Oecophorine Genera of Australia I: The Wingia Group (Lepidoptera: Oecophoridae)

Oecophorine Genera

of Australia

I

The Wingia Group

(Lepidoptera : Oecophoridae)

Monographs on Australian Lepidoptera

Editorial Board

Editor-in-Chief

E.S. Nielsen

Australian National Insect Collection

CSIRO Division of Entomology

GPO Box 1700

Canberra, ACT 2601, Australia

Editorial Panel

I.F.B. Common, Toowoomba, Australia

D.C. Ferguson, Washington D.C., USA

M, Horak, Canberra, Australia

R.L. Kitching, Brisbane, Australia

G.S. Robinson, London, UK

M.J. Scoble, London, UK

Myrascia bracteatella (Walker,1864) female

Drawn by Mr R. Nielsen, Copenhagen

Monographs on

Australian

Lepidoptera

Volume 3

Oecophorine Genera

of Australia

I

The Wingia Group

(Lepidoptera: Oecophoridae)

Ian F.B. Common

CSIRO Division of Entomology

Canberra, Australia

CSIRO Cataloguing-in-Publication Entry

Common, I. F. B. Ian Francis Bell.

Oecophorine genera of Australia. I. The Wingia group (Lepidoptera : Oecophoridae) / Ian F.B. Common.

(Monographs on Australian Lepidoptera; v. 3) Includes bibliograpical references and index. ISBN 0 643 05524 X.

1. Oecophoridae - Australia. 2. Oecophoridae - Classification. 3. Moths - Australia. I. CSIRO. II. Title. (Series: Monographs on Australian Lepidoptera; v. 3).

595.7810994

© 1994 CSIRO Australia

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Monographs on Australian Lepidoptera

Previous volumes in this series:

Volume 1, 1989

Primitive Ghost Moths

Morphology and taxonomy of the Australian genus Fraus Walker

(Lepidoptera : Hepialidae s. lat.)

E.S. Nielsen and N.P. Kristensen

Volume 2, 1993

Teneid Genera of Australia

(Lepidoptera)

