absent. Haustellum oped legs and 5 pairs of prolegs; ventral crochets
naked, usually reduced, loosely joined and un- usually in a uniserial circle, ellipse, or penellipse.
coiled. Maxillary palpi 5-segmented in many Tin- Pupa: Adecticous, incomplete, with appendages
eidae and in Crepidochares (Eriocottidae), 4-seg- variably appressed to body and with 3 - 6 mov-
mented to absent in all other genera. Labial palpi able abdominal segments. Maxillary palpi usu-
usually 3-segmented, with stout, erect, piliform ally present. Abdomen with tergal spines usually
scales (bristles) from apical and lateral surfaces in rows, sometimes in patches. Pupa extruded
of second segment (Fig. 7.3 B); bristles absent in partially from cocoon prior to adult eclosion.
Arrhenophanidae and most Acrolophidae and Six families are here recognized within Ti-
Psychidae. Thorax: Metafurca with apophyses neoidea. Family interrelationships were dis-
usually free, joined to secondary arms in Arrhen- cussed by Robinson (1988) and Robinson &
ophanidae and Psychidae. Forewings moderately Nielsen (1993), who tentatively presented them
broad, index (W/L) 0.25-0.47, with Rs4 usually as Tineidae + (Eriocottidae + (Acrolophidae +
terminating on costa or apex; Rs4 to termen in (Psychidae + Arrhenophanidae))). The Lypusi-
Psychidae, and most Arrhenophanidae and Erio- dae have subsequently been removed from syn-
cottidae. Chorda with accompanying accessory onymy with the Psychidae. Possible synapomor-
cell present or absent. Base of M usually present phies of the non-tineid families are the strong
and either simple or forked and forming interca- tendency of the antennae to bipectination, the
lary cell. 1A + 2 A usually with basal fork. Spe- reduction of the maxillary palp (which has at
cialized subanal interlocking microtrichia pre- most 3 segments except for some Eriocottidae),
sent. Hindwing index 0.20.55; base of M usu- forewing with Rs4 terminating on the termen
ally preserved, either simple or forked; discal cell rather than on the costa, male retinaculum aris-
closed. Abdomen: Tergal spines absent. A2 with- ing from wing membrane between C and Sc
out tergosternal connection. Tympanal organs rather than on Sc, female frenulum with more
absent except on S2 of Harmocloninae (Tine- than 4 bristles, apex of sacculus in male genitalia
idae). Anterior margin of S2 (Figs. 7.4 AE) with one or more thorn-like sensilla and spinose
with a pair of slender apodemes which usually membranous lobe ('pulvillus') on inner valve sur-
continue caudally into S2 as sternal rods (Kyrki face. Possible autapomorphies of the acrolophid/
92 Donald R. Davis & Gaden S. Robinson

psychid/arrhenophanid assemblage (to which the Tineidae. Very small to medium-sized moths,
Lypusidae apparently do not belong) are the ab- forewings 2.5 to more than 25 m m long (Figs.
sence of an antennal pecten and the fusion of the 71. AL). Features for rapid recognition of
metafurcal apophyses with the secondary furcal many species are: head vestiture of erect piliform
arms, forming a bridge-like structure; on the scales; labial palpi with stiff lateral bristles; pro-
other h a n d it is now k n o w n that distinct, albeit boscis very short or even absent, never strongly
short, galeae are indeed retained in the psychid sclerotized, the galeae only loosely associated
ground plan (Fig. 7.3 H). Separation of the Arr- (Fig. 7.2 A); hind tibiae with erect and elongate
henophanidae f r o m the Psychidae has been ques- scales on upper surface; adults resting with wings
tioned (Robinson 1988), but they are pheneti- raised tent-like over the body and with body par-
cally distinct (see account below) and the separa- allel to substrate, moving with characteristic
tion is here upheld. scuttling run.
Head vestiture of erect piliform scales on occi-
put, vertex and frons; scales grouped into tufts,
Key to the tineoid families (Lypusidae not
forming more or less conspicuous whorls; head
with semi-appressed forward-directed scales in
1 Forewings with Rs4 terminating on costa. (e. g.) Setomorpha, with appressed plate-like
Antennae filiform. Maxillary palpi 5-seg- scales in Hieroxestinae (Fig. 7.1 L). Head with-
mented in most species Tineidae out externally visible ocelli. C o m p o u n d eye un-
Forewings with Rs4 usually terminating at modified, with facets of regular size; interocular
termen or apex. Antennae often bipecti- index 0.50 (some Machaeropteris) to 2.20 ( M e t -
nate. Maxillary palpi usually 4-segmented apherna).
or less 2 M o u t h p a r t s (except galeae) fully developed in
2 Antennal pecten present. Metathoracic many species (Fig. 7.2 A), but reduction fre-
furca free; furcal apophyses arising f r o m quent. Pilifers large, or reduced; mandibles usu-
lateral ridges of metafurcal stem (Fig. ally large, together often as wide as frons, but
7.4 H). 9 with 1 (fused) or 2 short, pseuda- absent in some groups. Maxillary palpi more
pophyses within dorsal wall of A8 (Figs. strongly developed than in any other ditrysian
7.4 M - N ) Eriocottidae group, but reduction occurring frequently, corre-
Antennal pecten usually absent. Dorsal lated with reduction of the galeae. Fully devel-
pseudapophyses absent within A8 of oped maxillary palpi 5-segmented, folded; f o u r t h
Metathoracic furca usually joined as a and fifth segments with densely packed sensilla
bridge to secondary furcal arms (Figs. trichodea interspersed with microtrichia, fifth
7.4 J, K), or with only tendon of vestigial segment usually with conspicuous apical and
bridge arising f r o m secondary arms (Fig. subapical processes bearing sensilla basiconica.
7.41) 3 Galeae short, disassociated, not strongly sclero-
3 abdomen without corethrogyne. Oviposi- tized and, if rolled, rolled haphazardly; elongate
tor short, less than length of enlarged S8 and apparently forming a functional proboscis in
(Fig. 7.4 L); anterior apophyses absent. only a very few genera. Labial palpi invariably
Metathoracic bridge vestigial, broken (Fig. 3-segmented, in some Hapsiferinae and Hier-
7.41). Larva with lateral pinacula partially oxestinae distinctly recurved, drooping in some
separated f r o m p r o n o t u m (Fig. 7.5 M) . . Meessiinae; second segment in most genera with
Acrolophidae u p to 15 coarse lateral bristles arranged in irreg-
ular line and with terminal whorl of u p to 15
9 abdomen with corethrogyne on A7. Ovi-
bristles, usually with ventral brush of scales but
positor usually elongate, exceeding length
smooth-scaled in (e. g.) Opogona; terminal seg-
of more narrow S8; anterior apophyses
ment slightly to strongly flattened in m a n y taxa
usually present (sometimes absent and ovi-
and often swollen (e. g., Tiquadra), most genera
positor short in apterous ). Metathoracic
with conspicuous vom Rath's organ. Antenna 0.3
furca forming a bridge. Larva with lateral
to 1.3 length of forewing; scape smooth-scaled in
pinacula either separated or fused with
most genera, usually stout but distinctly slender
pronotum 4
and elongated in Erechthiinae and Hieroxesti-
4 $ genitalia with non-retractile vesica and
nae; pecten present in m a n y genera, consisting of
slender, tubular anellus. $ often with bipec-
two or three to about forty stout bristles; flagel-
tinate antennae. Larva with lateral pina-
lomeres cylindrical or approximately barrel-
cula widely separated f r o m p r o n o t u m (Fig.
shaped, conspicuously ciliated to finely pubes-
7.5 N ) Arrhenophanidae cent, females with fewer and shorter cilia than
$ genitalia with retractable vesica; anellus males, flagellomeres in most genera completely
usually m e m b r a n o u s or with separated lat- covered by one or, rarely, two annuii of overlap-
eral lobes. 9 antennae filiform or vestigial. ping scales.
Larva with lateral pinacula containing
spiracle and completely fused to p r o n o t u m Thorax: Legs usually smooth-scaled except for
(Figs. 7.5 J, O) Psychidae 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 Donald R. Davis & Gaden S. Robinson

on upper surface; foretibial epiphysis present in or specialised scaling on opposing wing surfaces
most genera; tibial spur pattern invariably 0-2-4. in Gerontha and Harmaclona.
Wings typically moderately broad (forewing Abdomen with S I vestigial; I a quadrangular
index 0.25-0.30, hindwing index 0.20-0.25), frame; without prespiracular bar connecting S II
rarely very narrow. Both wings generally sub- and I; S II with pair of anterior apodemes but
ovate. Forewing apex produced and upturned in usually lacking additional sclerotization (venu-
some Erechthiinae; termen usually convex; with lae) along margins; S II with tympanal organs in
a wide variety of forewing patterns, from uni- Harmaclona and Micrerethista, with hair-pencils
form dirty brown of some Tinea (Figs. 7.1 J) to in male Comodica and Dryadaula. Male abdo-
striking metallic transverse strigulae of some men in scattered genera with various types of
Moerarchis (Fig. 7.1 B); complicated patterns in scent-organs in various positions; many genera
some Erechthias make use of particoloured with shallow coremata in pleural membrane of
scales; pattern elements transverse, rarely longi- eighth segment; terminal segmentation highly
tudinal; bright coloration may provide mimetic modified in Setomorpha and Dryadaula. Func-
resemblance, disruptive patterning, or possibly tional and evolutionary aspects of male genital
aposematism. Marked sexual dimorphism rare, musculature are discussed by Kuznetsov & Ste-
e.g. Timaea (pattern), Gerontha (Fig. 7.1 E) kolnikov (1976, 1977), Robinson (1981) and Ste-
(shape), Setomorpha (venation). Hindwing un- kolnikov & Kuznetsov (1995). Female abdomen
patterned. Scales of forewing often in rough or with terminal corethrogyne in some genera (e. g.,
raised patches, some taxa with specialised sex- Tinissa); corethrogyne invaginated into three in-
tersegmental pouches in Perissomasticinae. Male
scales. Typically with full complement of free
genitalia of diverse form; vinculum and tegumen
forewing veins; Sc invariably present, but
not differentiated, saccus usually present, rarely
strongly displaced anteriorly in Dryadaula and
bilobed; uncus usually simple and bilobed, but
Eudarcia; all branches of Rs terminating anterior
articulated and hook-like in Tineinae; gnathos
to or at apex, with rare exceptions; pterostigma
usually present, absent in Scardiinae; valval apo-
developed sporadically; R absent in Crypsithyris,
demes elongate, robust; aedoeagus usually sim-
absent or vestigial in Opogona and Amphixystis;
ple and cylindrical, ductus and bulbus ejacula-
absence of any other branch of R/Rs rare, but
torius entering and connecting at base, thus with-
Rs3 occasionally lost; stalking of Rs3 + 4 or out coecum penis. Female with ovipositor typi-
Rs2 + 3 frequent, Rsl always free; more compli- cally elongate and telescopic with soft and setose
cated stalking sequences occur, involving Rs2, papillae anales, with ventral pair of sclerotized
Rs3, Rs4 and, in some cases, M l ; all three M rods set in membrane; armoured piercing ovipos-
branches usually present, but one may be absent; itor developed in a few genera (e. g., Tiquadra);
stalking of Ml + 2, M2 + 3, and M3 + CuAl oc- eight tergum and sternum typically separate, ter-
curs, also loss (or fusion) of either CuAl or gal margins extending anteriorly to articulate
CuA2; CuP present in all taxa, but tubular only with apophyses anteriores which invaginate from
proximally and distally; Al and A2 typically anteriorly extended corners of sternum; ostium
forming a broad basal loop but fused for distal bursae invariably within margins of eighth ster-
two-thirds; A2 weak in many genera, rarely un- num; bursa copulatrix usually differentiated into
developed; venational modification occurs in ductus and corpus bursae; ductus seminalis typi-
some genera (e. g., Monopis) to accommodate a cally connected to ductus bursae close to ostium;
hyaline spot. Hindwing venation typically com- signa present in a few genera.
plete, all veins free; Se + R and Rs separate and
subparallel; Sc + R displaced anteriorly in Eud- Egg of 'flat' type, with micropylar axis parallel
arcia; stalking of Ml + 2 common; M3 lost in or subparallel to oviposition substrate; chorion
some genera; full complement of posteriormost minutely wrinkled and in some taxa with coarser
veins CuP, 1A + 2A and 3 A expressed only ridging, either longitudinal ribbing or polygonal.
in large species with broad hindwings; 3 A absent Larva with few specializations (Figs. 7.5 AB,
in most groups with narrow hindwings. Female H, L), thorax not as strongly sclerotized as in
retinaculum consisting of a few semi-erect scales Psychidae, with only primary setae present; most
on Cu. Male retinaculum typically an elongately taxa with ecdysial lines of head extending be-
triangular tongue with a rolled apex arising from yond apex of frontoclypeus and terminating at
Se, but exhibiting considerable variation epicranial notch; frontoclypeus large, extending
broad and shallow lobe from Se or from mem- almost to epicranial notch; with six or fewer
brane between costa and Se, or triangular tongue pairs of stemmata and three pairs of epipha-
from spur from Se, or absent. Male frenulum a ryngeal setae; hypostoma either narrowly sepa-
single bristle, thickened and spatulate in a few rated or contiguous. Thorax with prespiracular
taxa; female frenulum typically consisting of two L sclerite typically separated from pronotum
or three bristles, but varying from one (e. g., which is usually distinct; prothoracic L-group
Coryptilum) to about 15 (Scardia). Supplemen- trisetose (bisetose in Scardiinae); three pairs of
tary wing-coupling devices involve microtrichia thoracic legs present, with precoxal plates rarely
fused; four pairs of ventral prolegs and one pair
The Tineoidea and Gracillarioidea 95

of anal prolegs; crochets invariably present on peratures close to the soil surface. Long tubes
prolegs, arranged in uniserial circle or ellipse that ramify through the lichen host are built by
(mesal penellipse in Oinophila); D1 and D2 pre- larvae of Infurcitinea. Short, fixed tubes are built
sent on all segments with D2 ventral; XD1, XD2 as shelters for ecdysis by certain Monopis and
and SD forming an almost straight vertical line; Tinea species. Fully portable tubular cases of
SV group bisetose but on second and third seg- varying degrees of sophistication are built by
ments it is unisetose in all groups except Tinei- many Tineinae and Meessiinae.
nae. Abdomen with D1 and D2 developed on Pre-pupation behaviour involves firstly move-
all segments, D2 caudal to DI; SD setae on all ment to the pupation site. Portable cases are sus-
segments but SD2 absent from A9; L group tri- pended well away from the food substrate prior
setose; SV group trisetose to A6, bisetose on re- to pupation. Case- or tube-dwellers pupate in the
maining segments (A9 unisetose in Tineinae); case or tube; free-living species spin a simple co-
anal plate with four pairs of setae; dermal cuticle coon; case- or tube-dwelling species then clean
with numerous fine microtrichia. Larvae of the out their refuge and seal it with a few strands of
termite commensal Acridotarsa are exceptional silk. Internal feeders partially cut out an opercu-
(see Harris, 1968). lum in the surface of the fungus or wood in
Pupa adecticous, obtect; at least two pairs of which they have fed and then spin a cocoon close
coxae visible; first and second abdominal seg- to this exit.
ments fused in both sexes; 7th, 8th and 9th seg- Eclosion involves considerable movement by
ments fused in females, 8th and 9th in males. the pharate adult; the pupa is wriggled and its
Typically with anterior band of posteriorly-di- abdomen contracted and expanded to bring the
rected spines on segments 39, additional pos- spines of the abdominal segments to bear against
terior band of spines on segments 36 in females the walls of the case or cocoon. Once the cocoon
and 3 7 in males; this pattern may be consider- is ruptured, the pupa is protruded clear for about
ably modified. Ventral abdominal surface with half its length and the adult emerges.
broad fields of posteriorly-directed microtrichia. Some Tineidae have been recorded as forming
Biology. Instead of occupying the typical di- mating swarms, notably Nemapogon and some
trysian leaf-feeding role, practically all tineid lar- Scardiinae; at least one Monopis needs several
vae are fungivorous in the sense that their food cubic metres of free space for successful court-
comprises a substantial proportion of fungal tis- ship. Gyratory dance has been recorded in sev-
sue. Tineid larvae have traditionally been catego- eral Tineidae, but this may not have a courtship
rised as detritophagous, lichenivorous or fungiv- function. Female pheromones of Tineidae have
orous. Detritophagy involves tineids in a wide been little studied but responses have been re-
range of niches in a wide range of biological sys- corded to octadecenyl acetate derivatives, nota-
tems with habits ranging from the mundane to bly E3Z13 18Ac; two pheromones have been
the bizarre, from feeding on fallen leaves to feed- isolated from Tineola, E2-18A1 and E2E13-
ing on guano in bat-caves, on mammal corpses, 18A1.
on insect remains in or below spiders' webs, and Oviposition may involve insertion of eggs into
on feathers in birds' nests. Some detritophagous crevices (by species with soft ovipositors) or pen-
Tineidae are commensals, sharing food resources etration of host tissue (by species with piercing
with social Hymenoptera, Isoptera, and with ovipositors); eggs may be protected with fine
mammals including man. Synanthropic species hair or scales from the corethrogyne or abdomi-
include pests of stored food products and of fur- nal apex glued to the pool of mucilage surround-
nishings and fabrics made from wool, fur or ing each egg.
feathers. There is strong correlation between the Larviparity is known in numerous Andean
higher taxonomic category to which a tineid be- Tinea and Indo-Australian Monopis species; fe-
longs and its larval food. cundity is traded off against egg mortality. Mo-
While many tineid larvae build portable cases nopis with conventional reproductive systems
from which they feed (Figs. 7.6 AC), in a man- carry 150200 mature and immature ova
ner analogous to that of Psychidae, case-building whereas larviparous species produce 4060
is by no means consistent within genera or even fully-developed first-instar larvae which are ma-
within species, and the definition of 'case' is tured in the grossly enlarged common oviduct.
blurred by the range of practices observed Classification and information sources. The
among Tineidae. The simplest structure is no family Tineidae encompasses some 3000 de-
more than a tubular cocoon constructed at the scribed species in 320 genera. The group may not
beginning rather than at the end of larval life. be monophyletic and represent a grade rather
The tube may be attached to the substrate or than a clade (Robinson 1988; Robinson & Niel-
form a tunnel through it. Trichophaga swinhoei sen, 1993). Fifteen subfamilies are currently rec-
feeds on weathered carnivore faeces in the Horn ognised. Of these, 13 are probably monophyletic.
of Africa and builds a 4 cm vertical tube within Relationships between subfamilies are still ob-
which it lives; the tube may provide not only scure. Some 150 genera have yet to be attributed
protection but a means of escaping extreme tem- 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 97

be synonyms of genera whose systematic posi- in the Old World; some New World groups may
tion is already known. For a broad and detailed also be attributable to the monophyletic 'core' of
review of the classification and biology of Tine- this subfamily. Larvae feeding externally, or
idae, see Robinson & Nielsen (1993). tube-dwelling, on plant detritus.
Present knowledge of Tineidae on a worldwide Harmacloninae. This group which was in-
scale is patchy, and most recent work has been cluded in the myrmecozeline assemblage by Rob-
restricted to particular geographical regions. inson & Nielsen (1993) has been given subfamily
There are checklists for the Nearctic (Davis status by Davis (in press a). Autapomorphies:
1983), Neotropical (Davis 1984), Oriental (Rob- Unique wing coupling system consisting of
inson & Tuck, 1996) and Australian (Robinson & curled, rigid scales along ventral hindmargin of
Nielsen 1993) regions; the checklist of New forewing interlocking into straight, rigid scales
Zealand Tineidae (Dugdale 1988) is cross-refer- along dorsal subcostal area of hindwing; pretar-
enced to the colour illustrations of Hudson sus lacking arolium and pseudempodial seta;
(1928; 1939). The last monographic treatment of paired tympanic organs present on abdominal
the Nearctic Tineidae was the unillustrated work sternum II; enlargement of sternal apophyses on
of Forbes (1923); this, in combination with the abdominal segment II; and aedoeagus with a
earlier monograph by Dietz (1905), is neverthe- basal, midventral keel.
less useful. More than 100 papers on the Tine- 2 genera (Harmaclona, Micrerethista) and 22
idae of the Palaearctic region have been pub- species. Pantropical distribution, most diverse in
lished since 1955 by A. K. Zagulajev, G. Petersen the Old World, especially southeast Asia and
and R. Gaedike (see Zoological Record, passim). Australia. Biology poorly known; a single rear-
The contents of very many of these papers were ing record reports larvae boring in logs of Bu-
summarized by Zagulajev (1960; 1964; 1973; chanania latifolia (Anacardiaceae; Fletcher
1975; 1979). Afrotropical Tineidae have been 1933). The Harmacloninae are the only group of
treated by Gozmny & Vri (1973); this work microlepidopterans (in a restricted sense, exclud-
should be used in conjunction with that of Janse ing cossuids and pyraloids) known to have devel-
(1968). For the Oriental region, figures of adults oped tympanic organs (discussion and illustra-
of some Indian species with descriptions of life tion in 211).
histories were given by Fletcher (1921; 1933); the Meessiinae. Autapomorphies: none. The group
Oriental Nemapogoninae, predominantly Indian was considered by Nielsen & Robinson (1993)
species, were reviewed by Robinson (1980 a). unlikely to be monophyletic.
Robinson & Nielsen (1993) provide a detailed Constituent species are small, often narrow-
treatment of the Australian Tineidae. Clarke winged; antennal pecten frequently lacking; fore-
(1971; 1986) and Zimmerman (1978) dealt with wing pattern simple, often consisting of large
spots or blotches, or transverse bands. Most spe-
the Tineidae of the islands of the eastern Pacific;
cies with reduced venation; forewing with Rs
the latter work contains detailed reviews of bio-
weak and with loss or stalking of Rs and M
logical information. Robinson (1979) revised the
branches, the cell narrow and open; hindwing
case-bearing clothes-moths, and the cave-dwell-
with Rs basally weak and one M branch lost, cell
ing species of Tineidae (Robinson, 1980 b). The
narrow and open, anal venation reduced and 3 A
same author reviewed the fungivorous group
rarely present. Male genitalia of many species
Scardiinae in a series of papers (Robinson 1976;
with uncus and gnathos reduced or absent.
1981; 1986). Robinson & Tuck (1997) have re-
Larva lichenivorous.
viewed the Hieroxestinae. Hinton (1956) gave
23 genera (Homosetia, Baeophylla, etc.) with
detailed descriptions of larvae of a number of
200 species; a further 2 0 - 2 5 genera, most mono-
species. G. Chauvin and collaborators (1960 to
basic, have been doubtfully attributed to this
date) have published numerous papers on tineid subfamily. Worldwide distribution but diversity
physiology (see Zoological Record, passim), dominated by the Palaearctic species of Eudarcia
other recent works are reviewed by Robinson & (= Meessia) and Infurcitinea. Larvae feeding ex-
Nielsen (1993). ternally or within tube galleries on lichen.
Myrmecozelinae (= Haplotineinae, Phthoro- Dryadaulinae (= Archimeessiinae). Autapo-
poeinae, Rhodobatinae). Autapomorphies: none. morphies: transfrontal suture of head reflexed
The group was considered by Robinson & Niel- dorsally, inverted V-shaped; labial palpus
sen (1993) to be polyphyletic but might contain broadly spatulate, lacking terminal bristles; fore-
a monophyletic core defined by: male genitalia wing venation with Sc displaced anteriorly, R
with aedoeagus strongly curved and basally arising close to base, A2 weak or absent; hind-
boot-shaped; anterior margin of vunculum re- wing with M3 or CuA2 absent, 3 A absent; male
curved dorsad. retinaculum a broad and very shallow lobe aris-
Constituent species are generally large and ro- ing from Sc; female frenulum a single bristle;
bust and cannot be included in any other sub- male frenular bristle sharply angled; abdominal
family. terga I and II entirely fused; male with terminal
36 genera (Myrmecozela, Cephimallota, Rho- abdominal segmentation strongly reduced and
dobates etc.) with 180 species. Widely distributed 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 99

ment incorporated into genitalia which are mandibles, prolegs with band of minute recurved
strongly modified and asymmetrical, gnathos ab- spines above crochets on A3 to A6.
sent; female with ovipositor strongly attenuated; Maxillary palpi and galeae are fully developed
apophyses anteriores rudimentary or absent. in Prosetomorpha (Davis, 1996) or vestigial in
One genus {Dryadaula) with 35 species. Lindera and Setomorpha (Robinson & Nielsen
Worldwide distribution but speciose only in Neo- 1993); the larva has six pairs of stemmata in
tropical region and Australia. Larva feeding ex- Prosetomorpha (Davis, 1996) but none in Lindera
ternally on lichen or fungus. and Setomorpha.
Scardiinae (= Tinissinae). Autapomorphies: 3 genera (Prosetomorpha, Setomorpha and
male genitalia invariably lacking gnathos; larva Lindera) with 9 species. Two cosmopolitan pest
with only two prothoracic L-group setae; D1 se- species with worldwide distributions, remaining
tae more widely separated than D2 setae on Al taxa Neotropical. Larvae detritophagous, feed-
to A8; L3 absent from A9; mandible with small ing externally or burrowing in loose substrates
ventral tooth and six or more distal teeth. or (Prosetomorpha) inhabiting borings in plant
24 genera (Scardia, Morophaga, Tinissa etc.) stems made by Coleoptera, on plant or arthro-
with 112 species (Fig. 7.1 A). Worldwide distribu- pod remains.
tion, most speciose in Indo-Australian region but Perissomasticinae. Autapomorphies: forewings
with greatest generic diversity in Neotropics. usually glossy and unicolorous; labial palpi
Larva boring in bracket-fungi or in bark or short; male genitalia with gnathos absent,
wood permeated by fungal mycelia. aedoeagus articulating with vinculum, bulbus
Nemapogoninae. Autapomorphies: male geni- ejaculatorius hypertrophied; female with cor-
talia with gnathos transverse and appressed to ethrogyne invariably present on intersegmental
diaphragma with membranous connection for membrane between A7 and A8, invaginated into
most of its length; valva with costa rolled inward one dorsal and a pair of lateroventral pouches.
and invariably cleft longitudinally, dorsal lobe 16 genera (Perissomastix, Edosa, Neoepiscar-
often a flexible, hirsute digitate process; valvae dia etc.) with 200 species. Distribution restricted
fused or articulated at ventral margins; juxta to the Old World tropics and subtropics, with
fused with inner surfaces of valvae and, with greatest diversity in the Afrotropical region. Bio-
anellus, forming complex support for aedoeagus; logy unknown.
female genitalia with ostium bursae in transverse Hapsiferinae. Autapomorphies: head vestiture
fold of S VIII; ventral lip of ostium with elongate consisting of lamellate scales appressed more or
setae; ductus bursae narrow, ornamented with less loosely to the head; epicranial suture absent,
microtrichia and with thorny internal ring close scale-bases of vertex forming an unbroken field;
to junction with corpus bursae. second segment of labial palpus with sparse lat-
eral bristles and no terminal whorl; male genita-
7 genera (Nemapogon, Nemaxera, Triaxomera,
lia with valvae fused medially at base, juxta
etc.) with 82 species (Fig. 7.1 C). Distribution
forming a bridge between faces of valvae,
primarily Holarctic but with one monobasic
transtilla present (most taxa), coremata invagi-
Australian genus (Vanna). Larva boring in
nated into ventral intersegmental membrane
bracket-fungi or in bark or wood permeated by
(most taxa); in female eighth segment and ovi-
fungal mycelia.
positor elongated, corethrogyne invaginated into
Tineinae. Autapomorphies: male genitalia with pair of ventral pouches.
uncus lobes fused and forming an articulated 11 genera (Hapsifera, Tiquadra, Scalidomia
hook; pupa with only anterior bands of abdomi- etc.) with 103 species (Fig. 7.1 D). Worldwide
nal spines; larva with bisetose SV-group on distribution but majority of genera and species
meso- and metathorax, stemmata reduced to a are Afrotropical. Larvae feeding externally or
pair or none. burrowing in soft substrates, on vegetable detri-
36 genera (Tinea, Tineola, Trichophaga, Mo- tus including sawdust used for cultivating mush-
nopis, Proterospastis etc.) with 320 species (Figs. rooms; also recorded from mammal burrows.
7.1 G - K ) . Worldwide distribution but poorly Hieroxestinae (= Oinophilidae). Autapomor-
represented in the Neotropical region. Larvae phies: head wedge-shaped in lateral view; head
feeding externally, or burrowing in loose sub- vestiture consisting of lamellate scales closely ap-
strates, on keratin and chitin (wool, feathers, ar- pressed to head, posterior scales of vertex and
thropod remains, guano, etc.), often in the pres- scales of occiput directed caudally; scale-bases
ence of fungal hyphae. on frons in regular transverse lines; antennal
Setomorphinae. Autapomorphies: head with scape elongated; antennal pecten absent; labial
lamellate semi-appressed scales; head sutures palpi strongly divergent; scale bases on abdomi-
fused; male eighth sternite a narrow, reflexed, al- nal sternites in transverse lines; female with sin-
most complete ring; male genitalia with dia- gle bobsled-shaped signum.
phragma microtrichiate; female with eighth seg- 6 genera (Opogona, Oinophila, Amphixystis,
ment membranous and lacking rami from the Phaeoses etc.) with 272 species (Fig. 7.1 L) (Rob-
apophyses anteriores; larva with unidentate inson & Tuck 1997). Pan tropical distribution, de-
100 Donald R. Davis & Gaden S. Robinson

