Zygaenid Moths of Australia: Revision of the Zygaenidae of Australia (Procridinae: Artonini) Monographs on Australian Lepidoptera, Volume 9 (Monographs on Australian Lepidoptera)

Zygaenid Moths of Australia A revision of the Australian Zygaenidae (Procridinae: Artonini) Monographs on Australian ...

Author: Gerhard M. Tarmann

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Zygaenid Moths of Australia A revision of the Australian Zygaenidae (Procridinae: Artonini)

Monographs on Australian Lepidoptera Editorial Board Editor-in-Chief M. Horak, Canberra, Australia

Editorial Panel I. F. B. Common, Toowoomba, Australia E. D. Edwards, Canberra, Australia R. L. Kitching, Brisbane, Australia S. E. Miller, Washington DC, USA G. S. Robinson, London, UK M. J. Scoble, London, UK M. S. Upton, Canberra, Australia

Hestiochora xanthocoma, male Drawing by R. Nielsen

M O N O G R A P H S O N AU S T R A L I A N L E P I D O P T E R A V O L U M E 9

Zygaenid Moths of Australia

A revision of the Australian Zygaenidae (Procridinae: Artonini)

Gerhard M. Tarmann ˇ

W I T H C O LO U R PA I N T I N G S B Y F R A N T I S E K G R E G O R

IV

ZYGAENID MOTHS OF AUSTRALIA

© Gerhard Tarmann 2004 All rights reserved. Except under the conditions described in the Australian Copyright Act 1968 and subsequent amendments, no part of this publication may be reproduced, stored in a retrieval system or transmitted in any form or by any means, electronic, mechanical, photocopying, recording, duplicating or otherwise, without the prior permission of the copyright owner. Contact CSIRO PUBLISHING for all permission requests. National Library of Australia Cataloguing-in-Publication entry Tarmann, Gerhard M. Zygaenid moths of Australia: A revision of the Australian Zygaenidae (Procridinae: Artonini). Includes index. ISBN 0 643 06798 1 (Hardback). ISBN 0 643 09219 6 (netLibrary eBook). 1. Zygaenidae – Australia. 2. Zygaenidae – Australia – Morphology. 3. Lepidoptera - Australia. I. Gregor, Frantiˇsek. II. Title. (Series: Monographs on Australian Lepidoptera; v. 9). 595.780994 Available from CSIRO PUBLISHING 150 Oxford Street (PO Box 1139) Collingwood VIC 3066 Australia Telephone: Local call: Fax: Email: Web site:

+61 3 9662 7666 1300 788 000 (Australia only) +61 3 9662 7555 [email protected] www.publish.csiro.au

Set in 9.5/11 Minion Cover and text design by James Kelly Typeset by James Kelly Printed in Australia by BPA Print Group 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 5, 1997 Oecophorine Genera of Australia II The Chezala, Philobota and Eulechria groups (Lepidoptera: Oecophoridae) I.F.B. Common

Volume 2, 1993 Tineid Genera of Australia (Lepidoptera) G.S. Robinson and E.S. Nielsen

Volume 6, 1999 Biology of Australian Butterflies R.L. Kitching, E. Scheermeyer, R.E. Jones and N.E. Pierce (Editors)

Volume 3, 1994 Oecophorine Genera of Australia. I. The Wingia Group (Lepidoptera: Oecophoridae) I.F.B. Common Volume 4, 1996 Checklist of the Lepidoptera of Australia E.S. Nielsen, E.D. Edwards and T.V. Rangsi (Editors)

Volume 7, 1999 Heliothine Moths of Australia A Guide to Pest Bollworms and Related Noctuid Groups M. Matthews Volume 8, 2000 Oecophorine Genera of Australia III The Barea Group and Unplaced Genera (Lepidoptera: Oecophoridae I.F.B. Common

Contents

Abstract

1

Introduction

3

Material and methods

5 5 5 5

Material studied Treatment of material Depositories and material examined

Acknowledgements Zygaenid morphology Imago Egg Larva Pupa and cocoon

Biology and life history Phenology Larval host-plants Parasitoids Cyanogenesis and resistance to hydrocyanic acid (HCN) Defensive biology Ecology, dispersal, diversity and distribution Pheromones and mating behaviour Genetics and individual variation Zygaenids as indicator species Pest zygaenids Conservation Fossil record

Phylogeny PAUP analysis

7 9 9 36 36 39 41 41 41 42 42 43 44 45 46 46 46 47 47 49 49

A possible history of Australian Zygaenidae

57

Checklist of Australian Zygaenidae

59

VI


Taxonomy Family Zygaenidae Latreille Subfamily Procridinae Boisduval Tribe Artonini Tarmann Key to Australian genera Genus Pollanisus Walker Key to Pollanisus species P. viridipulverulenta (Guérin-Méneville) P. cupreus Walker P. nielseni sp. n. P. modestus sp. n. P. lithopastus Turner Pollanisus sp. 1 P. empyrea (Meyrick) P. apicalis (Walker) P. trimacula (Walker) P. edwardsi sp. n. P. subdolosa (Walker) P. subdolosa subdolosa (Walker) P. subdolosa clara ssp. n. Male specimens probably conspecific with P. subdolosa Populations close to, but doubtfully conspecific with, P. subdolosa P. contrastus sp. n. Pollanisus cf. contrastus 1 Pollanisus cf. contrastus 2 Possible P. edwardsi, P. subdolosa or P. contrastus females P. eungellae sp. n. P. acharon (Fabricius) P. eumetopus Turner P. commoni sp. n. P. incertus sp. n. P. angustifrons sp. n. Pollanisus sp. 2 Pollanisus sp. 3 Pollanisus sp. 4 Pollanisus sp. 5 Pollanisus sp. 6 Pollanisus sp. 7 Pollanisus sp. 8 P. cyanota (Meyrick) P. isolatus sp. n. P. calliceros (Turner) P. calliceros calliceros (Turner) P. calliceros azurea ssp. n. Genus Onceropyga Turner Key to Onceropyga species O. anelia Turner O. pulchra sp. n.

61 61 61 61 61 63 65 67 72 75 76 77 79 79 81 84 86 87 87 89 91 91 91 93 93 93 94 95 97 98 99 100 102 102 102 102 103 103 104 104 105 106 106 107 133 134 134 135

CONTENTS

Genus Hestiochora Meyrick Key to Hestiochora species ‘Hestiochora tricolor auctt.’ – a complicated species-complex The ‘Hestiochora tricolor habitus’ – external appearance wasp-like Hestiochora erythrota-tricolor-group H. erythrota Meyrick H. tricolor (Walker) H. continentalis sp. n. H. furcata sp. n. Hestiochora queenslandensis-group H. queenslandensis sp. n. Hestiochora xanthocoma-occidentalis-group H. xanthocoma Meyrick H. occidentalis sp. n. H. intermixta sp. n. Genus Turneriprocris Bryk T. dolens (Walker) Genus Myrtartona gen. n. Key to Myrtartona species M. coronias (Meyrick) comb. n. M. leucopleura (Meyrick) comb. n. M. rufiventris (Walker) comb. n. M. mariannae sp. n. Genus Australartona gen. n. A. mirabilis sp. n. Genus Homophylotis Turner Key to Homophylotis species H. thyridota Turner H. pseudothyridota sp. n. H. artonoides sp. n. Genus Pseudoamuria gen. n. Key to Pseudoamuria species P. uptoni sp. n. P. neglecta sp. n. Genus Palmartona gen. n. P. catoxantha (Hampson) comb. n. Genus Thyrassia Butler T. inconcinna Swinhoe

139 141 141 143 143 143 144 145 148 149 149 150 150 151 152 163 164 169 170 170 172 174 176 185 186 191 192 192 194 195 201 202 202 203 207 208 215 216

Appendix I: List of localities

221

Appendix II: CryoSEM figures of eggs and first instar larvae of Pollanisus subdolosa clara ssp. n.

227

References

231

Index

243

VII

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Abstract The Zygaenidae, commonly known as burnets and foresters, are a group of aposematic moths that are usually diurnal. They have long attracted attention not only for their often beautiful appearance but also for their unusual biology. Zygaenids are capable of synthesising cyanogenic glucosides and releasing hydrocyanic acid. They can also detoxify hydrogen cyanide and are highly resistant to cyanides. Knowledge about the Australian zygaenid fauna has been very fragmentary until now, based on the original descriptions only. Together with an indepth revision of the Australian taxa, this study presents a comprehensive overview of the morphology and biology of the Zygaenidae, placing the Australian fauna into a world context and discussing the differences between subfamilies and genus groups. Of the four subfamilies currently recognised (i.e. Zygaeninae, Chalcosiinae, Callizygaeninae and Procridinae), only the Procridinae have a world-wide distribution. All Australian zygaenids belong to this subfamily which is divided into the two tribes Procridini and Artonini. All Australian species, 43 species in 10 genera, belong to the Artonini, and the monophyly of this tribe is discussed. Current knowledge of the Australian zygaenids, all foresters, is summarized and discussed. Nearly half the fauna, four genera, 21 species and two subspecies, are described as new and eight others recognised. All recognised species are endemic to Australia except Palmartona catoxantha (Hampson, [1893]), a Southeast Asian pest species on coconut palms not native to Australia but either already present or a very likely future incursion.

All species are described or redescribed, a diagnosis is given and the biology and distribution are summarised. The adults of all species are figured in colour, mostly upper as well as under side, from ˇ Gregor (Brno, Czech colour paintings by Frantisek Republic). Many zygaenid groups such as Pollanisus are taxonomically difficult, with few and unreliable diagnostic characters in genitalic structures. The fine details of colouration, even the brilliance of the shiny scales, are often the only means to separate species. The exquisite paintings, detailed portraits originally created at a ten-fold magnification of the moth, are crucial as they depict these subtle diagnostic details. These originals are reduced slightly in this book (see magnification of original size of moth given after the species name). Taxa recognised as possible distinct species but not described for lack of sufficient material have also been illustrated, anticipating that some will be named at a later date and a comparable illustration will then become important. Details of the morphology of adults and early stages are figured by SEM pictures. Keys to genera and species are provided and the genitalia of all species figured. A selection of colour photos illustrates biology and habitats. A distribution map for each species is given and the original data are listed, with an alphabetic list of all known Australian zygaenid localities. During field work, a conscious effort was made to find eggs and larvae of as many species as possible, resulting in the inclusion of life history information for five genera, namely Pollanisus, Hestiochora, Turneriprocris, Myrtartona gen. n., and Australartona gen. n.


Introduction The Zygaenidae are a family of small to large, mainly diurnal moths with a very variable habitus, biology and ecology. More than 1000 species are known world-wide and many more remain to be described. Four subfamilies are recognised: Zygaeninae, Chalcosiinae, Procridinae and Callizygaeninae (Epstein et al. 1998). The Zygaeninae are restricted to the Palaearctis, northwestern Oriental, and the Afrotropical regions. The Chalcosiinae (with the exception of the western Palaearctic genus Aglaope Latreille, 1809) occur in the eastern Palaearctic and Indo-Australian regions, including New Guinea and smaller islands eastwards to Fiji, but surprisingly do not reach Australia. The Procridinae have a world-wide distribution and are the only zygaenids represented in the Americas and Australia. Callizygaeninae are only known from south and Southeast Asia. Well-defined autapomorphies are known for most of the subfamilies, and there are significant differences in their appearance. Zygaeninae, Chalcosiinae and Procridinae seem to form a monophyletic group, whereas the taxonomic position of the Callizygaeninae is still uncertain (Yen 2003). Zygaenids can be recognised by their characteristic resting position with the wings laid back rooflike over the abdomen. When in this position, the wings are attached to the dorsal part of the metathorax by hook-like microtrichia. The antennae point forward and upward, with their distal ends slightly upturned or curved back. Their flight is often weak and linear but in the tropics some species fly rapidly, high over the forest canopy. As in all more primitive Lepidoptera, the wing-beat frequency is constant during flight. The larva is stout, with verrucae and setae. The head-capsule is retractile into the first thoracic segment (as in all Zygaenoidea). Representatives of the Zygaeninae, Chalcosiinae and Procridinae are cyanogenic, synthesizing the cyanoglucosides linamarin and lotaustralin in their body tissues. They are able to release hydrocyanic acid (HCN) by enzymatic breakdown of these two cyanoglucosides, and can also detoxify hydrocyanide using the enzyme β-cyanoalanine-synthetase (Witthohn and Naumann 1984b). In larvae of the Zygaeninae, two

types of cuticular cavities are present in the larval cuticle on most segments (Franzl 1980; Povolný and Weyda 1981; Franzl and Naumann 1984, 1985; Naumann and Povolný 1987; Fänger and Naumann 2001; Yen 2003); these cavities store a highly viscous defensive secretion consisting mainly of linamarin, lotaustralin and a large proteinaceous fraction (Witthohn and Naumann 1984a, 1984b). In the Chalcosiinae, only one type of cuticular cavity has been found (Fänger and Naumann 2001; Yen 2003). So far, no cuticular cavities have been found in the Procridinae. The pupa is stout and sometimes dorsoventrally flattened; the cuticle is moderately sclerotized, and the appendages are only weakly fused to each other and to the body. Abdominal segments 3–8 (male) or 3–7 (female) have a single row of dorsal spines; the cremaster is vestigial. Pupation takes place in weak, silken cocoons or in firm, parchment-like cocoons spun on leaves or stems, under rocks, in litter on the surface of the earth, or, rarely, in the stems of the host-plant. The wall of the cocoon is impregnated with microscopic whevellite crystals (calcium oxalate monohydrate) formed in the Malpighian tubules of the larva (Hepp 1920, 1922; Naumann 1977a; Hofmann 1994; Epstein et al. 1998; Naumann, Tarmann and Tremewan 1999; Yen 2003). All Australian zygaenids belong to the tribe Artonini of the Procridinae. The subfamily Procridinae with the two tribes Procridini and Artonini has an almost world-wide distribution but the Artonini are restricted to the Afrotropical and Indo-Australian regions (although a few species also occur in the eastern Palaearctis). The monophyly of the Artonini is well supported by the following three autapomorphies (Tarmann 1994): a chaetosema extending forward between the compound eye and the ocellus; small dorsolateral evaginations on abdominal segments 2 and 7 (not only in last instar larvae (as in most Procridinae) but also in the adults); and a fan-shaped valva. The family is represented in Australia by 43 species in 10 genera. Four genera, 21 species and two subspecies are described as new to science in this work. Many more species seem to exist, especially in the tropical parts of the continent. Eight of them are illustrated and discussed but not named

4


and formally described because more material and information is required. Alberti (1954: 296) recognised a close relationship between most Australian zygaenids and the Asian Artona-group (Balataea-group sensu Alberti) of the tribe Procridini. He referred the genus Hestiochora Meyrick, 1886, to the Callizygaenini that, in his opinion, was a tribe of the subfamily Procridinae. Tarmann (1994: 118) showed that the former Callizygaenini (sensu Alberti) lack the autapomorphies of the Procridinae and have to be excluded from that subfamily. They represent a distinct zygaenid subfamily. According to recent investigations by Yen (2003) they are closer to the Chalcosiinae than to the Procridinae. The Australian genus Hestiochora Meyrick, 1886, actually belongs to the procridine tribe Artonini and is closely related to the Australian genera Pollanisus Walker, 1854, and Onceropyga Turner, 1906. This is supported by a striking synapomorphy: the females of all species of the three above-mentioned genera have a characteristic hair-tuft at the end of the abdomen. This is combined with the lack of the pair of glands close to the oopore. These glands occur in all other known Procridinae and Zygaeninae. They appear to produce a poisonous liquid to protect the eggs from parasitoids and are described by Bode and Naumann (1987) as ‘Petersen’s glands’ in Zygaeninae. The females of these three Australian genera protect their eggs by touching them with their abdominal hair-tufts after oviposition and attaching long spiny scales to their surfaces, giving them a pincushion-like appearance. The tips of these scales (formerly being the bases that stuck in the abdominal skin of the female) seem to be poisonous: aphids die after having made contact

with them (Tarmann 1994: 118). Male genitalic characters indicate that most of the Australian zygaenids form a distinct group of Procridinae with an artonoid-type valva but without the autapomorphies of the Asian Artona-group and related genera. This suggests that most Australian zygaenids must have separated from the Asian group before the main autapomorphy of the Asian species developed (i.e. the development of a ventral ‘Artona-finger’ at the distal end of the valva). Only the genera Homophylotis Turner, 1904, Pseudoamuria gen. n., Australartona gen. n., and Palmartona gen. n. belong to that Asian group. Homophylotis is restricted to the tropical parts of northern Queensland (three species) and the Bismarck Archipelago (one species). Pseudoamuria gen. n., with two species in Australia, occurs in Queensland and is fairly closely related to the Asian Artona Walker, 1854, and Amuria Staudinger, 1887. Several species from New Guinea and surrounding islands also belong to this genus, e.g ‘Homophylotis’ melaleuca Jordan, 1908. Australartona gen. n., with the single included species A. mirabilis sp. n., is an interesting link between several Australian genera and the Asian genera close to Artona. It is distributed in New South Wales and southern Queensland. Palmartona gen. n., with only one species, P. catoxantha (Hampson, [1893]), is probably an introduction; outside Australia it is distributed from Myanmar (Burma) to New Guinea. Most Australian zygaenids are considered here to represent primitive Artonini displaying many plesiomorphies; however, certain subgroups that have evolved independently from the rest of the tribe display clear autapomorphies (see p. 57).

Material and methods Material studied In preparing this revision of the Australian zygaenids I have attempted to include all specimens available in collections. Most of the material is deposited in Australian museums and institutes and in The Natural History Museum in London. Field observations were made in November 1989 by G. M. Tarmann (assisted at times by the late E. S. Nielsen and by M. Shaffer), in September 1995 by G. M. and B. E. Tarmann, and in November 2001 by G. M. Tarmann and E. D. Edwards. N. McFarland made available a copy of his diary containing invaluable, mainly unpublished, field observations. A total of 3159 specimens have been examined during this study. Under the ‘Material examined’sections for each species, the localities are listed alphabetically within each state. The latitude and longitude of all localities is given in Appendix I.

Treatment of material At least one male and one female of each species, if ˇ Gregor in available, have been painted by Frantisek Brno, Czech Republic. Most figures are an abstraction and compiled from more than one specimen to show the most characteristic features of the population painted. Therefore, only superficial locality data and no collection data are provided (e.g. SA, Aldinga Scrub). The original paintings were 10× the size of the specimens. They have been reduced according to the breadth of the page of the book, but the same scale has been kept on each plate in a genus to avoid misinterpretations. The magnification in relation to the size of the original specimen is specified on each plate. Most species are illustrated with the upperside of the body and wings (right) and the underside of the wings (left); the underside of the body is also illustrated for some species. The genitalia were dissected by removing the whole abdomen and opening it laterally after maceration in potash. Abdomen and male genitalia were stained with Mercurochrome; female genitalia and the everted vesica of the male aedeagus were stained with Chlorazol Black. Sometimes, the entire male genitalia and abdomen were stained with Chlorazol Black after a first staining with Mercurochrome, especially in small specimens of

the genus Pollanisus, which have very translucent valva and abdomen. This double staining technique increases the contrast and leads to enhanced photographs. The genitalia and abdominal skins were mounted in Euparal on slides. In some groups, e.g. Hestiochora and Pollanisus, it was necessary to dissect parts of the male genitalia in order to see the characters satisfactorily (see p. 108–116, 154–157). Some genitalia were also mounted in their natural position to give a better impression of the function of the different parts. Parts of the head, body, and wings were prepared for SEM photomicrography. Small pieces were cut from the right forewing and hindwing and mounted on aluminium stubs with hot wax. The head, antennae, legs and abdomen were detached from the specimen and mounted on small cardboard points with conductive silver paint (silver particles suspended in methyl-iso-butyl-ketone). The mounted material was then gold-coated in the usual way. The SEM at the CSIRO (Canberra) was used for specimen examination and photography. The photos of the genitalia and abdomina were made in the Tiroler Landesmuseum Ferdinandeum (Innsbruck) using an Olympus SHZ Microscope with various filters, an Olympus OM2 camera and AGFA ORTHO film, and at the CSIRO Black Mt Microscopy Centre (Canberra) with a Leitz Diaplan Microscope and a ProgRes 3012 digital camera, using Adobe Photoshop 5.5 (later 6.0) and compiled with AutoMontage programme on computer. Eggs, larvae and pupae were preserved in 70% ethanol, the larvae first being fixed in KAA (1 part kerosene, 2 parts glacial acetic acid, 10 parts absolute ethylalcohol) for several days. CryoSEM Figs 436–448 were taken of non-fixed specimens using a Cambridge S360 at the ANU Electron Microscopy Unit.