G.S. Robinson and E.S. Nielsen

Contents

Abstract

Introduction

Material and Methods

Material Studied

Preparatory Techniques

Abbreviations, Definitions and Conventions

Acknowledgments

1. Phylogeny of the Oecophorinae

2. Morphology

Head

Thoracic Appendages

Pregenital Abdomen

Male Genitalia

Female Genitalia

Egg

Larva

Pupa

3. Biology

Life Histories

Host Relationships

Adult

Oviposition

Larva

Pupa

Pests and Synanthropes

Parasitoids and Predators

Collecting and Rearing Oecophorinae

4. Diversity and Distribution

5. Australian Genera of the Wingia Group of Oecophorinae

Key to Genera

Subgroup 1

1. Lamproxantha gen. n.

2. Polyeucta Turner

3. Ancistromorpha nom. n.

4. Hapaloteucha Meyrick

5. Leucorhabda gen. n.

6. Orthiastis Meyrick

7. Habroscopa Meyrick

Subgroup 2

8. Crepidosceles Meyrick

9. Lepidotarsa Meyrick

Subgroup 3

10. Ageletha gen. n.

11. Lophopepla Turner

12. Anthocoma Turner

13. Pycnozancla Turner

14. Elaphromorpha Turner

15. Eochrois Meyrick

16. Cosmaresta gen. n.

17. Plectobela gen. n.

18. Poliorhabda gen. n.

19. Tanyzancla Meyrick

20. Saphezona gen. n.

21. Heteroteucha gen. n.

22. Rhadinoloba gen. n.

23. Psaroxantha gen. n.

24. Coeranica Meyrick

25. Brachynemata Meyrick

26. Stictochila gen. n.

27. Zonopetala Meyrick

28. Piloprepes Meyrick

Subgroup 4

29. Euphiltra Meyrick

30. Hoplomorpha Turner

31. Euthictis Meyrick

32. Scotodryas Turner

33. Eremnotypa gen. n.

34. Placocosma Meyrick

Subgroup 5

35. Ironopolia gen. n.

36. Prodelaca gen. n.

Subgroup 6

37. Linosticha Meyrick

38. Psaltriodes Meyrick

39. Limothnes Turner

40. Basiplecta gen. n.

41. Idiozancla Turner

Subgroup 7

42. Parocystola Turner

43. Liocnema Turner

44. Arachnographa Meyrick

45. Epicurica Meyrick

46. Phytotrypa gen. n.

47. Echinobasis gen. n.

48. Acmotoma gen. n.

49. Baiocystis gen. n.

50. Paneutricha gen. n.

Subgroup 8

51. Heteroptolis Meyrick

52. Callimima Turner

53. Hybocrossa Turner

54. Acorotricha Meyrick

55. Trachyzancla Turner

Subgroup 9

56. Deigmoesta gen. n.

57. Tortricopsis Newman

58. Wingia Walsingham & Durrant

Subgroup 10

59. Platyphylla Turner

60. Ptyoptila Turner

61. Acanthodela gen. n.

62. Ericibdela gen. n.

Subgroup 11

63. Archaereta Meyrick

64. Enoplidia gen. n.

65. Euchaetis Meyrick

66. Heliocausta Meyrick

67. Prionocris gen. n.

68. Syringoseca gen. n.

Subgroup 12

69. Catacometes gen. n.

70. Chrysonoma Meyrick

71. Bathrosterra gen. n.

72. Garrha Walker

73. Sclerocheta gen. n.

74. Atelosticha Meyrick

75. Antipterna Meyrick

76. Coryphoscola gen. n.

77. Diaphorodes Turner

78. Endeolena gen. n.

79. Hemibela Turner

80. Idioxantha nom. n.

81. Mionolena gen. n.

82. Thalerotricha Meyrick

83. Zacorus Butler

84. Zelotechna Meyrick

85. Hadrocheta gen. n.

86. Ocystola Meyrick

87. Myrascia Common

88. Oligoloba gen. n.

89. Compsotropha Meyrick

90. Phyllophanes Turner

91. Stereoloba gen. n.

6. Unplaced Species

7. References

8. Index to Lepidoptera Names

9. Index to Plant Names

Abstract

The Oecophorinae represent at least 20% of the Australian moth fauna, with some 1850 valid named species in more than 250 genera, and with an estimated total fauna of more than 5000 species. Until now the classification of this large number of genera and species has been dependent on the classical work of Meyrick and, more recently, Turner, each of whom based his conclusions largely upon superficial characters of wing venation, scale vestiture, antennae and mouthparts. No attempt was made to examine and utilise structures which were not visible on an undamaged specimen, and little was known about the biology and larval food plants. The results reported here have revealed the chaotic state of the classification adopted in these earlier revisions.

The present volume presents a revision of the Wingia group of 91 genera, a group which appears to be almost entirely endemic to Australia. Detailed information is provided on the morphology, including the genitalia of both sexes, and up-to-date data on the distribution, biology and larval food plants of the species referred to each genus. A list of the species assigned to each genus is given, with full synonymy, references to the original descriptions, label data and repository of the primary type, lectotype designations, and abbreviated distribution data. Illustrations of the adults, wing venation, and male and female genitalia of the type species are included, as well as the heads and eggs of representative genera. A key to the genera is also provided.

Thirty-five new genera are proposed; three of these are based on new species, descriptions of which are added. A total of 500 valid named species are here referred to the 91 genera of the Wingia group, and an additional 268 undescribed species have been identified in collections, mainly the Australian National Insect Collection. The revision proposes 305 new generic combinations and 88 new specific synonyms.

A very tentative phylogenetic analysis of the Wingia group of genera has been attempted using the Hennig86 computer program. This utilised only 33 characters; the group was thus divided into 12 subgroups; within several subgroups the phylogeny remains unresolved and the position of certain genera appears to be anomalous.

The Wingia group is well represented in each of the Australian states and territories, but both species and genera are much more numerous in the southern half of the continent. The largest number of genera, 80 and 78 respectively, and named species, 274 and 270, have been recorded from southern Queensland and New South Wales.

It is clear that a high percentage of the Australian Oecophorinae depend for their larval food on plants belonging to the ‘dry-fruit’ genera of the family Myrtaceae, especially Eucalyptus, but also Lophostemon, Angophora, Syncarpia, Melaleuca, Leptospermum and Kunzea. It is thought that most of the Australian Oecophorinae evolved within Australia from Gondwanan ancestors, and were subjected to the same environmental restraints as the plant genera upon which they depend and, like their hosts, proliferated with the gradual onset of aridity. It is significant that a high percentage of the Australian species utilise dead leaves, especially of Eucalyptus, a material of low nutritive value which few other Australian Lepidoptera favour. It is reasonable to conclude that, in view of the large number of species and often relatively high populations of individuals, this group of moths must play an important role in breaking down eucalypt leaf litter to humus.

I.F.B. Common

CSIRO Division of Entomology

GPO Box 1700

Canberra, ACT 2601

Australia

Private mail address:

32 Katoomba Crescent

Toowoomba

Queensland 4350

Australia

Introduction

Anyone interested in Australian moths soon comes to recognise the disproportionate representation of the Oecophorinae. Although most species are small, with a wingspan of less than 50 mm, and many are relatively dull insects, a high proportion have distinctively patterned wings, often in orange-yellow and dark brown, and some are delicately coloured. As in other groups of the large superfamily Gelechioidea, the labial palpi are usually strongly recurved and the proboscis has overlapping scales near the base. Most species can be recognised by the nearly parallel condition of the veins Rs and M1 of the hind wing, and by the male genitalia in which the gnathos is either long and beak-like or is much shorter and rounded, in both instances with an area of scobinations or short fine spines above. The abdominal terga nearly always carry areas of sclerotised spine-like setae set in small sockets, as well as the scale vestiture.

The adults of most Australian Oecophorinae rest with their wings folded either roof-wise or flat above the body. They usually walk with a slow waddling gait and have a characteristic, fairly slow undulating flight. They come freely to lights at night, and most specimens may be captured either at a sheet illuminated by a bright light or in specially designed light traps. A few are diurnal, especially at high altitudes, and some tend to fly for a short period in early morning sunshine or towards dusk. The early stages are still not well known, but there are strong indications that a high percentage of the Australian species are associated with the dry-fruit section of the plant family Myrtaceae, especially Eucalyptus. Accordingly, they have been given the common name of mallee moths.

The Australian Oecophorinae include some 1850 valid named species referred to more than 250 genera, less than two-thirds of which are named. It is estimated that the fauna may well exceed 5000 species, and already more than 1500 undescribed species are represented in collections, especially the ANIC. As a detailed revision of such an enormous group of insects would be a daunting task, it was decided to provide generic descriptions, with figures of adults, wing venation, and genitalia of the type species, and provide a tentative classification based on a cladistic analysis. It has always been clear that the discrimination of genera in such a numerically large group would be difficult and, indeed, this was the experience of the only previous revisers of the subfamily, Edward Meyrick and and Alfred Jefferis Turner.