pauperate in the New World. Larvae feeding ex- of interfacetal microsetae. Antenna 0.4-0.7 the
ternally or burrowing in soft or loose substrates, length of forewing; pecten present; flagellum
on dead or decaying plant material, or grazing usually filiform, bipectinate in S Compsoctena,
fungi in termite mounds or birds' nests; Asym- with one row of slender bidentate scales per seg-
plecta are reported to be leaf-miners; Wegneria ment, either restricted to dorsal surface or com-
are reported to feed on guano. pletely encircling segment, more concentrated in
Erechthiinae. Autapomorphies: male genitalia Compsoctena. Pilifers variable, well developed to
with juxta forming a deep pouch that displaces absent. Mandibles vestigial to absent. Haustel-
aedoeagus dorsad; male retinaculum distinctly lum vestigial (Fig. 7.3 D) to elongate. Maxillary
broadened at base and comparatively shallow; palpi 2-3-segmented and extremely short or elon-
ovipositor lacking the ventral rods present in al- gate and 4-5-segmented. Labial palpi moderately
most all other Tineidae; larva with PI and P2 on long, 3-segmented, either upturned or porrect.
head much closer than in other Tineidae, labrum Thorax: Metafurca (Fig. 7.4 H) with apophyses
with only three pairs of marginal setae, abdomi- free. Wings moderately broad (Fig. 7.3 k) fore-
nal D2 displaced ventrad and roughly level with wing index 0.23-0.4; microtrichia sparsely scat-
SD-group (larval synapomorphies require con- tered over all wing surfaces; of Deuterotinea
firmation). Many species have the forewing apex brachypterous (Zagulayev 1988); all veins arising
upturned and/or one branch of R and one of M separate from cell; Rs4 terminating either on
missing in the forewing. costa or termen; forewing with both accessory
and intercalary cells preserved; $ retinaculum
Two genera (Erechthias, Comodica) with 140
consisting of a broad fold arising ventrally from
species. Indo-Pacific distribution, most speciose
costal margin and partially extending over base
in Australasia and the Pacific; isolated species on
of Sc: 1A + 2 A with basal fork. Hindwings
St. Paul's Rocks and Ascension I. Larvae feeding
about as broad as forewings; hindwing index
externally on plant detritus or, rarely, on living
0.32-4; $ frenulum a single stout seta; with
leaf tissue of tough-leaved plants such as palms
24 smaller frenular setae. Legs with tibial spur
and Pandanus; some larvae tunnel in soft sub-
pattern of 0-2-4; epiphysis relatively smooth
strates including galls on Acacia.
(without pecten) in Compsoctena. Abdomen:
Siloscinae. Autapomorphies: forewing with Paired sternal apodemes of A2 slender, continu-
erect spicular scales; male genitalia with articu- ing half way into sternum as sternal rods (Fig.
lated saccus. 7.4 B). Coremata and corethrogyne absent. Male
3 genera (Silosca, Organodesma, Autoch- genitalia. Uncus moderately bilobed, fused to
thonus) with 12 species. Afrotropical distribu- hoodlike tegumen. Vinculum usually short and
tion. Biology unknown. V-shaped, rarely Y-shaped (with an elongate sac-
Euplocaminae. Autapomorphies: male antenna cus). Gnathos slender, U- to V-shaped or re-
bipectinate, pectinations elongate and strongly duced and widely separated. Subscaphium mod-
ciliate; labial palpus lacking lateral bristles. erately sclerotized. Transtilla present. Juxta usu-
One genus (Euplocamus) with 11 species. Pa- ally indistinct, slender and weakly sclerotized in
laearctic distribution including the Himalayas. Eriocottis. Valvae typically with a prominent spi-
Larvae boring in rotten wood and bracket fungi. nose lobe (clasper) from medioventral margin, or
Stathmopolitinae. Autapomorphies: female with a more basal, variably developed process
eighth abdominal segment asymmetrical, with from sacculus (in Crepidochares). Aedoeagus a
process on left side. simple, slender cylinder without cornuti. Female
One genus (Stathmopolitis) with 1 species. Ca- genitalia: Segments 8 and 9 + 1 0 forming a slen-
nary Is. Biology unknown. der, elongate, telescoping ovipositor (Fig. 7.4 M)
Teichobiinae. Autapomorphies: head with semi- supported by typically 2 pairs of elongate apo-
appressed forward-directed lamellate scales, frons physes and 2 (rarely 3) pairs of pseudapophyses
smooth-scaled; mouthparts reduced; antenna at- (Davis 1990 a); anterior dorsal pseudapophyses
tenuated; male genitalia lacking gnathos, juxta located entirely within A8 and usually fused for
fused with inner faces of valvae; ovipositor of most of their length (Fig. 7.4 N). Ostium situated
female with papillae anales sclerotized and at caudal margin of membranous S8. Ductus
pointed to form piercing apex. bursae elongate, membranous. Corpus bursae
One genus (Psychoides) with three species. Pa- membranous; signum a single, irregular scleroti-
laearctic distribution. Larva leaf-mining in early zation, or absent. Morphology of immature
instars then feeding externally on fern sporangia. stages unknown. Some larvae known to con-
struct underground silken tubes (Zagulayev
Eriocottidae (Figs. 7.1 MN). Small to medium 1988) or to bore into decayed wood, leaves, or
size moths, with forewings ranging from 4.5 to stems (Dierl 1970). Two subfamilies are recog-
23 mm in length. Head (Figs. 7.3 CD): Vesti- nized (Nielsen 1978, Davis 1990 a).
ture rough consisting of mostly long piliform
scales with simple, acute apices. Ocelli present or Eriocottinae. Ocelli typically present, rarely re-
absent. Eye small to moderately large; interocu- duced or absent. Antennae filiform in both sexes.
lar index 0.61.1; cornea with sparse scattering 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 Donald R. Davis & Gaden S. Robinson

palpi 3-5-segmented. Forewing with Rs4 termi- duced, 3-segmented, or absent. Labial palpi mod-
nating usually on costa, or termen (in Deutero- erate to extremely long, curving over head in
tinea). Male genitalia with V- to U-shaped some S Acrolophus, 3-segmented; lateral bristles
gnathos. Mostly Old World from southern Pale- present or absent. Thorax: Metafurca (Fig. 7.41)
arctic or Spain and North Africa (Eriocottis, with furcal apophyses short, free; furcal bridge
10 sp., Deuterotinea, 5 sp.) to Australia (Eucryp- vestigial with paired tendons arising caudally
togona, 1 sp.), with a single genus in South from secondary arms. Forewings moderately
America (Crepidochares, 5 sp.). broad (Fig. 7.3 L); index 0.29-0.39, with veins
Compsocteninae. Ocelli absent. Antennae pec- arising separate from cell; Rs4 terminating usu-
tinate in $, filiform to serrate in $. Haustellum ally at apex; forewing with both accessory and
minute (Fig. 7.3 D). Maxillary palpi 2-3-seg- intercalary cells usually preserved; 1 A + 2 A
mented. Forewings with Rs4 terminating on ter- with moderately short basal fork; retinaculum
men. Male genitalia with gnathos reduced, consisting of a broad, triangular lobe arising
widely separated. A single genus ranging from from underside of costal membrane immediately
southern Africa to Taiwan (Compsoctena, ca. anterior to and curling over base of Sc. Hind-
50 sp.). wing index 0.3-0.4; $ frenulum a single stout
seta; with 2 - 7 frenular setae. Legs with tibial
Lypusidae. Monobasic. The Palaearctic Lypusa spur pattern of 0-2-4; epiphysis rarely reduced.
maurella is a unicolorous dark moth, forewing Abdomen: Paired sternal apodemes of A2 (Fig.
length some 67 mm; sexes overall similar. Head 7.4 C) relatively short with broad bases; sternal
dorsally with appressed lamellar scales; antennae rods absent. Coremata and corethrogyne absent.
simple, scapus and pedicellus with scale tufts; Male genitalia: Uncus either acute or bilobed.
proboscis and maxillary palpi rudimentary; la- Vinculum U- to V-shaped, a slender to broad
bial palpi very short. Forewing Rs4 to termen. In ventral ring. Gnathos either fused (V-shaped) or
male genitalia vinculum U-shaped; uncus lobes divided. Transtilla absent. Juxta usually absent,
slender; prominent, setose 'peniculus' arising sometimes present in Acrolophus. Valvae vari-
from tegumen/valve articulation area. Female in able, typically elongate and moderately slender,
A8 with dorsal strengthening ridges confluent with elongate costal apophysis at base; basal part
with base of anterior apophyses. Dierl (1996) of sacculus fused medially to opposite valva. Ae-
gives additional details on adult structure. doeagus variable, usually with one or more cor-
The affinities of Lypusa are unresolved. It was nuti. Female genitalia: Segments 9 + 1 0 ex-
previously associated with the Psychidae, either tremely short with posterior apophyses ca. equal
as a distinct subfamily (Torre and Strand 1929) in length to A 9 + 10 (Fig. 7.4L); sternum 8
or family (Tutt 1900), but this relationship was more elongate, much broader and almost com-
disputed by Sauter and Httenschwiler (1991). pletely sclerotized with ostium opening within
Indeed it is clearly not a psychid as evidenced by sternum or on caudal margin; ostium usually
the free arms of the adult metafurca, or even a flush, sometimes projected into a sclerotized
tineid (Sauter 1982) because of its unusual larval cone. Anterior apophyses absent (completely
case construction. The latter is formed by folding fused within eighth sternum). Ductus bursae usu-
a leaf. Lypusa was assigned to a family of its own ally short and membranous, rarely elongate and
by Heinemann (1870), and very recently again sclerotized. Corpus bursae usually a single mem-
by Dierl (1996). A further assessment of the rela- branous sac, sometimes partially divided and
tionships of the taxon is needed following a thor- with numerous spicules.
ough study of the immature stages.
Egg: Poorly known; white and smooth in
Acrolophidae (Figs. 7.1 OP): Small to medium Amydria effrentella (Johnson and Martin 1969),
size moths, with forewings 4 to 26 mm in length. averaging 0.42 mm long and 0.32 mm wide.
Head (Fig. 7.3 E): Vestiture usually rough with Larva (Figs. 7.5 C - G ) : Body small to large, to
dense tufts of piliform scales directed upward more than 50 mm in length, whitish with dark
from frons and forward from occiput, to rela- suffusion sometimes dorsally and brownish to
tively smooth with slender scales possessing black head, notai plates, and pinacula. Fron-
rounded apices. Ocelli absent. Eyes small to toclypeus broad, extending ca. half the distance
moderately large; interocular index 0.65-1.1; to epicranial notch (Fig. 7.5 E); ecdysial line ter-
cornea with sparse covering of interfacetal minating at epicranial notch. Usually 6 pairs of
microsetae to densely setose. Antennae 0.250.7 stemmata, rarely reduced to 3 pairs. Labrum
the length of forewing; scape smooth, pecten ab- with three pairs of epipharygeal setae (Fig.
sent; flagellum filiform to bipectinate in , fili- 7.5 G). Hypostoma variable (Figs. 7.5 CD),
form in all $; a single row of slender bidentate broadly joined at midventer in Acrolophus. Pre-
scales per segment. Pilifers and mandibles ab- spiracular plate partially separated from preno-
sent. Haustellum a minute, setose lobe (Fig. tai plate (Fig. 7.5 M). Coxae well separated.
7.3 E) usually shorter than first maxillary palpal Crochets arranged in a uniserial ellipse with sides
segment, or absent. Maxillary palpi greatly re- 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 ridge. Pupation usually in coarse cocoon or in
caudally. Larvae typically constructing long larval tube.
silken tubes (Fig. 7.6 D) underground or on/in The family is exclusively New World and pri-
plant detritus and feeding primarily on plant de- marily Neotropical with ca. 80 species occuring
bris as well as living plants, rarely coprophagous in North America north of Mexico. The largest
or mycetophagous (Hasbrouck 1964; Davis genus is Acrolophus (245 sp.) followed by Amy-
1984; Davis et al. 1986; Davis and Milstrey 1988; dria (17 sp.), Ptilopsaltis (4 sp.), Exoncotis (3 sp.)
Davis and Hogue 1992). Pupa: Head smooth, and Drastea (1 sp.). The most diagnostic features
without a frontal ridge. Wings relatively short, of the family include the incomplete metafurcal
seldom surpassing A4. Dorsum of A38 with a bridge and loss of anterior apophyses in the
single, anterior row of dense, ridgelike spines. adult $, and the partially separated prespiracular
Segments A3 7 movable in $ and . Cremaster plate in the larva (Robinson 1988, Davis 1990).
reduced, variable, sometimes with small, paired Adult Acrolophus have been examined from mid-
dorsal and ventral spines or with a dorsal furcate Tertiary Dominican amber.
104 Donald R. Davis & Gaden S. Robinson

Arrhenophanidae (Figs. 7.1 QR): Small to mod- Ductus bursae elongate, usually broader and
erately large moths, with forewings 4 to 33 mm more thickened over caudal third. Ductus semi-
in length. Head (Fig. 7.3 FG): Vestiture gen- nalis joining ductus bursae slightly anterior to its
erally rough, with slender scales over vertex middle, of variable length, extremely long in
longer and more erect than much shorter scales those species with $ possessing greatly length-
over frons. Ocelli absent. Eyes large, interocular ened vesica. Corpus bursae relatively small,
index ca. 1.5. Cornea naked. Antenna 0.30.6 membranous and without signa.
the length of forewing; scape smooth to rough; Egg: Undescribed. Larva: Body small to large,
pecten absent; flagellum piliform to serrate in $, up to 30 mm in length, whitish with yellowish
serrate to bipectinate in ; a single row of slender brown head, light brown to black notai plates
scales per segment over dorsal half. Pilifers and and pinacula. Frontoclypeus extending ca. half
mandibles absent. Haustellum (Fig. 7.3 G) ex- the distance to epicranial notch; ecdysial line ter-
tremely reduced to a minute setose lobe less than minating well before cranial notch. Six pairs of
length of first maxillary palpal segment. Maxil- stemmata. Labrum with 3 pairs of epipharyngeal
lary palpi reduced to two short, rounded seg- setae. Hypostoma narrowly joined at midventer.
ments. Labial palpi moderately long, upcurved, Prespiracular plate completely separate from
3-segmented; lateral bristles absent. Thorax: prenotai plate (Fig. 7.5 N). Coxae partially fused
Metafurca (Fig. 7.4 J) with furcal apophyses at midventer. Crochets arranged in a lateral pen-
joined as a bridge to secondary arms. Forewings ellipse; prolegs without peripheral spines; anal
moderately broad (Fig. 7.3 M); index 0.40.47, crochets in a semi-ellipse, opened caudally. Lar-
with Rsl usually completely fused to Rs2
vae feed on woody bracket fungus (Polyporus)
(stalked in Cnissostages) and Rsl + 2 usually
into which are constructed long, irregular silk-
stalked to Rs3 (separate in Cnissostages and in
lined tunnels; a tough, slightly flattened case
unnamed Taiwanese genus); Rs4 terminating
(Fig. 7.6 E) is attached to the outer opening of
either on apex or termen; accessory and interca-
the tunnel (Busck 1912). Pupa: Head smooth,
lary cells usually absent; 1 A + 2 A with basal
without frontal ridge. Wings relatively short, not
fork; S retinaculum consisting of a broad, trian-
surpassing A4. Dorsum of A3 - 8 in $ and A 3 - 7
gular lobe arising from underside of costal mem-
in 9 with a single anterior row of short spines;
brane immediately anterior to and curling over
base of Sc. Hindwing index 0.43-0.55; $ frenu- A 3 - 7 movable in $ and A 3 - 6 in Cremaster
lum a single stout seta; with 412 frenular se- reduced to a pair of small tubercules on venter
tae. Legs with tibial spur pattern of 0-2-4; epi- of AIO. Pupation occurs inside larval case with
physis present, inner surface densely spinose. Ab- the adult emerging from distal slit-like opening.
domen: Paired sternal apodemes of A2 (Fig. The Arrhenophanidae are mostly restricted to
7.4 D) short, stout to slender; sternal rods pre- the New World where four genera are recognized
sent; S8 of S with a bilateral pair of short, broad (Arrhenophanes, 2 sp.; Cnissostages, 2 sp.; Ecpa-
apodemes directed anteriorly. Coremata some- thophanes, 2 sp., Dysoptus, 2 sp.). An unnamed
times present on S8 of consisting of small, genus and two species have recently been re-
paired tufts of short, broad scales. Female with ported from Taiwan (Davis 1991). Closely allied
dense corethrogyne of long piliform scales encir- to the Psychidae, they differ from the latter by
cling A7. Male genitalia: Uncus fused to tegu- the antenna being serrate to bipectinate in the
men, usually tapering to a slender, rounded apex, $; genitalia with sclerotized anellar tube and
rarely bilobed. Vinculum usually Y-shaped, with aedoeagus often with non-retractible vesica that
a slender, elongate saccus, rarely V-shaped. is stored in an invaginated apotheca; oviposi-
Gnathos absent. Distinct juxta and transtilla ab- tor often with prominent lamella postvaginalis;
sent. Anellus a sclerotized tube tightly enclosing larva with three pairs of epipharyngeal setae and
aedoeagus, and broadening at base to fuse with the prespiracular plate separate from the prono-
inner surfaces of valvae. Valvae elongate, usually tum; and pupa with only a single anterior row of
with a prominent, strongly sclerotized lobe or tergal spines.
process from apex of sacculus. Diaphragma usu-
ally invaginated to form a large membranous sac Psychidae (Figs. 7.1 S - T , 7 . 2 A - G ) . Small to
(apotheca, Bradley 1951) dorsad to anellus. Ae- medium size moths, with forewings 4 to 28 mm
doeagus extremely slender and elongate, usually in length; fully winged, $ either fully winged,
with a tenuous, non-retractile, partially sclero- brachypterous, apterous, or vermiform (Fig.
tized vesica often exceeding length of aedoeagus; 7.2 F) with all body appendages vestigial or lost.
vesica normally coiled inside apotheca prior to Head (Figs. 7.3 HI): Vestiture generally rough,
mating, sometimes broken off inside ductus sem- consisting usually of moderately long, slender to
inalis of $ during copulation. Female genitalia: piliform scales directed mostly forward. Ocelli
Segments 8 and 9 + 1 0 forming a slender, elon- present in some primitive genera, usually absent.
gate telescoping ovipositor supported by 2 pairs Eyes small to large; interocular index 0.42.3;
of rodlike apophyses and 1 pair of pseudapo- cornea naked. Antennae 0.30.8 the length of
physes. Ostium located at caudal margin of S8. forewing; usually vestigial in vermiform 9; scape
rough, pecten present but often indistinct; flagel-
The Tineoidea and Gracillarioidea 105

lum filiform to strongly bipectinate in <?, usually Egg: Poorly known. Oval in shape. Chorion
filiform in (when developed) to rarely serrate; smooth to finely reticulated; micropylar area
vestiture variable, usually with scales scattered with faint reticulations (Davis 1990). Usually
over dorsal half, sometimes with one distinct row laid in masses of 1813,000 eggs (Kozhanchikov
per segment. Pilifers and mandibles absent. 1956; Davis 1975, 1987) inside larval case and
Haustellum usually absent, present in a few of mixed with piliform scales from body of ; de-
the most primitive genera (Davis in press b). posited on host substrate in some primitive
Maxillary palpi usually absent, rarely rudimen- forms. Larva (Figs. 7 . 5 1 - K , O): Body usually
tary to 3-segmented (in Kearfottia, Fig. 7.3 H, between 6 - 5 0 mm long, with head and thorax
Lamyristis). Labial palpi often lost, 1-3 seg- well sclerotized and variously pigmented; abdo-
mented in more primitive genera; lateral bristles men usually unicolorous, varying from white to
absent. Thorax: Metafurca (Fig. 7.4 E) with fur- fuscous, or with dark, dorsal plates. Frontoclyp-
cal apophyses joined as a bridge to secondary eus extending ca. 0.5 to epicranial notch; ecdysial
arms. Forewings (Figs. 7.3 N - O ) slender to lines converging at or just dorsad to apex of
broad; index 0.3-0.66, forewings normally with frontoclypeus. Six pairs of stemmata. Venter of
7 - 9 veins arising from cell; Rs4 terminating on labrum with 4 pairs of epipharyngeal setae (Fig.
termen; M present in cell; accessory and interca- 7.51). Thoracic legs well developed and directed
lary cells usually absent; 1 A + 2 A either with or anteriorly (Fig. 7.5 J); coxae partially to com-
without basal fork; S retinaculum varying from pletely fused at midline; lateral pinacula of Tl,
a broad fold to a more slender coiled flap from bearing 3 L setae and spiracle, fused to prono-
underside of costal membrane immediately ante- tum (Fig. 7.5 O). All prolegs present, moderately
rior to and extending under base of Sc. Hindw- short, with crochets of A36 uniordinal and ar-
ing index 0.33-0.76; $ frenulum a single stout ranged in a lateral penellipse; crochets of AIO
seta; alate ? with 3 8 frenular setae. Legs with in a uniordinal semi-ellipse. All larvae are case
tibial spur pattern highly variable, ranging from bearers throughout all instars and feed on a
0-2-4 to 0-0-0; epiphysis present or absent. Abdo- broad range of plant hosts, from lichens, grasses,
men: Paired sternal apodemes of S2 (Fig. 7.4 E) to leaves of evergreen and decidous trees and
elongate, straight to curved, continuing more shrubs; a few have also been reported as insec-
than half way into 2 S as sternal rods. A pair of tivorous (Clausen 1940). Structure of larval case
short coremata sometimes present on S8 of $ in highly divergent (Figs. 7.6 FN), from helical
more primitive genera; S8 usually sclerotized (Fig. 7.6 H) to greatly elongated, from less than
with a pair of prominent apodemes directed an- 6 mm long at maturity to 32.7 cm (Oiketicus aris-
teriorly. Female with dense corethrogyne of long tocosma Fig. 7.6 G). Number of instars not well
piliform scales encircling A7, often concentrated known, at least 4 or more (Jones and Parks 1928;
in clusters. Male genitalia. Uncus completely Davis 1964; Kozhanchikov 1956) the $ some-
fused, rounded caudally or represented by short, times possessing one more instar than $. Pupa:
setose lobes. Tegumen well developed, hoodlike. Vertex usually with short frontal process. Wings
Vinculum U- to Y-shaped; saccus often elongate. usually terminating on A3 or 4, variably reduced
Gnathos absent. Juxta sometimes present and in $ to absent. Tergal spines highly variable, typi-
fused with partially sclerotized anellus in lower cally present on at least some segments between
forms. Subscaphium sometimes sclerotized, usu- Al - 8, ranging from an irregular cluster in many
ally absent. Valvae relatively slender, elongate, primitive genera to 2 rows in most advanced
typically with a broad, setose pad at base (often forms with the posterior row usually more slen-
referred to as pulvillus, vallum penis, but proba- der, elongate, and recurved (Davis 1975). Seg-
bly derived from anellus) as well as an acute or ments A 2 - 7 movable in A 2 - 6 in . Cremas-
spinose process from ventral margin or apex of ter usually present as a single pair of small to
sacculus. Transtilla usually present. Seven pairs large ventral hooks. Pupation occurs inside lar-
of genital muscles present (Kuznetsov and Stek- val case. Prior to this the last instar larva atta-
ol'nikov 1977). Aedoeagus elongate, slender, ches the case to some support and then typically
usually without cornuti. Female genitalia. Highly inverts itself with the head pointing to the rear
variable depending upon degree of specializa- of the case; pupal extrusion and adult emergence
tion; segments 8 and 9 + 10 usually long and tel- thus occurring usually through the lower or cau-
escoping, with 2 pairs of apophyses and 1 pair of dal end.
pseudapophyses (within AIO) in more primitive Nearly 1000 species are known from all major
genera, to greatly shortened and with only 1 - 2 regions except Antarctica (Heppner 1991), with
pairs of apophyses in most specialized, vermi- more than 85% of these restricted to the Old
form $ (Davis 1964). Ostium located on S8 or at World. Although generally rare in Lepidoptera,
caudal margin. Ductus bursae and corpus bursae parthenogenesis has developed in at least five
usually short and membranous; signum absent; genera of Psychidae (Dahlica (= Solenobia),
ductus seminalis sometimes greatly elongated in Scoriodyta, Luffia, Apterona, and Acanthopsyche;
most advanced genera. 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 107