Depositories and material examined Below is a list of acronyms for the depositories in which the material examined is preserved. The data recorded for each species under ‘Material examined’ have been rationalised. The main locality is mentioned first, e.g. ‘Paluma, 2 mls W’ instead of ‘2 mls W of Paluma’ as written on the original label.

6


Only for type-material has the text of the original label been retained for most localities. Even if mentioned on the labels, latitude and longitude are not included under ‘Material examined’, except for type-material. Localities within states are given in alphabetical order, and Appendix I provides an alphabetical list of all localities with corresponding latitudes and longitudes (as used for the distribution maps). Collectors’ names are italicised and placed in parentheses. The acronyms of depositories are given in parentheses.

NHRM NMW NSWA OUM QM SAM TLMF

AMS ANIC BCRI BMNH BPBM DPIQ MV

Australian Museum, Sydney, Australia Australian National Insect Collection, CSIRO Entomology, Canberra, Australia Biological and Chemical Research Institute, Rydalmere, NSW, Australia The Natural History Museum, London, UK Bernice P. Bishop Museum, Honolulu, USA Department of Primary Industries, Queensland, Brisbane, Australia Museum Victoria, Melbourne, Australia

TMH UQIC WADA WAM ZMB

Naturhistoriska Riksmuseet, Stockholm, Sweden Naturhistorisches Museum Wien, Vienna, Austria New South Wales Department of Agriculture, Orange, Australia Oxford University Museum, Oxford, UK Queensland Museum, Brisbane, Queensland, Australia South Australian Museum, Adelaide, Australia Tiroler Landesmuseum Ferdinandeum, Innsbruck, Austria Tasmanian Museum and Art Gallery, Hobart, Tasmania, Australia University of Queensland, Insect Collection, Brisbane, Australia Western Australian Department of Agriculture, South Perth, Australia Western Australian Museum, Perth, Australia Zoologisches Museum der HumboldtUniversität, Berlin, Germany

Acknowledgements I acknowledge with thanks the financial support received from the Australian Biological Resources Study (ABRS), CSIRO, and the Österreichische Akademie der Wissenschaft/Royal Society exchange programme that enabled me to visit Australia twice and to study type-material at The Natural History Museum, London. For the loan of material I am indebted to the following people: Dr G. R. Brown (BCRI, NSW Department of Agriculture, Rydalmere), Dr B. K. Cantrell (DPI, Indooroopilly), Dr E. C. Dahms (Queensland Museum, Brisbane), the late W. Dierl (Zoologische Staatssammlung München, Munich), Ms A. J. A. Green (Tasmanian Museum, Hobart), Dr G. F. Gross (South Australian Museum, Adelaide), Mr M. R. Honey (The Natural History Museum, London), Dr T. F. Houston (WA Museum, Perth), Mr O. Karsholt (University Zoological Museum, Copenhagen), Dr M. Lödl (Naturhistorisches Museum Wien, Vienna), Dr D. McAlpine (Australian Museum, Sydney), Dr M. B. Malipatil (NT Museum of Arts and Sciences, Darwin), Dr. W. Mey (Zoologisches Museum der HumboldtUniversität, Berlin), Dr S. E. Miller (Bishop Museum, Honolulu), Prof. Dr C. M. Naumann (Zoologisches Forschungsinstitut und Museum Alexander Koenig, Bonn), Dr A. Neboiss (Museum Victoria, Melbourne), Mr K. T. Richards (WA Department of Agriculture, Perth), Ms M. A. Schneider (University of Queensland, St Lucia). For assistance in searching for specimens and literature, help in compiling computer distribution maps and the cladistic analyses, for identification of plants, help in the production of line drawings, photography and advice on field trips I thank the following people at CSIRO Entomology, Canberra: Mr C. Beaton, Mr M. Dominick, Ms H. Geier, Mr D. Giesemann, Mr J. Green, Mr E. Hines, Dr C. Lambkin, Ms S. Narain, Ms T. V. Rangsi, Dr S. O.

Shattuck, Ms K. Smith, Ms S. Wragg, Dr D. K. Yeates, and Mr A. Zwick; and at other institutions: Mr B. Dixon (Botanic Garden King’s Park, Perth), Ms J. Harvey (BMNH, London), Dr R. Heady (ANU Electron Microscopy Unit, Canberra), Mr S. Heim (TLMF, Innsbruck), Ms K. Keys (Botanic Garden King’s Park, Perth), Mr M. Shaffer (BMNH, London), and Mr. L. S. Willan (Sydney). Grateful thanks go to Mr Paul Reekie and his colleagues at CSIRO Publishing, in particular Ms Ann Crabb, for their advice and help for this volume. I thank my colleagues Dr I. F. B. Common (Toowoomba), Mr E. D. Edwards (CSIRO, Canberra), Prof Dr K.A. Efetov (Simferool, Crimea, Ukraine), Dr R. Hoare (Auckland, New Zealand), Dr J. D. Holloway (The Natural History Museum, London, U.K.), Dr M. Horak (CSIRO, Canberra), Dr M. Matthews (Canberra), the late E. S. Nielsen (Canberra), Dr W. G. Tremewan (Truro, UK), and Mr M. S. Upton (Canberra) for commenting on the manuscript and for checking the English text. I am especially indebted to the late Dr E. S. Nielsen (Canberra) who encouraged me to contribute to the series Monographs on Australian Lepidoptera and who channelled all his energy and influence to make this work technically and financially possible. This work would not have reached the high standard expected by CSIRO Publishing without the substantial assistance of Dr M. Horak (CSIRO, Canberra) as editor-in-chief, whose invaluable and time intensive comments to the typescript and help with the arrangement of figures, plates and captions gave the author every imaginable support. Last, but not least, I thank my friend Dr F. Gregor (Brno) for the magnificent paintings of Australian zygaenids that appear in this book, and encouraged me greatly during the preparation of this work.


Zygaenid morphology Imago Head (Figs 1–97) The head capsule of zygaenids is strongly sclerotized. The frons is projected in many groups, especially dorsally (Yen 2003: fig. 3) (e.g. Figs 2, 59). The adult mouthparts include short, but easily visible maxillary palps consisting of one or two segments (Epstein et al. 1999: 171, Yen 2003: 304) (e.g. Figs 1, 3, 15); a long and well-developed unscaled proboscis with a spiny surface is normally present (e.g. Figs 3, 25, 67 for artonine Procridinae, Figs 84–86, 75, 76, for procridine Procridinae). The proboscis is only reduced in a few groups: e.g. Pryeria Moore, 1877 in the Zygaeninae; Aglaope Latreille, 1809 and Boradia Moore, 1879 in the Chalcosiinae; and Theresimima Strand, 1917, and Rhagades Wallengren, 1863 (one of four species) in the Procridinae (Efetov and Tarmann 1995, 1999; Naumann, Tarmann and Tremewan 1999; Efetov 2001). The tip of the proboscis bears sensory hairs responsible for perceiving chemical and other stimuli during feeding (Naumann, Tarmann and Tremewan 1999: 25, text-fig. 22). The labial palps are porrect or upcurved and densely covered with scales (Figs 1, 4). Their tip bears sensory organs (Fig. 60). The ommatidia of the large compound eyes are large; inter-ommatidial setae (sensu Kristensen and Nielsen 1979: 73, fig. 4) are present in most Procridinae, but are absent in Zygaeninae (as far as examined) (Naumann, Tarmann and Tremewan 1999: 24, text-fig. 20) and Chalcosiinae (Yen 2003). Their length does not exceed the diameter of the ommatidia and they can therefore only be seen under high power magnification (e.g. Figs 18, 22, 23, 40, 53 for Artonini; Figs 90, 91 for a Procridini with inter-ommatidial setae and Figs 79, 80 for a Procridini without inter-ommatidial setae). This character is considered to be typical for some of the more primitive groups of Lepidoptera (Davis 1978: 9, figs 6, 8, 9) and has been overlooked by Yagi and Koyama (1963). In the more derived groups these interfacetal hairs (sensu Scoble 1992: 27) are long and clearly visible even under low power magnification. The diameter of the compound eye is larger in the male than in the female in most Procridinae

and Chalcosiinae, while the breadth of the frons is correspondingly smaller in the male than the female. This character is also useful for taxonomic purposes as it is very constant and there are sometimes clear differences between closely related species and species-groups in the Procridinae (Tarmann 1983, 1984; Naumann, Tarmann and Tremewan 1999) (e.g. Figs 1, 24, 29, 37, 41, 49, 57, 65). In Zygaeninae the compound eyes are usually of almost equal size in both the males and females. The inter-ocular index (sensu Davis 1975), a term also used by Kristensen and Nielsen (1979), is not used here. It is considered sufficient to estimate the relative breadth of the frons with respect to the compound eyes in frontal view (Fig. 1). For an accurate measurement for any index it would be necessary to remove the scales from parts of the head; this was avoided because of the rarity of much of the material. All zygaenid moths have well-developed ocelli and chaetosemata on the vertex of the adult head. The ocelli may be either small or large, and either pigmented or white (e.g. Figs 2, 6, 14, 17, 26, 33, 52). The chaetosemata are large and fused mediodorsally in Zygaeninae and Chalcosiinae (Yen 2003), but separated into two triangular parts in Procridinae. Jordan (1923) found two major types of chaetosemata in Lepidoptera, the first consisting of long upright sensilla trichodea arising from swollen plates dorso-posterior to the compound eye, the second with many additional erect specialised scales sitting together with sensilla trichodea on the ‘plates’. In Zygaenidae the second type is present (Yen 2003). The function of the chaetosemata is still unknown. In Procridinae the shape of the chaetosema is a character useful for separating the tribes Procridini and Artonini (Tarmann 1994: 120). For comparison see the two Palaearctic species Illiberis (Primilliberis) pruni Dyar, 1905 (Figs 77, 78), and Adscita (Adscita) statices (Linnaeus, 1758) (Figs 87, 88) for Procridini and the Australian species (e.g. Figs 2, 6, 14, 52, 61) for Artonini. Between the inner margin of the compound eye and the base of the proboscis there is an opening that allows zygaenid moths to secrete a vitreous white or yellow liquid or foam when disturbed (Epstein et al. 1999).

10


Zygaenid antennae display considerable variation between subfamilies and even between genera; moreover, specific characters occur in the Procridinae. Most zygaenid antennae, if bipectinate or biserrate, are densely covered with sensory hairs, the latter being generally longer in the male (e.g. Figs 48, 56). A pecten is absent. In most Zygaeninae the antennae are long, clavate and without any pectinations and serrations. The distal segments form a club with the tip either pointed or rounded. Only in some primitive Zygaeninae from Asia (Pryeria Moore, 1877) and the Afrotropical region (Orna Kirby, 1892) are the antennae bipectinate. The antennae of Chalcosiinae are usually strongly bipectinate to the apex in both males and females; the pectinations are much longer in the males. The distal segments are often very close together, forming a club. Some Chalcosiinae also have biserrate or simple, slightly clavate antennae. The Procridinae display all antennal types, but in most groups the male antenna is bipectinate and the female antenna biserrate, the apex is generally pointed or rounded, sometimes clubbed. In Zygaeninae the surface is spiny and densely covered with various types of short sensory hairs, many of which are specialised to perceive volatile stimuli, including those produced by sex pheromones and various compounds of flower scents (Naumann, Tarmann and Tremewan 1999: 38, fig. 40). In most Chalcosiinae the antennal sensory setae are very short and are generally only distributed on the rami (Yen 2003). In Procridinae the surfaces of both the antennal shaft and pectinations are built of irregular crests and a series of holes where the scales and sensory hairs are attached. There are significantly more of these hairs at the tips of the antennae. Only sensilla trichodea (sensu Kristensen and Nielsen 1979: 79, fig. 11) have been found so far (e.g. Figs 11, 28, 55, 56). For comparison with the antennae of Australian species, all Artonini, those of the two Palaearctic Procridini species Illiberis (Primilliberis) pruni Dyar, 1905 (Figs 81–83) and Adscita (Adscita) statices (Linnaeus, 1758) (Figs 92–97) are also figured. Thorax and wings (Figs 98–123) The prothorax is very short in all zygaenids. Its dorsal part forms the ‘patagia’ and ‘parapatagia’ and is often covered with coloured scales of taxonomic significance. The meso- and metathorax are large. The dorsolateral ‘tegulae’ on the mesothorax are often covered with coloured scales that are taxonomically important (Naumann, Tarmann and

Tremewan 1999: 26, text-fig. 23). The meso- and metathorax bear the wings and, as most zygaenids are strong fliers, the muscles needed for the movement of the wings are strongly developed. The wings are synchronized by a retinaculum on the underside of the forewing and a frenulum on the anterior margin of the hindwing basally. The retinaculum is a strong hook on the base of Sc in the male, and a row of upright, forward-pointing scales on the base of CuP in the female. The male frenulum consists of a strong, single spine, whereas the female frenulum consists of 1–5 smaller bristles, usually attached to the hindwing in a single row basally on the anterior margin (Naumann, Tarmann and Tremewan 1999: 26, text-fig. 24). These characters are often the only way to distinguish the sex of poorly preserved specimens, when the abdomen and head are missing. At rest, the wings are locked to the body by the interaction of a patch of small microtrichial hooks at the base of the underside of the forewing between CuP and 1A+2A, and a similar patch on the metathorax, dorsolaterally on both sides of the metascutum. The discovery of this mechanism is referred to Schneider (1878: 33) by various authors but, as Sattler (1991: 1) correctly remarks, Schneider did not describe a thorax-wing coupling mechanism but a mechanism fixing the fore- and hindwing together. It was Spuler (1895: 537) who first mentioned and described the structure later called a ‘Haftfeld’, a term also used in recent English literature (Scoble 1992: 62, figs 71–74). Detailed descriptions of the abovementioned structure were provided by Common (1969) and Kuijten (1974). The zygaenid thorax lacks sense organs such as tympanal organs and specialised eversible scent brushes, or coremata (Naumann, Tarmann and Tremewan 1999: 28). A striking trait of the Chalcosiinae is the capability of releasing a large amount of a fluid or foam (often with a hissing sound) containing cyanide from the membrane between the patagia and parapatagia on the prothorax (Yen 2003: 307). The shape of the wings is very variable in the different subfamilies and groups of zygaenids. Almost all Zygaeninae have elongate and rather slender wings, whereas the Chalcosiinae display nearly all wing-shapes known in the Lepidoptera; some are butterfly-like, some like skippers, some like tiger-moths, some resemble geometrids, and even tailed species occur. They seem to be involved in mimicry complexes but almost nothing is known about the possible interactions between these zygaenids and other insects.

ZYGAENID MORPHOLOGY

1

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3

4

5

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Figs 1–6. Head of Pollanisus viridipulverulenta, male. (C. Beaton.) 1: Frontal view: bce, breadth of compound eye in frontal view, bfr, breadth of frons; ce, compound eye; fr, frons; pb, proboscis; 2: lateral view: as, antennal shaft; ce, compound eye; ch, chaetosema; fr, frons; lp, labial palps; oc, ocellus; pc, pectinations; 3: frontal view, detail: el, eyelash; lp, labial palps; mp, maxillary palps; pb, proboscis; 4: lateral view, detail: ce, compound eye; el, eyelash; lp, labial palps; pb, proboscis; 5: lateral view, detail, eyelash; 6: lateral view, detail, chaetosema.

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Figs 7–11. Male antenna of Pollanisus viridipulverulenta. (C. Beaton.) 7: Entire view; 8–9: detail; 10: detail, distal segments; 11: detail, surface of antennal shaft. Figs 12–13. Head of Pollanisus viridipulverulenta, female. (C. Beaton.) 12: Lateral view; 13: antenna.

ZYGAENID MORPHOLOGY

14

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Figs 14–18. Head of Pollanisus calliceros, male. (E. Hines.) 14: Lateral view; 15: detail, maxillary palp; 16: detail, eyelash; 17: detail, chaetosema; 18: detail of compound eye: ommatidia, inter-ommatidial seta, ios.

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Figs 19–21. Male antenna of Pollanisus calliceros. (E. Hines.) 19: Entire view; 20–21: detail. Figs 22–23. Head of Onceropyga anelia, male, detail of compound eye. (E. Hines.) 22: Edge of chaetosema and ‘hairy’ surface of head at base of antenna; 23: inter-ommatidial seta. (E. Hines.)

ZYGAENID MORPHOLOGY

24

25

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Figs 24–28. Head of Hestiochora erythrota, male. (E. Hines.) 24: Frontal view; 25: detail, base of proboscis; 26: lateral detail; 27: detail, eyelash; 28: detail, antennal shaft.

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Fig 29–36. Head of Turneriprocris dolens, male. (C. Beaton.) 29: Frontal view; 30: frontal view, detail; 31: detail: clypeus, pilifer, maxillary palp, base of proboscis; 32: lateral view; 33: lateral view, detail, chaetosema; 34: lateral view, detail, antennal base; 35: antenna; 36: antenna, detail.

ZYGAENID MORPHOLOGY

37

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100 µm

Figs 37–40. Head of Myrtartona leucopleura, male.(C. Beaton.) 37: frontal view; 38: lateral view.; 39: lateral view, detail; 40: lateral view, detail of compound eye with inter-ommatidial setae. Figs 41–44. Head of Australartona mirabilis, male. (C. Beaton.) 41: Frontal view; 42: frontal view, detail of mouthparts; 43: lateral view; 44: lateral view, detail.

17

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45

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Figs 45–48. Male antenna of Australartona mirabilis. (C. Beaton.) 45: Distal part; 46: detail, distal end; 47: detail, pectinations; 48: detail, setae on pectinations. Figs 49–50. Head of Homophylotis thyridota, male. (E. Hines.) 49: Frontal view; 50: detail frontal view, mouthparts;

ZYGAENID MORPHOLOGY

51

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Figs 51–53. Head of Homophylotis thyridota, male. (E. Hines.) 51: lateral view; 52: lateral view, detail; 53: detail, ommatidia of compound eye with inter-ommatidial seta. Figs 54–56. Male antenna of Homophylotis thyridota. (E. Hines.) 54: Distal end; 55: detail, distal end; 56: detail, antennal pectination.