This revision of the Wingia group of the Australian Oecophorinae includes 91 genera, 35 of which are described as new. The number of available specific names was 620, of which 126 proved to be synonyms, including 88 new synonyms, with 500 valid described species and a further 268 undescribed species. There are 305 new generic combinations. Included is a detailed description of each genus, information on the distribution and biology, a synonymic list of the species referred to each genus, and a key to the genera.

The first Australian oecophorine to be named was Phalaena fascialis Fabricius (1775), based on a female specimen collected by Banks and now preserved in the Banks Collection in The Natural History Museum, London. This species was later designated as the type species of Meyrick’s (1914) genus Chrysonoma. Edward Donovan (1805) figured a specimen of fascialis as Tortrix bimaculana, and in the same work figured Tinea strigatella, now referred to Philobota Meyrick. Among 11 species of Oecophorinae described by Edward Newman (1856) from a small collection made on the Mount Alexander Range … about eighty miles from Melbourne, were specimens of fascialis which Newman identified as Oecophora bimaculella. One of his 11 new species was referred to Depressaria Haworth, one to a new genus Tortricopsis and the remainder to Oecophora Latreille.

Except for Newman, Walker and Meyrick, few Australian Oecophorinae were described between 1850 and 1890, no doubt because of the paucity of specimens in European collections. Wing (1850) named Palparia lambertella, Zeller (1855, 1877) named nine species, Wallengren (1861) named Gelechia antipodella, R. Felder and Rogenhofer (1874-75) named Symmoca herodiella and Tinea codrella, Butler (1882) named Conchylis auriceps, Latometus pilipes and Zacorus carus, and Rosenstock (1885) named Philobota athletica and Eulechria leucospina.

In 1863, 1864a, 1864b and 1869 Francis Walker proposed 97 names for Australian Oecophorinae, of which 15 prove to be synonyms and one, Oecophora divisella, a primary homonym. He proposed five new monotypic genera, Chezala, Barea, Thema, Tisobarica and Garrha, but referred most of his species to Oecophora Latreille, Cryptolechia Zeller, Depressaria Haworth, Psecadia Hübner, Incurvaria Haworth, Gelechia Haworth and Palparia Wing.

The first serious attempt to deal comprehensively with the taxonomy of the Australian Oecophoridae resulted in a series of papers by Edward Meyrick (1883a, 1883b, 1884a, 1884b, 1885, 1886, 1888, 1889). His interest in the microlepidoptera was stimulated during a period spent in Australia and New Zealand between 1877 and 1886, when he was introduced to an extensive, largely unknown oecophorid fauna. In these papers he gave a definition of the family, based on external characters of the adult, described the features he used to distinguish it from other related families, and proposed 66 new genera from Australia and 11 from New Zealand. Sixty-two of the Australian genera are now known to belong to the Oecophorinae and 21 of these to the Wingia group of genera, although four of the Wingia group are here treated as synonyms. He listed 756 species including 35 from New Zealand and 626 Australian species described as new. He also included a key to all of the then known genera and, for a few genera, keys to the known species. In the last paper of this series he added an appendix containing biological and other notes about the species described. He pointed out that the uniformity of structure throughout the whole [family] is extraordinary and that the points of structure on which it has been necessary to rely [in the generic classification] are in some cases slight, in others difficult of observation.

After his return to England in 1886 Meyrick continued to describe new genera and species of Australian microlepidoptera as well as many from other parts of the world. Three of his publications are of special importance to Australia. Meyrick (1902) described four new genera and 48 new species of Australian Oecophorinae. Between 1912 and 1923 in the first two volumes of Exotic Microlepidoptera, Meyrick described 227 new species of Australian Oecophorinae and 35 new genera. The specimens came from a variety of sources, including many in 1913 from George Lyell in Gisborne, Victoria, and in 1921 some from Oswald Lower in Adelaide, South Australia. In 1922 Meyrick published his revision of the world fauna of Oecophoridae (Meyrick 1922c). During his lifetime Meyrick described 108 genera of Australian Oecophorinae, of which 89 are currently accepted as valid, and about 850 species, of which 74 have been treated as synonyms.

Second in significance only to the enormous contribution made by Meyrick was that of Alfred Jefferis Turner, a medical practitioner in Queensland and a noted paediatrician. Turner dealt with most of the families of Australian Lepidoptera, but had a special interest in the microlepidoptera. He closely followed the taxonomic principles laid down by Meyrick and in his first paper in 1894 paid tribute to Meyrick’s work and stated that he had endeavoured to adhere faithfully to the genera [of Oecophoridae] laid down by Meyrick, but had to describe a few new genera which appear[ed] distinct from any of his. In that paper (Turner 1894) he proposed as new genera Eurypelta and Hemibela based on new species, and described other new species in Palparia Wing, Lepidotarsa Meyrick, Eomystis Meyrick and Euphiltra Meyrick. He published several additional papers containing new species of Oecophorinae between 1896 and 1917 (Turner 1896, 1898, 1914, 1915, 1916a, 1916b, 1917), and in two later papers (Turner 1927, 1939a) described new species and genera from Tasmania. Like Meyrick, Turner had a particular interest in the enormous Australian fauna of Oecophoridae and, in a series of papers between 1932 and 1947, the year of his death, published a detailed revision of the family. This included descriptions of many new genera and species and keys to the genera. Altogether Turner proposed 76 generic names of Oecophorinae, of which 49 are accepted as valid, and 1060 specific names, of which 225 are treated as synonyms. The contribution of Meyrick and Turner to our knowledge of this large group was enormous, although subsequent work has shown the necessity to alter considerably the concept of many of their genera.