netics of a few palearctic species of Dahlica and pupa (1) partially extruded from cocoon prior to
Talaeporia have been studied extensively by eclosion; (2) with abdominal tergal spines; adult
Seiler and his students (Seiler 1920-1963). Seiler (3) without pleural lobes on A8 of (4) with
discovered, for example, that Dahlica triquetrella smoothly scaled frons; (5) labial palpi without
has bisexual as well as diploid and tetraploid lateral bristles; (6) galeae usually elongate and
parthenogenetic races whose distributions were closely associated; (7) hindwings without interca-
closely associated with recent geological history lary cell; larva (8) with early instars mining or
and biotic changes. boring into plant host; and (9) crochets of later
Because of numerous and variously correlated instars usually biserial. The first three character-
morphological specializations which have ap- istics distinguish the Gracillarioidea from the
peared at different levels throughout this family, Yponomeutoidea (Kyrki 1984), and the last six
attempts to classify the Psychidae have varied separate them from the Tineoidea. The presence
considerably. Previous attempts to partition the of a naked haustellum and a second abdominal
family have resulted in as many as 10 families sternum with tineoid apophyses (Kyrki 1983 b),
(Tutt 1990) to as few as 2 subfamilies (Koz- along with the foregoing pupal characters,
hanchikov 1956). Gomez Bustillo (1979) superfi- closely associate the Gracillarioidea with the Ti-
cially divided the Psychidae into two paraphy- neoidea and serve to distinguish both groups
letic families: Micropsychidae (later emended by from the higher Ditrysia.
Gomez Bustillo (1984) to Micropsychiniidae) to Head: Frons smooth; vertex often smooth,
include the "tineid-like" forms and the true rough in Roeslerstammiidae, Bucculatricidae,
Psychidae, or "bombyx-like" forms. The most and some Gracillariidae. Ocelli usually absent,
recent classification (Httenschwiler 1989; Sauter present only in Douglasiidae. Antennae often as
and Httenschwiler 1991) lists 8 subfamilies, long as or longer than forewing, ranging from
mostly without accompanying keys or diagnoses. 0.61.75 the length of wing; scape with pecten
Until these are presented, hopefully within the present or absent; flagellum filiform; 1 - 2 rows
format of a cladistic analysis, it will not be pos- or annuii of scales per segment. Chaetosemata
sible to apply this classification on a global basis. absent. Pilifers present. Mandibles usually ab-
The subfamilies proposed by Sauter and Htten- sent. Haustellum naked, usually elongate, ca.
schwiler, with representative genera shown in pa- 12X length of labial palpi. Maxillary palpi re-
rentheses, are: Naryciinae (Dahlica (= Soleno- duced, 1 4 short segments. Labial palpi usually
bia), Diplodoma, Narycia, ?Kearfottia); Talepori- 3-segmented, reduced to 1 in most Bucculatrici-
nae (Bankesia, Pseudobankesia, Taleporia); Pla- dae (Fig. 7.7 F), without erect bristles. Thorax:
codominae (Placodoma); Typhoniidae (Dissoc- Metafurca with apophyses free. Forewings mod-
tena, Dissoctenioides, Melasma, Penestoglossa, erately broad to slender, index (W/L) 0.09-0.24;
and possibly the Neotropical genera Amiantastis, Rs4 terminating on costa or apex; chorda with
Plumana, Pterogyne); Scoriodytinae (Scorio- accompanying accessory cell usually absent,
dyta)', Psychinae (Bruandia, Luffia, Psyche sometimes present in Roeslerstammiidae and
(= Fumea)-, Epichnopteryginae (Bijugis, Epich- Gracillariidae; intercalary cell absent; 1 A + 2 A
nopterix, Rebelia, Whittleia), and Oiketicinae with or without basal fork; subanal interlocking
(Acanthopsyche, Amicta, Apterona, Brachycyt- microtrichia present. Hindwings often lanceo-
tarus, Cryptothelea, Eumeta (= Clania), Oiketi- late, index 0.1-0.4; intercalary cell absent; discal
coides, Oiketicus, Thyridopteryx). cell usually open (m-cu crossvein absent), closed
The Pseudarbela-group, previously given fam- in Roeslerstammiidae and most primitive Gracil-
ily status (Figs. 7.2 A - B ; Clench 1959) is cur- lariidae. Abdomen: Tergal spines absent. Anterior
rently included within Psychidae (Robinson 1988) margin of S2 with tineoid apodemes and usually
but some species are referable to the Limacodi- with sternal rods (Figs. 7.7 N - O , Q); rods absent
dae. Only two features - the elongate basal fork in Bucculatricidae (Fig. 7.7 P); a minute pair of
of 1 A + 2 A in the forewings and the spinose warts and 2 pairs of tuberculate plates usually
ridge of the S valvae define this taxon. These present as in Tineoidea. Male genitalia: Uncus
are of minor significance when compared to the usually absent, slightly bilobed in Roeslerstam-
major synapomorphies shared with Psychidae. miidae; genital muscles 3 - 7 pairs (Kuznetsov
The group includes 3 genera (Casana with 1 spe- and Stekol'nikov 1977, 1987). Female genitalia.
cies, Linggana with 1 species, Pseudarbela with Ovipositor moderately to only slightly extensi-
4 species) restricted to the Indo-Australian area. ble, with 2 pairs of moderate to short apophyses.
Females are fully winged, with forewings up to Ostium opening usually on S8 or at caudal mar-
20 mm in length. Immature stages are unknown; gin; opening between S7 and 8 in Roeslerstammi-
the larva is believed to be a casebearer, although idae.
this needs confirmation. Egg: Oval, fiat type, with micropylar axis par-
allel to oviposition substrate. Chorion relatively
smooth to finely reticulated; micropylar end with
GRACILLARIOIDEA reticulated ridges. Eggs laid singly and cemented
The Gracillarioidea are currently restricted to to plant host, with larva boring directly into tis-
four families that share the following features: 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 109

phic; early instars typically internal borers or nating on costa; accessory cell absent or weakly
miners, often apodal or with legs and prolegs re- present; intercalary cell absent; 1 A + 2 A with
duced; ventral crochets usually in a biserial circle basal fork; $ retinaculum a broad, triangular
with inner series incomplete (Fig. 7.81). Pupa: flap from costal cell coiling underneath base of
Adecticous, incomplete, with 4 - 5 movable abdo- Sc. Hindwing index 0.28-0.4; $ frenulum a sin-
minal segments. Maxillary palpi usually present. gle stout seta; $ with 2 frenular setae. Legs with
Abdomen with tergal spines usually in a single tibial spur pattern of 0-2-4; epiphysis relatively
anterior row, or in dense patches (Gracillarii- short, arising from distal third of tibia. Abdomen :
dae). Pupa extended partially from cocoon prior Paired sternal apodemes of A2 (Fig. 7.7 N) elon-
to adult eclosion. gate, slender, slightly curved inwards, continuing
caudad as well defined sternal rods 2/3 the length
of S2; a transverse, sclerotized bar extends par-
Key to the Families
tially across anterior margin of S2. Coremata
1 Ocelli present Douglasiidae and corethrogyne absent. Male genitalia: Uncus
Ocelli absent 2 slightly bilobed, setose. Tegumen moderately
2 Antennal scape broad, partially covering eye. broad dorsally. Vinculum usually Y-shaped with
Haustellum short, less than 1.5X eye diameter an elongate saccus, rarely U-shaped. Gnathos
(Fig. 7.7 E). Cocoon longitudinally ribbed slender, either fused medially or weakly united
(Fig. 7.6 P) Bucculatricidae by membrane. Subscaphium slender, weakly
Antennal scape slender, not covering eye. sclerotized. Valvae relatively simple, sometimes
Haustellum more than 2.5 X eye diameter. Co- with an acute basal or subapical process; an
coon smooth, not ribbed 3 elongate tuft of piliform scales arising from base.
3 Wings slender to lanceolate (Figs. 7.7 L - M ) ; 7 pairs of genital muscles present (Kuznetsov
hindwing index (W/L) 0.10.2; cilia longer and Stekol'nikov 1977). Aedoeagus a slender,
than hindwing width. Flagellomeres with sin- often curved cylinder without cornuti. Female
gle annulus of scales Gracillariidae genitalia: Ovipositor short, with 2 pairs of short
Wings broader (Fig. 7.71), hindwing index apophyses. Ostium opening in intersegmental
0.28-0.4; cilia less than hindwing width. Fla- membrane between S7 and 8. Antrum variously
gellomeres with 2 rows of dorsal scales . . . . sclerotized. Corpus bursae membranous with a
Roeslerstammiidae single, large, linear to cruciform signum.
Egg: Undescribed; deposited at the tip of host
Roeslerstammidae (Figs. 7.2 H - I ) . Small moths, leaf (Kyrki 1983a). Larva (Figs. 7.8 G - I ) : Body
with forewings 5 to 10 m m in length. Head (Fig. pale green, up to 13 mm long, with thoracic legs
7.7 AB): Vestiture of frons smooth with broad and 5 pairs of prolegs. Head brownish; pronotal
scales; vertex rough with dense covering of erect and anal plates paler brown to indistinct. Fron-
piliform scales. Ocelli absent. Eyes moderately toclypeus slender, extending 0.8 the distance to
large to greatly enlarged, sometimes indented epicranial notch (Fig. 7.8 G). Adfrontal sutures
posteriorly or completely divided by a slender terminate at epicranial notch. Six pairs of stem-
scaled band; interocular index 0.87-2.0; cornea mata present. Prothorax (Fig. 7.8 H) with 3 pre-
naked. Antennae 0.81.5 the length of forewing; spiracular (L) setae. Legs well developed; coxae
scape with dense pecten; flagellum piliform; two separated. Ventral prolegs with 2 4 - 3 4 crochets
rows of slender scales per segment dorsally, with arranged in a posteriorly uniserial or biserial lat-
distal row encircling segment. Pilifers well devel- eral penellipse (Fig. 7.81). Anal prolegs with ca.
oped. Mandibles absent. Haustellum moderate 15 uniordinal crochets in a semi-circle. First two
to elongate, usually equalling or exceeding length larval instars are leafminers (on Aceraceae, Betu-
of labial palpi. Maxillary palpi reduced to a sin- laceae, Elaeocarpaceae, and Tiliaceae) and the
gle ovoid segment (Fig. 7.7 B). Labial palpi elon- later instars feed externally, either exposed, or in
gate, 3-segmented, slender, and curved upwards; webbing on underside of leaf (Kyrki 1983 a), or
lateral bristles absent. Thorax: Metafurca (Figs. within a shelter formed by webbing several
7.8 - B ) with posterior apophyses slender, elon- leaves together (Moriuti 1978). Pupa: Frons
gate, free; anteromedial process elongate and smooth. Antennae extend to or exceed caudal tip
slender, equalling or surpassing length of furcal of abdomen, in which case they tend to coil
stem, and with a prominent ventral keel from around abdomen (Common 1990). Wings ex-
base. Forewings (Fig. 7.71) narrow, index 0 . 2 2 - tending to caudal margin of A6. An anterior row
0.31; Rs3 and 4 either stalked or fused and termi- of tergal spines present on A 3 - 7 . Segments

Bucculatrix solidaginiella', LM, Gracillariidae: L, Caloptilia stigmatella; M, Phyllocnistis insignis; NQ, Second
abdominal sterna: N , Roeslerstamiidae, Roeslerstammia erxlebella; O, Douglasiidae, Tinagma obscurofasciella; P,
Bucculatricidae, Bucculatrix solidaginiella; Q, Gracillariidae, Caloptilia stigmatella. (All scales = 0.5 mm, except
Fig. H.)
110 Donald R. Davis & Gaden S. Robinson

A 2 - 6 movable in $. AIO without sclerotized Aedoeagus a slender cylinder usually without

cremaster but with stout to hooked setae. Pupa- cornuti. Female genitalia. Ovipositor telescoping,
tion occurs inside a dense silken cocoon among moderately long; two pairs of apophyses of equal
leaf litter or within a folded leaf edge. length. Ostium a sclerotized ring of various out-
The family is widespread through the Old line at caudal margin of S8. Ductus bursae slen-
World in the Palearctic, Oriental, and Australian der, mostly membranous, sometimes with a few
regions where 10 genera and ca. 40 species are minute spicules or a distinct collar. Corpus bur-
known. A key to six of the Oriental-Australian sae with a single stellate signum.
genera (Agriothera, Amphithera, Chalcoteuches, Immature stages poorly known. Egg: Unde-
Enchoptila, Sphenograptis, Telethera) and one scribed. Larva (Fig. 7.8 J): Small; length ca.
Neotropical genus (Dasycarea) was compiled by 5 - 8 mm. Frontoclypeus higher than wide, ex-
Moriuti (1978). Common (1970) also included tending 0.6-0.75 to epicranial notch. Six pairs of
two additional Australian genera (Macarangela stemmata. Body white to yellow with pigmented
and Thereutis) within the family. Kyrki (1983) pronotal plate, pinacula, and anal plate. Prespi-
listed seven species ranging from western Europe racular (L) group of T1 trisetose. Primary setae
to Japan within the Palearctic genus Roeslers- extremely long. Thoracic legs present; coxae sep-
tammia. arated. Prolegs reduced; crochets apparently ab-
sent. Larvae have been reported as leaf miners
Douglasiidae (Fig. 7.2 J). Small moths, with fore- Klimeschia), flower petiole miners and stem bor-
wings 3.0 to 6.4 mm in length. Head (Fig. ers (Tinagma) on Boraginaceae, Rosaceae, and
7.7 CD): Vestiture of broad, smooth scales. Laminaceae (Heppner 1984). Pupa: Unde-
Ocelli well developed. Eyes small, interocular in- scribed, reportedly with abdominal tergal spines
dex 0.65-0.7; cornea naked. Antennae 0.6-0.7 (Kyrki 1984) and partially extruded from cocoon
the length of forewing; scape usually smooth, oc- prior to adult eclosion. The family is small with
casionally with rudimentary pecten of 2 - 3 pili- only 17 species represented in the Palaearctic re-
form scales; flagellum filiform, with a single row gion (Gaedike 1974), 7 for the Nearctic (Gaedike
of slender scales encircling each segment. Pilifers 1990), and one from Australia (Common 1990).
rudimentary. Mandibles reduced to a small lat- Two genera, Klimeschia and Tinagma (= Doug-
eral lobe. Haustellum elongate, more than 2X lasia) are recognized.
length of labial palpi. Maxillary palpi reduced to
a single lobate segment (Fig. 7.7 D). Labial palpi Bucculatricidae (Figs. 7.2 - M ) . Extremely to
relatively short, 3-segmented, drooping to moderately small moths, with forewings 2.5 to
slightly upcurved. Thorax: Metafurca (Fig. 7.8 C) 7 mm in length. Head (Figs. 7.7 E - F ) : Usually
with posterior apophyses short, free, curved elongate, with frons projecting well below eye;
ventrad; anteromedial process with prominent shortened, less specialized in Leucoedemia. Vesti-
ventral keel. Forewings (Fig. 7.7 J) narrow; index ture with frons smooth; vertex usually with large,
ca. 0.28; Rs with 3 or 4 branches; Rs4 terminat- erect tuft of piliform scales projecting in all di-
ing on costa, stalked with M l ; M either 2 or 3 rections, rarely smooth. Ocelli absent. Eyes mod-
branched; accessory and intercalary cells absent; erately large, interocular index ca. 1.0; cornea
A 1 + 2 A with or without basal fork; $ retinacu- naked. Antennae 0.6-0.9 the length of forewing;
lum a slender coiled flap under base of Sc. scape enlarged, flattened, with a dense row of
Hindwing index ca. 0.2; discal cell open; $ frenu- slender scales (pecten) partially covering eye; fla-
lum a single stout seta; ? with 1 2 frenular setae. gellum filiform, with 2 rows of short, slender
Legs with tibial spur pattern of 0-2-4; epiphysis scales encircling each segment; first flagellomere
present. Abdomen: Paired sternal apodemes of often notched or strongly curved in Pilifers
A2 (Fig. 7.7 O) elongate, slender; sternal rods usually reduced, well developed in Leucoedemia.
weak. Coremata and corethrogyne absent. Male Mandibles absent. Haustellum short, 1.5X eye
genitalia: Uncus absent. Tegumen a triangular diameter or less. Maxillary palpi vestigial, usu-
hood. Vinculum slender, often with a short, stout ally a minute, unsegmented lobe. Labial palpi
saccus which is lacking in some species. Gnathos drooping, without lateral bristles, usually ex-
and transtilla absent. Anellus a partially sclero- tremely short and one segmented (Fig. 7.7 F) or
tized ring. Valvae asymmetrical in Klimeschia or with three short segments in Leucoedemia. Tho-
symmetrical in Tinagma, relatively short and rax: Metafurca (Fig. 7.8 D) with apophyses re-
broad and often with a peculiar apical process. duced to a pair of low dorsal ridges. Forewings

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-
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 Donald R. Davis & Gaden S. Robinson

(Fig. 7.7 ) lanceolate, index 0.14-0.25, with nosperms and angiosperms have been reported
apex of forewing usually acuminate; forewing as hosts, with the Asteraceae, Betulaceae, and
with retinaculum a broad fold from underside of Fagaceae being particularly favored (Braun
fused C and Sc veins; Rs usually 4-branched, rar- 1963, Seksyayeva 1994 b). Pupa: Frontal ridge
ely 3; Rs4 to apex; Accessory and intercalary longitudinal, well developed. Wings and anten-
cells absent; Cu usually single branched, rarely nae usually extending to A7; A2 or 3 - 7 with
absent; 1A + 2 A without basal fork. Hindwing a single row of dorsal spines. Segments A37
index 0.180.21; $ frenulum a single stout seta; movable in S, A 3 - 6 in ? (Mosher 1916). AIO
$ with 2 frenular setae; discal cell open in hind- with cremaster reduced, consisting of minute,
wing. Epiphysis either present or absent; legs paired, dorsal spines and often a pair of larger,
with tibial spur pattern of 0-2-4. Abdomen: lateral spines.
Paired sternal apodemes of A2 (Fig. 7.7 P) elon- Approximately 250 species are known around
gate, slender; sternal rods absent. An eversible the world, with the greatest concentration
scale sac present in most $ Bucculatrix as a mid- (100 sp.) in the Nearctic region (Braun 1963,
dorsal invagination of membrane between A2 Heppner 1991, Seksyayeva 1994 a). With the no-
and 3; corythrogyne absent. Male genitalia. Un- table exception of the most plesiomorphic taxon,
cus usually absent, rarely well developed and the monotypic South African Leucoedemia,
acute. Tegumen and vinculum narrow to broad; nearly all the species are included in the single
saccus rarely developed. Gnathos usually absent; genus Bucculatrix . The Australian Ogmograptis
well developed, fused medially, and setose in
has recently been placed in this family (Nielsen &
Leucoedemia. Socii typically well developed and
Common 1991).
setose in Bucculatrix. Transtilla usually absent,
sometimes weakly developed. Anellus a membra-
Gracillariidae (Figs. 7 . 2 N - S ) . Small moths with
nous cone often with lateral sclerotizations. Dis-
forewings 210 mm in length. Head (Figs.
tinct juxta usually absent. Valvae usually elon-
gate and simple, rarely lobed. Three pairs of gen- 7.7 G - H ) : Vestiture variable, usually smooth (in
ital muscles present (Kuznetsov and Stekol'nikov Gracillariinae) with moderately broad scales
1987). Aedoeagus usually an elongate, sinuate from vertex directed forward and down over
cylinder; cornuti rarely present. Female genitalia: frons; similar in Lithocolletinae except with tufts
Ovipositor short, usually with only posterior of erect filiform and sometimes broader scales
apophyses present; anterior apophyses some- arising from occiput caudad to vertex. Ocelli ab-
times evident. Ostium opening either along cau- sent. Eyes moderately large; interocular index
dal margin of S8 or near center of S8. Ductus 0.9-1.0; cornea naked. Antennae 0.8-1.75 the
bursae usually sclerotized over caudal portion, length of forewing; scape usually smooth, with
usually elongate and slender. Corpus bursae or without pecten; flagellum filiform, with a sin-
abruptly enlarged; signum usually a single large gle row of slender scales completely encircling
plate with numerous rows of spines. each segment. Pilifers well developed. Mandibles
absent. Haustellum usually elongate, 1 - 2 X
Egg: Usually flattened and oval, with a reticu- length of labial palpi. Maxillary palpi rarely ab-
lated chorion (Braun 1963). Larva (Figs. sent, usually short to minute, 1-4-segmented. La-
7.8 K L): Hypermetamorphic, typically with 5 bial palpi 3-segmented, usually upturned, to
instars (Braun 1963); apodal in at least first two straight and drooping; segment 2 sometimes with
leafmining instars, with thoracic legs and 5 pairs ventral scale tuft. Thorax. Metafurca (Figs.
of prolegs in later externally feeding instars; pro- 7.8 E - F ) with anteromedial process elongate,
thoracic pretarsal claw highly modified in Buccu- relatively slender; furcal apophyses free, short to
latrix, greatly lengthened; crochets in an incom- moderately long and downcurved, arising well
plete circle. Frontoclypeus short, extending 0.5 caudad near secondary arms. Forewings (Figs.
0.6 to epicranial notch. Tentorial bridge very 7.7 L - M ) slender; index 0.9-0.25; venation
slender, deeply arched (Davis 1987). Stemmata variable with R/Rs 3- to 5-branched; Rs4 usually
5 - 6 pairs. Prespiracular (L) group of T1 triset- to costa, rarely to apex; M l - to 3-branched; Cu2
ose. Most larvae undergo two instars as leafmin- sometimes absent; discal cell elongate, extending
ers, with the third instar emerging to feed exter- 0.7-0.8 the length of forewing; accessory cell
nally. A flattened moulting cocoon (Fig. 7.8 L) is vestigial to absent; base of M vestigial to absent;
sometimes constructed underneath the leaf by intercalary cell absent; 1A + 2 A usually present,
the third instar larva as well as a second moult- sometimes vestigial, occasionally with minute
ing cocoon by the fourth instar; the fifth instar basal fork; $ retinaculum usually formed of
prior to pupation constructs a silken, ribbed co- curved scales from underneath slightly swollen
coon (Figs. 7.6 OP) typical for the family. base of Sc; Hindwing lanceolate; index 0.1-0.2;
Some species are stem borers or feed inside galls $ frenulum usually with 2 frenular setae, rarely 1 ;
(Braun 1963, Scoble and Scholtz 1984). The stout, composite pseudofrenular setae sometimes
larva of Ogmograptis scribula forms an elongate arising near apex of Sc in both sexes (Davis
mine in the bark of Eucalyptus (Nielsen and 1991); venation reduced, from 7 to 4 veins; discal
Common 1991). Nearly 30 families of gym- cell often open. Legs with tibial spur pattern usu-
The Tineoidea and Gracillarioidea 113

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 rotized plates. Hypermetamorphic development
(Fig. 7.7 Q) long and narrow, with long slender (Kumata 1978; Davis 1987) with at least 2 dis-
sternal apodemes continuing caudad as sternal tinct forms and habits: (1) an early, usually sap-
rods ca. 0.B the length of sternite. A8 and some- feeding (Trgardh 1913) or flattened (Chambers
times 7 of $ frequently with paired coremata; S8 1877) form, often comprising the first 25 in-
of $ often enlarged, particularly in Lithocolleti- stars, which possess a depressed, apodal body
nae; T8 sometimes with median sclerotization; with specialized, prognathous mouthparts and
corethrogyne absent. Male genitalia. Uncus ab- rudimentary spinneret (Figs. 7.8 N - O , P, R),
sent. Tegumen a relatively elongate, mostly and (2) a later hypognathous, tissue-feeding or
membranous dorsal hood. Vinculum U- to Y- cylindrical form possessing a more generalized,
shaped, with often a short, rarely elongate sac- eruciform body with unspecialized, chewing
cus. Subscaphium frequently distinct, variably mouthparts and legs (Figs. 7.8 M, Q); or a vari-
sclerotized. Gnathos absent. Transtilla present or ously modified nonfeeding spinning form, with
absent. Yalvae usually elongate and simple, rar- either a depressed or cylindrical body, mandibles
ely asymmetrical, sometimes lobed, spined, or and legs reduced or absent, but with a functional
with 1 - 2 pairs of pectinifers. Anellus usually spinneret (Figs. 7.8 S). An additional nonfeeding,
membranous. Juxta absent. Aedoeagus usually quiescent stage is also interposed between the fi-
elongate and slender; cornuti present or absent. nal sap-feeding and single spinning instars of
$ genitalia with 4 pairs of muscles, Ml absent Chrysaster, Dendrorycter, and Marmara (De-
(Kuznetsov and Stekol'nikov 1987). Female geni- Gryse 1916; Kumata 1978). Moreover, in the lat-
talia. Ovipositor short, with usually 2 pairs of ter two genera the quiescent instar exists in a
moderately short, rarely elongate apophyses; an- pharate condition within the cuticle of the last
terior apophyses sometimes reduced or absent. sap-feeding instar. Later instar larvae of Gracil-
Ostium on S8 or at caudal margin; sterigma lariidae are further characterized by 2 prespirac-
sometimes well sclerotized and variously modi- ular (L) setae on T1 and crochets, if present, on
fied. Ductus bursae usually elongate, slender, A35 and 10 and almost always absent on A6
either membranous or partially sclerotized, ex- (present on A 3 - 6 in Artifodina, Prophyllocnistis,
panding anteriorly into membranous corpus bur- and on A26 in Metriochroa psychotriella, Davis
sae; signa usually of 1 - 2 pairs, or sometimes ab- 1994). Number of instars varies from 4 - 1 1 , de-
sent. pending upon genus and sometimes species (Fitz-
Egg: Usually oval to elliptical, partially flat- gerald and Simeone 1971). Most species are leaf-
tened underneath and slightly convex above with miners (Figs. 7.9 - D ) ; others mine the subepi-
nearly smooth to finely sculptured chorion, dermal layers of new growth stems and fruits or
sometimes with upper and lower halves different bore inside stems and galls (Davis 1991). Early
in texture; less than 0.5 mm in length. Micropyle sap-feeding instars with slashing mandibles typi-
area finely reticulated, often reduced in size. cally feed in a horizontal plane, initiating slen-
Larva (Figs. 7.8 MS): Body usually not exceed- der, subepidermal serpentine mines while ingest-
ing 10 mm, variable in color depending upon ing relatively little solid tissue. In some genera
species and instar, from white to green, with (e. g. Phyllonorycter) the 'sap-feeding' instars in-
some becoming bright red prior to pupation, stead initiate a blotch mine by devouring mostly
with or without darkly pigmented head and scle- spongy parenchymal cells (Watson 1956). Tissue-
114 Donald R. Davis & Gaden S. Robinson

feeding instars possess typical chewing man- Phyllocnistinae: Adults rest with body parallel
dibles that enable these larvae to feed deeper into to surface. Head smooth. Hindwings with Rs
the host, usually ingesting the remainder of the closely parallel to costal margin. Abdomen of $
spongy parenchyma and most of the palisade with S8 membranous; coremata usually present.
cells within the blotch. Later tissue-feeding All feeding instars sap-feeding, creating elongate
instars of the most primitive genera (Gracilla- serpentine, subepidermal mines (Fig. 7.9 D); last
riinae) tend to exit the mine and feed externally, instar larva non-feeding with mandibles absent
often in a rolled-over leaf. Pupa: Head usually (Fig. 7.8 S), legs reduced to unsegmented stubs,
with a serrated or pointed process (cocoon cut- A9 with D bisetose. Pupation within slightly en-
ter) from vertex, or smoothly rounded. Antennae larged cavity at termination of mine. Representa-
extending to A6 to well beyond AIO. Wings to tive genera: Cryphiomystis, Metriochroa, Phylloc-
A56. Tergal spines variably developed, usually nistis, Prophyllocnistis).
small to minute in multiple, irregular rows or A fourth subfamily, Ornichinae, has been pro-
dense concentrations on A2 or 3-7 or 8; Phylloc- posed (Kuznetsov and Stekol'nikov 1987) for
nistis often with a few pairs of large tergal hooks Callisto and Parornix based on the presence of a
on A 2 - 7 . A 4 - 7 moveable in A 4 - 6 in sclerotized anellus and different genital muscula-
(Mosher 1916). Cremaster highly variable, usu- ture. Major characters in the larva, pupa, and
ally present as small paired spines, often with adult wing venation of these genera do not sup-
recurved apices. Pupation usually in a small, port subfamily distinction but instead place them
whitish silken cocoon either outside the host within Gracillariinae.
plant (most Gracillarinnae) or within the mine
(most Lithocolletinae and all Phyllocnistinae).
The Gracillariidae constitute the principal
family of plant mining Lepidoptera. Approxi- The authors wish to thank Vichai Malikul and
mately 2000 species grouped into ca. 75 genera Young Sohn of the Department of Entomology,
are known from all major regions of the world Smithsonian Institution, Washington, D. C.,
except Antarctica (Meyrick 1912 a, b; Vari 1961; USA and Biruta Akerbergs Hansen of Liverpool,
Kumata 1963, 1982; Kumata et al. 1988; Davis Pennsylvania and Andr del Campo Pizzini (de-
1983, 1984). Fossil leafmines of Gracillariidae ceased) for the line illustrations used in this
(Phyllocnistinae) are known from the early Cen- chapter. We also are grateful to Victor Kranz,
omanian of the Dakota Formation in Kansas formerly of the Smithsonian Photographic Labo-
and Nebraska (ca. 97 m. y. a.; Davis 1994; La- ratory and Susann Braden and Walter Brown of
bandeira et al. 1994) near the beginning of the the Smithsonian Scanning Electron Microscope
Angiosperm radiation. This currently represents Laboratory for photographic assistance. Peter
the earliest fossil record of any ditrysian moth Httenschwiler of Uster, Switzerland was helpful
assignable to family. Three subfamilies are recog- in reviewing the section on Psychidae.
Gracillariinae: Adults of most genera rest with References
anterior part of body raised at steep angle. Head
Arbogast, R. T. & Brower, J. H. (1989): External mor-
usually smooth. Hindwings with base of Rs
phology of the eggs of Tinea pallescentella Stainton,
arched toward costa and approaching apex of Sc Tinea occidentella Chambers, and Niditinea fuscello
(Ely 1918; Kumata 1978). Abdomen of $ usually (L.) (Lepidoptera: Tineidae). - Int. J. Insect Mor-
with S8 membranous; coremata usually present. phol. and Embryol., 18 (5/6): 3 2 1 - 3 2 8 .
Last instar larva with fully developed mandibles Bradley, J. D. (1951): Notes on the family Arrheno-
and legs, 5 coxal setae, and A9 with D group phanidae (Lepidoptera: Heteroneura), with special
bisetose. Pupation usually outside of mine. Rep- reference to the morphology of the genitalia, and
resentative genera: Acrocercops, Aristaea, Calop- descriptions of one new genus and two new species.
tilia, Epicephala, Macrostola, Parectopa, Paror- - Entomologist, Dorking, 84 (1059): 1 7 8 - 1 8 5 .
Braun, A. F. (1963): The genus Bucculatrix in America
nix, Philodoria, Stomphastis.
north of Mexico (Microlepidoptera). Mem. Amer.
Lithocolletinae: Adults rest with body parallel Ent. Soc., Philadelphia, 18: 208.
to surface or with head end lowered. Head usu- Clarke, J. F. G. (1971): The Lepidoptera of Rapa Is-
ally with occipital tufts of piliform scales. land. Smithson. Contr. Zool., Washington, 156:
Hindwings with Rs nearly parallel to costal mar- i-iv, 1-282.
gin. Abdomen of $ with S8 sclerotized, elongate; - (1986): Pyralidae and Microlepidoptera of the
coremata absent. Last instar larva (Fig. 7.8 M) Marquesas Archipelago. - Smithson. Contr. Zool.,
usually with fully developed mandibles and legs Washington, 416: 1 - 4 8 5 .
Clausen, C. P. (1962): Entomophagous Insects. -
(both reduced in Cameraria), 4 coxal setae, and
Hafner Pubi. Co., New York.
A9 with D unisetose. Pupation normally within
Clench, H. K. (1959): The Pseudarbelidae, a new fam-
blotch mine (Figs. 7.8 B - C ) . Representative gen- ily of psychoid moths, with description of a new
era: Cameraria, Chrysaster, Cremastobombycia, species from New Guinea. - Tijds. v. Ent., 102
Leucanthiza, Phyllonorycter (= Lithocolletis). (3-4): 223-229.
The Tineoidea and Gracillarioidea 115