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Figs 57–61. Head of Pseudoamuria uptoni, male. (E. Hines.) 57: Frontal view; 58: frontal view, detail, mouthparts; 59: lateral view; 60: detail, tip of labial palpus with sieve-like openings; 61: lateral view, detail. Figs 62–64. Male antenna of Pseudoamuria uptoni. (E. Hines.) 62: Distal part; 63: detail, distal end; 64: detail, pectination.

ZYGAENID MORPHOLOGY

65

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Figs 65–70. Head of Thyrassia inconcinna, male. (E. Hines.) 65: Frontal view; 66: frontal view, detail; 67: frontal view, detail, base of proboscis; 68: lateral view; 69: lateral view, detail, eyelash; 70: lateral view, detail, ‘hairy space’ ventrad compound eye.

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Figs 71–74. Male antenna of Thyrassia inconcinna. (E. Hines.) 71: Entire view; 72: detail, distal end; 73: detail, central part; 74: detail, pectination.;

ZYGAENID MORPHOLOGY

75

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Figs 75–80. Head of Illiberis (Primilliberis) pruni Dyar, 1905 (Procridini), male. (E. Hines.) 75: Frontal view; 76: frontal view, detail, base of proboscis; 77: lateral view; 78: lateral view, detail, typical short ‘procridine chaetosema’ with a free space between compound eye and ocellus; 79: detail, compound eye without interommatidial setae; 80: detail (high magnification), compound eye without inter-ommatidial setae.

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Figs 81–83. Male antenna of Illiberis (Primilliberis) pruni Dyar, 1905 (Procridini). (E. Hines.) 81: Entire view; 82: detail, distal end; 83: detail, distal end (high magnification). Figs 84–86. Head of Adscita (Adscita) statices (Linnaeus, 1758) (Procridini), male. (E. Hines.) 84: Frontal view; 85: frontal view, detail; 86: frontal view, detail, base of proboscis with ‘spiny wart’.

ZYGAENID MORPHOLOGY

87

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100 µm

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Figs 87–91. Head of Adscita (Adscita) statices (Linnaeus, 1758) (Procridini), male. (E. Hines.) 87: lateral view; 88: lateral view, detail, ‘procridine chaetosema’; 89: lateral view, detail, eyelash; 90: detail, compound eye; 91: detail, compound eye, with inter-ommatidial seta.

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Figs 92–97. Male antenna of Adscita (Adscita) statices (Linnaeus, 1758) (Procridini). (E. Hines.) 92: Entire view; 93: detail, distal part; 94: detail, distal part (high magnification); 95: detail, tip of antenna (high magnification); 96–97: detail, pectinations.

ZYGAENID MORPHOLOGY

The Procridinae are mainly smaller than the Zygaeninae and Chalcosiinae; they are uniform in appearance, the wing shape varying from elongaterounded to subtriangular or triangular. In several genera of Procridinae, shiny metallic scales cover parts of the body and wings, e.g. Zygaenoprocris Hampson, 1900 (partim), Jordanita Verity, 1946, and Adscita Retzius, 1783, in the Palaearctis, Neoprocris Jordan, 1915, in Central and South America, and Pollanisus Walker, 1854, in Australia. This character seems to have evolved several times independently or may otherwise be a procridine plesiomorphy. In Procridinae, mimicry appears to play a major role only in some tropical groups. In the tropical parts of the Indo-Australian region, species of Araeocera Hampson, 1892, Ephemeroidea Hampson, 1892, Lophosoma Swinhoe, 1892, Phacusa Walker, 1854, and Thyrassia Butler, 1876, seem to mimic ctenuchid arctiids or sesiids or even nonlepidopterous insects such as Hymenoptera. In the Americas, species of Pyromorpha Herrich-Schäffer, 1854, apparently mimic Lithosiinae (Arctiidae), Ctenuchinae (Arctiidae) and Tortricidae (Tarmann 1984). In Australia, the species of Hestiochora Meyrick, 1886, seem to mimic hymenopterous and dipterous insects (Pls 36–43) (Common 1970: fig. 6 ZC). The wing venation is rather constant in Zygaeninae, more variable in Chalcosiinae and extremely variable in Procridinae. All primitive zygaenids have a fully developed medial stem in both wings and a full complement of veins, the radial and medial veins in the forewing inserting at the posterior margin of the cell and regularly spaced from each other. CuP and 1A+2A are always fully developed (Fig. 98). The use of wing venation for classification is especially limited in Procridinae, and overestimation of its importance by earlier authors has led to some confusion (Tarmann 1984: 15). The ultrastructure of zygaenid wing scales is very variable. Primitive Zygaeninae, e.g. Pryeria Moore, 1877, Orna Kirby, 1892 and Praezygaena Alberti, 1954, exhibit ‘transverse striae’ (sensu Davis 1978: 17, figs 36–38) similar to those of Micropterigidae, Eriocranidae and primitive Incurvariidae (Naumann 1977b: 71, fig. 18c; 1977c: 16, fig. 4b). In the more advanced groups of Zygaeninae perforated scales are found (Naumann 1977b: 71, fig. 18d; 1977c: 16, fig. 4f). In Chalcosiinae and Procridinae the longitudinal ribs of the scales are connected by small ‘bridges’, the space between these being completely filled by a smooth plate in the more primitive forms, but in the more advanced groups

the plate is perforated by holes that may be large or small (Naumann, Tarmann and Tremewan 1999: 30, text-fig. 29). This kind of structure does not significantly differ from those found in other groups of higher Lepidoptera (Scoble 1992: 65, fig. 78). In Procridinae the ultrastructure of these plates is directly related to the appearance of the adult moth. A sieve-like ultrastructure produces the striking metallic sheen of many Procridinae, whereas large holes between the transverse bridges are always associated with a matt appearance. The most advanced character state is the complete reduction of the plate, leaving only a large square space between the transverse bridges. This produces a velvety, mainly dark (black) overall appearance. An attempt has been made to explain the evolution of a square hole from a smooth plate in the ultrastructure of the scales of Procridinae. On the basis of detailed studies of American Procridinae two pathways seem possible: one through the enlargement of a single central hole, the other through the increase 98

a

b

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d

e

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Fig. 98. Wing venation of male Zygaenidae, Procridinae: (a) Illiberis ulmivora (Graeser, 1888) (Procridini); (b) Pollanisus cupreus Walker, 1854 (Artonini); (c) Myrtartona rufiventris (Artonini); (d) Artona postvitta Moore, 1879 (Artonini); (e) Artona discivitta Walker, 1854; (f) Thyrassia inconcinna (Artonini).

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Figs 99–102. Forewing scales of Pollanisus viridipulverulenta, male. (C. Beaton). 99: Zone 1, overview; 100: Zone 1, detail; 101: Zone 2, overview; 102: Zone 2, detail. Figs 103–104. Hindwing scales of Pollanisus viridipulverulenta, male. (C. Beaton). 103: Zone 3, overview; 104: Zone 3, detail.

ZYGAENID MORPHOLOGY

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Figs 105–109. Forewing scales of Turneriprocris dolens, male. (C. Beaton.) 105: Zone 1. 106–107: Zone 1, detail; 108: Zone 2; 109: Zone 2, detail. Figs 110–111. Hindwing scales of Turneriprocris dolens, male. (C. Beaton.) 110: Zone 3; 111: Zone 3, detail.

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Figs 112–116. Forewing scales of Myrtartona leucopleura, male. (C. Beaton.) 112: Zone 1; 113–114: Zone 1, detail; 115: Zone 2; 116: Zone 2, detail. Figs 117–118. Hindwing scales of Myrtartona leucopleura, male. (C. Beaton.) 117: Zone 3; 118: Zone 3, detail.

ZYGAENID MORPHOLOGY

119

a

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121

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b

in size of the holes of a sieve-like structure (Tarmann 1984). Most of the characters demonstrated in that study can be seen in the Australian zygaenids, showing that the scales of this group have sieve-plate based structures only (Figs 99–118). The structure and ultrastructure of procridine scales are of taxonomic significance when compared in three areas of the upperside of the wings (‘3Zonen-Muster’ sensu Tarmann 1984: 17, fig. 219). Zygaenid legs are in general well developed, they vary (especially in the Procridinae) from short and stout, e.g. Thyrassia Butler 1876, to very long and slender, e.g. Palmartona gen. n., Pseudoamuria gen. n., Homophylotis Turner, 1904 (Figs 119–121). The foretibia may bear an epiphysis (Naumann, Tarmann and Tremewan 1999: 28, text-fig. 27); its presence or absence can be seen in all groups. Nevertheless, the epiphysis does sometimes characterise genera or groups of genera. It should be noted that the presence or absence of an epiphysis may vary within a single population, e.g. Illiberis pruni Dyar, 1905. On the whole, one pair of tibial spurs is developed on the midleg, and either one or two pairs on the hindleg. An interesting single medial spur on the hindtibia occurs in almost all Asian Artonini (Procridinae); it is on the central axis of the hindtibia and does not appear to be derived simply from the loss of one or other of the two medial spurs. This character is absent in the Australian genera except for Palmartona gen. n., which is of Asian origin and probably introduced to Australia. Abdomen (Figs 124–127) The sclerotized parts of the abdomen, consisting of eight free segments in the male and seven in the female, bear useful diagnostic characters in many Chalcosiinae and Procridinae, especially sternite VIII and tergite VIII in the males. In some Chalcosiinae, e.g. Chalcosiini, sternite VIII seems to play an important role in holding together the male and female abdomens during copulation (Yen 2003: 313).

Fig. 119. Examples of right forelegs of Zygaenidae: (a) Zygaena filipendulae (Linnaeus, 1758) (Zygaeninae); (b) Zygaenoprocris chalcochlora Hampson, 1900 (Procridinae, Procridini); (c) Adscita statices (Linnaeus, 1758) (Procridinae, Procridini). Ex Efetov (2001: figs 70, 72, 81). Fig. 120. Examples of right hindlegs of Zygaenidae: (a) Zygaena filipendulae (Linnaeus, 1758) (Zygaeninae); (b) Zygaenoprocris chalcochlora Hampson, 1900 (Procridinae, Procridini) ; (c) Adscita statices (Linnaeus, 1758) (Procridinae, Procridini). Ex Efetov (2001: figs 77, 79, 82). Fig. 121. Examples of right fore and hindlegs of Australian Zygaenidae, Procridinae, Artonini: (a) Palmartona catoxantha (with foretibial epiphysis and single medial spur on hindleg); (b) Pollanisus viridipulverulenta (without foretibial epiphysis and without medial spur on hindleg). (Ink drawing by Stefan Heim, TLMF, Innsbruck.)

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Figs 122–123. Foreleg, tarsus of Pollanisus viridipulverulenta. (C. Beaton.) 122: Entire view; 123. detail, areole. Fig. 124. Lateral gland on second abdominal segment on male of Australartona mirabilis. (J. Green, CSIRO, Canberra.) Fig. 125. Female abdomen of Pollanisus viridipulverulenta, abdominal hairtuft. (C. Beaton.) Figs 126–127. Female abdomen of Turneriprocris dolens, abdominal end. (C. Beaton.) 126: Overview; 127: papillae anales, detail.

ZYGAENID MORPHOLOGY

Coremata occur in the Zygaeninae (Zygaena Fabricius, 1775) in the 8th/9th intersegmental fold in males in the form of eversible hair pencils or brushes. These hair pencils are in close contact with dorsal and dorsolateral gland cells in the integument, which are believed to charge the coremata scales with scent molecules. The coremata are everted during courtship prior to copulation. The chemical structure of the main compound has been found to be the main constituent of scabious flowers (Dipsacaceae) frequently visited by burnet moths (Naumann, Tarmann and Tremewan 1999: 28, text-fig. 30). In Chalcosiinae a different system exists. An S-shaped hair brush is attached proximally to the posterior margin of the hindwing in both sexes, being much more strongly developed in the male; in some groups it is completely absent in the female. It inserts into a lateral fold of the 1st/2nd abdominal pleura when the insect is at rest; the epidermal cells of this fold have been transformed into gland cells (Tarmann 1992b: figs 42–45; Naumann, Tarmann and Tremewan 1999: 28; Yen 2003: 317, fig. 6). This organ exhibits group-specific differences within the Chalcosiinae as shown by Yen (2003). Coremata also occur at the end of the abdomen in the Procridinae e.g. Harrisina Packard, 1864 and Artona Walker, 1854, but their anatomy and function are still unknown. Hallberg and Subchev (1997) found a special pheromone-distributing system on abdominal tergites 3–5 of the female of Theresimima ampellophaga (Bayle-Barelle, 1808). When calling, the female exposes the abdomen by slightly spreading her wings. This habit has recently also been found in some other Palaearctic genera: Rhagades Wallengren, 1863; Jordanita Verity, 1946; Adscita Retzius, 1783, and Illiberis Walker, 1854 (Efetov and Tarmann 1999: 16, figs 337, 339, 346; Efetov 2001: pl. 53, figs 1–7; Nishihara and Wipking 2003: 111, fig. 3). It may be more widely distributed in Procridinae. In many species of Artonini (Procridinae) a pair of lateral, bulb-like, semi-eversible evaginations is present on abdominal segments 2 and 7, the surface of which is smooth and the function of which is unknown (Figs 124, 430, 431). A characteristic odour is produced by some species of Artonini, and it is possible that this and the lateral evaginations are in some way related. These structures can also be found in later instar larvae, and in the pupa (Efetov and Tarmann 2004). In some Australian Procridinae (Pollanisus Walker, 1854; Onceropyga Turner, 1906; Hestiochora Meyrick, 1886) the 7th/8th intersegmental fold in the female is enlarged and has a sticky surface.

Long, dart-like scales are attached to this area and form a large abdominal hair-tuft. A glandular structure seems to be developed just below the cuticular surface of this area, producing a poisonous compound that is transferred to the base of the scales of the hair-tuft. When these scales are attached to the surface of the eggs after oviposition the base becomes the poisonous tip. It has been observed that aphids die after contact with the tips of these scales (Tarmann 1994: 118) (e.g. Pl. 3, Fig. 1; Pl. 4, Fig. 2; Pl. 5, Fig. 2; Pl. 59, Figs 1, 2; Figs 125, 129, 130, 436–440). Male genitalia The external male genitalia of zygaenids, formed from the modified abdominal segments VIII, IX and X, are of extraordinarily different construction in the different subfamilies, tribes, groups and subgroups. In the Zygaeninae all structures are heavily sclerotized, dark brown, and maceration in hot potash takes a long time (up to an hour) until the parts can be separated easily. A double-lobed uncus is attached to a helmet-shaped tegumen that is connected in turn to a narrow, clasp-shaped vinculum. The valva is disc-like, densely covered with long, inward-pointing setae distally. The aedeagus is tube-like with a lamina dorsalis and lamina ventralis bearing spines of taxonomic significance. Spines of taxonomic importance are also often found on the vesica seminalis (for details see Alberti 1954, 1956 and 1958–1959; Naumann, Tarmann and Tremewan 1999: 34, text-figs 33–35). The Callizygaeninae have a similarly shaped, strongly sclerotized valva, a single uncus and a double-lobed vinculum. The aedeagus is small, tube-like and without a lamina dorsalis and a lamina ventralis (Tarmann 1994: 119, fig. 1). In the Chalcosiinae there are several different types of male genitalia. In the tribe Cyclosiini the male genitalia are not very strongly sclerotized, the uncus is long and single, often pointed distally and covered with setae, the valva is rectangular and rounded distally or slender and sharply pointed distally, and the tegumen is double-lobed. The aedeagus is small, mainly straight and tube-like. In the Chalcosiini the genitalia are very strongly sclerotized, the uncus is either double-lobed or helmetshaped, the valva is very short and rounded, the ventral end forming a crest with tooth-like structures medially. The aedeagus is very strongly sclerotized, narrow and hook-like. Abdominal sternum VIII is transformed into a pair of hook-like structures used for holding the female during copulation.

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In the Agalopini and Aglaopini, a generally helmetshaped uncus and sometimes a large gnathos occur, the valva is elongate, and rounded or pointed distally, and the aedeagus is in the form of a very small spine. The greatest variation in the genitalia of the Chalcosiinae occurs in the tribe Heteropanini, where almost all possible structures are developed. The aedeagus is mainly tube-like and straight (for the genitalia of the Chalcosiinae see Alberti 1954). Recently, an extended analysis of the subfamily has been undertaken by Yen (2003). In the Procridinae the uncus is very uniform, always single and without distal setae. It is usually long and slender with a pointed, sometimes hooklike apex. Among several hundred species worldwide, the uncus is only different in the two American genera Neoprocris Jordan, 1915, and Pyromorpha Herrich-Schäffer, 1854, where it may even be completely reduced (Tarmann 1984). The valva in Procridinae is markedly different in the two tribes Procridini and Artonini. In Procridini it is usually strongly sclerotized with broad dorsal and ventral sclerotization (at least proximally) and a translucent part mediodistally. If there are appendages, they are always fused with these sclerotized areas and are fixed, not mobile. In Artonini the valva is very translucent and weakly sclerotized. In its primitive form it is simple, with a narrow sclerotized area dorsally and ventrally and a broad, strongly folded, translucent central part. This simple artonoid valva is found in some Australian genera, e.g. Pollanisus Walker, 1854, and Onceropyga Turner, 1906, and in one genus from China, Thibetana Efetov and Tarmann, 1995 (e.g. Figs 136, 138–185, 274, 278). The valvae of most Artonini are very complex. In general, a finger-like structure (‘Artonafinger’ sensu Efetov and Tarmann 1995: 74) is inserted at the distal end of the valva ventrally. This ‘Artona-finger’ bears brush-like setae in many groups, which are of taxonomic significance (Figs 402–405, 412, 414, 415). All transitions between a fixed, immovable ‘finger’ and one with a complex jointed articulation occur. In some genera the ventral sclerotization of the valva is divided into ventral and dorsal parts, and both parts are connected to another joint, allowing the distal part to move independently, e.g. Amuria Staudinger, 1887 and Artona Walker, 1854. Many other additional structures exist. The aedeagus is tube-like in all Procridinae. The vesica seminalis very often bears spines and/or cornuti. In some genera of Artonini, e.g. Artona Walker, 1854 and Pseudoamuria gen. n., bundles of deciduous cornuti with a spiny surface occur; these remain in the

female praebursa after copulation (Figs 408, 409, 410). The internal male genitalia of zygaenids have not yet been examined sufficiently, but seem similar to most other Lepidoptera (Naumann, Tarmann and Tremewan 1999). Female genitalia Zygaenidae are ditrysian Lepidoptera with two separate genitalic openings: the gonopore, through which the male aedeagus is inserted during copulation, and the oopore that is used for oviposition and is situated near the tip of the abdomen. The gonopore (the ostium bursae of taxonomic literature) is situated on sternite VIII of the female abdomen. The ostium bursae, ductus bursae, corpus bursae and ductus seminalis are derived from embryonic ectoderm and are therefore provided with cuticular lining. These parts are more heavily sclerotized than the internal female genitalia and often display useful and constant taxonomic characters. The ostium is often reinforced by lip-shaped sclerotizations in the Zygaeninae and Procridinae, but is only a simple round opening in the Chalcosiinae. A ductus bursae of very different form leads into the corpus bursae, where the spermatophore is deposited. In the Zygaeninae and Callizygaeninae the ductus bursae is simple, tube-like and often rather strongly sclerotized. In Chalcosiinae it is very translucent and usually narrow; it may be either short or long. In many Procridinae the ductus bursae is transformed into a large, sometimes very strongly sclerotized, balloon- or bottle-shaped structure called a ‘praebursa’ by Alberti (1954: 156) (e.g. Fig. 265). Functionally this praebursa often replaces the corpus bursae and accommodates the spermatophore (Fig. 242). Strongly sclerotized rows of teeth and/or spine- or hook-like ‘cornuti’ may be found in the praebursa of many groups of Procridinae (Figs 373–377, 410, 411, 419–422). These structures are supposed to prevent successful multiple copulations and/or seem also to be able to rupture the relatively strong wall of the spermatophore. The praebursa sometimes has a lateral appendix to accommodate cornuti of the male vesica and/or to store the deciduous male cornuti that occur in some groups (Fig. 410). These deciduous cornuti seem to prevent the praebursa from receiving a second spermatophore. So far, very few studies have been undertaken to explain all these complicated structures. The presence of a praebursa is supposed to represent an autapomorphy for most, if not all, Procridinae. According to Alberti (1954: 155) the plesiomorphic character state is a simple,