Another amateur lepidopterist who contributed many specific names and a few generic names for Australian Oecophorinae was Oswald Lower, a pharmacist who lived in Adelaide, South Australia and Broken Hill in western New South Wales. Between 1892 and 1920 he published a series of 23 papers in the Transactions of the Royal Society of South Australia and the Proceedings of the Linnean Society of New South Wales in which he described 181 new species, of which 33 are treated as synonyms and 16 have not yet been identified because of major damage to, or loss of, the original specimens. He also proposed seven new genera, two of which are valid, three are synonyms, and two monobasic genera remain unknown because the original specimens have apparently been lost.

Until recently there has been little published information on Australian Oecophorinae requiring any considerable modification of Meyrick’s (1922c) classification of the world fauna, which relied on external characters, especially the wing venation, antennae, scale vestiture and mouthparts. Turner’s (1932-1947) classification of the Australian Oecophoridae closely followed Meyrick’s system, with few major changes except for the addition of numerous new species and genera. Gaede’s (1938, 1939) catalogue of the world Oecophoridae was based uncritically on Meyrick’s (1922c) revision, with the addition of the data in Turner’s papers which had been published up to 1938. However, both Meyrick and Turner failed to recognise the value of genitalia or other internal characters in distinguishing species or higher categories.

More recent studies of the family Oecophoridae have been mainly on regional faunas. Busck (1908) first used the genitalia, among other characters, in his classification of the North American Oecophoridae. Philpott (1927) figured the male genitalia of the known New Zealand species and Pierce and Metcalfe (1935) those of species of the British fauna, but neither of these two authors attempted to utilise the information gained for reclassifying the family. Clarke (1941) revised the classification of the North American oecophorids featuring the genitalia of both sexes and added new genera and species. Toll (1964) presented a taxonomic revision of the Polish Oecophoridae using genitalia as well as other characters, and the North American fauna was again revised by Hodges (1974). However, because of the paucity of the North American and European faunas of Oecophorinae, these revisions have had only a limited relevance to our understanding of the vast, largely endemic Australian fauna.

A first attempt at providing an insight into the important oecophorid fauna of southern South America was by Clarke (1978), who dealt with 34 genera from Chile, including 26 new genera and a total of 60 species, 39 of which were described as new. He presented descriptions and figures of the genitalia of most of the species, and a key to the genera. I have examined the genitalic preparations of this material in the National Museum of Natural History, Washington, hoping to identify any Gondwanan links with Australia. Some of the genera clearly belong to the Depressariidae but, judging by the genitalia, it is doubtful if any of the Oecophorinae have any near relationship to the Australian fauna.

Material and Methods

Material studied

The largest amount of Australian oecophorine material, including most of the types of A.J. Turner, is contained in the Australian National Insect Collection, although the collection of The Natural History Museum, London (BMNH), which includes most of the types of Edward Meyrick, Francis Walker and other authors, is also of vital importance for the study of the Australian fauna. The type and other material in these two collections provided the basic information for this revision. This was supplemented by detailed examination of the Oecophorinae in the W.B. Barnard collection in the Queensland Museum, Brisbane, the Goldfinch collection in the Australian Museum, Sydney, the George Lyell collection in the Museum of Victoria, Melbourne, and the collection of the South Australian Museum, Adelaide, which includes the majority of the O.B. Lower types and a few of T.P. Lucas.

Lectotypes for Meyrick’s and Walker’s Australian species in BMNH have been selected and designated, where necessary, the holotypes and lectotypes photographed in colour and the genitalia of a large proportion of the types of these and other authors have been dissected and photographed. The photographs of the adults and genitalia have been matched with specimens in the ANIC and examples dissected. The Turner types have likewise been examined, lectotypes selected, and the genitalia dissected. The Turner types in the Queensland Museum and the Australian Museum, and the Meyrick, Turner and a few Walker types in the Museum of Victoria have also been examined, colour transparencies made and the types of critically important species dissected and the genitalia photographed. The surviving types of Lower, Lucas, and a few Meyrick and Turner types in the South Australian Museum have also been studied and photographed, lectotypes designated, and the genitalia of doubtful or unrecognised species dissected and the genitalia photographed.

A checklist of all oecophorine species recorded from Australia has been compiled and their synonymy established. The species composition of previously proposed genera has been critically examined, using criteria provided by as many characters as possible including those provided by the genitalia of both sexes. Where necessary, to establish the monophyly of each established genus, species were added or removed, or new synonymy discovered, necessitating the description of several previously unrecognised genera. New species have been described only where it has been necessary to establish new genera. No doubt further refinement of the classification of the Australian Oecophorinae will be necessary as additional species are discovered or described.

Preparatory techniques

Pinned adult specimens were examined in artificial light using a Nikon or Wild M5 stereoscopic microscope. A camera lucida attached to the eye-piece of the Nikon microscope was used to draw the head, genitalia and pupae. Slide preparations of heads, wings, legs, abdomens and genitalia were examined at higher magnifications with a Leitz compound microscope. Size measurements were made with an eye-piece micrometer and converted to millimetres. The heads of adults were examined with the low-power microscope and scanning electron micrographs of the type species of a representative number of genera were made after the heads were sputter coated with gold.