Common, I. F. B. (1990): Moths of Australia. - Mel- Tineoidea). Proc. Ent. Soc. Washington, 88(1):
bourne University Press, Carlton, vi + 535 p. 98-109.
Davis, D. R. (1964): Bagworm moths of the Western - & Milstrey, E. G. (1988): Description and biology
Hemisphere (Lepidoptera: Psychidae). - U. S. Nat. of Acrolophus pholeter (Lepidoptera: Tineidae), a
Mus. Bull., Washington, 255: 1 - 2 3 3 . new moth commensal from gopher tortoise bur-
- (1975): A review of the West Indian moths of the rows in Florida. - Proc. Ent. Soc. Washington,
family Psychidae with descriptions of new taxa and 90(2): 164-178.
immature stages. - Smithson. Cont. Zool., Wash- - & Hogue, C. L. (1992): Neotropical Microlepidop-
ington, 188: iiii, 1 - 6 6 . tera XXV. A new arboreal species of Acrolophus
- (1983): Micropterigidae, Eriocraniidae, Acanthop- from Peru (Lepidoptera: Acrolophidae). - J. New
teroctetidae, Hepialidae, Opostegidae, Tischeriidae, York Ent. Soc., New York.
Incurvariidae, Heliozelidae, Tineidae, Psychidae, DeGryse, J. J. (1916): The hypermetamorphism of the
Ochsenheimeriidae, Lyonetiidae, Gracillariidae, lepidopterous sapfeeders. - Proc. Ent. Soc. Wash.
p. 2 - 1 1 . In Hodges, R. W. et al., eds., Check list of 18: 164-168.
the Lepidoptera of America north of Mexico. - Dierl, W. (1970): Compsoctenidae: Ein neues Taxon
E. W. Classey Ltd. and the Wedge Entomological von Familienstatus (Lepidoptera). - Verff. Zool.
Research Foundation, London. Staatssammlung Mnchen, 14: 141.
- (1984): Neopseustidae, Nepticulidae, Opostegidae, - (1996): Der Familienstatus von Lypusa maurella
Tischeriidae, Incurvariidae, Cecidosidae, Adelidae, Denis und Schiffermller, 1775 (Insecta, Lepidop-
Prodoxidae, Tineidae, Psychidae, Arrhenophani- tera). - Spixiana 19: 3 9 - 4 2 .
dae, p. 16-25. In Hepper, J. R., ed.., Atlas of Neo- Dietz, W. (1905): Revision of the genera and species
tropical Lepidoptera, 1 (Checklist, pt. 1). Dr. W. of the Tineid subfamilies Amydriinae and Tineinae
Junk, The Hague. inhabiting North America. - Trans. Am. Ent. Soc.,
- (1987): Micropterigidae, Eriocraniidae, Acanthop- 31: 1 - 9 6 .
teroctetidae, Nepticulidae, Opostegidae, Tischerii- Dugdale, J. S. (1988): Lepidoptera - annotated cata-
dae, Heliozelidae, Adelidae, Incurvariidae, Prodox- logue and keys of the family-group taxa. - Fauna
idae, Tineidae, Psychidae, Ochsenheimeriidae, Lyo- N. Zeal. 14: 1 - 2 6 2 .
netiidae, Gracillariidae, Epipyropidae, p. 341-378, Ely, C. R. (1918): A revision of the North American
456, 459-460. In F. W. Stehr, ed., Immature insects. Gracilariidae [sic] from the standpoint of venation.
- Proc. Ent. Soc. Washington, 19 ( 1 - 4 ) : 2 9 - 7 7 .
- Kendall/Hunt Pubi. Co., Dubuque, Iowa.
Fitzgerald, T. D. & Simeone, J. B. (1971): Description
- (1988): The tineoid complex: a review of the bio-
of the immature stages of the sap feeder Marmara
logy, composition, and phylogeny of the most prim-
fraxincola (Lepidoptera: Gracillariidae). - Ann.
itive ditrysian moths. - Proc. XVIII Int. Cong.
Ent. Soc. Amer. 64 (4): 765-770.
Ent., Vancouver, p. 79, [abstract],
Fletcher, T. B. (1921): Life histories of Indian insects.
- (1990a): Neotropical Microlepidoptera XXII. First
Microlepidoptera. VIII. Tineidae and Nepticulidae.
report of the family Eriocottidae from the New
Mem. Dep. Agrie. India, 6: 181-196.
World, with descriptions of new taxa. Proc. Ent.
- (1933): Life histories of Indian Microlepidoptera
Soc. Washington, 92 (1): 1 - 3 5 .
(Second Series). Cosmopterygidae to Neopseusti-
- (1990 b): Three new species of Acrolophus from the dae. Sci. Monogr., Imp. Coun. Agrie. Res. India,
southeastern United States with remarks on the 4: 1 - 8 5 .
status of the family Acrolophidae (Lepidoptera: Forbes, W. T. M. (1923): The Lepidoptera of New
Tineoidea. - Proc. Ent. Soc. Washington, 92 (4): York and neighboring states. - Mem. Cornell agrie.
694-704. Exp. Sta., 68: 1 - 7 2 9 .
- (1991): First Old World record of the moth family Gaedike, R. (1974): Revision der palaarktischen
Arrhenophanidae (Lepidoptera: Tineoidea). - Douglasiidae (Lepidoptera). - Acta faun. ent. Mus.
Trop. Lep., Gainesville, 2 (1): 4 1 - 4 2 . Nat. Pragae, Prague, 15 (176): 79-102.
- (1994): New leaf-mining moths from Chile, with re- - (1990): Revision der nearktischen Douglasiidae
marks on the history and composition of Phylloc- (Lepidoptera). - Beitr. Ent. Berlin, 2: 287-300.
nistinae (Lepidoptera: Gracillariidae). - Trop. Gomez Bustillo, M. R. (1978): Reestructuracin del
Lepid., Gainesville, 5 (1): 6 5 - 7 5 . complejo Psychidae (sensu lato) con el estableci-
- (1996): Neotropical Tineidae, VI: Prosetomorpha miento de un gnero y una familia y subsiguiente
falcata, a new genus and species of Setomorphinae revisin sistemtica de la fauna Ibrica (Lepidop-
(Lepidoptera: Tineoidea) from Colombia associ- tera-Tineoidea) (1 parte). - SHILAP, Revista
ated with curculionid galleries in stems of Solanum. Lepid., 6(24): 325-331.
- Proc. Ent. Soc. Washington, 98 (2): 173-187. - (1984): Revisin de los Micropsychiniidae (Gmz.-
- (in press a): A world classification of the Har- Btllo., [1979]) de la Pennsula Ibrica (Lep. Ti-
macloninae, a new subfamily of Tineidae (Lepidop- neoidea). - SHILAP, Revista Lepid., 12(47):
tera: Tineoidea). - Smithson. Cont. Zool., Wash- 183-193.
ington. Gozmany, L. A. & Vari, L. (1973): The Tineidae of
- (in press b): A revision of the genus Lamyristis the Ethiopian Region. Transv. Mus. Mem., 18:
Meyrick and proposal of a new genus Acoremata i-iv, 1-238.
from southeast Asia (Lepidoptera: Psychidae). - Httenschwiler, P. (1989): Genus Scoriodyta Meyrick,
Proc. Ent. Soc. Washington. 1888, a new subfamily and description of a new spe-
- , Clayton, D. H Janzen, D. H. & Brooke, A. P. cies and forms (Lepidoptera: Psychidae). New
(1986): Neotropical Lepidoptera, II: Biological Zealand J. Zool. 16: 5 1 - 6 3 .
notes and descriptions of two new moths phoretic Hasbrouck, F. F. (1964): Moths of the family Acro-
on spiny pocket mice in Costa Rica (Lepidoptera: lophidae in America north of Mexico (Microlepi-
116 Donald R. Davis & Gaden S. Robinson

doptera). - Proc. U. S. Nat. Mus., Washington, 114 (1990): Tentative reclassification of holarctic Ypo-
(3475): 4 8 7 - 7 0 6 . nomeutoidea (Lepidoptera). - Nota lepid., 13(1):
Henderickx, H. (1987): Morphologie et discussion du 28-42.
status spcimens mles de Luffia au sein d'une pop- Labandeira, C. C., Dilcher, D. L., Davis, D. R. &
ulation parthnogntique (Lep., Psychidae). - Wagner, D. L. (1994): Ninety-seven million years of
Alexanor, 15(3): 141-149. angiosperm-insect association: Paleobiological in-
Heinemann, H. (& Wocke, M. F) (1870(-1877)): Die sights into the meaning of coevolution. Proc.
Schmetterlinge Deutschlands und der Schweiz II/2. Natl. Acad. Sci. USA, Washington, 91: 12278-
- C. A. Schwetschke & Sohn, Braunschweig. 12282.
Heppner, J . B . (1987): Douglasiidae, p. 4 0 8 - 4 0 9 . In Martin, J. L. (1956): The bionomics of the aspen
F. W. Stehr, ed. Immature insects. - Kendall/Hunt blotch miner, Lithocolletis salicifoliella Cham. (Lep-
Pubi. Co., Dubuque, Iowa. idoptera: Gracillariidae). - Can. Ent. 88(4): 1 5 5 -
- (1991): Faunal regions and the diversity of Lepi- 168.
doptera. - Trop. Lepid., 2 (suppl. 2): 1 - 8 5 . Meyrick, E. (1912 a): Adelidae, Micropterygidae,
Hinton, H. E. (1956): The larvae of the species of Tine- Gracilariadae. Lepidopterorum Catalogus, pars 6,
idae of economic importance. - Bull. ent. Res., 47: p. 2 5 - 6 8 .
251-346. (1912 b): Lepidoptera Heterocera (Tineae), Fam.
Hudson, G. V. (1928): The butterflies and moths of New Gracilariadae. - Genera Insectorum, p. 1 - 3 6 ,
Zealand. - Wellington, 386 p., 52 pis. 1 col. pi.
- (1939): A supplement to the butterflies and moths of Moriuti, S. (1978): Amphitheridae (Lepidoptera): Four
New Zealand. - Wellington, [vi] + p. 3 8 7 - 4 8 1 , pis. new species from Asia, Telethera blepharacma
53-62. Meyrik new to Japan and Formosa, and Spheno-
Johnson, N. E. & Martin, P. (1969): Amydria effren- graptis Meyrick transferred to the family. - Bull.
tella from nests of mountain beaver, Aplodontia Univ. Osaka Prefl, Osaka, ser. , 30: 1 - 1 7 .
rufa. - Ann. Ent. Soc. Amer., 62: 396399. Mosher, E. (1916): A classification of the Lepidoptera
Kozhanchikov, I. V. (1956): Nasekomye tscheschuek- based on characters of the pupa. - Bull. Illinois
rylye. - Fauna SSSR, 3 (2): 1 - 5 1 6 . M o s k a u - L e - State Lab. Nat. Hist., Urbana (12 (2): 1 - 1 6 5 .
ningrad [In Russian] [English translation 1969. Narbel, M. (1946): La cytologie de la Parthenogenese
Smithsonian Institution and the National Science chez Apterona helix Sieb. - Rev. suisse Zool. 53:
Foundation, Washington, D. C.]. 625-681.
Kumata, T. (1963): Taxonomic studies on the Litho- Nielsen, E. S. (1978): On the systematic position of the
colletinae of Japan (Lepidoptera: Gracillariidae), genus Eriocottis Zeller, 1847, with remarks on the
pts. 1 - 2 . - Ins. Matsum. 25(2): 5 3 - 9 0 ; 26(1): phylogeny of primitive Tineoidea (Lepidoptera). -
1-48. Ent. scand., 9: 2 7 9 - 2 9 6 .
- (1978): A new stem-miner of alder in Japan, with a & Common, I. F. B. (1991): Lepidoptera (Moths
review of the larval transformation in the Gracilla- and butterflies). In CSIRO, The Insects of Australia,
riidae (Lepidoptera). Ins. Matsum. new ser. 13: Melbourne University Press, Carlton, Victoria, 2:
1-27. 817-915.
Robinson, G. S. (1976): A taxonomic revision of the
- (1982): A taxonomic revision of the Gracillaria
Tinissinae of the world (Lepidoptera: Tineidae).
group occurring in Japan (Lepidoptera: Gracillarri-
Bull. Br. Mus. nat. Hist. Ent., 32: 2 5 3 - 3 0 0 , pis.
idae). Ins. Matsum., new ser. 26: 1 186.
- , Kuroko, H. & Ermolaev, V. P. (1988): Japanese spe-
(1979): Clothes - moths of the Tinea pellionella
cies of the Acrocercops-gToup (Lepidoptera: Gracil-
complex: a revision of the world's species (Lepidop-
lariidae), pts. 1 - 2 . - Ins. Matsum., new ser. 38:
tera: Tineidae). - Bull. Br. Mus. nat. Hist. Ent., 38:
1 - 1 1 1 ; 40: 1 - 1 3 3 .
5 7 - 1 2 8 , figs. 1 - 1 0 3 .
Kuznetsov, V . l . & Stekol'nikov, A . A . (1976, 1977):
(1980 a): The Himalayan Nemapogoninae, with a
Phylogenetic relationship between the superfamilies
note on the female of Nemapogon levantinus Pe-
Psychoidea, Tineoidea, and Yponomeutoidea (Lep-
terson. - Entomologist's Gaz. 31: 129142.
idoptera), taking into account the functional mor-
(1980 b): Cave-dwelling tineid moths: a taxonomic
phology of the male genital apparatus. I, II. - Ent.
review of the world's species (Lepidoptera: Tine-
Rev. [English translation of Ent. Obozr.], 55 (3):
idae). Trans. Br. Cave Res. Ass., 7: 83120.
(1981): Remarks on the classification of the fungiv-
- (1987): Functional morphology of the male genita-
orous Tineidae with special reference to the Tinissi-
lia and notes on the classification and phylogenetic
nae (Lepidoptera). - Ent. scand., 12: 363-380.
relationships of mining moths of superfamily Grac-
(1986): Fungus moths: a review of the Scardiinae
illarioidea (Lepidoptera). - Ent. Rev. [English
(Lepidoptera: Tineidae). Bull. Br. Mus. nat. Hist.
translation of Ent. Obozr.], 66(3): 1 6 - 3 0 .
Ent., 52: 3 7 - 1 8 1 , figs. 1 - 2 0 0 .
Kyrki, J. (1983 a): Roeslerstammia Zeller assigned to (1988): A phylogeny for the Tineoidea. - Ent.
Amphitheridae, with notes on the nomenclature scand., 19(2): 117-129.
and systematics of that family (Lepidoptera). & Nielsen, E. S. (1993): Tineid genera of Australia.
Ent. scand., 14: 3 2 1 - 3 2 9 . - Monogr. Austr. Lepid. 2, xvi + 344 p. CSIRO,
- (1983 b): Adult abdominal sternum II in the ditry- Melbourne.
sian tineoid superfamilies morphology and phy- & Tuck, K. R. (1996): A revisionary checklist of the
logenetic significance (Lepidoptera). - Ann. Ent. Tineidae (Lepidoptera) of the Oriental region -
Fenn., 49: 8 9 - 9 4 . Occ. Pap. Syst. Ent., 9: 1 - 2 9 .
- (1984): The Yponomeutoidea: a reassessment of the (1977): Phylogeny and composition of the Hier-
superfamily and its suprageneric groups (Lepidop- oxestidae (Lepidoptera: Tineidae). Syst. Ent.
tera). - Ent. scand., 15: 7 1 - 8 4 . 22: 3 6 3 - 3 9 6 .
The Tineoidea and Gracillarioidea 117

Robinson, R. (1971): Lepidoptera genetics. Perga- Suomalainen, E. (1962): Significance of parthenogene-

mon Press, Oxford, New York. sis in the evolution of insects. - Ann. Rev. Ent. 7:
Sauter, W. (1982): ber das Weibchen von Meessia 349-366.
gallica Pet. nebst einigen systematischen Bemer- Torre, K. W. von Dalla & Strand, E. (1929): Psychidae
kungen zu den Tineidae (Lepidoptera). - Mitt. . Lepidop. Cat., Pars 34, 1211.
Schweiz. Entom. Ges., 55: 396-397. Trgardh, I. (1913): Contributions to the comparative
- & Httenschwiler, P. (1991): Zum System der pa- morphology of the trophi of the lepidopterous leaf
laearktischen Psychiden. (Lep. Psychidae) 1. Teil: miners. - Arkiv Zool 8 (9): 1 - 4 9 , 67 figs.
Liste der palaearktischen Arten. - Nota lepid., Tutt, J. W. (1900): A natural history of the British Lepi-
14(1): 6 9 - 8 9 . doptera. Swan Sonnenschein, London, etc., viii
Scoble, M. J. & Scholtz, C. H. (1984): A new gall-feed- + 584 p.
ing moth (Lyonetiidae: Bucculatricinae) from South Vari, L. (1961): Lithocolletidae, South African Lepi-
Africa with comments on larval habits and phyloge- doptera, vol. 1. - Transvaal Museum, Pretoria,
netic relationships. - Syst. Ent., 9: 8 3 - 9 4 . Mem. 12, xix + 238 p., 112 pis.
Seiler, J. (1920): Geschlechtschromosomen-Unter- Zagulajev, A. K. (1960): Tineidae; part 3 - subfamily
suchungen an Psychiden I. Experimentelle Beein- Tineidae. [In Russian]. - Fauna SSSR, 78: 1 - 2 6 7 ,
flussung der geschlechtsbestimmenden Reifeteilung 231 figs., 3 pis. [English translation, 1975, New
bei Talaeporia tubulosa. R Arch. Zellforsch. 15: Delhi.]
249-268. - (1964): Tineidae; part 2 - subfamily Nemapogoni-
- (1923): Geschlechtschromosomen-Untersuchungen nae. [In Russian], - Fauna SSSR, 86: 1 - 4 2 4 , 385
an Psychiden. IV. Die Parthenogenese der Psychi- figs., 1 pi. [English translation, 1968, Jerusalem.]
den. - Z. indukt. Abstamm. u. Vererb.-Lehre, 31: - (1973): Tineidae; part 4 - subfamily Scardiinae. [In
1-96. Russian.] Fauna SSSR, 104: 1 - 1 2 6 , 99 figs., 2 pis.
- (1949): Das Intersexualittsphnomen. Zusammen- - (1975): Tineidae; part 5 - subfamily Myrmecozeli-
fassende Darstellung und Deutung. - Experientia, nae. [In Russian.] - Fauna SSSR, 108: 1 - 4 2 6 , 319
Basel, 5: 425-438. figs., 8 pis. [English translation, 1988, New Delhi.]
- (1959): Untersuchungen ber die Entstehung der - (1979): Tineidae; part 6 - subfamily Meessiinae. -
Parthenogenese bei Solenobia triquetrella F. R. Fauna SSSR, 119: 1 - 4 0 8 .
(Lepid. Psychid.). I. Mitt. Die Zytologie der bi- - (1988): Grass stem moths, families Ochsenheimerii-
sexuellen Solenobia triquetrella, ihr Verhalten und dae and Eriocottidae. Lepidoptera. [In Russian.] -
ihr Sexualverhltnis. - Chromosoma, Berlin, 10: Fauna SSSR, 135: 1 - 3 0 2 .
- (1960): Untersuchungen ber die Entstehung der
Parthenogenese bei Solenobia triquetrella E R.
(Lepidoptera, Psychidae). II. Analyse der diploid Previous work on the skeleto-muscular structure
parthenogenetischen S. triquetrella. Verhalten, Auf- of tineid male genitalia by V. I. Kuznetsov and
zuchtresultate und Zytologie. - Chromosoma, Ber- A. A. Stekol'nikov has been supplemented by
lin, 11: 29-102. additional contributions from the same authors
- (1961): Untersuchungen ber die Entstehung der
in 1995 and 1997; both morphological and phy-
Parthenogenese bei Solenobia triquetrella F. R.
logenetic issues were addressed.
(Lepidoptera, Psychidae). III. Die geographische
Verbreitung der drei Rassen von Solenobia trique- Norman et al. (1995) listed the SE Asian
trella in der Schweiz und in angrenzenden Lndern. Psychidae and showed the taxonomic usefulness
- Zeits. Vererb.-lehre, 92: 261-316. of wing scale characters in the family.
- (1963): Untersuchungen ber die Entstehung der Norman, K., Robinson, G. S. & Basri, M. W. (1995):
Parthenogenese bei Solenobia triquetrella F. R. Common bagworm pests (Lepidoptera: Psychidae)
(Lepidoptera, Psychidae). IV. Mitteilung. Wie besa- of oil palm in Malaysia with notes on related
men begattete diploid und tetroploid parthenogene- South-east Asian species. Malay. Nat. J., 48:
tische Weibchen von S. triquetrella ihre Eier? - 93-123.
Zeits. Vererb.-lehre, 94: 2 9 - 6 6 . Kuznetsov, V. I. & Stekol'nikov, A. A. (1997): Phylo-
Seksjeva, S. V. (1994 a): Review of the mining moths genese relations of some subfamilies of true moths
(Lepidoptera, Bucculatricidae) of Russia. - Proc. (Lepidoptera, Tineidae) based on a new study of
Zool. Inst. Russian Acad. Sci. (St. Petersburgh), functional morphology of male genitalia. Ent.
255: 99-120. [In Russian], Rev. [English translation of Ent. Obozr.], 76:
- (1994 b): New intrageneric groups within the genus 362-372.
Bucculatrix (Lepidoptera, Bucculatricidae) desig- Stekol'nikov, A. A. & Kuznetsov, V. I. (1995): New ar-
nated on the basis of the morphology of male geni- chaic characters of the family Tineidae (Lepidop-
talia. - Zool. Zhurn. (Moscow), 73(2): 114-122. tera) and correction of the hypothesis concerning
[In Russian]. [English trans.: 1995, Ent. Rev. (New the origin of some male genital appendages. - Ent.
York), 74(2): 111-119], 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- jaeva (1988). Female anterior apophyses usually
semblage of relatively primitive micro-moths. continued into dorsal and ventral costae at base
Kyrki (1984) first recognized apomorphies indi- (Fig. 8.1 D); lamella antevaginalis unsclerotized.
cating the monophyly of this group but, as yet, Larva with prothoracic L usually trisetose (in
precise delimination of the superfamily remains post-1st. instars), occasionally (almost all Glyph-
debatable. ipterigidae, a few Yponomeutidae and Ypsoloph-
The principal yponomeutoid autapomorphy is idae) bisetose. Pupal abdomen lacking spine
the presence, in males, of posterior expansions of bands (single spines sometimes associated with
pleuron VIII (Figs. 8.1 F - G ) . The pleural lobes setae D1 and SD), segments I - I V immobile,
enclose the genital apparatus and their size is pupa not protruded prior to eclosion.
broadly correlated with that of the latter, the The controversial phylogenetic significance of
valves in particular; in the Plutellidae/Acrolepii- the tergal spinosity and pupal structure is dis-
dae and in the Heliodinidae the lobes are quite cussed in 1 - 2 . The yponomeutoids as now de-
small and narrow. In the Bedelliidae and Glyph- limited comprise the bulk of those non-apoditry-
ipterigidae (Fig. 8.1 H) the lobes are not present sian Ditrysia which have non-protruded pupae
as discrete formations, and in these extra ad hoc (conditions in Glyphipterigidae-Orthoteliinae
hypotheses are needed to justify their assignment may represent an autapomorphic character re-
to the superfamily. Another possible yponomeu- versal) and whose adults do not have the charac-
toid groundplan autapomorphy is the presence teristic gelechioid scaled proboscis.
of a transverse costa behind the anterior margin
of sternum II (Fig. 8.3 C); this costa is absent in Heppner (1987) corrected his earlier statements
Ypsolophidae-Ochsenheimeriinae and some Lyo- (1977) about the presence of pupal protrusion in ypo-
netiidae. nomeutoid families.