ZYGAENID MORPHOLOGY

tube-like ductus bursae found in some groups of Asian Procridini, e.g. subgenus Primilliberis Alberti, 1954, of Illiberis Walker, 1854. Careful re-examination of the female genitalia in the Procridinae from all parts of the world shows that this is doubtful. One finds a praebursa-like enlargement of the ductus in almost all Procridinae that, according to other characters, seem to represent primitive forms, e.g. most Illiberis Walker, 1854, in Asia; the subgenera Wiegelia Efetov and Tarmann, 1995, and Naufockia Alberti, 1954, of Rhagades Wallengren, 1863; or the subgenus Praviela Alberti, 1954, of Jordanita Verity, 1946, in the western Palaearctis; Acoloithus Clemens, 1861; Neoilliberis Tarmann, 1984, and Triprocris Grote, 1873 in America, and Pollanisus Walker, 1854 in Australia. The most primitive group of Procridinae is, according to Alberti (1954: 230), the subgenus Primilliberis of Illiberis with a large, funnel-shaped posterior part of the ductus bursae with ring-like sclerotizations near the ostium. Alberti concludes that this is the beginning of the development of a praebursa. If this is true, we must conclude that all other Procridinae worldwide are derived from forms similar to Primilliberis. If this is so, a praebursa was developed early, before the Procridinae spread over the rest of the world, and Primilliberis must be an old, relict group with the plesiomorphic character state of a simple ductus still present. If we look at the Australian genus Pollanisus with very simple and constant male genitalia, supposed almost to represent the groundplan of an artonoid valva, and we also accept the wing venation in this genus as rather primitive, Pollanisus must be seen as a primitive group of Artonini although it has several reductions (e.g. only one pair of abdominal evaginations) and other apomorphies (e.g. abdominal hair-tuft of female); this is also supported by larval characters. The praebursa is well-developed but very simple: a translucent enlargement of the ductus bursae, with a small appendix to accommodate the cornutus of the male vesica (e.g. Figs 240, 242, 244, 246, 252). Similar character combinations are found in the American genera Acoloithus Clemens, 1861; Neoalbertia Tarmann, 1984, and Triprocris Grote, 1873. These three genera are, according to their male genitalia and the ultrastructure of their wing scales, primitive Procridini (Tarmann 1984: 148). In the Asian genus Illiberis (Procridini), all possible combinations between a simple ductus and a very complicated praebursa with rows of spines, crests and teeth occur. It seems more likely that the present character state in Primilliberis is just a reduction of a former well-developed praebursa and therefore not

plesiomorphic but an autapomorphy of this subgenus. This corresponds with other newly discovered apomorphies in Primilliberis, e.g. the absence of interommatidial setae in the compound eye (Figs 79, 80). Although we cannot exclude the possibility that an enlargement of the ductus bursae has developed independently several times, the ability to form a praebursa seems to represent an autapomorphy of the entire subfamily Procridinae. The praebursa is connected to the corpus bursae by a short ductus intrabursalis (Alberti 1954: 155). The corpus bursae are translucent and spherical, or pocket- or bag-shaped. Signa are known in the Zygaeninae, Callizygaeninae and in some Chalcosiinae, e.g. Heteropanini, but are absent in the Procridini. The ductus seminalis inserts at the posterior end of the ductus bursae close to the ostium in the Zygaeninae, at the anterior part of the ductus bursae in the Callizygaeninae and many Chalcosiinae, and at the corpus bursae near its orifice into the ductus bursae in all Procridinae and some Chalcosiinae. An enlargement, or sometimes a lateral sac-like appendix of the ductus seminalis (‘bulla seminalis’ sensu Alberti 1954: 154), described as a ‘pseudobursa’ by Petersen (1900), is present in the Zygaeninae and Callizygaeninae but absent in the Chalcosiinae and Procridinae. The internal female genitalia of Zygaenidae have been studied by Naumann (1988). They consist of both mesodermal and ectodermal parts and they are difficult to study as they are very translucent and fragile. They provide excellent characters for phylogenetic analysis at the generic and higher systematic levels. There are four ovarioles on each side of the body, leading into two oviducti simplices that unite to form the oviductus communis. The receptaculum seminis is situated dorsad of the oviductus communis, opening into an enlargement of the oviductus communis, the infundibulum. The receptaculum seminis consists of a ductus receptaculi with a spiral interior structure and the spermatheca, consisting of the tube-like utriculus and the sac-like lagena that is attached to the utriculus laterally. The distal part of the utriculus is transformed into a long, tube-like accessory gland, the glandula receptaculi (Figs 342, 373). As in all Lepidoptera the spermatozoa must cross the infundibulum when leaving the ductus seminalis and are stored in the receptaculum seminis. The spermatozoa are ‘activated’ by components of the glandula receptaculi and descend towards the infundibulum through the spiral tube of the ductus receptaculi. Each egg is inseminated when it passes through the infundibulum. In the Zygaeninae, Callizygaeninae and Chalcosiinae, the

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receptaculum seminis is similar to that of most other Lepidoptera. In the Procridinae the lagena is reduced or fused with the utriculus, and this character is an important autapomorphy of the subfamily. The glandula receptaculi is long and very slender in most Zygaeninae, Callizygaeninae and Chalcosiinae but relatively short in Procridinae. After insemination at the infundibulum, the egg is coated with secretions from the glandula sebacea, which give the egg shell or chorion its consistency. The glandula sebacea in the Zygaenidae consists, as in most other Lepidoptera, of a pair of generally long and tube-like glands fused together basally and a slender ductus that inserts dorsally into the oviductus communis close to the ooporus. In the Zygaeninae, Callizygaeninae and Chalcosiinae, we find the glandula sebacea in the above-mentioned basic form. In the Procridinae a spherical ‘bulla’ is very often developed at the point where the two glandular parts are fused (Figs 342, 373). This ‘bulla sebaceae’ is sometimes large and round, e.g. in Illiberis and Rhagades, but may also be tube-like or heart-shaped, e.g. in Pollanisus. In the Zygaeninae a large pair of additional glands close to the oopore are connected to the oviduct; these were described as ‘Petersen’s glands’ by Bode and Naumann (1987). They are clearly separated into a secretory part and a reservoir where the secretions are stored. Examination of their ultrastructure shows that the gland cells have a characteristic extracellular ductus leading into the internal cytoplasm and ending in a curve, thus circumventing the nucleus. The cell also has additional characteristics peculiar to lipid-producing cells (Bode and Naumann 1987; Naumann 1988). The liquid released by these glands is highly proteinaceous and forms a thin coating on the eggs. Although it does not contain cyanoglucosides, it is thought to be poisonous and to protect the eggs from predators. ‘Petersen’s glands’, as described by Bode and Naumann (1987) and also figured by Naumann (1988: 87, fig. 1) and Naumann, Tarmann and Tremewan (1999: text-figs 38, 39), are considered to be an autapomorphy of the Zygaeninae. In the Procridinae we find a similar pair of glands at exactly the same place where the ‘Petersen’s glands’ occur in the Zygaeninae, and there is little doubt that both glands are homologous. The structure and ultrastructure of the glands are different, however. They form a simple sac in the Procridinae, and the existence of a pair of additional glands close to the oopore may represent a synapomorphy of the Zygaeninae and Procridinae. The further development into ‘Petersen’s glands’ in the Zygaeninae, and into a pair of different, simpler glands in the Procridinae are autapomorphies of

these two subfamilies. These glands are absent in the Chalcosiinae. They are also absent in three Australian genera of Procridinae (Artonini): Pollanisus Walker, 1854; Onceropyga Turner, 1906, and Hestiochora Meyrick, 1886. As described above, the females of these three genera protect their eggs with long dart-like scales from an abdominal hairtuft (Figs 129, 130).

Egg (Pl. 59, Figs 1, 2; Figs 128–130, 436–438) Zygaenid eggs are small, ovoid or slightly rectangular, with an almost smooth surface lacking any prominent structure. The chorion is slightly reticulate. No significant characters have so far been found around the micropyle (Naumann, Tarmann and Tremewan 1999: text-figs 4, 5). Their colour varies from pale yellowish green or whitish green to bright yellow or orange. The eggs are laid singly, in small rows, in single flat layers or in pyramidal or irregularly shaped batches. They are attached to the substrate by a secretion from the female glandula sebacea. In Zygaeninae and most Procridinae they are additionally coated with a sticky proteinaceous secretion or by urticating spiny scales. Oviposition of single eggs is considered to represent the ancestral egg-laying pattern, and pyramidal or irregular batches the derived pattern; the latter can be clearly seen in the Zygaeninae (Naumann, Tarmann and Tremewan 1999: text-fig. 5). Mainly flat egg batches are known in the Procridinae, but, in some groups at least, two layers of eggs can occur (e.g. Illiberis) (Nishihara and Wipking 2003: 115, fig. 4c). In many procridine groups, single eggs are laid or they are deposited in short rows or flat layers in which the eggs are spaced apart from each other (Efetov and Tarmann 1999, Efetov 2001).

Larva (Figs 131–133) One of the first and most comprehensive descriptions of a zygaenid larva ever published was that by Tothill, Taylor and Paine (1930) for the coconut moth in Fiji, Levuana iridescens Bethune-Baker, 1906. However, the combination of characters mentioned as typical for the family (Tothill, Taylor and Paine 1930: 84) refers only to one subfamily (i.e. Procridinae) and one tribe (i.e. Artonini). A good description of a Zygaeninae larva (i.e. Zygaena trifolii (Esper, 1783)) is given by Tremewan (1985) and of a Chalcosiinae larva (i.e. Aglaope infausta (Linnaeus, 1767)) by Fänger and Naumann (2001).

ZYGAENID MORPHOLOGY

128

129

130

Fig. 128. Egg batch of Australartona mirabilis, flat layer, eggs not touching each other and without protecting spine-like scales from female abdomen. (J. Green, CSIRO, Canberra.) Figs 129–130. Egg batch of Pollanisus viridipulverulenta on leaf of Hibbertia obtusifolia DC. (J. Green, CSIRO, Canberra.) 129: flat layer, eggs not or slightly touching each other, with protecting spine-like scales from the abdominal hairtuft of the female; 130: with protecting spine-like scales from the abdominal hairtuft of the female, lateral view.

All zygaenid larvae are prognathous (head horizontal and mouthparts directed forward), and the head capsule is retractile under the slightly extended prothoracic segment (Tothill, Taylor and Paine 1930: 91, fig. 31; Naumann, Tarmann and Tremewan 1999: 18, text-fig. 6). However, this has recently been challenged by Fänger and Naumann (1998) and Yen (2003). The ability to retract the head has been considered to represent an autapomorphy of the superfamily Zygaenoidea (Minet 1986: 300; Epstein et al. 1999: 159). The antennae are long, the mandibles are of normal size, and the well-developed spinneret is preceeded on both sides by labial and maxillary palps. Six stemmata are normally present on each side of the head, but one can be reduced (Tothill, Taylor and Paine 1930: 93, figs 34A–D; Tremewan 1985: 87: fig. 52). The head capsule bears primary setae only. The prothorax has mainly a well-sclerotized cervical shield dorsally and is enlarged to accommodate the retractile head capsule. The thoracic legs are well-developed. A well-developed circular spiracle is present on the prothorax. The ten-segmented abdomen has a pair of prolegs on segments 3–6 (ventral prolegs) and 10 (anal prolegs), the crochets uniordinal, uniserial, arranged in mesoseries, spiracles on segments 1–8 (Tothill, Taylor and Paine 1930: 94, figs 37, 38; Tremewan 1985: 97). The body of the zygaenid larva is mainly covered by a rough cuticle, sometimes bearing minute spines. In Zygaeninae and Procridinae, primary setae are found in first instar larvae only. The larvae of these two subfamilies develop complementary subprimary setae in the second and secondary setae in subsequent instars that are arranged in dorsal, subdorsal, lateral and ventral verrucae (sensu Scoble 1992: 120, fig. 135). In the Chalcosiinae (as far as examined) only primary setae can be found (in all larval instars) except in Aglaope Latreille, 1809 (Fänger and Naumann 2001) and Arbudas Moore, 1879 (Yen 2003). In Procridinae, an anal comb is normally developed dorsad of the anus and two smaller comb-like aggregations of setae are present at the posterior base of the anal prolegs (Tothill, Taylor and Paine 1930: 95, figs 39, 40). Under low magnification they appear together with the anal comb as three dark spots. These combs are frequently reduced. They are present in all Australian genera so far examined except Australartona gen. n. (i.e. Pollanisus, Hestiochora, Onceropyga and Myrtartona gen. n.). The anal comb aids the ejection of faecal pellets or frass. The chaetotaxy of zygaenid larvae has been studied by various authors (Tothill, Taylor and Paine

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1930; Tremewan 1985; Stehr 1987; Yen and Fan 1995; Efetov and Tarmann 1999; Efetov et al. 2000; Efetov 2001, 2003a, 2003b, 2004; Efetov and Drouet 2003; Efetov, Mollet and Tarmann 2003; Efetov and Tarmann 2003a). So far, significant differences have only been found in the first instar larvae of Procridinae. In later instars the number of setae is variable and of little significance (Tothill, Taylor and Paine 1930: 86). Unlike Hinton (1946), who examined the pro-, mesothorax and 6th abdominal segment of the larva, Efetov and Tarmann (1999) and subsequent authors used the first abdominal segment for their studies. This segment is more easily accessible owing to the lack of legs and shows sufficiently reliable and constant characters. No differences have been found in setal combination and setal colour between closely related species but

131

there are differences between species of different subgenera (Fig. 131). Zygaenid larvae are either uniform or brightly coloured. The larvae of Zygaeninae very often show a typical pattern consisting of black and yellow spots, but they can also be very uniform. The larvae of Chalcosiinae are often very conspicuous and extremely colourful (e.g. Cyclosia Hübner, 1820; Erasmia Hope, 1841; Agalope Walker, 1854; Aglaope Latreille, 1809; Gynautocera Guérin-Méneville, 1831; Histia Hübner, 1820), but uniformly coloured larvae are also known (e.g. Arbudas Moore, 1879; Soritia Walker, 1854). Most larvae of western Palaearctic and American Procridinae are uniform in coloration but some of the eastern Palaearctic, Indo-Australian, and a few of the American species also have colourful larvae. In some regions,

132

a

b

c

d

e

f

133

g

h

i

Fig. 131. Diagrams of chaetotaxy of the first abdominal segment of first instar larvae of Zygaenidae, Procridinae. Anterior end to the left: (a) Jordanita (Solaniterna) subsolana (Staudinger, 1862) (Procridini); (b) Adscita (Adscita) statices (Linnaeus, 1758) (Procridini); (c) A. (Tarmannita) mannii (Lederer, 1853) (Procridini); (d) Jordanita budensis (Speyer & Speyer, 1858) (Procridini); (e) Illiberis (Primilliberis) rotundata Jordan, 1907 (Procridini); (f) Rhagades (Rhagades) pruni ([Denis & Schiffermüller], 1775) (Procridini); (g) Pollanisus viridipulverulenta (Artonini); (h) P. subdolosa clara (Artonini); (i) Australartona mirabilis (Artonini). Figs 131 a–d, f after Efetov (2001: figs 14, 17, 21, 27, 39); Figure 131e after Efetov (2003a: Figure 1); Figs 131g–i (ink drawings by Stefan Heim, TLMF, Innsbruck). Scale 0.1 mm. Figs 132–133. Adult larva of Pollanisus apicalis on Hibbertia obtusifolia DC. ACT, Black Mountain, November 1989. (J. Green, CSIRO, Canberra.) 132: Dorsal view; 133: lateral view.

ZYGAENID MORPHOLOGY

zygaenid larvae resemble lycaenid larvae and are also found on the same host-plants. Either group might gain from the other’s chemical protection or even from the ants that are very often symbiotic with the larvae of lycaenids (Naumann, Tarmann and Tremewan 1999: 18). Some colourful chalcosiine larvae also resemble limacodid larvae that are also well protected by toxins. For an account of the larval defence system of zygaenids see p. 43. The larvae of the Australian Zygaenidae are still poorly known. There is some material stored in alcohol in the ANIC and a few photos and descriptions are published (e.g. Common 1990: 295, fig. 52.10). However, the existing preserved material consists almost exclusively of larvae of the later instars. These larvae are extremely uniform, and no differences in setal combinations have been found, even at generic level. It seems that only the colour is significant (but the colour disappears in alcohol). It would be most useful to further examine first instar larvae. A remarkable autapomorphy for the tribe Artonini of the Procridinae is the presence of a pair of elliptical, semi-eversible, lip-shaped glands on abdominal segments 2 and 7 in the larva, and bulblike structures in the same position in the pupa and the adult (Pl. 60, Fig. 5; Figs 430, 431). In other Procridinae these glands can only be found in late instar larvae (Tothill, Taylor and Paine 1930; Stehr 1987; Efetov and Tarmann 2003b, 2003c, 2003d, 2004) but not in the pupa or the adult. This trait (in the adults), together with the specific form of the chaetosemata and the valvae, has been used as an important argument to separate the tribe Artonini and consider it to be monophyletic (Tarmann 1994: 121, fig. 3). However, these abdominal glands had already been discovered, described and figured by Tothill, Taylor and Paine (1930: 95, fig. 36 (larva), 107 (pupa)), who surpisingly did not mention its presence in the abdomen of adult specimens. The function of these glands is still unknown. They seem to be reduced in most (or all?) adult Procridini and can be reduced in a few Artonini (e.g. Hestiochora).

Tremewan 1985; Common 1990: 296, fig. 97.4; Naumann, Tarmann and Tremewan 1999: text-figs 12, 13). The second abdominal spiracle is covered by the wings, a character that is considered to represent an autapomorphy of the whole superfamily Zygaenoidea (Minet 1986: 300; Epstein et al. 1998: 159). The integument of the pupa is sometimes almost translucent whitish yellow, lacking pigment, but may be pigmented, varying from light brown to black and may even have a pattern of stripes, e.g. Theresimima ampellophaga (Bayle-Barelle, 1808) (Procridini) (Efetov and Tarmann 1999: fig. 328). In all species where the adults have abdominal lateral bulb-like structures, these structures can also be observed in the pupa. However, as the spiracle is covered by the edge of the wing on the second abdominal segment, they can only be seen on this segment if the wing is lifted up (Pl. 60, Fig. 5; Fig. 134). The cocoons of zygaenids consist of silken threads of seroproteins produced by the labial glands of the larva (Procridinae) in which microscopic crystals of calcium oxalate monohydrate (whewellite) produced by the Malpighian tubules of the last instar larvae are incorporated, giving the cocoon a parchment-like surface (Zygaeninae, Chalcosiinae) (Hofmann 1994: 204; Naumann, Tarmann and Tremewan 1999: 22, text-figs 14–16; Efetov and Shiryayev 2000, 2003). The cocoons of Zygaeninae are sometimes completely exposed on grass stems, twigs or leaves but usually they are hidden under plant litter, rocks, or on or under the surface of the earth (Tremewan 1985; Hofmann 1994; Naumann, Tarmann and Tremewan 1999; Efetov and Shiryayev 2003). The Chalcosiinae very often attach their cocoon along the midrib of a leaf of the larval foodplant, partially folding the leaf to enclose the cocoon. The cocoons can be rather hard 134

Pupa and cocoon The pupa of zygaenids is ‘incomplete’, the abdominal segments 3–7 in the male and 3–6 in the female are freely movable, the appendages are loosely fused to the body, articulated mandibles are absent, and tergites 3–8 in the male and 3–7 in the female bear a transverse row of hooked spines anteriorly (Tothill, Taylor and Paine 1930: 105, fig. 41, 196, fig. 42;

Fig. 134. Pupa and cocoon of Pollanisus apicalis. (J. Green, CSIRO, Canberra.)