Microscope slide preparations of wings were made using the method described by Common (1987, 1990). The two wings on one side were detached, wetted with 70% ethanol, and bleached with a solution of sodium hypochlorite or household bleach. After thorough washing in distilled water, they were immersed in an acid fuchsin stain for about 24 hours. They were then transferred briefly to 95% ethanol and dehydrated in absolute ethanol before mounting on a microscope slide in Euparal.

Genitalia were dissected, stained and mounted on microscope slides using the techniques described by Robinson (1976). They were stained with Chlorazol Black E (1% in 70% ethanol). Abdominal pelts were also stained with Chlorazol Black E and, after cleaning and removal of scales with a water-colour brush, either mounted flat on the slide, ventral surface uppermost (males), or cut along the left-hand side and spread out flat (females). The preparations were mounted in Euparal. At first difficulty was experienced in transferring dissected female genitalia having a coiled ductus bursae from one solution to the next or on to a slide, without completely displacing the coils. This problem was overcome by constructing a small scoop (a miniature ‘egg-lifter’ or ‘egg-slice’) from a right-angled piece of stiff transparent plastic attached with ‘Superglue’ to the handle of a water-colour brush. The whole genitalia could then be carefully lifted on the horizontal part of the plastic and transferred, with the coils undisturbed, from one solution to the next, or floated on to the slide in absolute ethanol for mounting in Euparal.

Abbreviations, Definitions and Conventions

The terminology used for the external and genital morphology follows that of Nielsen and Common (1991) and Klots (1956).

Acknowledgments

The work which has culminated in this monograph has been in progress intermittently over many years during my service with the CSIRO Division of Entomology in Canberra, and could not have been undertaken without the generous support of the Division’s administration, especially that of Dr Douglas Waterhouse and Dr Max Whitten. The most recent work on the generic revision and the preparation of the manuscript and figures have been greatly facilitated by grants from the Australian Biological Resources Study (ABRS) of the Australian Nature Conservation Agency (ANCA), which provided for valuable technical assistance, the provision of a new PC computer and a Bubblejet printer.

Special thanks for technical assistance are due to Ms Leane Regan, who provided immeasurable help with dissecting, wing preparations, curation, photocopying, reducing line drawings, inserting numbers and scale lines on the figures, and the preparation of the references and index for this monograph. I am extremely grateful to Ms Vanna Rangsi for mounting and labelling specimens, and other valuable assistance over many years. I thank Mr Colin Beaton for the scanning electron micrographs, Mr John Green for photographs of mounted specimens and genitalia, and for preparing half-tone negatives from my colour transparencies, and Ms Jenny Alyta for printing photographs. The majority of the line drawings of wing venation were prepared by Ms Leane Regan and Mrs Sybil Monteith. The phylogenetic analysis was carried out by Dr Ebbe Nielsen and Mr Eric Zurcher, and the cladogram was drawn by Ms Sandy Smith. The colour transparencies used in Figs 77, 232, 261 and 518 were by the late Mr Bob Jessop, and that used in Fig. 681 was by Mr Ted Edwards. Permission for republishing Figs 3, 24-26, 27, 610, 671, 672, and 681 was given by the editor of the Journal of the Australian Entomological Society, and for Figs 28-31 and 610 by Melbourne University Press.

The value of this project would have been greatly reduced without the generous help provided by the various institutions holding significant collections of Australian Oecophoridae. Thanks are due particularly to the authorities of The Natural History Museum, London, the Queensland Museum, Brisbane, the Australian Museum, Sydney, the Museum of Victoria, Melbourne and the South Australian Museum, Adelaide, and the Department of Primary Industry, Hobart, for providing facilities to work on their collections and for loans of valuable material. I am grateful to the many colleagues of these institutions for their ready assistance, including Dr Gaden Robinson, Dr Klaus Sattler and Mr Kevin Tuck of The Natural History Museum, London, Mr Ted Dahms, Dr Geoff Monteith and Ms Gudrun Sarnes of the Queensland Museum, Mr Max Moulds of the Australian Museum, Dr Artur Neboiss and Mr Ken Walker of the Museum of Victoria, and Dr Gordon Gross, Dr Eric Matthews and Ms Jan Forrest of the South Australian Museum. For their continued support, advice and help in numerous ways my sincere thanks go to my colleagues of the CSIRO Division of Entomology, especially Dr Max Whitten, Dr Ebbe Nielsen, Dr Marianne Horak, Mr Ted Edwards and Mr Murray Upton.

Chapter 1

Phylogeny of the Oecophorinae

Recent studies by Nielsen (1989), Minet (1986, 1991) and others have demonstrated that the four superfamilies Tineoidea, Gracillarioidea, Yponomeutoidea and Gelechioidea represent the most primitive ditrysian Lepidoptera. All the remaining Ditrysia, termed the Apoditrysia, may be distinguished by the presence of tortricoid-type apodemes of abdominal sternum 2, instead of tineoid-type apodemes. The phylogenetic inter-relationship of the four primitive superfamilies, however, is still not clear and has been discussed by Robinson and Nielsen (1993). The Gelechioidea appear to be a monophyletic group, defined by at least three apomorphies: the overlapping scaling on the base of the proboscis, the up-curved three-segmented labial palpi, in which the apical segment is usually slender and acute, and the four-segmented maxillary palpi, which are folded over the base of the proboscis (Minet 1986).