Ocelli and chaetosemata present or absent. The formal classification of the superfamily
Antenna simple, scape with or without pecten or has so far varied considerably. The family Ypo-
scale awning. Proboscis unsealed, usually well nomeutidae has at times comprised the taxa now
developed; maxillary palp small, 1 4 segmented, assigned to the Plutellidae and Ypsolophidae
sometimes appressed to proboscis base. Labial Ypsolophinae, but thus delimited it is almost cer-
palp drooping, porrect or ascending, not mark- tainly paraphyletic. A phylogenetic analysis of
edly elongate and recurved above head; apical the superfamily by Kyrki (published posthu-
segment with vom Rath's organ basally or near mously 1990) led to a reclassification, which is
midlength. Epiphysis present, spurs usually 0-2- followed by Scoble (1992) and in the present
4, occasionally reduced, hind tibiae usually work (except that the Acrolepia-group is treated
smooth-scaled. Forewing with pterostigma and as a distinct family). The tentative nature of this
chorda present or absent, Rs4 nearly always arrangement must be emphasized, and the mo-
postapical, M-stem in cell usually vestigial or ab- nophyly of the redefined Yponomeutidae is still
sent. Female frenulum usually of only two bris- in obvious need of further scrutiny. The family/
tles. Sternum II of tineoid type. Abdominal ter- subfamily position of a number of extra-Hol-
gal spinosity variably developed, often absent. arctic genera (listed by Kyrki 1984) which have
Male postabdomen usually with paired ventro- been assigned to the Yponomeutidae s. lat. re-
lateral coremata between pleural lobes and IX; mains in doubt.
X almost never forming well developed median
uncus (one known exception is the New Zealand
Cadmogenes, which may not belong in the Ypo- Key to yponomeutoid families
nomeutoidea); socii often well developed, flank- (based on Holarctic taxa)
ing anal tube, or absent (Glyphipterigidae, also 1 Ocelli present, rarely rudimentary (if rudi-
the Australian/New Zealand 'plutellid' Or- mentary, maxillary palp normally 34 seg-
thenches); gnathos present or absent, sometimes mented, porrect, and forewing pterostigma
fused medially with socii; subscaphium strongly extending to Rs2) 2
developed and strap-like, or weakly developed.
Ocelli absent 6
Valve base with strong apophysis for extrinsic
2(1) Wings pronouncedly narrow, forewing
muscle. The male genital musculature in ypono-
with only 5 postapical veins arising from
meutoids and its (phenetic-)systematic implica-
cell (Fig. 8.3 D) Heliodinidae
tions are discussed by Kuznetzov & Stekolnikov
Wings less narrow, forewing with 6 posta-
(1976-77, 1993) and Kuznetzov, Kozlov & Seks-
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 moreover the Scythropiinae have a long pteros-
palp often with transverse light and dark tigma, while the short kind also occurs within
bars; forewing often with metallic mark- the Yposolophidae. A difficulty with the inter-
ings; male genitalia partially enclosed in a pretation of the particularly strong melanization
posteriorly expanded tergopleural 'cone', of the male sternum VIII as a family autapomor-
i. e. distinct lateral lobes not developed phy is that the unspecialized state of this plate
(Fig. 8.1 H) in the subfamily Yponomeutinae must, then, be
. . . . Glyphipterigidae (Glypiphteriginae) secondary. The abrupt narrowing of the apical
Head rough-scaled (Fig. 8.1 B); labial palp hindwing area in front of Se + R (Fig. 8.1 L), a
without transverse bar pattern; forewings specialization characteristic of many yponomeu-
usually without metallic markings (such tids (Minet 1986, Scoble 1992), also occurs else-
markings do occur in some S. Hemisphere where in the superfamily (Fig. 8.1 MN).
taxa); male pleural area VIII/IX with well Second segment of labial palp usually without
developed lateral and/or ventral lobe . . 4 scale tuft. Larva with prolegs shorter than broad;
4 (3) Hindwing Rs and M l stalked or coincident on segment IX SV bisetose.
(Fig. 8.1 M) Ypsolophidae The taxa brought together in this family are
Hindwing Rs and M separate 5 diverse in facies, structure and bionomics. Six
5 (4) Hindwing with M3 and CuAl separate . . groups are recognized and given subfamily status
Plutellidae in the present classification; their phylogenetic
Hindwing with M3 and CuAl stalked . . . interrelationships remain unclarified. This classi-
Acrolepiidae fication, as that of the Yponomeutoidea as a
6(1) Wings greatly narrowed; hindwing vena- whole, is primarily based on a study of the Hol-
tion markedly simplified, with at most 4 arctic genera (Moriuti 1977, Kyrki 1984, 1990),
pre-CuP veins 7 and its applicability to tropical/south temperate
Wings less narrowed; hindwing venation faunas remains to be tested.
more nearly complete, with at least 6 pre- Friese (1960), Kuznetzov & Stekolnikov
CuP veins 8 (1976-77), Moriuti (1977), Hannemann (1977),
7 (6) Forewings simply lanceolate, R s 2 - 4 and Gershenzon (1990).
an M branch on common stalk from
pointed apex of discal cell; scape with
dense awning, but not expanded into genu- Key to yponomeutid subfamilies
ine eye cap Bedelliidae
Forewing apex narrowly produced, Rs2 1 Chaetosema present; male hind tibia with-
arising before apex (which is not a single out spurs, reduced (like the tarsus)
point) of discal cell, or forewing without Attevinae
closed discal cell; scape expanded into eye Chaetosema absent; male hind tibia with
cap (Fig. 8.4 J) Lyonetiidae spurs 2
8 (6) Rs3 apical/postapical; monobasic W. Pa- 2 (1) Antennal scape without pecten Praydinae
laearctic taxon, large micro-moth with Antennal scape usually with pecten . . . 3
broad, angular wings (Fig. 8.2 N), associ- 3 (2) Proboscis reduced, short and soft; pteros-
ated with freshwater plants tigma extending distad beyond R
Glyphipterigidae (Orthoteliinae) Scythropiinae
Rs3 apical/preapical . . . . Yponomeutidae Proboscis well developed; pterostigma,
when present, not extending distad beyond
Yponomeutidae. Autapomorphies few and obvi- R 4
ously homoplasious. Ocelli absent. Pretarsus 4 (3) Abdominal terga with normal scales only
('claw') of larval thoracic legs unusually long and Argyresthiinae
straight; absence of this trait in the endophagous Abdominal terga with spines (modified
Argyresthiinae interpreted by Kyrki (1990) as scales (Fig. 8.1 F) 5
secondary. Larval proleg crochets usually (the 5 (4) Male sternum VIII more strongly mela-
endophagous argyresthiines and a few ypono- nized than VII; segment X processes with
meutines excepted) bi-/multiserial; polarizing this specialized scales Saridoscelinae
character is difficult, however, since while Male sternum VIII not more strongly mel-
crochets are uniserial in tineoids a bi-/multiserial anized than VII; segment X processes un-
arrangement is predominant in those non-ditry- sealed Yponomeutinae
sians and gracillarioids which do have crochets.
Other possible groundplan autapomorphies of Yponomeutinae. Perhaps a paraphyletic as-
the family enumerated by Kyrki are even more semblage. Uniform melanization of male abdo-
questionable: there is no obvious support for the minal sterna VII and VIII diagnostic, but proba-
interpretation of the relative shortness of the bly plesiomorphic; this applies also to spinose
commonplace yponomeutoid pterostigma (which abdominal terga (Fig. 8.1 F) shared only with the
does not extend distad beyond R) as derived; 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- ration, and in having the socii set with fringed
ten. Maxillary and labial palps variable. Spurs sensilla recalling those of the Argyresthiinae.
0-2-4. In male genitalia (Fig. 8.1 E) saccus and Hindwing with all three M branches.
segment X processes ('socii' in principal taxo- Saridoscelis (4 described species and at least
nomic treatments) usually prominent, the latter twice as many undescribed, Asia; Fig. 8.2 C).
unsealed. As in the Saridoscelinae the phallus Larvae in communal webs on Ericaceae.
usually has a basal 'scape'. Larva with cranial Scythropiinae. Monogeneric. Probable autapo-
seta VI long, PI below A f 2 - P 2 line. morphies are the reduced proboscis and dorsally
A small genus group is characterized by plesi- hairy hind tibia.
omorphic traits including 4-segmented maxillary Scape with pecten. Maxillary palp 4-seg-
palp (Fig. 8.1 C; basal segment synscleritous with mented. Pterostigma extending distad from R.
stipes, hence palp described as 3-segmented by Abdominal terga without spines. Phallus without
Moriuti 1977), long 'basal fork' of forewing anal basal 'scape'. Socii unsealed.
veins, and hindwing with all three M-branches Larva with cranial seta VI long, PI below
present. This group has been given subtribe rank A f 2 - P 2 line.
(by Moriuti 1988 whose tribe Yponomeutini Scythropia (2 species, European; Fig. 8.2 D).
equals the present subfamily Yponomeutinae), Larva on woody Rosaceae, initially leafmining,
but there is no evidence for its monophyly. At eventually feeding externaly in a communal web.
least the genera Niphonympha (3 species, Eura- Attevinae. Probable autapomorphies are the
sian, Mexico) and Thecobathra (13 species, S. E. presence of chaetosema, the reduction of hindleg
Asian) pertain to this grade; the known larvae tibia and tarsus (particularly in males where tib-
feed on leaves of Fagaceae and Hamamelida- ial spurs are absent), the presence of two sub-
ceae. ventral setae in larval meso- and metathorax,
In the bulk of the Yponomeutinae the maxil- and the concealment of the labial palp in the
lary palp is only 2-segmented, the forewing anal pupa.
fork is very short, and the hindwing has only two Large, colourful (markings often red/orange/
M veins (Fig. 8.1 L). The near-cosmopolitan (not yellow) and probably aposematic micro-moths.
New Zealand) Yponomeuta ('ermine moths', 80 + Scape without pecten. Maxillary palp with 3 seg-
species; Fig. 8.1 P, 8.2 B) mostly have larvae liv- ments. No pterostigma. Abdominal terga with-
ing in communal webs on (botanically diverse) out spines. Phallus without basal 'scape'. Tegu-
trees and scrubs. Mass occurrence, with forma- men bearing an unsealed prong (probably an-
tion of highly voluminous and conspicuous webs other subfamily autapomorphy); socii complex;
occurs in a number of species, and some have gnathos W-shaped.
economical importance. Recent concerted inves- Larva with cranial seta VI short, PI not below
tigations of the ecology, behaviour, physiology A 2 - P 2 line.
and evolutionary biology of European species Atteva (50+ species, pantropical; Fig. 8.2 A)
have been reviewed by Menken, Herrebout & larvae live in communal webs recalling those of
Wiebes (1992). The more narrow-winged Zelleria Yponomeuta, but looser; host families include Fa-
(50 + species) have a characteristic resting posi- baceae, Simaroubaceae, Lauraceae and Aralia-
tion with the head end close to the substrate and ceae.
the abdomen lifted (as in argyresthiines, Fig. Praydinae. Probable autapomorphy an en-
8.1 K); ecology diverse, comprising webmakers larged (broader than long) male sternum VIII
on broad-leaved trees/scrubs as well as pine nee- (Fig. 8.1 G). Female anterior apophyses greatly
dle miners. The closely related Xyrosaris (Aus- reduced, without branched costa at base (Mori-
tralasian/Neotropical) is aberrant in having a uti 1977). Loss of forewing 'chorda' (mentioned
prominent ventral scale tuft on labial palp seg- by Kyrki 1984) parallelled in several Yponomeu-
ment 2 (recalling that present in some other ypo- tinae.
nomeutoid families). Other principal genera in-
Scape without pecten. Maxillary palp 1-seg-
clude Kessleria (Huemer & Tarmann 1991), Cal-
mented. Pterostigma not extending distad from
lithrinca and Swammerdammia. Ocnerostoma
R. Abdominal terga without spines. Phallus
(Holarctic) is exceptional in that the larvae (nee-
without basal 'scape'. Socii unsealed. Larva with
dle miners) have bisetose prothoracic L (proba-
cranial seta VI short, PI not below Af2P2 line.
bly related to the endophagy) and uniserial pro-
Prays (25 + species, Old World; Fig. 8.2 E).
leg crochets; it belongs to a small group with re-
Larvae on diverse woody plants, endophytic
duced proboscis. Altogether the subfamily com-
(leaf miners or borers in flower buds or shoots)
prises 360 + species worldwide.
in all instars, or older larvae feed externally in
In the following subfamilies the male sternum light webs; P. citri and P. oleae are pests on citrus
VIII is more strongly sclerotized than VII (Fig. and olives respectively.
8.1 G), and often of a characteristic V- or Y-like Argyresthiinae. Principal autapomorphies are
shape. in male genitalia (Fig. 8.11J): vinculum more
Saridoscelinae. Monogeneric, differing from or less produced laterally and lacking a median
preceding mainly in male sternum VIII configu- 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 Fig. 8.2 F) are narrow-winged moths with di-
(shorter stalks occur in a few Yponomeutinae). verse, often metallic forewing patterns, larvae
Larvae usually endophagous, rarely in loose bud-, fruit-, leaf- and twig-miners on a variety of
webs; with simple pretarsi of thoracic legs (auta- conifers and woody dicots, some attaining pest
phomorphic character reversal?) and uniserial status.
crochets on prolegs. Resting position of adult Friese (1969), Gershenzon (1990).
characteristic (Fig. 8.1 K), with head end lowered
and abdomen lifted (as in Yponomeutinae-Ze/-
leria). Putative clade Ypsolophidae +
(Plutellidae + Acrolepiidae + Glyphipterigidae +
Scape with pecten. Maxillary palp 1-seg-
(Heliodinidae + Bedelliidae + Lyonetiidae))
mented. Pterostigma not extending distad from
R. Abdominal terga without spines. Phallus The monophyly of this entity is weakly sup-
without basal 'scape'. ported. Kyrki (1990) enumerated three possible
Larva with cranial seta VI short, PI below autapomorphies: Ventral scale tuft on second la-
A f 2 - P 2 line. 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- complement on the larval prolegs (a single row,
bers of the heliodinid/bedelliid/lyonetiid assem- or complete absence).
blage). Larval prolegs (Fig. 8.10) longer than Larvae initially leaf miners in Poaceae
wide (not applicable to endophagous glyphipteri- (mainly), Cyperaceae and Juncaceae; later in-
gids and lyonetiids with reduced prolegs, nor to stars are stem borers.
certain genera currently placed in Plutellidae). 10+ valid species. Ochsenheimeria and the
Larva with only one SV seta on IX, a trait recur- closely related Aridomeria are Palaearctic;
rent in several endophytic micro-moth larvae. O. vacculella, occasionally a serious cereal crop
(Note: two setae are present in some ypsolophids pest in the Old World, has been casually intro-
and heliodinids; it remains unclear whether this duced into N. America.
is a genuine plesiomorphy or a character re- Davis (1975, 1987), Karsholt & Nielsen (1984),
versal.) Zagulyaev (1988, 1990), Kuznetznov & Stekolni-
kov (1993).
Ypsolophidae. Probable autapomorphies (Kyrki
1990) include the stalked (or coincident) hind-
wing Rs and Ml (Fig. 8.1 M), the anteriorly Putative clade Plutellidae + Acrolepiidae +
deeply bilobed male tegumen, the membranous Glyphipterigidae + (Heliodinidae + Bedelliidae +
and strongly spinose tubular anellus, the 2 cor- Lyonetiidae)
nuti/cornutus groups in phallus, the markedly
The monophyly of this assemblage was suggested
elongate female apophyses, the exit site of the
by Kyrki (1990) on the following: loss of the
ductus seminalis on ductus bursae close to os-
tium, the signum configuration (elongate plate gnathos and reduction of the tegumen (which
with two transverse ridges), and loss of pupal dorsally at most forms a narrow band) in the
cremaster setae. male genitalia; protruded pupal thoracic spira-
cle; larval feeding on herbaceous plants. How-
Ocelli usually well developed. Pterostigma,
ever, there are serious difficulties with this
when present, not extending distad beyond vein
scheme: contrary to the statements of Kyrki &
Itmies (1986) a gnathos is actually present in
Larval proleg crochets, when present, uni-
members of the Plutellidae, Heliodinidae and Ly-
onetiidae (e. g. Common 1990, Kuroko 1964).
The apparently close relationship between the
And as noted by Kyrki himself, the lyonetiids do
two constituent subfamilies has only recently
not have protruded pupal spiracles, and their
been recognized. The Ypsolophinae have mostly
host spectrum includes many woody plants; ac-
been associated with the phenetically similar Plu-
ceptance of the suggested phylogeny therefore
tellidae; the family distinction is based on the as-
requires, that these traits are interpreted as char-
sumption (which needs corroboration, see be-
acter reversals.
low), that a taxon centred on Plutella is cladisti-
cally closer to the Glyphipterigidae and the Heli-
odinid/Bedelliid/Lyonetiid assemblage. The Och- Plutellidae. As delimited by Kyrki (1990) the
senheimeriinae were consistently associated with family included the Acrolepia group of genera
the tineoid families until Kyrki's 1984 study, and which has often been ranked as a separate fam-
their status deserves continued scrutiny; it can- ily. The alleged autapomorphy of the family thus
not be ruled out that they are actually cladisti- circumscribed is the structure of the female post-
cally subordinate in the ypsolophines. vaginal lamella, which consists of two setose
Ypsolophinae. Monophyly not well supported. lobes; however, a somewhat similar lamella (with
Kyrki (1984) suggested the phallic coecum to be the lobes more or less distinctly united by mela-
an autapomorphy. nized cuticle, see Moriuti 1977) is widespread in
Ypsolopha (Holarctic) is the principal genus Yponomeutidae and Ypsolophidae, and it may
with 120 + species, some of which have the fore- be plesiomorphic at this level. Kyrki further
wing termen sinuate or even markedly falcate mentioned the large-meshed cocoon structure,
(Fig. 8.2 G); larvae in open webs on leaves of but the latter also occurs in some yponomeutids
diverse (usually woody) plants. (Heppner 1987). Both traits are absent in some
Kuznetzov & Stekolnikov (1976-77), Moriuti S. Hemisphere taxa currently included in the Plu-
(1977), Zagulyaev (1990). tellidae. In the absence of good synapomorphies
Ochsenheimeriinae. A compact group of moths of the Plutella and Acrolepia groups they are here
with characteristic habitus (Fig. 8.2 H). Possible treated as separate families.
autapomorphies include the 'shaggy' vestiture of According to Kyrki (1984) the Plutella group
the head and labial palp, the small size of the in the restricted sense is characterized by male
compound eyes (probably reflecting the moths' genitalia with 'teguminal processes' curved, sur-
diurnal activity, as do the well developed ocelli), rounding anal tube; the processes in question are
absence of pterostigma, the strongly sclerotized actually gnathal.
anal cone of the male, the asymmetrical ostium Ocelli usually distinct, rudimentary in some
bursae of the female, and the reduced crochet S. hemisphere genera currently placed in the
The Yponomeutoidea 125

family. Pterostigma extending beyond vein R. including choreutoids and immoids. Heppner
Female retinaculum a subcostal scale-patch. (1977, 1985, 1987, 1991) assigned the Glyphip-
Larval prolegs usually slender (Fig. 8), with terigidae s. str. to the Copromorphoidea with
uniserial crochets. which they share prominent protuberances bear-
Plutella (40 + species, cosmopolitan) larvae ing the larval spiracles (particularly on thorax
mostly feed on Brassicaceae, living in loose webs and VIII) and the dorsocaudal displacement of
and skeltonizing leaves. The genus includes the the larval spiracle VIII. However, the different
diamondback moth P. xylostella (= maculi- abdominal sternum II configurations (Kyrki
pennis) (Fig. 8.2 K), an ubiquitous pest species 1983, Minet 1983, contra Heppner 1977) and the
which is one of the few micro-moths renowned non-apical vom Rath's organ contradict a close
for long-range migrations; its biology and con- relationship between Glyphipterigidae s. str. and
trol are reviewed by Talekar & Shelton (1993). copromorphoids. The placement of the former
Other principal genera include Leuroperna, Rhi- within the Yponomeutoidea received additional
gognostis (Kyrki 1988), Eidophasia (Fig. 8.2 K). support with Kyrki & Itmies' arguments (1986)
The monobasic Embryonopsis from Indian for their close affinities to Orthotelia, whose male
Ocean subantarctic islands is brachypterous in has retained typical yponomeutoid pleural lobes.
both sexes. Proditrix (Fig. 8.21; New Zealand, Probable autapomorphies of the family as now
also known from Tasmania) species are note- delimited (Kyrki 1990) include the absence of an
worthy for their large size (some exceeding antennal pecten/scale flap, absence of segment X
50 mm wingspan) and their larval adaptations ('teguminal') processes; larvae endophagous
for feeding in intermittently waterfilled sites in (hence with shortened prolegs) and primarily
crowns of large monocots belonging to several monocot feeders, their cranial setae F l , Afl, Af2,
families (Dugdale 1987), or, in the case of the A2, P2, Ol, G2 (= 03 auct.) and S02 short, and
Tasmanian taxon, in a dicot (Richea, Epacri- the abovementioned larval spiracular traits.
dacreae) with a similar growth form. Their pro- Kyrki & Itmies (1986) mentioned two addi-
legs are blunt, with partially biserial crochets and
tional possible family autapomorphies in larval
the cocoon is non-meshed, so the systematic
structure, viz. the absence of an adfrontal 'punc-
placement is tentative. In this and other New
ture' and the presence of sclerotized shields on
Zealand 'Plutellidae' non-meshed cocoons have
segments VIII and IX. The larval prothoracic L
anterior and posterior openings, and the larval
group is almost always bisetose, but a New
skin is protruded from the latter; this is most
Zealand glyphipterigine has retained a vestigial
probably an autapomorphic condition.
third seta (Dugdale, in Heppner 1987). The pu-
Kuznetzov & Stekolnikov (1976-77), Moriuti pal thoracic spiracle is borne on a prominent
(1977), Zagulyaev (1990). process.
Orthoteliinae. Monobasic. Orthotelia sparga-
Acrolepiidae. As noted above this taxon has re-
nola (Fig. 8.2 N) is a rather large and broad-
cently been included in the Plutellidae; like the
winged, crepuscular micro-moth without ocelli.
bulk of the latter they have retained ocelli. Prob-
Vertex with tufts of raised narrow scales. Labial
able autapomorphies (Kyrki 1984) include details
palpi prominent, upcurved. Larva stem- and
in the male genitalia (reduction of the tegumen,
leafboring in freshwater plants belonging to sev-
teguminal processes and gnathos, basal widening
eral monocot families; prolegs with crochets in
of the phallus) and stalking of hindwing
two transverse rows (on anal pair a single). Pupa
Ml + M2 and M3 + CuAl (Fig. 8.1 N). Females
lack the ventral costa at the base of the anterior (Fig. 8.1Q) with prominent, bifid frontal process
apophysis, but this apomorphy is recurrent in and segments V - V I I I with spiny transverse
some S. Hemisphere taxa currently placed in Plu- ridges; pupal skin protruded from foodplant as
tellidae. adult emerges, an unusual condition within the
Adults are crepuscular/nocturnal. Larvae are Yponomeutoidea. W. Palaearctic.
leaf miners or borers in stems, flower buds, and Kyrki & Itmies (1986).
seeds. While the main pest species (including the Glyphipteriginae. A probable autapomorphy is
leek moth Acrolepiopsis assectella, Fig. 8.2 L, the more or less conical male segment VIII (Fig.
studied in detail by Frediani, 1954) are Liliaceae 8.1 H) with an apparently unusually enlarged ter-
feeders, the host spectrum also includes a variety gum, now interpreted (Kyrki & Itmies 1986) as
of dicot families such as Asteraceae, Lamiaceae being fused with the pleural lobes, which are not
and Solanaceae. distinguishable as such (Kyrki 1984). Kyrki & It-
95 + species; worldwide, but more than half mies also considered the vestigial forewing M-
are Palaearctic. Acrolepia, Acrolepiopsis and Dig- stem and CuP, as well as the approximation of
itivalva. the hindwing M3 and CuA (Fig. 8.3 B), to be
Kuznetzov & Stekolnikov (1976-77), Gaedike probable subfamily autapomorphies; these spe-
(1970, 1983, 1986), Zagulyaev (1990). cializations are commonplace elsewhere among
yponomeutoids, but Orthotelia exhibits the plesi-
Glyphipterigidae. This family long comprised an omorphic states. The diurnal activity of adult
assemblage of unrelated diurnal micro-moths, 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- cot families including Urticaceae and Crassula-
folded wings in repose, may be further derived ceae. The mesh cocoon is spun in the cavity made
groundplan traits. by the larva, or away from the food plant.
Moths mostly small, wingspan range 4,5 383 species (Heppner 1991). Many of the 20 +
30 mm (Heppner 1985), usually ca. 10 mm. genera are monobasic, and about two-thirds of
Ocelli prominent (Sericostola excepted, Heppner the species are placed in the cosmopolitan genus
1990), chaetosemata absent; head smooth-scaled Glyphipterix.
(Fig. 8.3 A). Labial palpi usually upturned, Arita (1987), Arita & Heppner (1992), Diako-
sometimes with prominent ventral scale tuft on noff (1986), Kuznetzov & Stekolnikov (1976-
segment 2. Forewing outline variable, in many 77), Heppner (1982, 1985), Kuznetsov (1990).
Glyphipterix termen concave; pattern (Fig.
8.2 M) usually with light transverse lines or bars,
often with prominent metallic markings. Hind- Clade Heliodinidae + Bedelliidae + Lyonetiidae
wings narrower than forewings. Males often with The monophyly of this assemblage was suggested
coremata on VIII, sometimes on valvae (not in by Kyrki (1990) on the basis of the following
both locations at the same time, Heppner 1985); probable synapomorphies of its constituent fami-
anal cone long, prominent. lies: maxillary palp reduced, at most with 2 seg-
Larval proleg crochets sparse, in lateral penel- ments (parallelled in subordinate taxa elsewhere
lipse or transverse band, or completely absent. in the superfamily). At most 8 branches present
Larvae mostly borers/miners in leaves or flow- in forewing Rs/M/CuA group (the identity of in-
ering stems of Poaceae, Cyperaceae, Juncaceae, dividual branches cannot be identified, but it is
Zingiberaceae and Araceae, but also of some di- assumed that one and the same branch has been
The Yponomeutoidea 127

lost in all families; Kyrki's inference that there Adults are mostly diurnal. Larvae are leaf
are "at most 4 M and CuA veins" may well be miners, borers in stems, buds/fruits (even in gall
correct, but it cannot be excluded that it is an midge galls) or feeding, sometimes in a commu-
Rs branch, which has been lost). Pupa without nal web, externally on leaves. Hosts include a
distinct maxillary palp, with strong lateral ridges, variety of dicot families (Chenopodiaceae, Ny-
and with immovable abdominal segments. The taginaceae, Araliaceae, Onagraceae and Ai-
consistent absence of a ventral scale tuft on labial zoaceae). Pupation in larval shelter or externally,
palp segment 2 may also be seen as a groundplan in some taxa in a loose silk webbing.
autapomorphy (parallelled in Plutellidae-Acro- 50 + species, the great majority of which are
lepiinae and many Glyphipterigidae-Glyphipteri- in the New World; stated by Heppner (1991) to
ginae). Kyrki also considered an open discal cell be absent from the Afrotropical region, but rep-
in the hindwing to be synapomorphic of these resented here by Hierophanes. About a dozen
families, but some Lyonetiidae-Cemiostominae genera, Heliodines being the largest and most
do have the cell closed (Kuroko 1964). All mem- widespread.
bers of this assemblage have very narrow wings, Meyrick (1914), Heppner (1987), Fal'kovich
and the simplifications of the venation (which (1990), Common (1990).
may be very pronounced in subordinate taxa) are
evidently related to this; similar modifications Bedelliidae. A monogeneric taxon previously in-
are recurrent in narrow-winged members of cluded in the Lyonetiidae, but as noted above the
other micro-moth superfamilies. Kyrki noted evidence concerning its affinities is ambiguous.
that particularly the assignment of the Lyonetii- Probable autapomorphies (Kyrki 1984): Fore-
dae to this assemblage is problematical, since wing Rs2, Rs3, Rs4 and ?M1 + 2 on common
they have (retained?) a non-protruded pupal tho- stem from pointed apex of discal cell. Male seg-
racic spiracle and at least the lack of ocelli could ment VIII without pleural lobes. Larva with only
be taken as indication that they are most closely 2 lateral setae on abdominal segments IVII.
related to, or even subordinate in, the Ypono- Pupa with conical process on frons. The non-spi-
meutidae. nose adult abdomen (Kuroko 1962) and the
Relationships within this assemblage remain shortening of larval prolegs III and VI relative
unresolved. As noted by Kyrki a sistergroup rela- to those of I V - V (Common 1990) may be con-
tionship between bedelliids and heliodinids is sidered additional family autapomorphies, the
suggested by characters in the larval spinneret former parallelled elsewhere among yponomeu-
and thoracic chaetotaxy as well as pupation toids.
mode; on the other hand the head scaling, ab- Small greyish moths. Head with tuft of pili-
sence of ocelli and the hind tibial vestiture sug- form scales on vertex, face smooth-scaled; ocelli
gest a sistergroup relationship between bedelliids and chaetosemata absent; scales on scape form-
and lyonetiids. ing dense pecten, but no true eye cap. Hind tibia
Kyrki also mentioned a very prominent hairy on dorsal side.
"fringe" on the underside of the larval labrum as Larva with long spinneret and undeveloped
a possible bedelliid/lyonetiid synapomorphy, but thoracic VI setae (similarities with Heliodinidae),
the epipharyngeal armature is actually somewhat and prominent epipharyngeal fringe (similarity
differently developed in these taxa (Jayewick- with Lyonetiidae) (Kyrki 1990).
reme 1940). Larvae leafminers, feeding on several plant
families, e. g. Poaceae, Liliaceae, Urticaceae and
Heliodinidae. Probable autapomorphies (Kyrki Convolvulaceae. Initial mine a narrow gallery.
1984): entire head scaling smooth, and forewing Later instars blotch miners and eroders, which
CuP vestigial (Fig. 8.3 D) (states also encoun- may make several mines and even change leaf;
tered in Glyphipterigidae-Glyphipteriginae); frass is ejected from the mines. An open silk net-
abdomen non-spinose; pupa with stiff lateral and work is spun outside the mined leaf. Pupation in
dorsal bristles. Many taxa previously included in a loose web below or above the mine, without
the family (Meyrick 1914) have no close relation- formation of a genuine cocoon.
ships to Heliodines and are now placed in various Bedellia (18 described species; Fig. 8.4 B) is
families in the Gelechioidea, or even (Schrecken- distributed in all regions except the Neotropical;
steinia) in the Apoditrysia. there is a small radiation of the genus in Hawaii,
Small moths with ocelli but no chaetosemata; where at least 14 species occur. The very wide-
male antennae typically thickened, scape without spread B. somnulentella is occasionally a pest on
pecten/scale flap. Hind tibia smooth scaled on Ipomoea.
dorsal side. Forewings with metallic markings Zimmerman (1978), Davis (1987), Kuznetzov,
(Fig. 8.4 A). Female frenulum a single bristle, ret- Kozlov & Seksjaeva (1988).
inaculum subcostal, of short recurved bristles.
Larval spinneret unusually long and VI unde- Lyonetiidae. Probable autapomorphies (Kyrki
veloped on thorax (similarities with Bedeliidae, 1990): scape slightly flattened, scales forming
Kyrki 1990); prolegs in uniordinal circle. 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 somnulentella\
C, Lyonetia clerckella; D, Leucoptera laburnella. Uniform magnification; scale (milimetres) in A.