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(Naumann, Tarmann and Tremewan 1999: 23). The cocoons of Procridinae are fragile (Fig. 134). They can be well concealed in or on the earth by small soil or plant particles attached to their outer surface (many Procridini), but may also be spun on bark, twigs and leaves (Naumann, Tarmann and Tremewan 1999: 23; Efetov and Tarmann 1999: 157, fig. 329). Many Artonini spin their cocoons on twigs or on bark but at times also on the leaves of the foodplant (Tothill, Taylor and Paine 1930: pl. XVII; Common 1990: 121, fig. 52.11). Before the adult moth emerges, the pupa ruptures the anterior part of the cocoon, using the

vertex of the head to break the silken threads, and moving its abdomen with a circular motion. Using the transverse rows of hooked spines on the abdomen, the pupa moves forward and protrudes its anterior part, the abdomen remaining in the cocoon. After the emergence of the moth, the pupal exuvia usually remains protruding from the cocoon. In Zygaena, the movements of the pupal abdomen in the cocoon can produce a distinct rasping sound (Tremewan 1985: 93). The question as to whether this sound has any defensive or other function has not been answered so far.

Biology and life history Phenology Most Australian zygaenids have two or more generations per year. Species occurring in the more temperate southern parts of the Australian mainland are usually bivoltine, with one generation in late spring or early summer (October to December) and another in autumn (February and March). In the colder mountainous regions of New South Wales and Victoria, and in Tasmania, the same species are usually univoltine. All subtropical and tropical zygaenids of Australia seem to be multivoltine. As in most Palaearctic species, diapause takes place in the larval stage. Real hibernation in a specialised larval instar or even in a hibernaculum, as is known for some groups of Palaearctic Procridinae, e.g. Rhagades Wallengren, 1863, is not known in the Australian species. A quiescent period in larval development observed in Australian species may be caused by dryness and lack of food. However, the possibility that larvae of some exclusively mountain species, e.g. Pollanisus calliceros Turner, 1926, have a genuine diapause cannot be excluded.

Larval host-plants The larvae of zygaenids are usually oligophagous or even monophagous. Polyphagy is only known in a few species, e.g. Zygaena exulans (Hohenwarth, 1792) (Zygaeninae) and Rhagades pruni ([Denis and Schiffermüller], 1775) (Procridinae). Zygaena exulans is a Palaearctic oreal species and has an arcto-alpine distribution. Hofmann and Tremewan (1996: 132) mention species of 12 plant families that are accepted by the larvae. The Palaearctic R. pruni is distributed from Spain to Japan and feeds on different host-plants according to the region where it occurs. Efetov (2001: 70) mentions various species belonging to six plant families. So far, more than 100 different foodplants belonging to more than 40 families are known for the entire family Zygaenidae (Tarmann 1992a). The larvae of primitive Zygaeninae feed exclusively on Celastraceae, and the more derived groups of this subfamily feed on Fabaceae, Apiaceae, Asteraceae and Lamiaceae (Hofmann and Tremewan 1996; Naumann, Tarmann and Tremewan 1999: 55). The larvae of Chalcosiinae

feed on a wide range of different Palaearctic and tropical plants but some groups seem to be rather specialised, e.g. Aglaope Latreille, 1809, and Agalope Walker, 1854, on Rosaceae; Eterusia Hope, 1841, and Soritia Walker, 1854, on Theaceae, and Campylotes Westwood, 1839, on Pinus spp. (Pinaceae) (Tarmann, 1992a). The larvae of Procridinae feed on several different plant families but within each group they are specialised. Some Palaearctic Procridinae are leaf-miners, e.g. Jordanita Verity, 1946, and Adscita Retzius, 1783, many American species are leaf-skeletonizers, e.g. Harrisina Packard, 1864, and a few Palaearctic species are stem-borers of Asteraceae, e.g. Jordanita subsolana (Staudinger, 1862); however, most Procridinae larvae feed externally on the leaves of their foodplants. Sometimes the early instar larva is a leaf-miner and the later instars are external feeders (e.g. many Palaearctic Adscita species) (Efetov and Tarmann 1999; Naumann, Tarmann and Tremewan 1999; Efetov 2001). The following foodplants are used by Australian zygaenids (Procridinae, Artonini). The large genus Pollanisus Walker, 1854, uses Dillenia, Hibbertia, Tetracera (Dilleniaceae), Pipturus (Urticaceae) and Kennedia (Fabaceae). Hestiochora Meyrick, 1886, Turneriprocris Bryk, 1936, and Myrtartona gen. n. use Eucalyptus, Melaleuca, Kunzea and Leptospermum (Myrtaceae). Onceropyga Turner, 1906, and Thyrassia Butler, 1876, use Vitis (Vitaceae), Palmartona gen. n. uses palms (Arecaceae) and Australartona gen. n. uses grass (Poaceae). Except for the tropical genera Palmartona gen. n. (on Arecaceae), Thyrassia (Vitaceae) and the subtropical genus Onceropyga (Vitaceae), the foodplants of most of the Australian zygaenids belong to plant genera endemic to, or showing their greatest diversity in, Australia (e.g. Eucalyptus, Melaleuca, Hibbertia). Dilleniaceae and Myrtaceae are not known as larval foodplants of Procridinae outside Australia (except for a single record of an unidentified Phacusa sp. feeding on Dillenia pentagyna Roxb. from India). Most Artonini (Procridinae) from Southeast Asia, New Guinea and the islands eastwards to Fiji feed on monocotyledons such as Poaceae, Arecaceae, Zingiberaceae and Musaceae (Tarmann, 1992a).

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Parasitoids Parasitoids of zygaenids are usually rather specialised as they must be able to detoxify hydrocyanic acid (for detoxification of cyanide see p. 43). However, large numbers of Hymenoptera, e.g. Chalcididae, Ichneumonidae, Braconidae, and Diptera (Tachinidae) are known to be able to cope with such detoxification. Moreover, the toxicity of zygaenids seems not to be a major problem for many spiders and Heteroptera (especially Reduviidae), and even some beetles have been recorded feeding on zygaenids. Parasitoids have been used successfully to control pest zygaenids. One of the first large biological control programmes ever undertaken was to control the coconut moth, Levuana iridescens Bethune-Baker, 1906 (Procridinae, Artonini), in Fiji (Tothill, Taylor and Paine 1930). Further successful programmes to control the Southeast Asian coconut moth, Palmartona catoxantha (Hampson, [1893]), are known from Malaysia and Indonesia (see p. 210). A few years ago, Artona chorista Jordan, 1907, caused major problems in cardamom plantations in Sikkim and biological control has recently been started there (Bhowmik 1963; Thakur and Sachen 1987; Yadav et al. 1992; Chatterjee, Kumar and Naidu 1993; Tarmann 2003). Major effort has been put into the biological control of the American grape-leaf skeletonizers Harrisina americana (Guérin-Méneville, [1844]), Harrisina metallica Stretch, 1885 (= H. brillians Barnes & McDunnough, 1910), Acoloithus falsarius Clemens, 1861, and Acoloithus novaricus Barnes & McDunnough, 1913 (all Procridinae, Procridini) (Robinson 1950; Langston and Smith 1953; Smith 1953, 1954; Clausen 1961; Mead 1970; Corette McGiffen and Neunzig 1985; Landolt, Heath and Tarmann 1991; Tarmann 2000a, 2003). The Mediterranean grape-leaf skeletonizer Theresimima ampellophaga (Bayle-Barelle, 1808) has been controlled almost exclusively with chemical pesticides (Efetov 1998; Tarmann 1998, 2000a; Efetov and Tarmann 1999, Tarmann 2003). The most important parasitoids known for the control of zygaenids are braconid Hymenoptera of the genera Apanteles Förster, 1862 and Cotesia Cameron, 1891, ichneumonid Hymenoptera of the genus Charops Holmgren, 1859 and tachinid Diptera of the genus Ptychomyia Brauer & Bergenstamm, 1889 (Naumann, Tarmann and Tremewan 1999: 73ff.) Almost nothing is known about the parasitoids of Australian zygaenids and so far only a few species

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Fig. 135. Tachinidae pupa ex larva and in cocoon of Pollanisus apicalis. (J. Green, CSIRO, Canberra.)

have been reared, among them a tachinid fly from Pollanisus apicalis Walker, 1954 (Fig. 135).

Cyanogenesis and resistance to hydrocyanic acid (HCN) All instars of zygaenid moths release hydrocyanic acid (HCN), an extremely toxic gas known to block the respiratory pathways of most living organisms owing to its extremely high affinity for alcohol-dehydrogenase, a component essential for energy production in living cells, and cyclooxigenase. Cyanogenesis of zygaenids was first reported by Rothschild (1961) and the highest concentration of HCN was found in the eggs. Davis and Nahrstedt (1979) and Nahrstedt and Davis (1981) found that HCN in zygaenids is derived from the two cyanoglucosides linamarin (2β-D-glucopyranosyloxy-2-methylpropionitrile) and lotaustralin (2-β-glucopyranosyloxy-2-butyronitrile). These compounds combine a sugar molecule with a cyanogenic aglucon. Linamarin and lotaustralin are present in many plant families, e.g. Fabaceae. As Fabaceae are one of the main foodplant families for the western Palaearctic burnets (Zygaena species), it was initially hypothesised that zygaenids are dependent on the acquisition of cyanoglucosides from plants and synthesize HCN through their enzymatic breakdown. This was refuted by the fact that, although many zygaenids world-wide feed on noncyanogenic plants (e.g. Arecaceae), all of them are cyanogenic. It has since been shown that zygaenids and their larvae are in fact able to synthesize linamarin and lotaustralin from the amino acid precursors valin and isoleucin by the same pathway as found in plants (Wray, Davis and Nahrstedt 1985; Witthohn and Naumann 1987a). According to

BIOLOGY AND LIFE HISTORY

Witthohn and Naumann (1987a: 1789) cyanogenesis in animals is restricted to a few instances in arthropods. Examples are known from Myriapoda, Diplopoda and Insecta (Heteroptera, Coleoptera, Lepidoptera). In Myriapoda, Diplopoda, and in Insecta in Coleoptera (Cicindelidae, Chrysomelidae), HCN is produced by catabolic decomposition of mandelonitrile (benzaldehyde-αhydroxynitrile). In the Lepidoptera so far studied, e.g. in Acraea and Heliconius butterflies and zygaenid moths, it is synthesized from linamarin and lotaustralin. Zygaenids not only release HCN but their haemolymph also contains large amounts of βcyano-L-alanine (BCA) (Witthohn and Naumann 1984a, 1984b). This substance was first discovered in larvae of the noctuids Helicoverpa zea (Boddie, 1850) (Heliothinae) and Spodoptera exigua (Hübner, 1808) (Hadeninae). BCA is synthesised when HCN is detoxified through BCA-synthase. The presence of this substance in an insect is therefore an indicator that the species concerned is able to detoxify HCN and is therefore resistant to it (for more details and figures see Naumann, Tarmann and Tremewan 1999: 56ff.). Cyanogenesis occurs in 14 lepidopterous families. In most of them BCA is only present in small amounts. Large quantities of BCA are found in all Zygaenidae, Anomoeotidae, and Heterogynidae, in some Megalopygidae and Yponomeutidae, and in some species-groups of Nymphalidae, e.g. most Heliconiinae, some Clossiana, Issoria and Acraea species, and in the geometrid Lomographa temerata (Linnaeus, 1758) (Witthohn and Naumann 1987a). Witthohn and Naumann (1987a: 1806) conclude that cyanogenesis is a widespread phenomenon in the Lepidoptera. They suggest that cyanogenesis is a basic character of Lepidoptera but it cannot be confirmed that it is based on linamarin and lotaustralin, and that detoxification via BCA is an autapomorphy of this order.

Defensive biology In the larvae of the Zygaeninae and most Chalcosiinae a highly complicated defensive system has been developed in conjunction with the synthesis of cyanide (Franzl 1980, 1986; Franzl and Naumann 1984, 1985; Franzl, Nahrstedt and Naumann 1986; Franzl, Naumann and Nahrstedt 1988). On the third thoracic segment and on abdominal segments 1 to 8 the larval integument has four cavities per segment where a highly viscu-

ous, sticky fluid is stored. This fluid moves directly through the unmodified cells of the larval epidermis from the body cavity to the storage chambers and is not, as one might assume, secreted by specialised glands. The dry weight of this secretion consists of about 50% cyanoglucosides. These cyanoglucosides are not lost with the exuvia during ecdysis as are most defensive substances in other insects but are transported into the body and reabsorbed before moulting. Immediately after moulting the cyanoglucosides are transported back into the new cuticular cavities. Four cavities are developed on each segment with different opening mechanisms in the Zygaeninae and Chalcosiinae (Naumann, Tarmann and Tremewan 1999: 21, text-fig. 11). When disturbed, the larva releases drops of a clear, toxic liquid by constriction of the circular segmental muscles (Naumann, Tarmann and Tremewan 1999: 58, text-figs 57, 58). This liquid has not yet been analyzed in detail but it is known to contain the cyanoglucosides linamarin and lotaustralin. Larvae of the Procridinae, and therefore all Australian zygaenids, seem to lack the abovementioned opening mechanisms and storage chambers in their larval cuticle. Nevertheless, these larvae are also highly toxic and some of them, if distrubed, also release small drops at the dorsal verrucae that are visible on the dorsal setae. Moreover, they have in their later instars eversible lip-shaped glands on abdominal segments 2 and 7 that are of unknown function but which may also play a role in defensive biology (see p. 39). Many adult zygaenids and their larvae are aposematic, some of them very conspicuously so. All larvae of Zygaeninae and Chalcosiinae, including those that are not aposematic, release drops of a clear, yellow-brown liquid from their integument when disturbed (see above). Another defensive system is well-developed in Zygaeninae and Chalcosiinae, in which a specialised opening beside the compound eye can be found in the adults. Here the toxic liquid is released as conspicuous foam, the release of the liquid producing a hissing sound. In Chalcosiinae, an additional pair of openings with the same function is developed between the patagia and parapatagia on the prothorax (Yen 2003). So far, active release of a defensive liquid has not been observed in adult Procridinae. The biological significance of linamarin, lotaustralin, β-cyanoglucosidase, and β-cyano-L-alanine (BCA) in zygaenids has been tested experimentally; specimens containing these compounds were fed to ants, frogs, toads, lizards, chameleons, birds and

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shrews (Rocci 1915, 1916). All animals tested showed strong reactions such as spitting, mouth cleaning, spasms and the refusal of food for a long time after ingesting zygaenids. These reactions were the same whether zygaenid larvae or adults were tested. Lane (1959) reports on an experiment in that his mother, Miriam Rothschild, vaccinated herself with the clear yellow fluid from the cervical ‘gland’ region of a burnet moth. Two drops of this fluid were placed inside the right forearm and the skin was scratched with a sterilized needle, the point being drawn quickly through the centre of each drop of fluid. The result was startling. Within a few minutes an oval area of dusky red formed around the site of each scratch about 8 × 9 cm in extent but there was no trace of a swelling or local pain. Almost immediately sensations of breathlessness occurred, associated with sweating and a rapid pulse rate that increased to 120–130 beats per minute. The main symptom was one of great restlessness and agitation. These symptoms lasted several hours. Most females of the Zygaeninae and Procridinae protect their eggs by covering them with the secretion from Petersen’s gland (Zygaeninae) or the pair of homologous glands (Procridinae) situated close to the oopore, or, in some Australian Procridinae genera, by urticating scales (see p. 37, Figs 129, 130). Whether or not the urticating tips of the scales from the abdominal hair-tuft also contain cyanide has not so far been investigated.

Ecology, dispersal, diversity and distribution Zygaenids are found in almost all habitats where their larval foodplants are found. Most Zygaeninae, Chalcosiinae and Procridinae are clearly derived from ancestral species inhabiting arboreal habitats, even if some of them are nowadays secondarily restricted to treeless steppe, semi-desert, oreal, or mountain habitats, e.g. many Zygaena species. Only in the Procridinae are species-groups of primary steppe or semi-desert origin known, e.g. Jordanita species feeding on Artemisia spp. from the treeless Asian steppe (e.g. Jordanita (Roccia) paupera (Christoph, 1887)) or Zygaenoprocris species feeding on Atraphaxis spp. and Polygonum spp. restricted to the Middle East semi-deserts (e.g. Zygaenoprocris duskei (Grum-Grshimailo, 1902) and Zygpr. persepolis (Alberti, 1938)). Zygaenids are stenoecious and usually restricted to small, isolated areas even if the biotopes in which they occur appear to be large and uniform. In general they develop only small colonies and are therefore difficult to observe. This is not the case,

however, for most species of the Palaearctic genus Zygaena, which are mainly also stenoecious but are often abundant in their habitats. They are well known and observations are usually easy because the moths are so conspicuous. In the Zygaeninae (Zygaena) most new colonies are started by windborne gravid females (Tremewan 1985: 96; Naumann, Tarmann and Tremewan 1999: 78). If a habitat is suitable, they may be very abundant for a limited period (usually 2–3 years). The abundance then decreases, mainly because of parasitoids. The ability to increase rapidly in numbers seems to be highly developed in many zygaenid species, especially in the Procridinae (see pest species p. 46, 210). At present, the highest species diversity is known from the interchange of centres of generic species richness between the Oriental and the eastern Palaearctic regions (northern India – Burma – south-western China). Very little is known about zygaenids occuring in tropical rainforests. Almost every zygaenid newly collected in the rainforests of the Neotropis represents an undescribed species, suggesting their diversity may be very high. As most of these species are very small, day flying, and canopy-living with the larva feeding mainly on climbing Vitaceae, they have never been collected systematically. A similar situation probably exists in Southeast Asia, New Guinea and adjacent islands, and in northern Australia. Many of the smaller species known from those areas have only been collected in very low numbers. Their larvae feed mainly on monocotyledons (Arecaceae, Musaceae, Zingiberaceae) or on Vitaceae, and we can assume that some of them are also subcanopy or canopy species. Zygaenids are usually diurnal. Some species are regularly attracted to light, especially m.v. light, but most of them are mainly active during the day. Females are always less active than males and usually never fly very far. In many of the smaller species, especially in the Procridinae, the females stay close to where they have emerged from the pupa, with the males sometimes swarming around the plants in large numbers. Usually adult activity starts in the late morning. The adults search for flowers to obtain nectar, except in those species with a reduced proboscis that do not feed at all (see p. 9). It is interesting to note that different plants seem to be preferred as nectar sources in different parts of the world. Palaearctic zygaenids (Zygaeninae and Procridinae) are strongly attracted to the pink, violet or white flowers of Dipsacaceae, e.g. Knautia, Scabiosa, Cephalaria, or Asteraceae, e.g. Carduus,


Centaurea, and Cirsium. Chemical analysis of the flower-scent compounds has shown that at least the main substances of some Dipsacaceae and the orchid Anacamptis pyramidalis (L.) Rich. are partly also present in the sex pheromones of some European Zygaeninae. The component found was 6,10,14-trimethyl-pentadecan-2-ol and this is the corresponding ketone to the alcohol 6,10,14trimethyl-pentadecan-2-ol, the main constituent of the male scent brushes (coremata) (Naumann, Tarmann and Tremewan 1999: 60, text-fig. 61). In America, the preferred plants have white flowers, e.g. Mimosaceae (Acacia) or Oleaceae (Ligustrum). In Australia there seems to be a preference for the yellow flowers of Asteraceae (Helichrysum, Arctotheca). Mainly in the afternoon, the females start to release pheromones to attract males. It is at this time that males can most easily be seen on the wing and waiting for this time of day is sometimes the only way of observing a rare species. Mating usually begins in the late afternoon and lasts until the following morning. Males are able to copulate several times. Many females have specialised structures in their genitalia to prevent successful multiple inseminations (see e.g. Figs 373–377, 410, 419–422). Females begin ovipositing soon after copulation has ceased, and many different modes of oviposition have developed (p. 36). Oviposition is often interrupted for feeding, and there are many species where the number of eggs laid depends on the availability of nectar resources during the oviposition period. Most females are slow fliers, and their biology (particularly for the Procridinae) has been elucidated by following females, watching them oviposit and then rear the larvae.