The status of the 18 families included in the Gelechioidea by Common (1970) has been greatly modified by more recent authors such as Bradley (1972), Hodges (1978), Common (1990), Minet (1986, 1990), Nielsen and Common (1991) and Leraut (1992). Bradley (1972) adapted Common’s (1970) system to the limited British fauna, including only nine families, but made relatively minor changes. Hodges (1978) proposed a total of 10 families, including the Ethmiidae, Xyloryctidae and Stenomatidae as subfamilies of the Oecophoridae, along with the Oecophorinae, Depressariinae, Peleopodinae, Autostichinae, Chimabachinae, Deuterogoniinae and Hypertrophinae. He treated the Stathmopodidae as a tribe within the Oecophorinae, Batrachedrinae as a subfamily of the Coleophoridae, Symmocinae as a subfamily of the Blastobasidae, and Blastodacninae as a subfamily of the Agonoxenidae. Common (1990) and Nielsen and Common (1991) accepted 17 families in the Gelechioidea, giving family status to the Batrachedridae, Hypertrophidae, Depressariidae, Ethmiidae, Symmocidae, Blastodacnidae and Holcopogonidae. They recognised that the spinose gnathos represents an apomorphy shared by the Depressariidae, Elachistidae, Agonoxenidae and Coleophoridae, necessitating the removal of the Depressariidae from the Oecophoridae. They also considered that the upright, exposed pupae of the Hypertrophidae and some genera of the Depressariidae, represents an apomorphy distinguishing the Hypertrophidae from the Oecophoridae. However, they followed Hodges (1978) by retaining the Xyloryctinae, Autostichinae, Stenomatinae and Stathmopodinae as subfamilies of the Oecophoridae.

Major modifications to the family relationships within the Gelechioidea were proposed by Minet (1990), who listed a series of 32 characters of the immature and adult stages which he considered provided significant apomorphies for his proposed 17 families. Of special note is his new concept of the family Elachistidae, which he enlarged to include as subfamilies the Agonoxeninae, Elachistinae, Stenomatinae, Cryptolechiinae, Hypertrophinae, Ethmiinae and Depressariinae. He proposed as an apomorphy defining the Hypertrophinae the erect, exposed pupa, attached by numerous hooked setae to a pad of silk spun by the larva on the substrate. This redefined subfamily thus included not only the genera listed by Common (1980) but also those genera with a more or less upright pupa, such as Scorpiopsis Turner, Pedois Turner, Peritornenta Turner, Octasphales Meyrick and Tonica Walker, previously referred to the Depressariinae. Based principally on larval apomorphies, especially the very close proximity of setae SD1 and SD2 on abdominal segments 1 to 8, Minet considered that nine families (Xyloryctidae, Batrachedridae, Oecophoridae, Symmocidae, Lecithoceridae, Scythrididae, Epimarptidae, Blastobasidae and Stathmopodidae) form a monophyletic group. Of special note in the context of my revision of the Australian Oecophorinae, Minet suggested that the Oecophoridae (sensu stricto) may be paraphyletic. He also maintained that the Stathmopodidae are most closely related to the Blastobasidae, giving as evidence (Minet 1986) the scaling of the head, the proximity of R1 to R2 and the short CuA1 and CuA2 in the fore wing, and the female retinaculum, which he considered resembled that of the Blastobasidae.

In a study of the occurrence and phylogenetic significance of tineoid-type apodemes and venulae (sternal rods) of sternum 2 in the primitive Ditrysia, Kyrki (1983) showed that the development of these structures varied considerably in families of the Tineoidea, Yponomeutoidea and Gelechioidea. Leraut (1992) made a more extensive examination of the apodemes and venulae of a wide range of families and subfamilies in the Gelechioidea and, as a result, redefined most of these taxa including the Oecophoridae, in which he recognised two subfamilies, the Oecophorinae and the Philobotinae. The latter subfamily, he said, is without doubt the more archaic. Unfortunately his conclusions show some inconsistencies which detract from their value. In his definition of the family Oecophoridae, for example, he stated that the apodemes of sternum 2 are long in the female and reduced in the male, whereas in the subfamily Philobotinae he claimed that the apodemes are well developed in the male. Nevertheless, he figured (his fig. 1) sternum 2 of the male of Philobota interlineatella showing the apodemes and venulae reduced or absent. In fact, this species belongs to Myrascia Common, one of many Australian genera in the Wingia group of Oecophoridae in which the apodemes and venulae of the male are reduced or absent.

Our knowledge of the vast Australian fauna of Oecophoridae is still not sufficient to determine with any degree of confidence if Leraut’s Oecophorinae or Philobotinae are monophyletic or, indeed, if there is an apomorphy defining the family Oecophoridae. Certainly all the Australian species that I have examined have an unarticulated gnathos, the arms of which unite to form a short, dorsally concave lobe, usually produced distally into either a short drooping, apically acute process or a much longer beak-like structure. Dorsally all have an area of scobinations or short thorns or, less frequently, an area of minute, fine, proximally directed spines. This type of gnathos may prove to be an apomorphy defining the Oecophoridae. The Stathmopodidae also have a beak-like gnathos, not unlike many Oecophoridae and, in addition, have an undivided valva, and are without the specialised, setose anellus and inner longitudinal sclerite of the aedeagus characteristic of all known Blastobasidae (Adamski and Brown 1989). The pterostigma of the fore wing between veins Sc and R1 found in Blastobasidae is absent in Stathmopodidae and, although CuA2 is a short vein in the latter, it does not form a near right angle with CuA as in Blastobasidae. There seems to be little evidence therefore for associating the Stathmopodidae with the Blastobasidae and, following Common (1990), Nielsen and Common (1991) and Hodges (in press), they are here accepted as a subfamily of the Oecophoridae.