produced. Larva with head (Fig. 8.3 F) flattened, Cemiostominae. Probable autapomorphies
cranial setae F l , Afl, A , LI and G2 (= 03) (Kyrki 1984): forewing with tornai patch of
short, labrum at most with four pairs of dorsal raised metallic scales. Larval head with adfrontal
setae, and abdominal setae D1 and D2 approxi- ridges subparallel, reaching epicranial notch
mated, D1 more laterad than D2. Two first lar- (Fig. 8.3 F). The adult's spinose abdomen is not
val instars without prolegs, and frequently also here considered apomorphic.
without thoracic legs (Davis 1987, Common Vertex with tuft of piliform scales, or smooth-
1990). Pupal thoracic spiracle non-protruded scaled like the face. Antenna not longer than
(plesiomorphy or autapomorphic character re- forewing. Forewing mostly with uniform white/
versal?). greyish ground colour and a characteristic
Scaling on vertex variable, on face consistently pattern (Fig. 8.4 D) of brownish and black
smooth; ocelli and chaetosemata absent. Hind bands/bars in the apical half (strikingly similar
tibia hairy on dorsal side. to that of GracillariidaePhyllocnistinae). In
Larval epipharynx with prominent fringe resting moths the apical black bars in the fringes
(similarity with Bedelliidae, Kyrki 1990). are believed to mimic the antennae of an insect
Larvae leaf- (occasionally twig-) miners. Pupa- head (Emmet 1985; this phenomenon is recurrent
tion outside the mine.
in several other groups of micro-moths).
Kuroko (1964), Bradley & Carter (1982), Em-
met (1985), Davis (1987), Kuznetsov, Kozlov & Larval leaf mines are blotches or gallery-
Seksjaeva (1988), Seksyaeva (1990), Mey (1994). blotches, twig mines narrow galleries through-
Lyonetiinae. Probable autapomorphies (Kyrki out. Some blotch miners have a characteristic
1984): extreme narrowing of the wings and abdo- habit of smearing the viscous faeces in concentric
minal terga. Female 'ovipositor' (Fig. 8.3 G) rings on the upper epidermal wall of the mine
piercing, strongly sclerotized, pointed and (Fig. 8.31). Pupation in a usually fusiform co-
equipped with small lateral/ventral teeth. The coon, spun on the hostplant or among leaf litter.
non-spinose abdomen may also be considered Principal foodplants belong to Fabaceae, fol-
autapomorphic (but very homplasious, cp. Be- lowed by Rubiaceae, but about a dozen dicot
delliidae above). host families have been recorded, and even the
Vertex always with tuft of piliform scales (Fig. monocot family Liliales.
8.3 J) Antennae long, exceeding forewings. Fore- 120+ species, in all regions. Main genera in-
wing often with dark markings in basal half. clude Paraleucoptera, Perileucoptera (with coffee
Larval mines may be narrow galleries pests: P. coffeella in the Neotropics and Masca-
throughout, or gallery-blotch mines. Pupa usu- rene Isles, P. meyricki in Africa) and Leucoptera
ally in a flimsy cocoon suspended by silk threads, (L. malifoliella is a Eurasian orchard pest,
but occasionally naked (Davis 1987). Hosts are L. psophocarpella a pest of Psophocarpus in
woody dicots belonging to a dozen (partly unre- New Guinea).
lated) families.
Cosmopolitan; Heppner (1991) counted 89 de-
scribed species (only 1 Afrotropical). Lyonetia is Acknowledgments
the principal genus; L. clerckella (Fig. 8.4 C) is
occasionally a pest on rosaceous fruit-trees in the Comments from Mr. Frank Hsu (Heliodinidae)
Palaearctic. Stegommata in the Australian region and Mr. Ole Karsholt on our manuscript drafts
are Proteaceae miners. are gratefully acknowledged.
The Yponomeutoidea 129

References dae), Gespinstmotten (Yponomeutidae), Echte

Motten (Tineidae). 273 pp. in Die Tierwelt Deutsch-
Arita, Y. (1987): Taxonomic studies of the Glyphipteri- lands, 63. Teil.
gidae and Choreutidae (Lepidoptera) of Japan. - Heppner, J. B. (1977): The status of the Glyphipterigi-
Trans. Shikoku ent. Soc. 18: 1 - 2 4 4 . dae and a reassessement of relationships in Ypono-
- & Heppner, J. B. (1992): Sedge moths of Taiwan meutoid families and Ditrysian superfamilies. J.
(Lepidoptera: Glyphipterigidae). - Tropical Lepi- Lepid. Soc. 31: 124-134.
doptera 3 (suppl. 2): 1 - 4 0 . - (1981): A world catalog of genera associated with
Bradley, J. D. & Carter, D. J. (1982): A new lyonetiid the Glyphipterigidae auctorum (Lepidoptera). -
moth, a pest of winged-bean. - Syst. Ent. 7: 1 - 9 . J. N. Y. Ent. Soc. 89: 220-294.
Common, I. F. B. (1990): Moths of Australia, vi + - (1982): Synopsis of the Glyphipterigidae (Lepidop-
535 pp., 32 pis. Leiden, E. J. Brill. tera: Copromorphoidea) of the world. - Proc. ent.
Davis, D. R. (1975): A review of Ochsenheimeriidae soc. Wash. 84: 3 8 - 6 6 .
and the introduction of the Cereal Stem Moth Och- - (1985): The sedge moths of North America (Lepi-
senheimeria vacculella into the United States (Lepi- doptera: Glyphipterigidae). - Flora & Fauna Hand-
doptera: Tineoidea). - Smithson. Contr. Zool. 192: book No. 1, 254 pp.
1-20. - (1987): Glyphipterigidae, Plutellidae, Yponomeuti-
- (1987): Lyonetiidae. Pp. 370-372 in F. Stehr (Ed.) dae, Argyresthiidae, Acrolepiidae, Heliodinidae
Immature insects. Kendall/Hunt, Dubuque. (Yponomeutoidea). Pp. 403-411 in Stehr (Ed.) Im-
Diakonoff, A. (1986): Glyphipterygidae auctorum mature insects of North America, Kendall/Hunt,
sensu lato: (Glyphipterygidae sensu Meyrick, 1913); Dubuque.
Tortricidae: Hilarographini, Choreutidae, Brachod- - (1991): Faunal regions and the diversity of Lepi-
idae (partim), Immidae and Glyphiperigidae in doptera. - Tropical Lepidoptera 2 (Suppl. 1): 1 - 8 5 .
Microlepidoptera Palaearctica 7: 436 pp. + Plates Huemer, P. & Tarmann, G. (1991): Westpalarktische
Volume (175 pis.). Gespinstmotten der Gattung Kessleria Nowicki:
Dugdale, J. S. (1987): Proditrix, a new genus for Tita- Taxonomie, kologie, Verbreitung (Lepidoptera,
nomis tetragona Hudson and Plutella megalynta Yponomeutidae). Mitt. Mnch. Ent. Ges. 81:
Meyrick and allies (Lepidoptera: Yponomeutidae), 5-110.
shoot destroyers of Monocotyledones. - N. Z. Karsholt, O. & Nielsen, E. S. (1984): A taxonomic re-
Journal of Zoology 14: 99 - 1 1 2 . view of the stem moths, Ochsenheimeria Hbner, of
Emmet, A. M. (1985): Lyonetiidae. Pp. 2 1 2 - 2 3 9 in J. northern Europe (Lepidoptera: Ochsenheimerii-
Heath & A. M. Emmet (eds.) The moths and butter- dae). - Ent. scand. 15: 233-247.
flies of Great Britain and Ireland 2. Kodama, T. (1961): The larvae of Glyphipterigidae
Fal'kovich, M. I. (1990): 38. Family Heliodinidae. (Lepidoptera) in Japan (1). - Pubis ent. Lab. Univ.
Pp. 6 9 9 - 7 0 0 in G. S. Medvedev (ed.) - Keys to the Osaka Prefect. 6: 3 5 - 4 5 .
Insects of the European Part of the USSR, II. Kuroko, H. (1964): Revisionai studies on the female
Leiden, E. J. Brill. Lyonetiidae of Japan (Lepidoptera). - Esakia 4:
Frediani, D. (1954): Ricerche morfo-biologiche sul- 1 - 1 6 1 , 17 pis.
YAcrolepia assectella Zell. (Lep. Plutellidae) nel- Kuznetsov, V. I., Kozlov, M. & Seksjaeva, S. V. (1988):
l'Italia centrale. - Redia 39(2): 187-249. To the systematics and phylogeny of mining moths
Friese, G. (1960): Revision der palarktischen Ypono- Gracillariidae, Bucculatricidae and Lyonetiidae
meutidae unter besonderer Bercksichtigung der (Lepidoptera) with consideration of functional and
Genitalien (Lepidoptera). - Beitr. Ent. 10: 1 - 1 3 1 . compa[ra]tive morphology of male genitalia. - Tr.
- (1969): Beitrge zur Insekten-Fauna der D D R : Zool. Inst. Akad. Nauk SSSR 1767: 1 4 - 5 1 [In Rus-
Lepidoptera-Argyresthiidae. - Beitr. Ent. (Berlin) sian].
19: 693-752. Kuznetsov, V. I. & Stekolnikov, A. A. (1976-77): Phy-
Gaedike, R. (1967): Zur systematischen Stellung ei- logenese relationships between the superfamilies
niger Gattungen der Heliodinidae/Schreckensteini- Psychoidea, Tineoidea and Yponomeutoidea in the
idae sowie Revision der palarktischen Arten der light of the functional morphology of the male geni-
Gattung Pancalia Curtis, 1830 (Lepidoptera). - talia 1 - 2 . - Ent. Rev., Wash. 55: 19-29; 56: 1 4 -
Beitr. Ent. 17: 363-374. 22.
- (1970): Revision der palarktischen Acrolepiidae - (1993): Comparative morphology of the male geni-
(Lepidoptera). - Ent. Abh. 38: 4 - 5 4 . talia and systematic status of some little-known lep-
- (1978): Versuch der phylogenetischen Gliederung idopterous families (Lepidoptera: Pterolonchidae,
der Epermeniidae der Welt (Lepidoptera). Beitr. Ochsenheimeriidae, Carposinidae, Callidulidae,
Ent. 28: 201 - 2 0 9 . Epicopeiidae) from the fauna of Russia. - Trudy
- (1984): Revision der nearktischen und neotro- Zool. Inst. Acad. Sci. 255: 321 [In Russian].
pischen Acrolepiidae (Lepidoptera). - Ent. Abh. Kyrki, J. (1983): Adult abdominal sternum II in ditry-
Mus. Tierk. Dresden 47, 10: 179-194. sian tineoid superfamilies - morphology and phy-
- (1986): Die Typen der orientalischen, australischen logenetic significance (Lepidoptera). - Annls ent.
und thiopischen Acrolepiidae (Lepidoptera). fenn. 49: 8 9 - 9 4 .
Beitr. Ent., Berlin 36: 6 3 - 6 8 . - (1984): The Yponomeutoidea: a reassessment of the
Gershenzon, Z. S. (1990): 28. Family Yponomeutidae. superfamily and its suprageneric groups (Lepidop-
29. Family Argyresthiidae. Pp. 4 3 6 - 4 7 2 in G. S. tera). - Ent. scand. 15: 7 1 - 8 4 .
Medvedev (ed.), Keys to the Insects of the European - (1989): Reassessment of the genus Rhigognostis
Part of the USSR, IV, II. Leiden, E. J. Brill. Zeller, with descriptions of two new and notes on
Hannemann, H.-J. (1977): Kleinschmetterlinge oder further seven Palaearctic species (Lepidoptera: Plu-
Microlepidoptera III. Federmotten (Pterophori- tellidae). - Ent. scand. 19(4): 437-453.
130 John S. Dugdale, Niels P. Kristensen, Gaden S. Robinson & Malcolm J. Scoble

- (1990): Tentative reclassification of holarctic Ypo- Talekar, N. S. & Shelton, A. M. (1993): Biology, ecol-
nomeutoidea (Lepidoptera). - Nota, lepid. 13(1): ogy, and management of the diamondback moth.
28-42. - A. Rev. Ent. 38: 275-301.
Kyrki, J. & Itmies, J. (1986): Immature stages and the Zagulyaev, A. K. (1988): Cereal stem moth: families
systematic position of Orthotelia sparganella Ochsenheimeriidae and Eriocottidae. Fauna
(Thunberg) (Lepidoptera: Yponomeutoidae). - SSSR, 135: 302 pp. [In Russian],
- (1990): 14. Family Ochsenheimeriidae. 30. Family
Syst. Ent. 11: 93-105.
Plutellidae. 31. Family Acrolepiidae. Pp. 130-136,
Mey, W. (1994): Taxonomische Bearbeitung der west-
4 7 3 - 5 5 1 in G. S. Medvedev (ed.) Keys to the Insects
palarktischen Arten der Gattung Leucoptera of the European Part of the USSR, IV, II. Leiden,
Hbner, [1825], s. 1. (Lepidoptera, Lyonetiidae). - E. J. Brill.
Dtsch. ent. ., . E 41 (1): 173-234. Zimmerman, E. C. (1978): Insects of Hawaii. 9. Micro-
Minet, J. (1983): Etude morphologique et phylogn- lepidoptera. ( 1 - 2 ) . 1903 pp., Honolulu.
tique des organes tympaniques des Pyraloidea. I -
gnralits et homologies. (Lep. Glossata). Annls
Soc. ent. Fr. (N. S.) 19: 175-207. Addendum
- (1986): Ebauche d'une classification moderne de Agassiz (1996) monographed the British Ypono-
l'ordre des Lpidoptres. - Alexanor 14: 291-313. meutoidea.
Menken, S. B. J., Herrebout, W. M. & Wiebes, J. T. The exclusively Southern Hemisphere Or-
(1992): Small ermine moths (Yponomeuta): Their thenches g r o u p of plutellids w a s discussed b y
host relations and evolution. - A. Rev. Ent. 37: Dugdale (1996). It is distinctive by the slender
41-66. pedunculi (tegumen arms). Most members
Meyrick, E. (1914): Lepidoptera Heterocera. Fam Hel- browse or mine angiosperms, with one genus
iodinidae. Gen. Ins. 165, 29 pp. (Chrysorthenches) specific t o conifers.
Moriuti, S. (1977): Fauna Japnica. Yponomeutidae A detailed a c c o u n t of the lyonetiid Lyonetia
s. lat. (Insecta: Lepidoptera). 327 pp., 95 pis., prunifoliella w a s given b y S c h m i t t et al. (1996).
Robinson, G. S Tuck, K. R. & Shaffer, M. (1994): A Agassiz, D. J. L. (1996): Yponomeutidae. In Emmet,
Field Guide to the Smaller Moths of South-east Asia. A. M. (ed.) The Moths and Butterflies of Great Brit-
308 pp. Natural History Museum, London & Ma- ain and Ireland 3, 39 - 1 1 4 .
Dugdale, J. S. (1996): Chrysorthenches new genus, co-
laysian Nature Society, Kuala Lumpur.
nifer associated plutellid moths (Yponomeutoidea,
Scoble, M. J. (1992): The Lepidoptera. Form, function Lepidoptera) in New Zealand and Australia. -
and diversity. 404 pp. Oxford Univ. Press. N. Z. J. Zool. 23: 3 3 - 5 9 .
Seksyaeva, S. V. (1990): Family Lyonetiidae (Leucop- Schmitt, J. J., Brown, M. W. & Davis, D. R. (1996):
teridae). Pp. 5 5 2 - 5 6 6 in G. S. Medvedev (ed.) Keys Taxonomy, morphology and biology of Lyonetia
to the European Part of the USSR, IV, II. Leiden, prunifoliella (Lepidoptera: Lyonetiidae), a leafminer
E. J. Brill. of apple. - Ann. ent. Soc. Am. 89: 334-345.
9. The Gelechioidea
Ronald W. Hodges

Gelechioidea are a large group with 1,425 genera lected larvae or pupae. Few species have been
and 16,250 described species. They are generally reared through a complete cycle, and very few
distributed throughout the world, becoming less voucher specimens document this information.
numerous at higher latitudes. Representation of One apomorphy defines Gelechioidea: the
families and subfamilies within geographic re- haustellum has overlapping scales on the dorsal
gions varies greatly: Cosmopteriginae are ex- surface from the base extending variously to 1/2
tremely speciose in one of four genera in Hawaii its length (Common 1970, 1990; Hodges 1986;
(Zimmerman 1978); Oecophorinae are the pre- Minet 1990); however, this state also occurs in
dominant Lepidoptera in Australia (Common Pyraloidea and in Choreutoidea among ditry-
1990), the second most species numerous taxon sians. Pyraloidea have paired tympanic organs at
in New Zealand (Dugdale 1988), but absent from the base of the abdomen that never are present
Hawaii before the advent of man (Zimmerman in Gelechioidea; and Choreutoidea have naked,
1978); Gelechiidae far outnumber Oecophori- minute 1/2-segmented maxillary palpi, a state
dae-Elachistidae in America north of Mexico that does not occur in Gelechioidea. The pupa
(Hodges 1983); and Stenomatinae are extremely remains in its chamber or in place on adult eclo-
abundant, 1,095 species, in the Neotropical re- sion, a behavioral character shared with Ypono-
gion (Becker 1984), but have only nine species in meutoidea (Common 1990).
the Antilles (Duckworth 1969). Gelechioidea
Characters common to most Gelechioidea are
have been very poorly collected except for very
1) maxillary palpus 4-segmented, scaled and
few areas. They may be the least well described
folded over/parallel with base of the haustellum
large superfamily of Lepidoptera. In the Nearctic
(probable apomorphy); 2) labial palpus upturned
region approximately 30% of Gelechiidae are de-
and third segment long and acute; 3) head
scribed, 10% of Scythridinae (Landry 1991),
smooth scaled; 4) chaetosemata absent; 5) dorsal
20% of Coleophorinae, 25% of Momphinae,
surface of hind tarsus with long, slender scales;
6 0 - 7 0 % of Cosmopterigidae, 30% of Elachisti-
6) pupal antennae meeting mesially before their
dae (unpublished); for the Neotropical region
apexes (probable apomorphy); and 7) larval ab-
Stenomatinae may be 50% described, Gelechi-
dominal segments 1 - 8 with setae L1/L2 closely
idae < 20% described (V. O. Becker, unpub-
approximated or on the same pinaculum (Com-
lished); for all of Subsaharan Africa only Gelech-
mon 1970, 1990; Hodges 1978, 1986; Minet 1990,
iidae and Lecithoceridae of South Africa have
1991; Nielsen 1989).
been studied in any detail (Janse 1949-63), sug-
gesting that the 1,027 species of Gelechioidea Gelechioidea have the following characters:
(Vri & Kroon 1986) are but a small part of the Head smooth scaled; frons evenly convex, occa-
total. The oecophorine fauna of Australia is sionally with processes; haustellum usually de-
about 50% described (Common, unpublished). veloped, rarely absent, scaled on anterior sur-
face, at least at base; labial palpus three seg-
Adults have forewing lengths of 2.0 mm mented, usually upturned, rarely porrect with
(Siskiwitia falcata) to 35 mm (Cryptophasa setio- third segment slightly upturned, third segment
tricha) with those of most species 5 15 mm. normally acute; maxillary palpus four segmented
Most are active at dusk or night; many genera (rarely one-three segmented), scaled, folded over
are diurnal, and they often are very brightly col- base of haustellum or downturned and parallel
ored. Alpine and arctic species may be diurnal with base of haustellum; ocellus present or ab-
but are very somberly colored. Larvae feed on sent; antenna simple or ciliate, rarely pectinate
dead/decaying plant or animal tissue, virtually or bipectinate, pecten present or absent; chaeto-
any living plant tissue, and sometimes on living semata absent. Forewing broad to narrowly lan-
invertebrates. Most species are concealed exter- ceolate, pterostigma present or absent, cell open
nal feeders; they usually construct a silk shelter or closed, 10 to 12 veins present, CuP present or
or tie leaves. Some are root or stem gall makers, absent, 1 A and 2 A forked basally; retinaculum
leaf miners, or case bearers (Bradley 1987; Pow- subcostal in male, subcostal to subcubital in fe-
ell 1980). Pupation usually occurs in the larval male, sometimes with secondary wing coupling
chamber, but the pupa may be fully exposed. It structures. Hindwing broad to linear, fringe
remains in situ on eclosion of the adult. Eggs short to over five times width of wing; cell closed
normally are laid singly (Common 1990). or open; six to nine veins present; frenulum a
Except for the Palearctic region very little is single acanthus in male, usually three acanthi in
published on the immature stages. Most reports female; cubital pecten sometimes present; base of
in the literature are from rearings of field-col- wing sometimes with hair pencils. Wings rarely
132 Ronald W. Hodges

reduced or non-functional in , then lanceolate/ ing at their distal ends to expose the metathora-
linear or short lobes. Legs: epiphysis present; tib- cic legs." She included Coleophoridae in Ypono-
ial spurs 0, 2, 4; hindtibia usually with numerous meutoidea.
long, slender scales on dorsal surface. Abdomen Subsequently, Forbes (1923), McDunnough
frequently with spiniform dorsal setae; some- (1939), Common (1970, 1990), Bradley (1972),
times with secondary hair pencils from lateral or Hodges (1978), Kuznetsov and Stekol'nikov
ventral surface in male; segments seven and eight (1978, 1984), Minet (1986, 1990), and Nielsen
sometimes modified in male; S2 with tineoid ven- and Common (1991) have proposed classifica-
ulae and apodemes, sometimes venulae or apo- tions; however, either no characters were given
demes, infrequently neither. Male genitalia sim- or each taxon was descriptively characterized
ple and symmetrical to highly specialized by re- without an indication of relationship across the
duction, modification, and asymmetry; aedeagus superfamily.
sometimes ankylosed, rarely unsclerotized; val- A phylogenetic analysis using Hennig 86
vae sometimes reduced and replaced functionally (Farris 1988) based on 38 characters provides the
by modifications of preceding segments. Female basis for my discussion concerning relationships
ovipositor usually adapted for external place- among gelechioid higher taxa. No previous clas-
ment of eggs but sometimes for piercing and in- sification of Gelechioidea has been based on an
ternal placement of eggs; ostium bursae ventro- analysis of characters and character states using
mesial, rarely lateral or dorsal. Egg normally parsimony. Other cladograms resulting from dif-
flat, rarely upright in Coleophoridae and ferent interpretations of characters (different
Scythridinae (Common 1990); chorion ridges character state matrixes) and methods of analy-
frequently present; texture rough (Peterson 1967) sis (Mickevich & Mitter 1981, Mickevich and
or smooth. Larva sometimes with submental pit, Weiler 1990), are possible; however, most clades
almost always with six stemmata, rarely three or are stable among the many trees examined dur-
none; prothorax with three L setae, rarely two; ing the analysis.
setae LI and L2 usually approximate on abdo- Within Gelechioidea several characters are ho-
men, L2 anterior to dorsal of LI; A8 usually moplasious; a relatively large number of charac-
with SD anterodorsad to dorsad of spiracle; A9 ters are polymorphic in the sense that a character
with D2 setae on separate pinacula, SD setae has more than one state within a taxon (Micke-
often hairlike; anal comb sometimes present in vich & Mitter 1981) (appendix 3); and homology
Gelechiidae; crochets uniserial, in a circle, el- has not been determined for several genital struc-
lipse, or two transverse parallel rows, rarely very tures. These facts likely are the basis for continu-
few, rarely a mesoseries. Pupa obtect, frontocly- ing problems in classification of parts of the su-
peal suture present or absent, almost always perfamily and make the resulting classification
without dorsal spines, maxillary palpus usually tentative. Discovery of additional characters and
visible, labial palpus and prothoracic femur reinterpretation of existing ones should result in
sometimes exposed, antennae usually touching a more nearly stable classification. Concurrently,
mesially before apexes, abdominal segments five- Gelechioidea exhibit an extremely wide array of
seven movable in male, five and six in female, morphological diversity. Likely, much additional
lateral condyles sometimes present on segments diversity will be discovered as new species and
5/6 and 6/7; segment nine sometimes with paired, the immature stages of nearly all taxa are col-
ventrolateral, anteriorly directed lobes; segment lected and studied.
10 with recurved setae or cremaster.
Analytical procedure. Characters and states detailed
Classification. Fracker (1915) proposed Gelechi- in appendix 1 and coded in appendix 2 were used in the
oidea, but he did not indicate characters to de- analysis. Terminology follows Braun (1924), Common
(1990), Heinrich (1920), Klots (1970), Kyrki (1983),
fine them. In his key to families the included taxa
Minet (1982), Mosher (1916), Stehr (1987). Plesiomor-
arise in different places; Elachistidae and Col-
phic states are indicated by " O . "
eophoridae were associated with the tineoid Characters were polarized on the basis of knowl-
series but without superfamily assignment. He edge of the Gelechioidea and other Lepidoptera; a hy-
stated (1915: 45) "The grouping and the division pothetical outgroup consisting of the plesiomorphous
into families is principally a compilation from character states was used. Three characters were
Walsingham and from published and unpub- weighted: 1) valva divided or not (character 1), 2) pupa
lished work of August Busck. At the same time with/without lateral condyles (character 31), and 3)
we believe that nothing in the larval structure pupa with/without paired ventromesial lobes on A9
precludes this arrangement and that it is a more (character 32). An "mh*" run yielded one tree with a
natural classification than any published more length of 201, consistency index of 37, and retention
index of 65. The "bb*" command yielded 2,456 trees
than a decade ago."
with the same length and consistency and retention in-
Mosher (1916) recognized the pupae of Gel- dexes as for the "mh*" run.
echioidea as having "... a distinct epicranial su- Taxa shifted association among the trees because
ture, with the caudal portions of the antennae polymorphic characters were coded as "?," several
lying adjacent on the meson and usually separat- characters are homoplasious, and states for several
The Gelechioidea 133