Pheromones and mating behaviour Females of all Zygaenidae release sex pheromones. The chemistry of these pheromones has only been identified for a few western Palaearctic and American species. They consist of unsaturated carbohydrates with double-bonds. For Zygaeninae they are rather simple alcohols or aldehydes varying in chain length and position of the single doublebond (Zagatti and Renou 1984; Arn, Toth and Priesner 1992; for formulae see Naumann, Tarmann and Tremewan 1999: text-fig. 63). It has been proved that a mixture of at least three different components is most effective in attracting males. The chemical structure of the pheromones of Procridinae is more complicated. 2-butyl-(Z)-7-

tetradecenoate has been identified as a main component of the sexual pheromone of the American western grape leaf skeletonizer, Harrisina metallica Stretch, 1885 (= brillians Barnes and McDunnough, 1910) and some other American species (Myerson, Haddon and Soderstrom 1982; Soderstrom et al. 1985; Landolt et al. 1986; Landolt, Heath and Tarmann 1991). The same chemical substance has been identified for the Mediterranean vine-bud moth Theresimima ampellophaga (BayleBarelle, 1808) by Subchev and co-authors (Subchev and Harizanov 1990; Subchev et al. 1998; for formulae see Efetov 2001: text-fig. 43). Moreover, Subchev and co-authors observed an unusual mating behaviour in Theresimima ampellophaga (Subchev 1996; Hallberg and Subchev 1997; Subchev 2000; Subchev 2003). The female’s sex pheromone glands are not situated at the terminal end of the abdomen as usual but located on the anterior part of abdominal tergites 3–5. To attract males, the female moves into a specific calling position with the wings slightly spread and lowered, thus exposing the down-flexed abdomen to release the pheromone (Subchev 2003: 146, fig. 1). It has been shown that this habit is not restricted to Theresimima but seems to occur throughout at least the Palaearctic Procridinae. Since its discovery by Subchev for Theresimima, it has also been found in the genera Rhagades Wallengren, 1863; Zygaenoprocris Hampson, 1900; Adscita Retzius, 1783; Jordanita Verity, 1946 and Illiberis Walker, 1854 (see summarized photo documentation in Efetov 2001: pl. 53, figs 1–7, and for Illiberis in Nishihara and Wipking 2003: fig. 3). All the above-mentioned genera belong to the tribe Procridini and not to Artonini, in which all the Australian genera are found. So far, we have no records on calling Artonini females either from Asia or Australia. As we already know that this habit does not occur in Zygaeninae, where the pheromone glands are in a ‘normal’ position at the end of the abdomen, it would be most interesting to know whether this character combination (unusual location of pheromone glands in female combined with unusual calling habit) is an autapomorphy of the whole subfamily Procridinae, of the tribe Procridini, or only a synapomorphy of a group of related genera. Usually calling can only be observed with reared virgin females as in nature the latter are very difficult to find. Interspecific attraction and copulation between species of Procridinae is well known and documented (e.g. Efetov 2001: pl. 53, figs 7, 8), but no hybrids have been observed so far. Females of

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closely allied but allopatric species seem to have almost identical pheromones as they attract males of the other species as easily as conspecific females. But sympatric species also have a strong affinity to each other, and attempts by males to mate with females of another species are frequent (observations made by the author together with K. A. Efetov in Spain in 2002 and W. G. Tremewan in Italy in 2003).

Genetics and individual variation To date the genetics of only a few species of Zygaena Fabricius, 1775 (Zygaeninae) have been studied. A summary of these results is given in Naumann, Tarmann and Tremewan (1999) and a comparative study has been provided by Tremewan (2002). They are based on extended rearing and crossing experiments, mainly studying red and yellow individuals, or populations with five or six spots. Similar variation between red and yellow individuals within one population of a single species, as in Zygaena, can be observed in other zygaenid groups. In the Chalcosiinae, red/yellow variation is known in Aglaope labasi Oberthür, 1922, and Aglaope infausta (Linnaeus, 1967) (Aglaopini). Striking examples also occur in the Procridinae (Procridini), e.g. in the Central American species Neoilliberis ignorata (Hering, 1925); Neoalbertia variabilis Tarmann, 1984, and Pyromorpha brueckneri (Hering, 1926), in which the red and yellow markings form a large number of different wing patterns against the black ground colour (Tarmann 1984). So far nothing is known of the genetics of Australian zygaenids. Individual variation is not very great in the group except for size and colour in a few species. Size is most probably not a result of genetic factors but dependent on the foodplant availability during larval development. Variation in the colour of the shiny metallic scales is probably controlled genetically; only metallic green and blue are variable in the Procridinae, other metallic colours in the subfamily do not vary. However, it is known from Palaearctic species of Jordanita and Adscita that the variability of the green and blue metallic colour is related to the humidity of the locality where the population occurs, and may even vary within one population in different years, once again depending on the humidity in a particular year.

Zygaenids as indicator species Being stenoecious insects, zygaenids are good indicator species for particular plant-animal associations. Only in Europe have species of the genera

Zygaena (Zygaeninae), Rhagades, Jordanita and Adscita (Procridinae) recently been used for environmental monitoring. In northern Italy it could be shown that zygaenids are perfect indicators for monitoring destructive events in the environment caused by moulting inhibitors (e.g. Dimilin, Insegar). For years these chemicals have been used in apple plantations against the codling moth (Cydia pomonella (Linnaeus, 1758) (Tortricidae)) and have killed almost the entire fauna of holometabolous insects in these southern Alpine valleys (Tarmann 2000b, 2000c). Zygaenids proved to be more sensitive to these chemicals than most other Lepidoptera species and, being conspicuous diurnal species, they are easy to observe. As a result of these studies the plantations are now treated with pheromones and the zygaenids are used as key species to monitor the recovery of the devastated areas. In regions outside of Europe, zygaenids are usually too difficult to observe to be used for this purpose. However, in Australia many common Pollanisus species are restricted to undisturbed native bushland, their larvae feed on Hibbertia (Dilleniaceae), they are easy to observe, and so could well be used for environmental monitoring.

Pest zygaenids Some species of zygaenids are crop pests, most of them belonging to the Procridinae (16 species) and Chalcosiinae (nine species) and only one, i.e. Pryeria sinica Moore, 1877, to Zygaeninae (for a summary and bibliography see Tarmann 2003). The oldest records known are for the Mediterranean species Theresimima ampellophaga (Bayle-Barelle, 1808), an important pest in vineyards since Roman times (Tarmann 1998: 58). The use of insecticides during the last 50 years has brought about a rapid decline in the number of populations as well as individuals, and a few years ago this species was assumed to be extinct over most of its former range. On the Crimean Peninsula, however, some populations have recently managed to exploit the widespread naturalized American decorative vines Parthenocissus quinquefolia (L.) Planch., P. inserta (A. Kern.) Fritsch, P. inserta × P. quinquefolia (hybrid) and the Asian decorative vine P. tricuspidata (Siebold and Zucc.) Planch (Vitaceae) as larval host-plants. Such populations have recently begun to spread rapidly over south-eastern Europe and should be monitored carefully (Efetov 1990, 1996, 1998, 2001; Efetov and Tarmann 1999; Tarmann 1998, 2000a, 2003; Embacher and Tarmann 2001).


In North American vineyards most of the problems have been caused by the western grape leaf skeletonizer Harrisina metallica Stretch, 1885 (= brillians Barnes and McDunnough, 1910). Biological control has been used successfully for many years (Robinson 1950; Langston and Smith 1953; Smith 1953, 1954; Clausen 1961). In Asia, infestations of Illiberis pruni Dyar, 1905, are reported on apple trees in north-eastern China (Zhang 1994) and in Japan (Nishihara 2000; Wipking 2000; Nishihara and Wipking 2003). Illiberis tenuis (Butler, 1878) is known as a pest in vineyards in Japan (Inoue 1976, 1982). Artona chorista Jordan, 1907 has been recorded as a serious pest on cardamom in Sikkim (Bhowmik 1963 (as Clelea plumbeola); Thakur and Sachan 1987 (as Clelea plumbeola); Yadav et al. 1992; Chatterjee, Kumar and Naidu 1993). In the Sundaland region, major damage to coconut palms has been caused by Palmartona catoxantha (Hampson, [1892]) (see p. 208 for Palmartona catoxantha). One of the first successful examples of biological control was that of the coconut moth in Fiji (Levuana iridescens Bethune-Baker, 1906) (Tothill, Taylor and Paine 1930). This species is now thought to be extinct (Robinson 1975), although this is very doubtful. My own experience shows that tropical Procridinae may be unobserved for years, well hidden in the canopy of their host-plant trees or bushes when the population density is low. The best recent example of this is Artona chorista Jordan 1907. This species was only known from its two type specimens collected a century ago. Suddenly it appeared in huge numbers destroying large cardamom plantations in Sikkim. Even the well known Southeast Asian coconut palm pest Palmartona catoxantha (Hampson, 1892) disappears completely for years between periods of infestation. Another Artonini mentioned as a pest is the grassfeeding Artona (Balataea) funeralis (Butler, 1879). Efetov (1997) has shown that the type of this species is different from what has been generally understood as A. funeralis in the literature. The common East Asian Artona species that feeds on bamboo and bamboo-grasses and frequently causes damage, differs in genitalic structure and is easily separated from the rare A. funeralis by its dark proboscis (yellow in A. funeralis!). This common pest species was described as Artona (Balataea) martini Efetov, 1997. In March 1996 A. martini was discovered in New Zealand where it must have been introduced from east Asia. It has spread and is now well established (Gill 2000; Hoare 2001; Tarmann 2003).

Conservation Whether or not it is still necessary to collect zygaenids in Europe is debatable. Most of the life histories, ecologies and bionomics of the European burnets and foresters are already known. Several populations are seriously threatened and some have already become extinct although not through overcollecting. In most other parts of the world zygaenids are rarely encountered except for some pest species (see above). Australian zygaenids are not endangered although some of them have rather restricted distributions, e.g. Hestiochora occidentalis, Pollanisus calliceros. A sufficient network of natural native areas in the various Australian National Parks, Conservation Parks and Nature Reserves exists and is a reasonable guarantee for the preservation of the habitat of this group. There is so little information on the habitat preferences, early stages, foodplants, adult habits and their role as pollinators available for Australian zygaenids that any effort to obtain such information would be most worthwhile. Many amateur lepidopterists have made major contributions to the present knowledge of this group and they should be encouraged to continue to undertake research, especially on the habitat, distribution, ecology, ethology, mimicry, life histories, larval foodplants and insect/plant relationships.

Fossil record Very few fossil zygaenids are known; moreover, little is known about those that have been discovered. All fossils found so far originate from the middle Cenozoic (Tertiary) and belong to the subfamily Zygaeninae. The first fossil moth recognised as a zygaenid was ‘Zygaena’ miocaenica Reiss, 1936, a perfectly preserved specimen from bituminous deposits in southern Germany (Naumann 1987b: 301, figs 1–2, 303, figs 5–6; Hofmann 1994: 196, fig. xx; Naumann, Tarmann and Tremewan 1999: 54, text-fig. 54). A second specimen was described as Zygaena controversus Burgeff, 1951 (Naumann 1987b: 301, figs 3–4, 303, figs 7–8; Hofmann 1994: 197, fig. xx). Further species were recently found in Spain and France and described as Zygaena? turolensis Fernández-Rubio, Peñalver and MartinezDelclos, 1992 and Neurosymploca? oligocenica Fernández-Rubio and Nel, 2000 (Fernández-Rubio, Peñalver and Martinez-Delclos 1992; FernándezRubio and Peñalver 1994; Férnandez-Rubio 1998; Fernández-Rubio and Nel 2000). No fossil Callizygaeninae, Chalcosiinae or Procridinae have been discovered.

47

48



Phylogeny The Zygaenidae form part of the Zygaenoidea. There is still uncertainty over the monophyly of this superfamily and the relationships among the 12 included families. A summary of current concepts and controversy is given by Epstein et al. (1998) and Yen (2003). Really convincing autapomorphies for Zygaenidae have not yet been found. It has been assumed that the capacity to synthesise the two cyanogenic glucosides linamarin and lotaustralin from the amino acids valin and isoleucin, the presence of a specialised muscle in the mid gut of the larva, and the presence of a pair of accessory glands (viz. Petersen’s glands or homologous structures) in the internal female genitalia represent possible autapomorphies supporting monophyly of this family (Naumann, Tarmann and Tremewan 1999: 14). Yen (2003: 323) summarises the following potential apomorphies for Zygaenidae: (1) adults secrete cyanogenic foam from the intersegmental membrane between the mandibular lobe and the subgena; (2) the breadth of the area around the eyes occupied by the chaetosemata exceeds the length of the antennal base; (3) paired spatulate setae are present on the tarsus; (4) a row of additional sclerites is present below the crochets; (5) the front of the pupa has two pairs of short setae. I am not able to discuss these proposals in detail in this book as comparative studies for all these characters on Procridinae from all regions in the world would be necessary in order to comment on them. All characters that separate Zygaenidae easily from other families of Zygaenoidea are plesiomorphies (i.e. ocelli present, chaetosemata present, and proboscis well-developed). In Australia, only Procridinae of the tribe Artonini have been found so far. Procridinae are clearly monophyletic, an assumption at least supported by the following two characters: 1, female genitalia with ductus seminalis lacking a pseudobursa (‘bulla seminalis’ sensu Alberti, 1954); 2, lagena absent in the receptaculum seminis. Moreover, there are two more characters that seem to represent autapomorphies of the subfamily: 3, presence of a pair of accessory glands, homologous to ‘Petersen’s gland’ in the Zygaeninae, close to the oopore, but of different structure; 4, larva with an

unspecialised chemical defense system (Tarmann 1994: 117; Efetov and Tarmann 1999: 8). While the remaining Procridini are paraphyletic, the Artonini are considered to be monophyletic, based on the following autapomorphies: 1, abdomen of pupa and imago with a pair of dorsolateral evaginations on segments 2 and 7 (Pl. 60, Fig. 5; Figs 430, 431); 2, chaetosemata triangular, extending forward between the compound eye and the ocellus (e.g. Figs 6, 61); 3, valva fan-shaped with a more strongly sclerotized dorsal and ventral margin and a very translucent and strongly folded central part (Tarmann 1994: 120) (e.g. Fig. 136).

PAUP analysis To elucidate the relationship between the Australian and Asian genera of Artonini and produce a more definitive cladistic analysis, further investigations are necessary. Careful examination of character variation is sometimes difficult, as more material of many of the very rare tropical species is needed. However, an attempt has been made below to list and discuss characters and character combinations based on the available material. A cladistic analysis has been based on 31 characters for the 10 genera of the Artonini present in Australia. As outgroup taxa the genera Adscita (Procridini), Artona (Artonini) and Amuria (Artonini) have been selected. Adscita Retzius, 1783 (= Procris Fabricius, 1807) is a typical genus for the tribe Procridini, Artona Walker, 1854, and Amuria Staudinger, 1887, are the dominant genera in the Artonini outside of Australia with much information available. The data matrix was prepared using MacClade version 4.03 (Maddison & Maddison 2001) and PAUP* version 4.0b10 (Swofford 2002), and the consensus tree figured was prepared using PAUP*. The data matrix for the 13 taxa was analysed using PAUP* on a Power Macintosh 7100/66. Branch and bound analysis yielded eight trees of length 52, CI = 0.71 (CI excluding uninformatives = 0.67), RI = 0.73. The strict and semistrict consensus trees have the same topology, and only the first is illustrated.