As a first step towards a thorough species-level revision of the Australian Oecophoridae, it was decided to undertake the present generic revision. Of course, generic concepts must be based on a study of the species, but this need not be as exhaustive as a species-level revision and, for such an enormous number of species, a generic revision is much more practical and should lead to a more rapid understanding of the generic relationships and the diversity of the Australian fauna. In preparing my oecophorid part of the Checklist of the Lepidoptera of Australia (Nielsen, Edwards and Rangsi in preparation), it was possible to recognise several major groups of genera, most of which are probably monophyletic, although some may prove to be paraphyletic. However, I consider it would be premature to give these groups formal names. This first monograph deals with the Wingia group of genera.

The Wingia group of genera may be defined by three synapomorphies: (a) the sexually dimorphic apodemes of abdominal sternum 2, greatly reduced or absent in males (Figs 4, 6), but elongate in females (Fig. 5); correlated with this the venulae in both sexes are reduced or vestigial; (b) the presence of a pair of narrow, oblique sclerites of abdominal sternum 2 (Figs 4-6); these could represent a vestige of an inner arm of a bifid venula, dissociated from the normal venula and the base of the apodeme; and (c) the presence of a slender, sclerotised rod or spicule in a swollen section of the ductus ejaculatorius near the base of the male aedeagus (Fig. 14); this is usually short, elliptical in longitudinal section, occasionally very long and tortuous or rarely with one end truncate or slightly expanded. Whereas synapomorphy (a) appears throughout the genera of this group, synapomorphies (b) or (c) are absent in some genera.

A very tentative phylogenetic analysis, using the Hennig86 computer program, version 1.5, designed and written by J.S. Farris, has been attempted based on 33 characters (Tables 1, 2). Polarisation of the character states has been carried out manually, using as outgroup a hypothetical ancestor with plesiomorphic character states based on general information on the Gelechioidea and other Lepidoptera. All characters have been treated as additive. Treating particular sets of characters as additive versus non-additive (unordered) was examined; trees generated by treating all characters as non-additive contained almost no resolution and, running the analysis with subsets non-additive, provided different tree configurations, although some of the principal patterns were congruent. Because of the low number of characters and the overall lack of resolution, the option generating the most resolution was adopted and, in the analysis presented, all characters have been treated as additive.

Fig. 1 Consensus cladogram of the Wingia group of genera.

The complete dataset contains 91 taxa and 34 characters, one of which is scored 0 for all taxa in order to permit ready interpretation of the Hennig86 results. The command m*;bb*; was used; m* constructs an initial cladogram and extended branch-swapping bb* was undertaken generating as many trees as will fit the memory available. This analysis yielded 695 trees (with overflow) with length 685, a consistency index of 0.13 and a retention index of 0.97. From this the nelsen option was used to generate the consensus cladogram shown in Fig. 1. The characters (Tables 3 and 4) supporting the nodes and taxa in the cladogram were listed by loading the nelsen-tree into PAUP Version 3.1.1, and generating lists of apomorphies using command describel/apolist.

The tree indicates that the Wingia group of genera can be treated in 12 subgroups, although subgroups 1 and 3 consist of clusters of genera not included in any of the other monophyletic subgroups. Nine genera in subgroups 1 and 2 form the sister-group of the whole of the remaining genera, with a sister-group, subgroup 2, of two genera, Crepidosceles and Lepidotarsa arising at node 2. Node 3 gives rise to the bulk of the Wingia group comprising subgroups 3 to 12. Of these, there is little doubt that subgroup 3, and possibly 4 and 5, are well supported and monophyletic, but anomalies arise in the groupings of some genera in the remaining subgroups. Comments will be made about these problems in the appropriate sections to follow. From Table 4 it will be seen that the genera Scotodryas, Heteroptolis and Wingia are not supported by unique autapomorphies in their current positions in the cladogram.

Table 1 Characters and character states, Oecophorinae, Wingia Group

Table 2 Character matrix, Oecophorinae, Wingia Group

Table 3 List of characters and character states supporting the nodes in the cladogram shown in Fig. 1

Table 4 List of characters and character states supporting each genus in the cladogram in Fig. 1

Chapter 2

Morphology

Head (Fig. 2).

The cranium or head capsule of Oecophorinae, and indeed of Gelechioidea, lacks clearly defined sutures, although there is usually a weak occipital suture and a scale-free line representing the transfrontal suture between the bases of the antennae. These separate the scale-fields of the vertex and occiput on the one hand, and the vertex and frontoclypeus on the other. Except for these scale-free zones, and narrow scale-free areas adjacent to the antennal-bases and the eyes, the scale sockets of the cranium are more or less evenly distributed. Ocelli and chaetosemata are absent.

The compound eyes of the Oecophorinae lack interfacetal hairs and are either nearly circular, or oval with a vertical long axis; a few have the margin adjacent to the antennal base slightly flattened, but not emarginate. The size of the eyes varies considerably, as indicated by the interocular index. Large eyes with an interocular index exceeding 1.46 frequently occur in rainforest species and others which are normally active at night, whereas the smallest eyes, with an interocular index of less than 1.10, are characteristic of species which are normally active during the day. Some of the latter, such as Orthiastis (Fig. 68), frequent areas of heath and shrubs at the higher altitudes, especially in the south-east and in Tasmania, but some such as Deigmoesta (Fig. 423) occur in heathlands at lower altitudes.