characters of immature stages are unknown. The somesial surface usually finely scobinate . .
following discussion of alternative associations of (Oecophoridae) 2
some taxa caused by homoplasious or polymorphic
- Gnathos present and fused or articulated
characters led to selection of a single tree. 1. Depressa-
riinae/Elachistinae: a) sister-groups to each other, b)
with tegumen or absent; when present and
Depressariinae the sister-group to clade Elachistinae/ fused with tegumen then a sclerotized band
Agonoxeniae/Hypertrophinae/Deuterogoniiae/Aeolan- (fig. 9.4.B), mesial region downturned and
thinae, or c) forming a trichotomy among Depressarii- laterally compressed (fig. 9.4 D), or with a
nae, Elachstinae, and Agonoxeninae/Hypertrophinae/ mesial spinose bulb (fig. 9.4 I) 3
Deuterogoniinae/Aeolanthinae. " a " was selected be- 2 Abdominal terga with spiniform setae on
cause of the synapomorphies in the gnathos and uncus
posterior margin of each segment (fig.
shared by Depressariinae and Elachistinae. 2. Xyloryc-
tinae/Scythridinae: Because they share three apomor- 9.5 H) Stathmopodinae
phies (larva with secondary SV setae on Al8, larva - Abdominal terga usually without spiniform
with a pore posterad of SVI on A l - 8 , and larva with setae, spiniform setae in broad band when
a pinaculum ring around SVI on A l - 8 ) , they may present (fig. 9.5 G) Oecophorinae
associate in four diverse ways, a) forming a trichotomy 3 Pupa with lateral condyles between abdomi-
with Stenomatinae/Aeolanthinae and Chimabachidae/
nal segments 5/6 and 6/7 (fig. 9.7 F)
/Pexicopiinae, b) associated with Ethmiinae in clade
Stenomatinae//Aeolanthinae, c) Xyloryctinae/Scy- (most Elachistidae) 4
thridinae sister group to clade Chimabachidae/- /Pexi- - Pupa without lateral condyles between ab-
copiinae. d) derived taxa within clade Chimabachidae/ dominal segments 12
Scythridinae/Xyloryctinae. " a " was discarded because 4 Valva almost always with some setae that
Xyloryctinae/Scythridinae share many character states are thickened and have bifid or multifid
with the clade ending with Pexicopiinae; " b " was dis-
apexes Stenomatinae
carded because Xyloryctinae/Scythridinae lack lateral
condyles in the pupa, an apomorphy of clade Steno- - Valva without setae having such modified
matinae/-/Aeolanthinae; and " d " was discarded be- apexes 5
cause the apomorphy defining the clade is weak, "c" 5 Gnathos fused broadly with tegumen, apex
was selected to emphasize the significance of the three with stout, toothlike projections
apomorphies. 3. Physoptilinae: a) sister-group to Gel- Ethmiinae, part
echiinae/-/Pexicopiinae, b) sister-group to any one of
- Gnathos articulated with tegumen, with
Gelechiinae, Dichomeridinae, or Pexicopiinae. " a " was
selected because the retinaculum of Physoptilinae fe- finely spinose mesial bulb (fig. 9.4 I), some-
males is state 7, whereas that of Gelechiinae/-/Pexi- times arms separate and each arm ending in
copiinae is state 4, which I construe as the most signifi- finely spinose bulb (fig. 9.4 K) or toothlike
cant apomorphy of the clade. 4. Gelechiinae/Dichom- projections (fig. 9.4 L), or gnathos absent 6
eridinae/Pexicopiinae: a) Gelechiinae sister-group to 6 Gnathos present 7
Dichomeridinae + Pexicopiinae, b) Dichomeridinae
- Gnathos absent 9
sister-group to Gelechiinae/Pexicopiinae, c) Pexicopii-
nae sister-group to Gelechiinae/Dichomeridinae. " a " 7 Larva with secondary setae in SV group on
was selected because Gelechiinae have character states A46 (fig. 9.7 I); pupa with paired leglike
that could give rise to Dichomeridinae - Pexicopiinae structures arising from ventral surface of
through loss or modification. Dichomeridinae have segment 9 or 9/10 (fig. 9.7 G) Agonoxeninae
two apomorphies, 1) 9 with secondary corpus bursae - Larva without secondary setae in SV group
and 2) pupal terga 1/2 and 2/3 with setose, raised, me-
on A 4 - 6 (SVI, SV2, SV3 only); pupa with-
sial/submesial prominences on opposing margins; and
Pexicopiinae usually have only one pair of apodemes
out paired leglike structures arising from
on sternum 2 (apomorphy within clade), often have an ventral surface of segment 9 or 9/10 . . . .8
antennal pecten, and never have an anal fork. 8 Hindwing narrowly lanceolate, cell open, Rs
and Mi long stalked (fig. 9.6 N)
The proposed classification (fig. 9.1) is a major - Hindwing subquadrate, cell usually closed,
rearrangment of family and subfamily-group Rs and Mi arising separately from cell (fig.
concepts. My use of family names reflects some 9.6 K) Depressariinae
conservatism based on the tentativeness of this 9 Hindwing with Rs and Mi stalked
analysis but recognizes what probably are reli- Aeolanthinae
able associations. For this reason all taxa in clade - Hindwing with Rs and Mi separate . . . 10
2 are included in Elachistidae and the taxa in 10 Abdominal sternum 2 with paired venulae
clade 24 in Coleophoridae. The list of valid and
and apodemes (fig. 9.5 E) Deuterogoniinae
synonymic family/subfamily-group names to-
- Abdominal sternum 2 with paired venulae
gether with authors and dates of publication will
only (fig. 9.5 D) 11
help associate information in previous literature
11 Valva with free sclerite at apex of sacculus,
with recognized taxa (appendix 4).
socii present Hypertrophinae
- Valva without free sclerite at apex of saccu-
Key to Families and Subfamilies lus, socii absent Ethmiinae, part
1 Gnathos present, fused with tegumen later- 12 Juxta present 13
ally, expanded mesially (fig. 9.4 C), dor- - Juxta absent 36
134 Ronald W. Hodges

& <
, & cT &


f f j/i
f /- ff j
/ / / //
-V #

*&4 & M &&

" /I

10-<3.10-0 :13-1 4-6.7-J-15. 10-1 6-1 1-1 ^36-1 3-1 4 - 1 6 - 1 ^13-5 3-1 0.1
. .11-1 11-3 4-2
92 :20-1 . . 5-0.1 - I S 0. 12-1 8-1-1 31-0.1 1,0-1. 17-1 : 30-1 i 10-2 23--0.1 23-3 : =14-1
:27-1 13 l ' I S 22-1
) -l : 10-:
-2 36-0 -12-0. 22-0 19-1 ! s 171
: :15 .22-0.1 24-1 : 16-1 : : 28-0.1 : :24-1 ! 24-1
. .21 : :27-1 20-1 -37-1 ; :35-1
: 23 37-1

4 9

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

/ /

y. /
/ P >
< i $f
/ <8* /
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
> 9-0. 5-0.1 ^ 7-1 8-0
10-2 10-0.1 4-6 6 1
15-0 20-0 11-1.2 5-1 5-1 12-1 8-0.1:: 10-2 137-0 10-2.- 7-0.1 ! 8-0.1 T10-(1).2
21-2 22-2 13-1.5 10-3. 8-1 ;13-1 12-0.1 12-0.1 91 I 9-1 123-3 " " 10-0 29-0.1
22-0 :17-1 ?2 12-2 13-2.4!113-0 20-1 25-(0).1 * tc 16-1
28-1 :24-1 13-0 13-6 :18-1 I122-0 '27-0.1 i 22-1 .1!
20-0 16-1 :22-1 ::30-1 29-0.1 s 29-1
21-1 28-0.1
34-1 : 34-1

_ 28 _

27-1 13-2


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 Ronald W. Hodges

Gnathos present, without spinose bulb(s), or without patches of raised scales

absent 33 Cosmopteriginae
32 Hindwing with Rs and stalked beyond Male genitalia symmetrical or asymmetrical,
end of cell Peleopodidae uncus present and symmetrical or absent,
Hindwing with Rs and Mi arising separately aedeagus often ankylosed with saccus; labial
from cell Amphisbatidae palpus never extremely slender, often stout;
33 Abdominal sternum 2 with paired venulae forewing often monochromatic gray, black,
and apodemes (fig. 9.5 E) 34 or brown, often with patches of raised scales
Abdominal sternum 2 with paired apodemes Chrysopeleiinae
only (fig. 9.5 F) 35
34 Forewing with CuP present
Gelechiidae, Physoptilinae Family-group statements
Forewing with CuP absent . Glyphidoceridae Apomorphic characters, possible apomorphic
35 Transtilla present Chimabachidae characters, and untreated characters (often of
Transtilla absent Xyloryctidae, Scythridinae larvae and pupae) are provided for each taxon.
36 Female retinaculum consisting of anteriorly Appendices 2 and 3 give the states of characters
directed scales on R (fig. 9.6 F), gnathos used in the parsimony analysis. When the family-
usually present (absent in Isophrictis group) group concept is new, no literature may exist for
(Gelechiidae, part) 37 the entity.
Female retinaculum consisting of anteriorly
directed scales on CuA (fig. 9.6 I), anteriorly Elachistidae. Defined by the apomorphy of pupa
directed scales between CuA and R in a row, with lateral condyles on abdominal segments 5/6
or posteriorly directed scales/setae arising on and 6/7, which character occurs as a polymorphy
Sc + anteriorly directed scales between Sc in Elachistinae; ventral surface of A9 often with
and R (fig. 9.6 C); gnathos absent two submesial, slightly swollen zones of recurved
(Cosmopterigidae) 39 setae. Highly diverse, worldwide taxon; 8 sub-
37 Abdominal sternum 2 with a pair of venu- families, 165+ genera, 3270+ species. (Minet
lae; forewing usually with CuA] und CuA2 1990).
stalked and directed posteriorly from end of Stenomatinae (fig. 9.2 A). All species, except
cell; female with secondary bursa arising some Agriophara, have some setae on valva with
from corpus bursae; pupa with raised, se- prominent, multilobed apexes (only some Anar-
tose, mesial or submesial prominences on sia (Gelechiidae) have such setae elsewhere in
opposing margins of terga 1/2, 2/3, 3/4 . . . Gelechioidea); uncus uni-, bi-, trilobed, or ab-
Dichomeridinae sent; abdominal support diffuse/diffuse with very
Abdominal sternum 2 with a pair of venulae short apodeme/sclerotized ridge at 45 degree an-
+ a pair of apodemes or a pair of apodemes gle with anterolateral margin of second sternum
(if a pair of venulae only, then forewing with (), well-developed apdeme, rarely sclerotized
CuAi and CuA2 separate); female lacking area with lateral ridge ($); hindwing with Rs bent
secondary bursa from corpus bursae; pupal toward Sc near end of cell; antenna long cili-
terga without prominences 38 ated. Larva: frons < , = , > 1/2 distance to epicra-
38 Abdominal sternum 2 with a pair of venulae nial notch; thoracic legs approximate or distant;
+ a pair of apodemes (apodemes very short Al with 2(3) SV setae; A 3 - 6 : SD setae without
in Anacampsis group); antenna without pec- pinaculum rings; crochets complete, biordinal el-
ten (some Aristotelia, Bryotropha, Metzneria, lipse, numerous. Pupa: beginning development
Monochroa with pecten usually a single, of "legs" Rectiostoma xanthobasis (Duckworth
slender scale, Apatetris group with full pec- 1971: 31), R. fernaldella (Keifer 1936: 356), Cer-
ten of broader scales) Gelechiinae conota monella, C. palliata (USNM), fully devel-
Abdominal sternum 2 with a pair of apo- oped in Timocratica melanocosta (Becker 1982:
demes; antenna often with a pecten of sev- 262); antennal scape usually with prominent
eral slender scales Pexicopiinae ridges. Larvae feed on turgid, living (16 families,
39 Female retinaculum with series of posteri- 35% Myrtaceae) and wilted plant tissue. Mainly
orly directed slender scales or setae arising New World, esp. Neotropics; Madagascar, India,
on base of Sc 40 Ceylon, New Guinea, Australia, New Zealand;
Female retinaculum without series of poste- 30 + genera, 1,200 + species (Anadasmus, An-
riorly directed slender scales or setae arising taeotricha, Chlamydastis, Phylomictis, Stenoma,
on base of Sc Antequerinae Timocratica).
40 Male genitalia asymmetrical, uncus two Becker (1982, 1984 a, 1984 b), Busck (1934),
separate lobes, right lobe almost always Common (1990), Duckworth (1964, 1967, 1970,
larger than left lobe, aedeagus ankylosed 1971), Hodges (1978), Minet (1990), Powell
with diaphragma; labial palpus usually very (1980), Stehr (1987).
slender and sickle shaped; forewing often Ethmiinae (fig. 9.2 B). Variable in most charac-
brightly colored with well-defined pattern, 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- Depressariinae (fig. 9.2 C). Maxillary palpus
tached to tegumen, when present. Larva: A 3 - 6 usually four segmented; hindwing with Rs and
or A l , 2, 79 with secondary SV setae in New Mi separate; fore- and hindwings broad. Larva:
World taxa, but assamensis and nigroapicella stemmata with gap between 5/6, T1 normally
groups in Palearctic with secondary setae on with 3 L setae (rarely 2), Al with 2/3 SV setae,
Al9. Pupa with "legs" in section I of Ethmia, A9 with SD usually hairlike. Larvae leaf tiers,
absent in section II, extremely poorly developed seed feeders, stem borers on dicots (17 families,
in sphaerosticha and in undescribed Pyramido- including Apiceae, Asteraceae, Betulaceae, Cory-
bela from Mexico (USNM). Larval hosts mainly laceae, Fabaceae, Fagaceae, Malvaceae, Rosa-
Boraginaceae and Hydrophyllaceae (70%), but ceae, Rutaceae, Salicaceae, Urticaceae). World-
also Papaveraceae, Ranunculaceae, Rosaceae, wide, but absent from many island groups; 80 +
Sabiaceae, Scrophulariaceae. Most numerous in genera, 600 + species (Agonopterix, Depressaria,
tropical-subtropical, seasonally arid areas, esp. Exaeretia, Semioscopis, Tonica).
northern Neotropics; 35 genera, 250 species Clarke (1941, 1963, 1978), Common (1964,
(Pseudethmia). 1990), Hannemann (1953), Hodges (1974),
Common (1990), Keifer (1936), Minet (1990), L'vovskii (1981), Minet (1990), Palm (1989), Pa-
Powell (1973, 1980, 1985), Sattler (1967), Stehr tocka (1989), Powell (1980), Stehr (1987), Toll
(1987). (1964).
The Gelechioidea 139

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

gnathos uncus


costal lobe of valva




gnathos uncus


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 graceae, Poaceae, Polygonaceae, Portulacaceae,
leafminers, on 20 plant families including Apia- Solanaceae. Worldwide; 26 genera, 700 + spe-
ceae, Asteraceae, Cactaceae, Caryophyllaceae, cies; many undescribed taxa (Areniscythris, Apos-
Chenopodiaceae, Convolvulaceae, Fabaceae, tibes, Aratura, Eretmocera, Mapsidius, Neo-
Hydrophyllaceae, Lamiaceae, Malvaceae, Ona- scythris, Rhamphura, Scythris).
The Gelechioidea 141

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

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 143

Garrha, Hofmannophila, Inga, Orophia, Oxystola, with Mi and M2 stalked. Worldwide; 6 genera,
Philobota, Pleurota, Tanyzancla, Wingia). 100 + species.
Clarke (1941, 1963), Common (1990), Hodges Epimarptinae (fig. 9.2 R). Defined by three
(1974), Leraut (1989, 1993), L'vovskii (1981), parallelisms: 1) abdominal support a pair of apo-
Meyrick (1922), Patocka (1989), Stehr (1987), demes, 2) hindwing with Mi and M2 fused, 3)
Toll (1964). $ antenna notched. Immature stages unknown.
Stathmopodinae (fig. 9.2 P). Defined by auta- Western India; 1 genus, 1 species (Epimarptis).
pomorphy, abdominal terga with spiniform setae Meyrick (1914).
on posterior margins. Wall of aedeagus with Batrachedrinae (fig. 9.2 S). Defined by abdo-
ventrodistal, sclerotized projection (possible apo- minal second sternum with venula or venula +
morphy); signa paired, sclerotized plates with apdeme (polymorphy); hindwing with Rs/Mi
mesial, inwardly directed flange, often secondary separate (reversal); and hindwing with M3 and
signa consisting of rows of inwardly directed Cu fused (polymorphy). Mi and M2 often
conical projections. Most species very slender fused and arising from Rs; tegumen often with
winged, with scales closely appressed to surface strong, mesial sclerotized ridge; larva with sub-
of head. Larvae feed in fruits and flowers, on mental pit present or absent, T1 with 2 or 3 L
dead plant tissue, predatory on scale insects and setae, Al with 1 - 3 SV setae; A9 usually with
aphids. Mainly pan tropical/subtropical; 26 gen- SD hairlike. Larvae feed on plant tissue (Amar-
era, 150 species (Hieromantis, Lactistica, Oede- yllidaceae, Arecaceae, Bromeliaceae, Cypera-
matopoda, Pseudaegeria, Snellenia, Stathmo- ceae, Juncaceae, Pinaceae, Salicaceae), fern spo-
poda, Tor tilia). rangia, scale insects, or scavenge. Worldwide; 5
Common (1990), Hodges (1978), Kasy (1973), genera, 100 + species (Batrachedra, Chedra, Ho-
Walsingham (1889). maledra).
Clarke (1963), Hodges (1966), Zimmerman
Lecithoceridae (fig. 9.2 Q). Sister-group to clades (1978).
18 36; defined by apomorphy, gnathos fused to
tegumen, mesial region downturned and laterally Deoclonidae, new family. Type genus: Deoclona
compressed, and five parallelisms: 1) frenum pre- Busck 1903. Sister-group to clades 2236; de-
sent, 2) forewing with CuP absent (rarely pre- fined by gnathos an articulated band with a me-
sent), 3) hindwing with outer margin slightly ex- sial bulb bearing parallel rows of short spines
cavated posterad of apex, 4) larva with second- (parallelism), and female retinaculum with ante-
ary setae, 5) larva with 2 L setae on prothorax. riorly directed scales on CuA or with anteriorly
Head usually smooth scaled, sometimes tufted or directed scales between CuA and R (poly-
loosely scaled. Larvae feed on detritus. Southern morphy). New World and Palearctic; 3 genera,
Palearctic, Africa; mainly Indoaustralia; 90 gen- 4 species.
era, 500 + species (Atrichozancla, Ceuthoma- Deocloninae, new subfamily. Type genus:
darus, Crocanthes, Dragmatucha, Eridachtha, Deoclona Busck 1903 (fig. 9.2 T). Defined by
Frisilia, Homaloxestis, Idiopteryx, Lecithocera, gnathos articulated with mesial, spinose bulb(s)
Odites, Tisis, Torodo ra). Torodorinae [defined by (parallelism), abdominal terga with spiniform se-
apomorphy, uncus thorn shaped, tapering to tae in band or two submesial patches (poly-
slender, acute apex (fig. 9.4 D)]; Lecithocerinae morphy), retinaculum consisting of anteriorly
[defined by uncus with narrow base and broad, directed scales on CuA or on R (polymorphy),
bilobed apex (fig. 9.4 F) and slender sclerite ex- forewing with CuP absent (parallelism), hind-
tending from base of tegumen to mesial surface wing with outer margin slightly excavated post-
of valva]; and Ceuthomadarinae (defined by ab- erad of apex (parallelism). Larva: T1 with 3 L
sence of haustellum, apophyses anteriores usu- setae, Al with 2 SV setae, A9 with SD setiform.
ally as long as bursa copulatrix, and slender scle- Pupa with paired, submesial, ridged protuber-
rite extending from base of tegumen to mesial ances on T l . Larva in dry seed pods of Fabaceae,
surface of valva) proposed for Palearctic fauna, Liliaceae, or fruit mummies of Rosaceae. New
but additional diversity extant in Australasian World, California to northern Argentina; 2 gen-
region. era, 3 species (Proclesis).
Clarke (1955 a, 1965), Common (1990), Goz- Busck (1939), Clarke (1969 b), Keifer (1936).
mny (1978), Janse (1949-54, 1958-63), Kana- Syringopainae, new subfamily. Type genus
zawa (1986), Moriuti (1982), Piskunov (1981), Syringopais Hering 1919 (fig. 9.2 U). Defined by
Sattler (1973). 4 parallelisms 1) gnathos absent, 2) vinculum
weakly sclerotized mesially, 3) hindwing with
Batrachedridae. Sister-group to clades 20-36; M1/M2 stalked, 4) larva with a submental pit and
defined by one polymorphy, abdominal support a reversal, hindwing with cell open. Larva: fron-
either pair of venulae or pair of apodemes, and toclypeus reaching epicranial notch, Al with 2
two parallelisms: 1) abdominal terga with spini- SV setae, A 3 - 6 SD with adjacent, very small
form setae in two submesial patches, 2) hindwing socket, A9 with SD setiform. Larva leaf miner
144 Ronald W. Hodges

on Triticum (Poaceae). Palearctic region, Medi- ha, Mompha, Moriloma, Synallagma, Urodela,
terranean to India; 1 genus, 1 species. Zapyrasta.
Hering (1919). Clarke (1963), Riedl (1969), Stehr (1987), Za-
gulyaev & Sinev (1981).
Coleophoridae. Sister group to clades 26-36. De- Blastobasinae (fig. 9.2 X). Defined by two par-
fined by hindwing with Rs/Mi separate (rever- allelisms: 1) forewing with pterostigma, 2) larva
sal), hindwing with M3/CUA1 separate or con- with submental pit. Aedeagus nearly membra-
nate or stalked (polymorphy), antennal pecten nous with sclerotized, ventral band; anellus
present (parallelism). Males of Blastobasinae + finely or coarsely setose before aedeagus. Larva:
Momphinae + Coleophorinae have the valva di- frontoclypeus 1/2-3/5 distance to epicranial
vided with the costa developed as a free lobe that notch; A l - 6 with 3 SV setae; A l - 8 , SD setae
often appears to arise from the sacculus (synapo- often with pinaculum rings, L2, LI vertical; A9,
morphy). Worldwide; 43 genera, 1418 species. SD seta hairlike or setiform. Pupa: T1 femur
Coleophorinae (fig. 9.2 V). Defined by gnathos visible; AIO without cremaster or recurved setae.
fused with tegumen laterally and bearing a me- Larvae mainly scavengers, sometimes on living
sial, spinose bulb (apomorphy), and five paral- plant tissue, sometimes opportunistic predators.
lelisms: 1) uncus absent; 2) Aedeagus ankylosed; Worldwide, best developed in New World; 22
3) $ frenulum with two acanthi fused distally, ap- genera, 300 species (Auximobasis, Asaphocrita,
pearing as a single acanthus; 4) retinaculum Blastobasis, Calosima, Holcocera, Hypatopa, Pi-
consisting of anteriorly directed scales between gritia, Tecmerium).
Sc and R + posteriorly directed scales/setae on Adamski and Brown (1989), Clarke (1963),
Sc; 5) forewing with CUAI/CUA2 stalked and di- Common (1990), Hodges (1978), Piskunov
rected toward posterior margin from end of cell. (1981), Powell (1976 b), Stehr (1987).
Aedeagus membranous; valva sometimes ap- Pterolonchinae (fig. 9.3 A). Defined by three
pearing like Oecophorinae with well-developed parallelisms: 1) abdominal support a pair of apo-
sacculus and prominent costal area not a discrete demes, 2) abdominal terga with spiniform setae
lobe to the base. Larva: frontoclypeus nearly to in band across segments, 3) haustellum absent
epicranial suture; A l , A2 with 2 SV setae; A9 and two reversals: 1) $ retinaculum consisting of
with SD setiform; prolegs highly reduced, diffuse, anteriorly directed, slightly upturned
crochets usually two transverse, uniordinal rows. scales, 2) hindwing with M3/CUA1 separate.
Pupa: labial palpus present, antennae often ap- Larva with no/few crochets; pupa with antennae
proximate mesially (sometimes meet), diverge approximate but separated mesially. Pterolonche
distally; segments 9/10 blunt distally, with short a root borer in Centaurea (Asteraceae). Palearc-
lateral projection. Larvae mostly leaf miners or tic (circum Mediterranean) and South Africa, in-
seed feeders on 34 plant families in Coniferae troduced into Nearctic; 2 genera, 8 species (Ana-
(rare), Monocotyledonae, and Dicotyledonae; thyrsa).
those of most species case makers. Worlwide, but Hodges (1978), Minet (1988), Vives Moreno
mainly Holarctic; 13 genera, 1,050 species; (1986).
Nearctic fauna extremely rich but poorly de-
scribed (Augasma, Coleophora, Goniodoma, Met- Autostichidae. Sister-group to clades 2936. De-
riotes). fined by gnathos an articulated band with an un-
Capuse (1971, 1972, 1973, 1975), Common articulated mesial hook (parallelism) and abdo-
(1990), Falkovitsch (1972), Powell (1980), Ra- minal terga with spiniform setae in band across
zowski (1990), Sattler & Tremewan (1974, 1978), segment (parallelism). Larvae feed on dead/de-
Stehr (1987), Toll (1952, 1962), Vives Moreno caying plant/animal material. Palearctic, Aus-
(1988). tralasia, Polynesia, Nearctic; 67 genera, 308 spe-
Momphinae (fig. 9.2 W). Defined by paral- cies.
lelism gnathos absent. Juxta ending in two small Holcopogoninae (fig. 9.3 B). Defined by $ reti-
lobes closely associated with aedeagus; signa naculum a series of anteriorly directed scales be-
paired with lightly sclerotized, circular base and tween Sc and R and posteriorly directed scales/
heavily sclerotized rod. Larva: T1 with 2 L setae, setae on Sc (parallelism). Hindwing with Rs and
tarsus sometimes with scalelike seta; Al with Mi long stalked (possible autapomorphy), lan-
1 - 3 SV setae; A9 with SD setiform; plantae ceolate; spiniform setae on abdominal terga
separate, distant or mesial, almost fused, prominent. Larva with stemmata reduced or un-
crochets uniordinal transverse circle, open el- developed, Al 8 without pinaculum ring
lipse, two parallel rows. Pupa with labial palpus around SD or pore posterad of SD, A9 with 2
present, antennae touching mesially. Larvae feed extra D setae. Larva of Holcopogon bubulcellus
on living tissue, mainly Onagraceae, also Astera- feeds on cattle dung. Palearctic region, mainly
ceae, Fabaceae, Fagaceae, Halogoridaceae, Rub- Mediterranean and eastward in xeric areas; 7
iaceae. Mainly Holarctic, very few in Neotropics, genera, 18 species (Arragonia, Charadraula, He-
New Zealand; 6 genera, 60 species (Anchimom- 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 Ronald W. Hodges