50


Characters used in phylogenetic analysis 1. Presence of an epiphysis on fore tibia (Fig. 119) The presence of a fore tibial epiphysis is a plesiomorphic character. It is present in most primitive Lepidoptera but is reduced in many groups. This reduction has evolved independently many times in Lepidoptera beginning with the most primitive groups (e.g. Micropterigoidea) (Kristensen 1998: 42). A well-developed foretibial epiphysis is supposed to belong to the ground plan of Zygaenidae (Naumann 1977c: 22). In Procridinae it is reduced in many genera, but it is present in almost all tropical Artonini. In the Australian genera it is present in Palmartona gen. n. (Fig. 121a), Pseudoamuria gen. n., Homophylotis and Australartona gen. n., but absent in Myrtartona gen. n., Turneriprocris, Pollanisus (Fig. 121b), Onceropyga and Hestiochora. No character variations have been observed in the examined Australian and Southeast Asian material, but those variations are known for some Procridini (e.g. Illiberis pruni Dyar, 1905; see Alberti 1954). 2. Presence of a single medial spur on the hindtibia (Fig. 121a) This character is found in many genera of Artonini and may even be an autapomorphy of the whole tribe. This spur seems not to have evolved just from the loss of one of the two medial spurs of primitive Lepidoptera, as its position is completely central. However, the spur is absent in some genera and it is not yet clear whether this is a secondary loss or a plesiomorphy. In the Australian genera this medial spur is missing except in Palmartona gen. n. This genus is considered to originate from Southeast Asia where almost all species of various genera have a well-developed medial spur. No infrageneric variation of this character is known. 3. The lateral ‘glands’ on abdominal segments 2 and 7 in pupa and adult (Pl. 60, Fig. 5;Figs 430, 431) Eversible lip-shaped evaginations, presumably glands that produce a scent, are present in the later instar larva of Procridinae. In the same position, round evaginations with a smooth surface can be found in the pupae and the adults of many species. They are situated at the dorsolateral part of the second and seventh abdominal segments just below the spiracles. They can be found in most tropical Artonini, but are partly or completely reduced in some Australian genera. In Myrtartona gen. n., Turneriprocris, Pollanisus and Onceropyga these glands can only be found on the second abdominal

segment and they are completely absent in Hestiochora and Thyrassia. Australartona gen. n. and Homophylotis, however, not only have the full set of evaginations, but these evaginations are also significantly enlarged on the second abdominal segments in the male (see character 4). 4. Lateral evaginations enlarged on second abdominal segment in the male (see comment above and Figs 430, 431) 5. Asymmetry in the length of the male antennal pectinations In some genera of Artonini the antennae are strongly asymmetrical owing to different lengths of the anterior and posterior antennal pectinations. In Australia, Homophylotis and Palmartona gen. n. have asymmetrical male antennae. No infrageneric variability has been found. 6. The white spot on the distal part of the antenna (Pls 50–55) This is a very strange character. Some segments of the darkly scaled antenna are shiny white in some groups of Artonini. This spot is very conspicuous and easily visible in resting specimens that present their antennae in the characteristic upward-pointing position of zygaenids. The spots move with the antennae and seem to have some kind of signalling function. They are present in both sexes. The density and the number of the white scales are variable and in worn specimens they may even be absent. In most cases the presence or absence of these spots is clearly recognisable, though their function is unknown. However, in many Hymenoptera and Diptera similar characters can be found. 7. Presence or absence of the medial stem in the venation of the forewing (Fig. 98) The presence of a medial stem in the wing venation is supposed to be plesiomoprhic. In most Australian species a medial stem is present in the hindwing but only some genera have the medial stem also developed in the forewing (i.e. Turneriprocris, Pollanisus, Onceropyga, Hestiochora and Thyrassia). 8. Shiny, metallic scales on wings and body (Pls 1–32) Shiny metallic scales are found in several genera of Procridinae. The shiny metallic effect is caused by scales with an encaved plate between the transversal bridges with a sieve-like ultrastructure (Tarmann 1984). There are only three genera in the western Palaearctis (i.e. Zygaenoprocris (partim), Adscita and

PHYLOGENY

Jordanita) and one genus in Australia (i.e. Pollanisus) where, at least in some species, the whole forewing upperside and parts of the body are covered with such scales. This led to the conclusion by earlier authors (e.g. Jordan 1907) that the Australian Pollanisus are closely related to European Adscita (=Procris of earlier authors) or even that some of the Australian species are congeneric (Meyrick 1886, 1888). Now we know that Pollanisus is the sister-group of Onceropyga and is also monophyletic with Hestiochora (see p. 55), both genera lack shiny scales, and that none of the other Artonini of Asia and the Indo-Australian region have shiny scales. The presence of such scales in Pollanisus must have developed independently and is an autapomorphy of Pollanisus. 9. Wasp-like habitus (Pls 35–43) The peculiar wasp-like habitus of most Hestiochora is an autapomorphy of this group. 10. Ctenuchid-like coloration (Pl. 57) The ctenuchid-like coloration of Thyrassia is an autapomorphy of this genus. 11. Black and yellow wing pattern Artona species are very conspicuous owing to a characteristic black and yellow wing pattern and body. This character is supposed to represent a synapomorphy of Artona Walker, 1854 and Thibetana Efetov and Tarmann, 1995. 12. Hindwing strongly reduced in size (Pl. 57) Thyrassia species have a significantly reduced hindwing. The habitus of all species of that genus is very uniform. 13. Translucent hindwings (Pls 51–54) Translucent, almost clear hindwing areas occur in Homophylotis. They arise from a reduction of scale density and an upright position of the scales. This is thought to represent an autapomophy of Homophylotis. 14. The abdominal hair-tuft of the females (For description and discussion see p. 33) 15. The abdominal glands close to oopore in the female (For description and discussion see p. 36) 16. Female abdomen with long and slender urticating scales posteriorly (For description and discussion see p. 33, 36)

17. Female with enlarged abdominal segments 7 and 8 (For description and discussion see p. 33) 18. Eggs covered with pincushion-like scales from female abdomen (For description and discussion see p. 36, Figs 129, 130, 436, 437) 19. Female genitalia with praebursa This is a very unreliable character. In many Procridinae the ductus bursae is swollen to form a so-called praebursa (Alberti 1954). This praebursa is often large, well-sclerotized and bears numerous crests and spines. These characters are very useful for taxonomic purposes. In some groups the praebursa takes over the function of the corpus bursae and accommodates the spermatophore. However, it has to be concluded that a swelling of the ductus bursae could have occurred several times during evolution. Therefore, similar structures in different groups may not be homologous. However, the principal ability to swell and form a praebursa is here considered to represent an autapomorphy of Procridinae. For Artonini it is assumed that the presence of a praebursa is plesiomorphic and its absence is a secondary reduction and therefore apomorphic. In the Australian genera a well-developed praebursa is present in Palmartona gen. n., Pseudoamuria gen. n., Myrtartona gen. n., Onceropyga and Pollanisus. In Homophylotis the praebursa is secondarily fused with the corpus bursae (see comments p. 192 and Figs 397–400). In Australartona gen. n., Turneriprocris, Hestiochora and Thyrassia there is no praebursa developed. 20. Ductus intrabursalis inserting into praebursa at special hook-like or dentate sclerotisation In Myrtartona gen. n. and Pollanisus the large spherical praebursa has a strong sclerotisation developed at the point where the ductus intrabursalis is inserted. This structure has a rounded distal end and a dentate proximal end. The proximal dentate part is used to rupture the wall of the spermatophore that is always stored within the praebursa and never reaches the corpus bursae. The function of this structure is very easily visible and understood if the genitalia preparation is done carefully (e.g. Figs 242, 243, 373–377). By squeezing the round distal part of the sclerotisation, small portions of sperm material come out of the spermatophore and enter the ductus intrabursalis. This material is then stored in the corpus bursae from where it proceeds into the ductus seminalis and beyond. A very similar and

51

52


possibly homologous structure is also found in the monotypic genus Leptozygaena Jordan, 1907 (typespecies Leptozygaena gracilis Jordan, 1907) from New Guinea.

may have become fused with the wall of the aedeagus during the evolution of the genus (Figs 310–327). Moreover, in Hestiochora the vesica is not eversible in dissection.

21. Spiny crests in the praebursa (Figs 410, 411, 419–422) Spines and spiny crests are frequently found in the praebursa of Procridinae. A fully developed spiny crest may have two principal functions: 1, to rub off the deciduous cornuti from the vesica; and 2, to rupture the wall of the spermatophore and help the spermatozoa to enter the ductus intrabursalis and proceed into the corpus bursae and beyond. As stated earlier (see p. 51) the praebursa very often replaces the corpus bursae functionally. The spermatophore cannot proceed into the corpus bursae. In most cases, special mechanisms are developed to rupture the wall of the spermatophore. Species without a spiny crest in the praebursa but with strong spine-like sclerotizations never have deciduous cornuti in the male. The spiny crest is therefore almost certainly a mechanism to support the accommodation of the deciduous cornuti by ‘catching’ them during copulation. These crests are fairly constant in shape and number of spines and can be used for taxonomic purposes. In Australian species, spiny crests in the praebursa are only found in Pseudoamuria gen. n. and Palmartona gen. n. They are present in almost all Southeast Asian species of Procridinae.

23. Vesica with a bundle of slender, eversible (shootable) cornuti In many Artonini, bundles of deciduous spiny cornuti are developed on the more rounded smaller lobe of the vesica. They are packed together and become loose and detached (‘shooting out’) when the vesica is everted during copulation (Fig. 409). They are caught by the spines of the crest of the female’s praebursa, rubbed off and stored in a special pocket on the praebursa (Fig. 410). In the Australian genera we find this character combination only in Pseudoamuria gen. n.

22. Aedeagus with or without fixed cornutus The vesica of most Artonini is double-lobed, consisting of a longer more tube-like part and a smaller more spherical part (e.g. Fig. 224). So far no one has succeeded in everting a vesica of an Artonini by blowing so that the shape of these lobes is usually very difficult to describe. The longer lobe often bears a well-developed single cornutus that is attached to the vesica and situated at its distal end. This can be seen if the vesica is pulled out of the aedeagus using pins and a pair of forceps (e.g. Figs 137, 218). There can be more than one cornutus and the cornuti can also be lacking. In the Australian genera Pollanisus, Onceropyga and Myrtartona gen. n. there is one cornutus, in Palmartona gen. n. there are three (Figs 416–418); in Hestiochora, Turneriprocris, Australartona gen. n., Homophylotis and Pseudoamuria gen. n. fixed cornuti are absent. In Hestiochora the dorsal wall of the aedeagus is very strongly sclerotized and enlarged, somewhat spinelike distally, a character combination that leads to the conclusion that what was formerly a cornutus

24. Vesica with large triangular or rectangular spines The vesica of many zygaenids has a slightly spiny surface. However, really large spines in numbers are rare. In Homophylotis large spines are well-developed (Fig. 393). 25. The presence of an ‘Artona’-finger on the valva Most Artonini have a well-developed ‘Artona’-finger (for description see p. 34 and Figs 378, 380, 402, 403, 405, 412). This character is very striking, and many varieties of finger-like appendages inserted on the ventral posterior edge of the male valva are known. Because of the many different types of fingers and considerable variability within closely related groups, it is difficult to comment on these characters, particularly because their function is largely unknown. In the Australian genera the finger is well-developed in Palmartona gen. n., Pseudoamuria gen. n. and Australartona gen. n. In Homophylotis this finger is found in only one species in a considerably reduced form (H. artonoides sp. n.) (Figs 390, 391). In all other species of Homophylotis, as well as in all other Australian zygaenid genera, it is absent. 26. ‘Artona-finger’ with distal hair-brush Many ‘Artona-fingers’ have significant distal hairbrushes. Their function is still unknown. 27. The ‘Myrtartona-finger’ (e.g. Figs 344, 346, 347, 349, 350, 352) In Myrtartona gen. n. and Turneriprocris, a characteristic finger-like sclerotization is developed. Its basal part is forked and one of the forks is connected to the dorsobasal part of the valva by a small translucent connection. The other fork seems

PHYLOGENY

to be connected to a special muscle that moves the finger. The finger is bluntly pointed distally with a group of short spiny setae. Proximad of these setae several longer setae can be found pointing backwards (anteriorly). This characteristic pair of fingers points posteriorly in its natural position and can be moved like a pair of pliers. This character is presumed to represent a synapomorphy of the two above-mentioned genera. 28. Larval thorax ‘swollen’ The larvae of most Artonini examined so far are very uniform in morphological characters and are only different in coloration. However, Palmartona larvae have a significantly swollen thorax. This character is thought to represent an autapomoprphy for this genus. 29. Larva with long secondary hairs and an arctiidlike appearance The larvae of Thyrassia are hairy and look like arctiid larvae. They look remarkably different from other zygaenid larvae (see Piepers and Snellen 1903: pl. 13, fig. 12).

30. Larval host-plant families At least at generic level, Artonini seem to be monophagous or oligophagous. The groups examined so far are clearly associated with special plant families. Which families represent the ancestral state is still not clear. 31. Monocotyledons or dicotyledons as larval hostplants It is remarkable that all Australian genera bearing an ‘Artona’-finger and the Asian genera Artona and Amuria (also with ‘Artona’-finger) have larvae that feed on monocotyledons whereas all other genera feed on dicotyledons. For Homophylotis the larval host-plant is still not confirmed but, according to the relationship of Homophylotis to Pseudoamuria gen. n. and Levuana Bethune-Baker, 1906 (from Fiji), it is almost certain that Homophylotis larvae feed on monocotyledons. In the matrix below this genus has been coded as if the larval host-plants were already known.

Hestiochora

Thyrassia

11.

Onceropyga

10.

Pollanisus

9.

Turneriprocris

8.

Myrtartona

7.

Australartona

6.

Homophylotis

5.

Pseudoamuria

4.

Palmartona

3.

Amuria

2.

Foretibia epiphysis 0, present; 1, absent Medial spur on hindtibia 0, present; 1, absent Lateral evaginations (LEV) 0, present on abdomen on segments 2+7; 1, on segment 2 only; 2; absent LEV2 0, normal; 1, enlarged in male Antennal pectinations in the male 0, symmetrical; 1, asymmetrical White spot at distal end of antenna 0, absent; 1, present Wings with medial stem 0, at least partly present in forewing; 1, absent Wings: shiny, almost metallic scales 0, absent; 1, present Habitus 0, not wasp-like; 1, wasp like Habitus 0, not ctenuchid like; 1, ctenuchid-like Wing pattern 0, black and yellow; 1, other

Artona

1.

Adscita

Data matrix for PAUP analysis

1

0

0

0

0

0

0

1

1

1

1

1

1

1

0

0

0

1

1

1

1

1

1

1

1

1

2

0

0

0

0

0

0

1

1

1

1

2

2

0

0

0

0

0

1

1

0

0

0

0

0

0

0

0

0

1

0

1

0

0

0

0

0

0

0

0

0

0

0

1

1

0

0

0

0

0

0

0

0

1

1

1

1

1

1

1

0

0

0

0

0

1

0

0

0

0

0

0

0

0

1

0

0

0

0

0

0

0

0

0

0

0

0

0

0

1

0

0

0

0

0

0

0

0

0

0

0

0

0

1

1

0

1

1

1

1

1

1

1

1

1

1

0

53

23. 24. 25. 26.

27.

28. 29.

30.

31.

Thyrassia

22.

Hestiochora

21.

Onceropyga

20.

Pollanisus

19.

Turneriprocris

18.

Myrtartona

17.

Australartona

16.

Homophylotis

15.

Pseudoamuria

14.

Palmartona

13.

Hindwings 0, not strongly reduced in size; 1, strongly reduced in size Hindwings 0, not translucent; 1, partly translucent Female abdominal hair-tuft 0, absent; 1, present Female abdominal gland 0, close to ooporus; 1, absent Female urticating abdominal scales 0, absent; 1, present Female enlarged abdominal segments 7 and 8 0, absent; 1, present Eggs 0, not covered with pincushion-like scales; 1, covered with pincushion-like scales from female abdomen Female genitalia 0, with a well-developed praebursa; 1, praebursa fused with corpus bursae; 2, without praebursa Ductus intrabursalis 0, not inserting into praebursa at special hook-like or dentate sclerotization; 1, inserting into praebursa at special hook-like or dentate sclerotization Praebursa: rows of spiny crests 0, present; 1 absent Aedeagus 0, with one cornutus; 1, without cornutus; 2, with more than one cornutus Vesica: bundle of slender, deciduous cornuti 0, present; 1, absent Vesica: large triangular or quadrangular spines 0, absent; 1, present Valva ‘Artona’-finger 0, absent; 1, present ‘Artona’-finger 0, with significant distal hair-brush; 1, with a few hairs; 2, without hairs Proximal part of valva 0, without finger-like dorsal connection; 1, connected with a characteristic, finger-like process dorsally Larva 0, normal; 1, with ‘swollen’ thorax Larva 0, without long secondary hairs and arctiid-like appearance; 1, with long secondary hairs and arctiid-like appearance Larval host-plant 0, Myrtaceae; 1, Vitaceae; 2, Arecaceae; 3, Zingiberaceae; 4, Poaceae; 5, Dilleniaceae; 6, other Larval host-plant 0, monocotyledon; 1, dicotyledon

Amuria

12.

Artona


Adscita

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PHYLOGENY

Adscita Artona Amuria Palmartona Pseudoamuria Homophylotis Australartona Myrtartona Turneriprocris Pollanisus Onceropyga Hestiochora Thyrassia Strict consensus tree

Comments Not all questions can be answered from the results of the phylogenetic analysis presented here. It is more or less satisfactory according to morphological criteria but it is somewhat contradictory with respect to larval host-plants. The genera with larvae feeding on monocotyledons form a monophyletic grouping but the Myrtaceae feeders appear to be paraphyletic. This is not too disturbing if we consider Hestiochora to be plesiomorphic in that respect and Pollanisus and Onceropyga apomorphic. Thyrassia is a unique and highly derived genus with most species occurring outside Australia. In this group we find many morphological reductions and striking apomorphies in habitus and larval characters. At present no closely allied genus is known and the systematic and phylogenetic position of Thyrassia is preliminary.

55


A possible history of Australian Zygaenidae It is a most fascinating task to develop theories about the origin and dispersal of Zygaenidae on the Australian continent and in Tasmania. Alberti (1954: 296) stated that in his opinion all Australian zygaenids have come from Asia and are related to Artona Walker, 1854. He based this opinion on the following: 1. male: uncus, valva and aedeagus with similar characters; 2. female: praebursa present in most genera and similar to Artona; 3. wing venation with various reductions in the Australian groups. Alberti was correct in recognising a relationship between the Australian zygaenids and the Asian genera around Artona. No relationship to the American, and only a weak relationship to some African genera, has been found so far, and all known Australian species belong to the tribe Artonini of Procridinae (see p. 3). However, many Australian genera lack the characteristic ‘Artonafinger’ in the male genitalia. An ‘Artona-finger’ is present in almost all Asian and some African Artonini (exceptions in Asia are, for example, Thyrassia and Thibetana Efetov & Tarmann, 1995). This indicates that if this absence is not a secondary reduction (as it possibly is in Thyrassia and Thibetana), it must be plesiomorphic. Therefore, the Australian genera without an ‘Artona-finger’ in the male genitalia (i.e. Pollanisus, Onceropyga, Hestiochora, Turneriprocris and Myrtartona gen. n.) must have developed independently from the rest of Artonini, from ancestors that had split off from the Old World groups before the ‘Artona-finger’ was developed. Moreover, their vesica seminalis lacks deciduous cornuti, another important synapomorphy for most Asian genera. This corresponds closely with the female genitalia. There is a praebursa in Pollanisus, Onceropyga and Myrtartona with all the main Artonini characters (i.e. ductus intrabursalis inserting in centre of praebursa at a groove-like structure and the praebursa with appendix to accommodate the large cornutus of vesica) but still without the characteristic spiny crest that is developed to rub off the deciduous cornuti on the vesica. Of course, this crest is not necessary if there are no deciduous cornuti present in the male vesica.