The vestiture of the cranium in Oecophorinae includes a narrow area of long, slender scales of the occiput, with rounded or bifid apices, which usually curve posteriorly, a pair of dense lateral groups of similar scales, called side-tufts, above the eyes and curving posteriorly, mesally and anteriorly, and the central area of the vertex. The scales of the central part of the vertex are in two layers, a lower sometimes sparse layer of small lamellar scales directed anteriorly, and an upper layer of much longer lamellar scales, which are also directed anteriorly and usually project between the bases of the antennae. The long scales of the vertex may be appressed, loosely arranged or roughened. They may be partly or even completely obscured by the scales of the side-tufts.

Fig. 2 Oecophorinae, Wingia lambertella (Wing), frontal view, head terminology.

The vestiture of the frontoclypeus (frons) includes a central area, in which the scales are in two layers, and narrow lateral areas. The scales of the lower layer are small and fairly broad, while the scales of the upper layer are longer and usually broader; the scales of both layers may be appressed or the upper layer may be arranged more or less loosely or rarely erect; both layers are directed ventrally. The scales of the lateral areas are usually narrower than the median scales and are usually directed mesoventrally. The scale vestiture of the frontoclypeus is overhung above by projecting scales of the vertex and side-tufts.

The mouthparts of Oecophorinae are usually well developed, although reduction or loss of the galea has occurred in a few genera. Pilifers are always present. The maxillary palpi have four short segments, and are usually folded over the base of the proboscis. The terminal segment is the longest and is the only one bearing scales. Although the scales are nearly always small and appressed, they are much longer and loosely arranged in a few genera, especially Elaphromorpha (Fig. 132). The galeae are usually well developed, with the two elements hooked together to form a functional proboscis. In repose the proboscis is coiled. Occasionally it is reduced in length and is not coiled (e.g. Coeranica), and rarely it is vestigial or absent (e.g. Limothnes).

The labial palpi of Oecophorinae are three-segmented and are usually fairly long or long, recurved and sickle-shaped. The basal segment is short and the second is usually the longest, in length frequently exceeding the height of the eye and extending to or beyond the base of the antenna. Occasionally the second segment is shorter and fails to reach the base of the antenna. In a few genera it is porrect or rises obliquely, but the apical segment is then seldom porrect, but slightly recurved or vertical (e.g. Deigmoesta, Fig. 424). The second segment is usually fairly slender, but is occasionally stout (e.g. Trachyzancla, Fig. 418). The third segment is nearly always slender and shorter than the second, and the apex is acute; vom Rath’s organ has not been recognised.

The scaling of the labial palpi provides useful taxonomic criteria. A clothing of appressed, overlapping scales appears to be the plesiomorphic condition, with the anterior scales being sometimes slightly roughened. Modification of the scaling, particularly of the second segment, has occurred in many genera. Distally the anterior scales may be lengthened and slightly expanded (e.g. Ocystola, Fig. 664), the distal posterior and anterior scales may be lengthened and more or less expanded, or the anterior scales may be greatly lengthened to form a strong triangular scale-tuft (e.g. Wingia, Fig. 449). The apical segment usually has appressed scales, but is sometimes thickened or laterally compressed; rarely the scaling may be slightly thickened medially. In a few genera the labial palpi are sexually dimorphic. In Basiplecta, for example, they are longer in the female, with the second segment in males being 1.2x the height of the eye and extending to the base of the antenna, whereas in females it is 1.5x the height of the eye and extends beyond the base of the antenna.

The antennae in Oecophorinae vary considerably, in length ranging from less than two-thirds to more than the length of the fore wings. However, in the Wingia group of genera they are more usually between two-thirds and three-fourths the length of the fore wing. The scape is short and slender, and the pedicel is very small; they both have appressed scales. The scape often has a pecten of slender, scale-like setae: sometimes the setae are numerous and extend along most of the anteroventral side of the scape, and occasionally they are very dense. However, the number of setae may be reduced and their distribution may be restricted to the proximal half or one-third of the scape. In a few genera (e.g. Callimima, Fig. 404) the pecten is represented by only one or two setae at the base of the scape, while in many genera the scape is devoid of a pecten.

The flagellum is sexually dimorphic. Occasionally in males the flagellum is thickened with scales (e.g. Idioxantha, Fig. 621), but in most it is slender. In the male slender sensilla (ciliations) arise from each flagellomere, sometimes in small clusters on each segment. The ciliations are longest near the base of the antenna and diminish in length distally, usually to about two-thirds the length of the antenna, or even to near the apex. Near the base they vary greatly in length, ranging from less than the diameter of the flagellum to as much as twelve times the diameter (e.g. Acorotricha, Fig. 412). However, within some genera the length of the ciliations is variable. Females normally have very short antennal sensilla, an exception being Polyeucta callimorpha (Fig. 43), in which the ciliations near the base of the antenna are about twice the diameter of the flagellum. Each flagellomere has two rows of overlapping scales dorsally, but ventrally scales are absent in males. In the females of some genera the ventral surface of the flagellum is without scales, as in males, but in others two rows of ventral scales may be present from the base to near the apex of the flagellum, or the ventral scales may be restricted to a basal section of the flagellum, where only one row may be present. In the females of some genera ventral scales are lacking only in a narrow ventral strip.

Thoracic appendages

In Oecophorinae the fore legs always have a tibial epiphysis, and usually have overlapping, appressed scales. In a few genera, however, the fore tibiae, and sometimes also the basal segments of the tarsi, are dilated with scales. In these genera the fore tibiae and tarsi are pressed together and conspicuously extended in front when the insect is at