Amsel (1942), Benander (1937), Gozmny Bourquin (1957), Duckworth (1970), Hodges
(1964, 1967), Hodges (1978), Vives Moreno (1974), Kuznetsov & Stekol'nikov (1984, as Acri-
(1987). inae), Minet (1986, 1990).
Autostichinae (fig. 9.3 C). Defined by 1) $ reti-
naculum consisting of anteriorly directed setae/ Amphisbatidae (fig. 9.3 F). Sister-group to clades
scales starting in cell between base of CuA and 31-36; defined by 1) gnathos an articulated
R and extending to area between Sc and R (par- band with mesial, spined bulb (parallelism), 2)
allelism), 2) larva with pore posterad of SD on second abdominal sternum with venula only,
Al 8 (parallelism), and 3) larva with pinaculum venula and apdeme, or apdeme only (poly-
ring around SD on Al 8 (parallelism). Hind- morphy), 3) 9 frenulum three acanthi (reversal),
wing with Rs/Mi stalked/connate/separate; 4) $ retinaculum diffuse, anteriorly directed
broadly lanceolate to subquadrate; abdominal scales on CuA, or anteriorly directed scales be-
terga with fine, slender, sometimes deciduous se- tween CuA and R (polymorphy). Uncus broad
tae. Larva: frontoclypeus 2/5 distance to epicra- based, tapering to broadly rounded or acute
nial notch, Al with 2 SV setae, A l - 8 with LI/ apex; signum absent or a linear band with al-
L2 vertical, A9 with SD setiform. Pupa: labial ternating, lateral, toothed extensions; larva with
palpi minutely visible, antennae approximate/ SD hairlike on A9; antennae meet mesially on
meet mesially, cremaster present. Larvae on li- pupa. Larva (Amphisbatis) case bearer on Aster-
chens, decaying vegetable matter. Eastern Pale- aceae and Ericaceae; few others leaf tiers/rollers
arctic, Australasia, Polynesia; 18 genera, 120 + on Aceraceae, Betulaceae, Cornaceae, Fabaceae,
species (Apethistis, Autosticha, Pachnistis, Pro- Fagaceae, Juglandaceae, Myricaceae, Oleaceae,
cometis, Ptochoryctis, Stoeberhinus). Rosaceae, Salicaceae, Tiliaceae, Ulmaceae; most
Common (1990), Hodges (1978), Le Mar- unknown. Mainly New World and Palearctic; 11
chand (1947), Zimmerman (1978). genera, 65 species (Amphisbatis, Doina, Eupragia,
Symmocinae (fig. 9.3 D). Defined by forewing Machimia, Psilocorsis).
with CuP absent (parallelism), $ with retinacu- Hodges (1974), Minet (1990), Stehr (1987),
lum a series of anteriorly directed scales between Toll (1964).
CuA and R (parallelism), second abdominal ster-
num with venula, venula + apdeme, or ap- Cosmopterigidae. Sister-group to clade 34. De-
deme (polymorphy), forewing with R5 terminat- fined by 1) gnathos absent (parallelism), 2) ae-
ing on outer margin or on costa (polymorphy). deagus ankylosed (parallelism), 3) $ retinaculum
Gnathos a slender lateral band becoming a me- series of anteriorly directed scales between CuA
sial hook, distinctive; spiniform setae on abdomi- and R or series of anteriorly directed scales on
nal terga sometimes deciduous, slender. Larva: R (polymorphy), 4) hindwing with M3 and CuAi
Frons less than 1/2 distance to notch; Al - 8 with almost always separate (reversal). Worldwide;
L1/L2 vertical, SD followed by a pit, pinaculum 106 genera, 1628 species.
ring absent; A9 with SD setiform. Pupa: labial Common (1990), Fletcher (1933), Hodges
palpus and T1 femur exposed; antennae meet (1978), Kuroko (1982), Riedl (1969), Stehr
mesially, separate distally. Larvae scavengers on (1987), Zagulyaev & Sinev (1981), Zimmerman
dead plant tissue. Palearctic, circum Mediterra- (1978).
nean east to China in xeric areas, Nearctic; 42 Chrysopeleiinae (fig. 9.3 G). Defined by hind-
genera, 170 species (Amselina, Aprominta, Con- wing with cell open (reversal). Uncus present or
quassata, Dysspastus, Kurtomesis, Mylothia, Oe- absent (not two large, asymmetric lobes separate
goconia, Symmoca.) to base); larva: Al with 1/2 SV setae, Al - 8 with
Gozmny (1957, 1959, 1963, 1964), Hodges L1/L2 nearly horizontal, A9 with SVI setiform,
(1978), Stehr (1987). crochets uniordinal circle/ellipse (sometimes
open laterally); pupa: extremely compact, anten-
Peleopodidae (fig. 9.3 E). Sister-group to clades nae meet mesially, no cremaster, usually no
30-36; defined by gnathos with mesial, spinose cremastral setae. Larvae on living plant tissue,
bulb (sometimes paired) (parallelism), and abdo- leaf and bark miners, stem/root gall makers,
minal terga with scales only or with spiniform flower/seed feeders. Worldwide except Oceania;
setae in band (polymorphy). genitalia often 22 genera, 270 species (Ascalenia, Cholotis,
asymmetrical; male second sternum with ap- Ithome, Leptozestis, Perimede, Periploca, Sorha-
deme only, with venula and apdeme; forewing genia, Stilbosis, Walshia).
with CuP present; larva: T1 with 3 L setae, L pi- Clarke (1965), Hodges (1964, 1969, 1978),
naculum fused with shield, Al with 2 SV setae, Kasy (1968 a, 1968 b, 1969), Mosher (1916),
A9 with SD setiform. Immature stages poorly Riedl (1969), Zagulyaev & Sinev (1981).
known, feed on Arecaceae, Malpighiaceae, Plat- Cosmopteriginae (fig. 9.3 I). Defined by 3 par-
anaceae, Polygonaceae, Verbenaceae. New allelisms 1) vinculum weakly sclerotized or open
World and Oriental region; 6 genera, about 25 mesially, 2) $ genitalia asymmetric, 3) antennal
species (Acria, Antoloea, Durrantia, Letogenes, pecten present and one polymorphy, hindwing
Peleopoda, Pseuderotis). with Rs and Mi stalked or separate. Larva: Al
The Gelechioidea 147

with 2/3 SV setae, A 3 - 6 usually with 3 SV setae and extending to area between Sc and R, 3) pupa
(rarely 4-10), A9 with SD setiform or hairlike, with two rows of spines on terga 47 (USNM).
A l - 8 with L1/L2 nearly horizontal to nearly $$ and 9 have area of hindwing between Rs
vertical, crochets uni/biordinal circle (sometimes and CuA lacking broad scales, except on veins,
open laterally); pupa: extremely compact to very scales very slender, wing appears transparent
slender, T3 legs usually not exposed distally. Lar- (possible autapomorphy). Characters of larva
vae leaf miners/tiers, stem and root borers/gall unknown; larva stem borer in Lecythidaceae. In-
makers, seed feeders, aquatic, scavengers on at dia, Java, Borneo, Australia; 1 genus, 7 species
least 26 plant families (including Asteraceae, (Physoptila).
Cyperaceae, Fabaceae, Pandanaceae, Poaceae, Clarke (1963), Common (1974), Meyrick
Rosaceae, Typhaceae, Zamiaceae, see Zimmer- (1914).
man, 1978: 1129). Worldwide; 80 genera, 1350 Gelechiinae (fig. 9.3 K). Defined by abdominal
species (Anoncia, Asymphorodes, Cosmopterix, support a pair of venulae and a pair of apodemes
Hyposmocoma, Labdia, Leptozestis, Limnaecia, (Anacampsis and allies often with very short,
Macrobathra, Persicoptila, Pyroderces, Stagma- broad venulae) (parallelism). Several different S
tophora, Trachydora, Triclonella, Trissodoris). genital types typified by Anarsia, Aristotelia, Chi-
Clarke (1965), Common (1990), Gaedike onodes, Gelechia, Gnorimoschema, Hypatima,
(1967), Hodges (1962, 1978), Riedl (1969), Zagu- Isophrictis, Recurvara. Larva: frontoclypeus 1/2
lyaev & Sinev (1981), Zimmerman (1978). distance-nearly to epicranial notch, Al usually
Antequerinae (fig. 9.3 H). Defined by 9 frenu- with 2 SV setae, A9 with SD setiform on hair-
lum with three acanthi (reversal), retinaculum like, anal fork usually absent, crochets absent/
usually a row of anteriorly directed scales be- few/many in circle or 2 rows; pupa: antennae
tween CuA and R (parallelism), and hindwing usually touch mesially, diverge distally, A7 often
with closure of cell directed at 45 0 angle toward with fringe of hairs on posterior margin, cremas-
base of wing from M2 (parallelism). Aedeagus ter usually absent. Larval habits and hosts nearly
free, valva a single lobe, costal/saccular margins as for family. Worldwide; 480 genera, 3,400 spe-
not developed nor differentiated; larva: Al with cies (Anarsia, Aproaerema, Apatetris, Aroga, Bat-
one SV seta, A9 with SD setiform, crochets taristis, Bryotropha, Caryocolum, Coleotechnites,
sparse, a uniordinal circle; pupa unknown. Compsolechia, Deltophora, Exoteleia, Filatima,
Larva of Euclemensia parasitoids of armored Keiferia, Kiwaia, Lita, Megacraspedus, Metzne-
scale insects. North America, England; 4 genera, ria, Mirificarma, Ornativalva, Palumbina,
8 species (Antequera, Euclemensia, Panealia, Phthorimaea, Polyhymno, Protolechia, Pycnos-
Ramphis). tola, Rhynchopacha, Schizovalva, Scrobipalpa,
Hodges (1978). Scrobipalpula, Stegasta, Stomopteryx, Telphusa).
Benander (1937), Common (1990), Hodges
Gelechiidae. Defined by gnathos a pair of lateral, (1986), Janse (1949-1954, 1958-1963), Meyrick
articulated, symmetric sclerites with an articu- (1925), Minet (1979), Piskunov (1981), Po-
lated, mesial hook (synapomorphy); forewing volny & Sustek (1987), Sattler (1973).
with CuP absent (parallelism); and three poly- Dichomeridinae (fig. 9.3 L). Defined by 1)
morphies: 1) hindwing with R1 and Sc united pupa with raised, setose mesial/submesial promi-
from base of wing or R1 running into Sc beyond nences on terga 1/2, 2/3, 3/4 (apomorphy), 2) cor-
base of wing, 2) hindwing with closure of cell pus bursae with prominent secondary bursa
perpendicular with long axis of wing or directed (apomorphy), 3) abdominal support a pair of
at 45 angle toward base of wing from M2, 3) venulae (reversal), 4) frenum absent (paral-
appendix appendicular absent or present. lelism), 5) forewing with CuAi/CuA2 stalked and
Hindwing usually with outer margin excavated downcurved from end of cell, 6) hindwing with
posterad of apex. Larvae usually leaf rollers/ R1 running into Sc beyond base of wing (paral-
tiers, also within seeds, stem/root galls, mining lelism), 7) appendix appendicular present (paral-
leaves/needles, recorded from 82 plant families lelism). Larva: frontoclypeus 1/2-3/4 distance to
(Gymnospermae and Angiospermae). World- epicranial notch, Al with 2 SV setae, A9 with
wide; 507 genera, 4,530 + species. SD hairlike, anal fork usually present, prolegs
Benander (1937), Clarke (1969 a, 1969 b), noticeably long, slender, crochets usually biordi-
Common (1990), Hodges (1986), Janse (1949- nal in two sparse rows; pupa: antennae usually
1954, 1958-1963), Meyrick (1925), Moriuti touch mesially, cremaster usually absent. Larvae
(1982), Mosher (1916), Patocka (1987 a, 1987 b, leaf tiers on many plant families. Worldwide, but
1987c, 1987d, 1987e, 1988), Piskunov (1981), not endemic to New Zealand or Hawaii; 4 gen-
Powell (1980), Sattler (1973, 1979), Stehr (1987), era, 1,000 + species (Brachmia, Dichomeris, Hel-
Weber (1948). cystogramma, Scodes).
Physoptilinae (fig. 9.3 J). Sister-group to clade Hodges (1986), Povolny (1980, 1983).
35. Defined by 3 parallelisms 1) aedeagus anky- Pexicopiinae (fig. 9.3 M). Defined by abdomi-
losed, 2) 9 retinaculum of anteriorly directed se- nal support a pair of apodemes (rarely venulae
tae starting in cell between base of CuA and R + apodemes) (polymorphy). Ocellus usually pre-
148 Ronald W. Hodges




submental pit

pinaculum pore

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- meet mesially, usually diverge apically, cremaster
toclypeus 3/5 distance to epicranial notch, Al absent, cremastral spines usually present. Larvae
with 2 (1) SV setae, crochets 1, 514 in uniordi- internal feeders in seeds/developing seed pods
nal lateral penellipse, A9 with SD usually seti- (Bombacaceae, Malvaceae, 1 on Poaceae), most
form, AIO without anal fork; pupa: antennae life histories unknown. Mainly pantropical, but
The Gelechioidea 149

also Palearctic, New Zealand; 21 genera, 110 spe- - (1987): The ditrysian 'microlepidoptera' superfami-
cies (Pectinophora, Pexicopia, Sitotroga, Thio- lies. In Holloway, J. D., J. D. Bradley, and D. J.
trie ha). Carter. CIE Guides to Insects of Importance to Man.
1. Lepidoptera. [iv] + 262 pages. Wallingford [Eng-
land]: C. A. B. International.
Common (1958), Hodges (1986). Braun, A. F. (1924): The frenulum and its retinaculum
in the Lepidoptera. Ann. ent. Soc. Am., 17 (3):
Acknowledgments 234-256, pi. 23.
- (1948): Elachistidae of North America (Microlepi-
I thank Drs. V. O. Becker, I. F. B. Common, J.- doptera). - Mem. Am. ent. Soc., 13: 1 - 1 1 0 , + i, ii,
F. Landry, J. Minet, S. Passoa, J. A. Powell, K. pl. 1 - 2 6 .
Sattler, . A. Sols, and . Vandenberg for re- Bruand, T. (1850): Catalogue systmatique et syn-
view of the manuscript and my wife, E. R. S. onymique des Lpidoptres du dpartment du
Hodges, for the line drawings. Doubs. (suite). Tinides. - Mm. Soc. mul. Doubs,
3(5, 6): 2 3 - 5 8 .
References - (1859): Classification des Tinides. - Ann. Soc. ent.
Adamski, D. & Brown, R. L. (1987): A new Nearctic Fr., (3)6: 601-702.
Glyphidocera with descriptions of all stages (Lepi- Busck, A. (1903): A revision of the American moths
doptera: Blastobasidae: Symmocinae). - Proc. ent. of the family Gelechiidae, with descriptions of new
Soc. Wash., 89(2): 329-343. species. - Proc. U. S. natn. Mus., 25 (1304): 767-
- (1989): Morphology and systematics of North 938, p. 2 8 - 3 2 .
American Blastobasidae (Lepidoptera: Gelechi- - (1909): Notes on microlepidoptera, with descrip-
oidea). - Mississippi Agrie. For. Exp. Stn. Tech. tions of new North American species. - Proc. ent.
Bull., 165 (Mississippi Ent. Mus., 1): 1 - 7 0 , fig. Soc. Wash., 11: 87-103.
1-126. - (1934): Stenomidae. - Lepid. Cat., 67: 1 - 7 3 .
Amsel, H. G. (1942): Revision der Gattung Holcopo- - (1939): Un Lepidoptero nuevo que perjudica el
gon Stgr. (Lepidoptera: Scythrididae). Verff. dt. nspero en la Argentina (Familia Gelechiidae).
Kolon, u. bersee-Mus. Bremen, 3: 224-237, An. Soc. cient. argent., 127: 361-365.
pl. 16-18. Cpuse, I. (1971): Recherches morphologiques et v-
- (1977): Vier neue Kleinschmetterlingsarten aus Iran tematiques sur la famille de Coleophoridae ( Lepidop-
(Lepidoptera). Beitr. naturk. Forsch. SdwDtl, tera). - Bucarest: l'Institut International de Tech-
36: 227-236. nologie et Economie apicole d'Apimondia. 66 + [2]
Barnes, W. & McDunnough, J. (1917): Check List of pages, 46 plates.
the Lepidoptera of Boreal America, viii + 392 pp. - (1972): Contributions l'tude de la famille des
Decatur, Illinois: Herald Press. Coleophoridae (7). - Trav. Inst. Spol. Emile Ra-
Becker, V. O. (1982): Stenomine moths of the neotropi- covitza, 11: 265-270.
cal genus Timocratica (Oecophoridae). - Bull. Brit. - (1973): Sur la taxonomie de la famille des Coleopho-
Mus. nat. Hist. Ent. ser., 45: 211-306. ridae (Cls de dtermination des taxa superspci-
- (1984a): Gelechioidea. In Heppner, J. B. (ed.) Atlas fiques). Bucareti. 24 pages.
of neotropical Lepidoptera. Checklist . Part 1 Micro- - (1975): Complment systmatique la famille des
pterigoidea-Immoidea. xxvii + 112 pp. The Coleophoridae (Lepidoptera). Fragm. ent., 11 :
Hague: W. Junk. 1-64.
- (1984 b): Taxonomic notes on the neotropical Chapman, T. A. (1902): The classification of Gracilaria
microlepidoptera. - Revta. braz. Ent., 28 (2): and allied genera. - Entomologist, 35: 1159-1164.
129-201. Clarke, J. F. G. (1941): Revision of the North Ameri-
Benander, P. (1937): Die Gelechiiden-Raupen. Opus. can moths of the family Oecophoridae, with de-
ent., 2: 4 9 - 1 0 9 . scriptions of new genera and species. Proc. U. S.
Bengtsson, B. (1984): The Scythrididae (Lepidop- Natl. Mus., 90: 33-286, iviii, pl. 1 - 4 8 .
tera) of northern Europe. Fauna ent. scand., 13: - (1955 a): Catalogue of the type specimens of micro-
1 - 1 3 7 , fig. 1 - 1 3 6 . lepidoptera in the British Mus. (Natural History) de-
Brner, C. (1939): Die Grundlagen meines Lepidopte- scribed by Edward Meyrick. Vol. I 332 pages. Lon-
rensystems. - Verh. VII Int. Congr. Ent., 2: 1372 don: Trustees of the British Museum (Natural His-
1424. tory).
Bottimer, L. J. (1926): Notes on some Lepidoptera - (1955 b): Catalogue of the type specimens of micro-
from eastern Texas. - Jour. Agrie. Res., 33: 7 9 7 - lepidoptera in the British Mus. (Natural History) de-
819. scribed by Edward Meyrick. Vol. II. 531 pages. Lon-
Bourquin, F. (1957): Notas sobre la metamorfosis de don: Trustees of the British Museum (Natural His-
Pseuderotis cannescens Clarke, 1956 (Lepidop. Oe- tory).
cophoridae). Revta. soc. ent. argent., 19: 4 7 - 4 9 . - (1962): Neotropical microlepidoptera, I and II. -
Bradley, J. D. (1966): A comparative study of the coco- Proc. U. S. natn. Mus., 113 (3457): 373-388.
nut flat moth (Agonoxena argaula Meyr.) and its - (1963): Catalogue of the type specimens of microlepi-
allies, including a new species (Lepidoptera, Ago- doptera in the British Mus. (Natural History) de-
noxenidae). - Bull. Ent. Res., 56: 453-472, fig. scribed by Edward Meyrick. Vol. IV. 521 pages.
1-38. London: Trustees of the British Museum (Natural
- (1972): In Bradley, J. D D. S. Fletcher & P. E. S. History).
Whalley. A check list of the British insects, second - (1965): Catalogue of the type specimens of microlepi-
edition (revised). - Handbks. Identification Br. In- doptera in the British Mus. (Natural History) de-
sects, 11 (2): [i]viii, 1 - 1 5 3 . scribed by Edward Meyrick. Vol. V. 581 pages. Lon-
150 Ronald W. Hodges

don: Trustees of the British Museum (Natural His- Falkovitsh, M. I. (1972): New genera of palaearctic
tory). casebearers (Lepidoptera, Coleophoridae). [in Rus-
- (1969 a): Catalogue of the type specimens of micro- sian], - Ent. Obozr., 51: 369-386.
lepidoptera in the British Mus. (Natural History) de- - (1981): Families Scythrididae, Elachistidae, Stath-
scribed by Edward Meyrick. Vol. VI. 537 pages. mopodidae. - Keys Insects European Part USSR,
London: Trustees of the British Museum (Natural 4(2): 584-696. [English translation, 1989, New
History). Delhi: Oxonian Press Pvt. Ltd.]
- (1969 b): Catalogue of the type specimens of micro- Farris, J. S. (1988): Hennig 86. Version 1.5.
lepidoptera in the British Mus. (Natural History) de- Fletcher, T. B. (1929): A list of the generic names used
scribed by Edward Meyrick. Vol. VII. 531 pages. for microlepidoptera. - Mem. Dep. Agrie. India, 11 :
London: Trustees of the British Museum (Natural [i]-ix, 1 -[246].
History). - (1933): Life histories of Indian microlepidoptera
- (1978): Neotropical microlepidoptera XXI: new (second series) Cosmopterygidae [sic] to Neopseus-
genera and species of Oecophoridae from Chile. tidae. Imperial Council Agrie. Res., Scient. Mono-
Smithson. Contr. Zoology, 273: 180. gr., 4: 1 - 8 5 , pi. 1 - 7 7 .
Common, I. F. B. (1958): A revision of the pink boll- Forbes, W. T. M. (1923): The Lepidoptera of New
worms of cotton (Pectinophora Busck (Lepidop- York and neighboring states. - Mem. Cornell Univ.
tera: Gelechiidae) and related genera in Australia. agrie. Exp. Stn., 68: 1 - 7 2 9 .
- Aust. J. Zool., 6: 268-306.
- (1931): Supplementary report on the heterocera or
- (1964): The systematic status of the family Thala-
moths of Porto Rico. J. Dept. Agrie. Porto Rico,
marchellidae (Lepidoptera). J. ent. Soc. Qn., 3:
4(4): 339-394, pi. 4 3 - 4 7 .
Fracker, S. B. (1915): The classification of Lepidopter-
- (1970): Lepidoptera (moths and butterflies). In
ous larvae. Illinois biol. Monogr., 2(1): [1]161,
CSIRO Division of Entomology (Ed.), The insects
pi. 1 - 1 0 .
of Australia. Melbourne: Melbourne University
Press: 765-866. Fujisawa, K. (1991): The genus Deuterogonia (Oeco-
- (1974): Lepidoptera (moths and butterflies). In phoridae) from Japan. - Tinea, 13(13): 113-120.
CSIRO Division of Entomology (Ed.), The Insects Gaedike, R. (1967): Zur systematischen Stellung ei-
of Australia. Supplement 1974. Melbourne: Mel- niger Gattungen der Heliodinidae/Schreckensteinii-
bourne University Press: 98-107. dae sowie Revision der palarktischen Arten der
- (1980): The systematic position of Hypertropha Gattung Pancalia Curtis, 1830. - Beitr. Ent., 17:
(Lepidoptera) and related Australian genera. 363-374.
Ent. scand., 11: 17-31, fig. 1 - 4 3 . Gozmny, L. A. (1957): Notes on the generic group
- (1990): Moths of Australia, vi + 585 pages. Victoria: Symmoca Hbn. (Lep. Gelechiidae). - Annls. hist.-
Melbourne University Press. nat. Mus. natn. hung., 8 (n. s.): 325346.
Diakonoff, A. (1954): Revision of the Hypertrophinae (1959): Some new considerations on the generic
(Lepidoptera, Glyphipterygidae). I - I V . - Proc. K. group Symmoca Hbn. (Lep., Gelechiidae). Acta
ned. Akad. Wet., section C, 57: 273 - 258, 471-490, zool. hung., 5: 4 1 - 4 8 .
686-695, pi. [1], (1963): The family Symmocidae and the description
- (1954): Microlepidoptera of New Guinea, part IV. of new taxa mainly from the Near East (Lepidop-
- Verh. K. Akad. Wet., Amst., Tweede reeks, tera). Acta zool. hung., 9: 67134.
50(1): 1 - 1 9 1 . - (1964): On the generic groups Eremica Walsingham
Dietz, W. G. (1910): Revision of the Blastobasidae of and Symmocoides Amsel (Lepidoptera: Symmoci-
North America. - Trans, am. ent. Soc., 36: 1 - 7 2 , dae). - Acta zool. hung., 10: 101-129.
pi. 1 - 4 . (1967): The family Holcopogonidae fam. nov. (Lep-
Duckworth, W. D. (1964): North American Stenomi- idoptera) and its constituent taxa. - Acta zool.
dae (Lepidoptera: Gelechioidea). - Proc. U. S. hung., 13: 271-278.
natn. Mus., 116 (3495): 2 3 - 7 1 , pl. 1 - 4 . - (1975): New symmocid species (Lepidoptera) and
- (1967): Neotropical microlepidoptera XIII. Review the description of an unknown abdominal organ. -
of genus Loxotoma (Lepidoptera: Stenomidae). Acta zool. hung., 21: 263-269.
Proc. U. S. natn. Mus., 122 (3590): 1 - 8 , pl. 1. (1978): Lecithoceridae. Microlepidopt. Palaearctica,
- (1969): Bredin-Archbold-Smithsonian Biological
5: [i] xxviii + 1 - 3 0 6 , pi. 1 - 9 3 .
Survey of Dominica: West Indian Stenomidae (Lep-
Guene, A. (1845): Essai sur une nouvelle classification
idoptera: Gelechioidea). - Smithson. Contr. Zool-
des microlpidoptres. - Ann. Soc. ent. Fr., (2) 3):
ogy, 4: 1 - 2 1 , fig. 1 - 3 0 .
- (1970): Neotropical microlepidoptera XVIII: revi-
Hampson, G. F. [1893], (1892): The Fauna of British
sion of the genus Peleopoda (Lepidoptera: Oeco-
India, including Ceylon and Burma. Moths, vol. 1.
phoridae). Smithson. Contr. Zoology, 48: 1 - 3 0 .
xxiii + 527 pages. London: Taylor & Francis.
- (1971): Neotropical microlepidoptera XX: revision
of the genus Setiostoma (Lepidoptera: Stenomidae). (1918): Some small families of the Lepidoptera
- Smithson. Contr. Zoology, 106: 1 - 4 5 . which are not included in the key to the families in
- (1973): The Old World Stenomidae: a preliminary the Catalogue of Lepidoptera Phalanae, a list of the
survey of the fauna, notes on relationships, and re- families and subfamilies of the Lepidoptera with
vision of the genus Eriogenes (Lepidoptera: Ge- their types, and a key to the families. Novit, zool.,
lechioidea). - Smithson. Contr. Zoology, 147: 1 - 2 1 , 25: 366-394.