The wing venation is rather underived in Pollanisus but is reduced in most other genera. This is also the case in the Asian Artonini if compared with Procridini. Based on this information the following scenario is suggested. After the splitting of the Artonini from the Procridini somewhere in the south-eastern Palaearctis or the Oriental region, only the Artonini invaded the Australian continent. The main characters of the Artonini had already been developed: 1, chaetosemata triangular, extending forward between compound eyes (e.g. Figs 6, 61); 2, valva fan-shaped, without processes (e.g. Fig. 136). These traits are present in a very basic form in Pollanisus and can also be easily recognized in Onceropyga (e.g. Fig. 278). However, both genera have developed various autapomorphies, the most important being the abdominal hair-tuft on the abdominal end of the females combined with a special habit to protect the eggs from predators (Figs 125, 129, 130). The wasp-like Hestiochora share this character as a synapomorphy but they have also developed many autapomorphies. Pollanisus, Onceropyga and Hestiochora, although clearly monophyletic (see also p. 55), are very different in habitus and biology. Pollanisus and Hestiochora have a wide distribution in all temperate parts of Australia, including Tasmania, their larvae feeding on plants endemic to Australia (e.g. Australian Dilleniaceae in Pollanisus and Eucalyptus spp. in Hestiochora). Moreover, many characters are very uniform in these two genera (e.g. genitalia in Pollanisus and habitus in Hestiochora). This indicates a fairly long development in isolation. Early in the development of Zygaenidae in Australia the genera Turneriprocris and Myrtartona must have split off from the rest of the group. This must have been before the transformation of the female abdomen into the ‘hair-tuft type’ started. The larvae of both genera feed on Myrtaceae (i.e. Leptospermum, Melaleuca, Kunzea). Myrtartona species have a praebursa with all the basic characters of Artonini but they have also developed striking autapomorphies in males and females (e.g. Figs 352, 360, 362, 363, 367, 368, 374–375, 377). Turneriprocris must be an early offshoot from ancestral Myrtartona with a very primitive wing venation

58


but with reductions in the male and female genitalia (Figs 336–343). Later, further invasions of the Australian continent by zygaenids of the Artonini type must have taken place; Artonini with a well developed ‘Artonafinger’ began to reach Australia. The earliest invasion may have resulted in one interesting relict species (i.e. Australartona mirabilis sp. n.) (Figs 378–384). Australartona mirabilis is confined to temperate mountainous rainforest biotopes, its larva feeding on a climbing grass (see p. 187, and Pl. 62, Fig. 6; Pl. 64, Fig. 4). Poaceae are common larval host-plants of east and Southeast Asian Artonini and many of these species also prefer a humid but more temperate climate. Subsequent invasions brought a further group of species to the northernmost rainforest areas of

Australia that are closely related to the tropical zygaenid fauna of Southeast Asia and New Guinea. These genera (i.e. Pseudoamuria gen. n., Homophylotis and Thyrassia) reach their southern distributional limit in northern Queensland and have more species outside Australia. The important coconut palm pest, Palmartona catoxantha, has been collected only once in Australia and may now have disappeared or the specimen may be mislabelled. It belongs to the Southeast Asian species-group and it is not clear whether this species is an indigenous element of the Australian fauna. However, it is possible for wind-assisted invasions of zygaenids from New Guinea, or accidental introductions by human transport, to occur at any time.

Checklist of Australian Zygaenidae

ZYGAENIDAE Latreille, 1809: 189 PROCRIDINAE Boisduval, [1829]: 38 ARTONINI Tarmann, 1994: 120 [Type-genus: Artona Walker, 1854: 439]

Pollanisus Walker, 1854: 114 [Type-species: Procris viridipulverulenta Guérin-Méneville, 1839: pl. 11]

angustifrons sp. n. Pollanisus sp. 2 Pollanisus sp. 3 Pollanisus sp. 4 Pollanisus sp. 5 Pollanisus sp. 6 Pollanisus sp. 7 Pollanisus sp. 8

viridipulverulenta (Guérin-Méneville, 1839: pl. 11) (Procris) cupreus Walker, 1854: 115 nielseni sp. n. modestus sp. n. lithopastus Turner, 1926b: 443

cyanota (Meyrick, 1886: 793) (Procris) cyanotas [sic] Aurivillius, 1920: 43, misspelling. isolatus sp. n. calliceros Turner, 1926a: 117 calliceros calliceros Turner, 1926a: 117 calliceros azurea ssp. n.

Pollanisus sp. 1 empyrea (Meyrick, 1888: 927) (Procris) amethystina (Meyrick, 1888: 927) (Procris)

Onceropyga Turner, 1906: 137 [Type-species: Onceropyga anelia Turner, 1906: 137] Ocneropyga [sic] Bryk, 1936: 259, misspelling.

apicalis (Walker, 1854: 111) (Procris) sequens Walker, 1854: 115 novaehollandiae (Wallengren, 1860: 39) (Procris)

anelia Turner, 1906: 137

trimacula (Walker, 1854: 110) (Procris)

pulchra sp. n.

edwardsi sp. n. subdolosa (Walker, [1865]: 32) (Procris) subdolosa subdolosa (Walker, [1865]: 32) (Procris) subdolosa clara ssp. n. contrastus sp. n. Pollanisus cf. contrastus 1 Pollanisus cf. contrastus 2 eungellae sp. n. acharon (Fabricius, 1775: 556) (Zygaena) acheron [sic] Bryk, 1936: 117, misspelling. eumetopus Turner, 1926: 443 commoni sp. n. incertus sp. n.

Hestiochora Meyrick, 1886: 788 [Type-species: Procris tricolor Walker, 1854: 111] Histiochora [sic] Pagenstecher, 1900: 44, misspelling. erythrota Meyrick, 1886: 789 tricolor (Walker, 1854: 111) (Procris) continentalis sp. n. furcata sp. n. queenslandensis sp. n. xanthocoma Meyrick, 1886: 788 xanthosoma [sic] Kirby, 1892: 86, misspelling. occidentalis sp. n. intermixta sp. n.

60


Turneriprocris Bryk, 1936: 304 [Type-species: Procris dolens Walker, 1854: 112] Neoprocris Turner, 1926b: 445, praeocc. Turneriptocris [sic] Tarmann, 1994: 121, fig. 3, misspelling. dolens (Walker, 1854: 112) (Procris)

Myrtartona gen. n. [Type-species: Procris rufiventris Walker, 1854: 110] coronias (Meyrick, 1886: 792) (Procris), comb. n. leucopleura (Meyrick, 1886: 792) (Procris), comb. n. rufiventris (Walker, 1854: 110) (Procris), comb. n. mariannae sp. n.

Australartona gen.n. [Type-species: Australartona mirabilis sp. n.] mirabilis sp. n.

Homophylotis Turner, 1904: 243 [Type-species: Homophylotis thyridota Turner, 1904: 243] thyridota Turner, 1904: 243 pseudothyridota sp. n. artonoides sp. n.

Pseudoamuria gen. n. [Type-species: Pseudoamuria uptoni sp. n.] uptoni sp. n. neglecta sp. n.

Palmartona gen. n. [Type-species: Brachartona catoxantha Hampson, [1893]: 233] catoxantha (Hampson, [1893]: 233) (Brachartona), comb. n.

Thyrassia Butler, 1876: 355 [Type-species: Syntomis subcordata Walker, 1854: 132] inconcinna Swinhoe, 1892: 55 mimetica (Turner, 1902: 200) (Monoschalis)

Taxonomy Family ZYGAENIDAE Latreille, 1809

Procridini are distributed world-wide, with the exception of Australasia. The Artonini are restricted to the Afrotropical Region, south, southeast and east Asia, and the Australian Region.

Subfamily PROCRIDINAE Boisduval, [1829]

Tribe Artonini Tarmann, 1994

Procridinae Boisduval, [1829], Europaeorum Lepidopterorum Index methodicus: 38 (as Procridae).Type genus: Procris Fabricius, 1807.

Artonini Tarmann, 1994, Nota lepid., Suppl. 5: 120. Type genus: Artona Walker, 1854, List Specimens lepid. Insects Colln Br. Mus 2: 439.

Diagnosis

Diagnosis

Small to medium-sized, usually day-flying moths. Wing shape and coloration variable. Most species are unicolorous but richly coloured species are also known. A very brilliant, sometimes metallic sheen is typical for some genera. Procridinae are separated from other zygaenids by the following characters (Tarmann 1994: 117; Efetov and Tarmann 1995: 63, 1999: 8; Efetov 2001: 11): 1. Female genitalia with ductus seminalis lacking a pseudobursa (‘bulla seminalis’ sensu Alberti 1954); 2. lagena absent in the receptaculum seminis; 3. a pair of accessory glands present, homologous to ‘Petersen’s gland’ in the Zygaeninae, close to the oopore; 4. larva with an unspecialised chemical defense system (without structurally developed opening mechanisms distributed over large parts of the integument). Characters 2 and 3 are supposed to represent good autapomorphies for the subfamily, supporting their monophyletic origin. Larvae of the Procridinae are leaf-miners, leafskeletonizers, borers, or external feeders. The Procridinae are the only subfamily of zygaenids with a world-wide distribution, whereas all other subfamilies have more restricted distributions. On the American and Australian continents they are the only representatives of the family. The subfamily Procridinae is divided into the tribes Procridini Boisduval, [1829] and Artonini Tarmann, 1994. The Procridini are a paraphyletic group containing a mixture of several, possibly monophyletic, subgroups that may have to be described as separate tribes in the future. The Artonini are monophyletic (see below). The

The Artonini are monophyletic, based on the following autapomorphies: 1. Chaetosema triangular, strongly extending forward between the compound eye and the ocellus (Figs 6, 61). In all other Procridinae the space between the compound eye and the ocellus is naked or covered with smooth, flat scales and the chaetosema is not extended (Fig. 88). However, this character is slightly variable; some American Procridinae and species of the Asian genus Clelea Walker, 1854 show intermediate characters. 2. Valva fan-shaped with a more strongly sclerotized dorsal and ventral margin and a very translucent and strongly folded central part. In its basic, simple, form this ‘artonoid valva’ is found in some Australian genera, e.g. Pollanisus (Fig. 136), but complex structures are found in many tropical genera, e.g. Artona, Palmartona, Pseudoamuria (Figs 402–406, 412–415).

Key to Australian genera 1.

– 2.

– 3.

–

With reduced wings (female) (Pl. 32, Fig. 2) . . . Pollanisus (Pollanisus calliceros Turner only) (p. 106) With normal wings . . . . . . . . . . . . . . . . . . . . . . 2 Parts of body and wings with shiny, metallic scales, females with abdominal hair-tuft (Pls 1–32) . . . . . . . . . . . . . . . . . Pollanisus (p. 63) Without shiny, metallic scales . . . . . . . . . . . . . 3 Ctenuchid-like habitus with large forewings and very small hindwings (Pl. 57) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Thyrassia (p. 215) Other habitus . . . . . . . . . . . . . . . . . . . . . . . . . . 4

62


4. – 5. – 6.

–

Hindtibia with three spurs, a single medial spur present (Fig. 121a) . . . .Palmartona (p. 207) Hindtibia with two spurs (Fig. 121b) . . . . . . . 5 Antenna with a conspicuous white spot distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .6 Antenna without white spot distally . . . . . . . . 7 Male antenna with asymmetrical pectinations. Male genitalia: valva very simple, without or with only a very short ‘Artona-finger’ and with a long and slender finger-like prolongation of the dorsal edge proximally bearing some setae distally that point backwards; vesica seminalis without cornuti (Figs 386–396). Female genitalia: without praebursa (Figs 397–401). Hindwing translucent centrally (scales upright) in the Australian species (Pls 51–54) . .Homophylotis (p. 191) Male antenna with symmetrical pectinations. Male genitalia: valva with well-developed, moveable ‘Artona-finger’, finger-like prolongation of the proximal dorsal edge of valva replaced by a short rounded process that is covered with many setae distally pointing upwards; vesica seminalis with a bundle of deciduous, eversible cornuti (Figs 402–409). Female genitalia: praebursa well developed with rows of spines and a pocket to accommodate the spiny, deciduous cornuti of the male vesica (Fig. 410). Hindwing not translucent centrally (Pl. 50, Fig. 2; Pl. 55) . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudoamuria (p. 201)

7. – 8. – 9. – 10. –

11. – 12. – 13. – 14. –

Wasp-like habitus, females with abdominal hairtuft (Pls 35–43) . . . . . . Hestiochora (p. 139) Not wasp-like . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Male (antenna bipectinate, frenulum represented by one single spine) . . . . . . . . . . . . . 9 Female (antenna biserrate, frenulum represented by three small bristles). . . . . . . . . . 12 Antenna with long pectinations (e.g. Pl. 48, Fig. 2) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 Antenna with short pectinations (e.g. Pl. 48, Fig. 1) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 Forewing matt, brown, with darker spot at end of cell (Pls 33, 34) . . . . . Onceropyga (p. 133) Forewing shiny varnish-like, olive-green to blue-green with a purple tinge (Pl. 48, Fig. 2; Pl. 49, Fig. 1; Pl. 50, Fig. 1) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Australartona (p. 185) Proboscis dark brown . . Turneriprocris (p. 163) Proboscis yellow . . . . . . . . Myrtartona (p. 169) Abdominal hair-tuft present (Pl. 33, Fig. 2; Pl. 34, Fig. 2) . . . . . . . . . Onceropyga (p. 133) Abdominal hair-tuft absent . . . . . . . . . . . . . . 13 Proboscis dark brown . . Turneriprocris (p. 163) Proboscis yellow . . . . . . . . . . . . . . . . . . . . . . . 14 Female genitalia without praebursa (Fig. 385) . . . . . . . . . . . . . . . . . . . Australartona (p. 185) Female genitalia with large praebursa (Figs 373, 376) . . . . . . . . . . . . . . . . Myrtartona (p. 169)

TAXONOMY

Genus Pollanisus Walker Pollanisus Walker, 1854, List Specimens lepid. Insects Colln Br. Mus. 1: 114. Type species: Procris viridipulverulenta Guérin-Méneville, 1839: pl. 11, by subsequent designation by Kirby (1892: 87). [Australia: ‘Nova Hollandia’.] Pollanisus Walker; Meyrick 1886: 788; Kirby 1892: 15; Turner 1906: 137; Jordan 1907: 10; Strand 1915: 118; Hering 1922: 13; Turner 1926b: 442; Bryk 1936: 117; Turner 1939: 110; Alberti 1954: 297; Common 1970: 766, fig. 36.1.A. (head), 829, fig. 36.35.A. (wing venation); Fletcher & Nye 1982: 129; Common 1990: 297 (bionomics); Moore, Brown and Rothschild 1990: 45 (biochemistry); Tarmann 1994: 121 (bionomics); Efetov and Tarmann 1995: 64; Tarmann 1996: 143; Epstein et al. 1998: 173; Efetov and Tarmann 1999: 18.

Diagnosis Species of different size and with variable wing shape (Pls 1–32). Body and wings sometimes partly, sometimes completely covered with blue, green, golden or coppery metallic scales. Some species with upperside of forewings densely scaled, shiny metallic blue or green, strongly resembling Palaearctic Jordanita species.

Redescription Head (Figs 1–13) Frons projected, its breadth varying in the different species and representing a good diagnostic character, usually slightly broader in the female; maxillary palps very small; proboscis brown or yellow; labial palps short, slightly upturned; pilifer short; compound eye of varying size, usually significantly smaller in the female; ocelli small to large, white or black; chaetosemata strongly developed, artonoid; male antenna bipectinate proximally, biserrate or simple distally, pointed apically; female antenna biserrate, dentations close together, giving the antenna a groove-like appearance ventrally. Thorax Foreleg without tibial epiphysis; tibial spurs 0–2–2, of equal length on midleg, the outer spur slightly longer on hindleg, no single medial spur on hindtibia as in many other genera of the Artonini. Wings broad or narrow, triangular or slightly rounded, length of hindwing approximately 70–90% that of forewing. Forewing with all veins

free, medial stem present or absent; hindwing with Sc fused with cell for a short distance, M3+CuA1 fused, medial stem present, sometimes slightly reduced proximally. Male frenulum a strong spine, retinaculum a well developed hook at base of Sc; female frenulum consisting of three spines in one row, inserted into a retinaculum consisting of a row of anteriorly pointing scales at base of CuP. One species, P. calliceros Turner, has a brachypterous female. It is the only known zygaenid species that has the wings reduced. Pregenital abdomen Abdomen with lateral evaginations on segment II only, the evaginations being very small and almost invisible. This has been interpreted as a secondary loss of the evaginations on segment VII (Tarmann, 1994: 121). Female with striking yellow or orange abdominal hairtuft (except for P. cyanota and P. calliceros that have a brownish green or greyish ochreous abdominal hairtuft). Sternum II (Fig. 426) with a pair of short, almost straight apodemes, lateral sternal rod and longitudinal rod weakly developed; sclerotized part of sternum II with broad lateral prolongations anteriorly. Tergum II with approximately 50–70% of segment sclerotized posteriorly. Sternum VII and VIII in the female weakly sclerotized and taxonomically significant. The translucent parts of segments VII and VIII enlarged in the female, the epidermis with smooth and sticky surface bearing long, needle-like scales that form the abdominal hairtuft. Genitalia The genitalic differences in Pollanisus are usually insufficient to recognise species. There is a remarkable variability within species in the shape of the valva, length and shape of the cornutus, and in the sclerotizations of the praebursa. Most structures are very translucent and their form is flexible and variable, depending on their treatment or state, for example, a valva when completely spread is almost twice as broad as when not spread, and similar changes occur with the presence or absence of a spermatophore in the praebursa. Very careful dissections of a series using constant methods are therefore essential to show the few characters that may be used for recognising species-groups. This is very unusual in the Procridinae, as normally there is uniformity in their external appearance but striking differences in their genitalia. Only two genera are known world-wide where genitalic characters seem to be inadequate for determination: Harrisina

63

64


Packard, 1864, in the Americas and Pollanisus in Australia. The following treatment of the genus is therefore based on external characters. The discovery of Pollanisus nielseni sp. n. in Western Australia by serendipity only (see p. 76) shows that even more species may still exist than we are able to recognise at present. In this work, only those species that possess clear diagnostic characters, and where enough material is available, are described as new. Some species, in which a formal description would have to be based on insufficient material, are figured and discussed but not named (Pollanisus sp. 1 to sp. 8). Male genitalia (Figs 136–239) Very uniform, with small specific differences. Uncus long and slender, with two groups of long setae basally, tegumen and vinculum weakly sclerotized, slender, saccus not pronounced. Valva long and slender, fan-shaped with the dorsal and ventral margin more strongly sclerotized, and with a very translucent, strongly folded central part, without processes, sacculus well developed, with long setae. Pulvinus well developed, oval, with long setae. Juxta rounded basally, with a slender medial prolongation dorsally. Aedeagus stout, everted vesica double-lobed, with a strongly sclerotized, almost straight, long, slender cornutus, a group of weakly sclerotized, needleshaped spines, and sometimes with a lateral plate-like sclerotization. Female genitalia (Figs 240–272) All parts very translucent. Ostium and ductus bursae not sclerotized. Praebursa well developed, very translucent, posterior part in some species slightly sclerotized or with ring-like structures, with a small, translucent, lateral appendix. The ductus intrabursalis inserts into the praebursa at a point where a strongly sclerotized thorn- or hook-like, sometimes dentate structure is developed. This sclerotization is very variable in shape and size even within populations (e.g. Figs 250, 251). It is used for rupturing the wall of the spermatophore that is stored in the praebursa; from there the spermatozoa are transported through the ductus intrabursalis into the very translucent corpus bursae where they can frequently be seen forming cloud-like aggregations. The pair of abdominal glands (homologous to Petersen’s gland in Zygaeninae) close to oopore absent.

Bionomics The larvae of Pollanisus species are brightly coloured (Pl. 59, Fig. 6; Pl. 60, Figs 1–3). Adult

larvae have the normal two pairs of lateral glands on abdominal segments 2 and 7 (the posterior pair on segment 7 is reduced in pupa and imago). As far as is known, most Pollanisus species feed on Dilleniaceae. Various Hibbertia spp., Dillenia alata (DC.) Martelli, and Tetrocera nordtiana F. Muell. are confirmed food sources. One species has been reared from larvae found on Pipturus argenteus (G. Forster) Wedd. (Urticaceae) and another from Kennedia rubicunda Vent. (Fabaceae). The eggs are deposited singly, or in small, flat, layers on the leaves of the foodplant and are covered with the deciduous and possibly poisonous scales of the abdominal hairtuft of the females, giving them a pincushion-like appearance (Pl. 59, Figs 1, 2; Figs 125, 436–438). The brightly coloured larvae are usually nocturnal (Pl. 59, Fig. 6; Pl. 60, Figs 1–3; Figs 132, 133). Pupation takes place in a silken cocoon either on the ground between leaves and litter or on stems, twigs and leaves of the foodplant (Fig. 134). The adults are diurnal and are most active in bright sunshine. Several specimens have been collected at mercury vapour light, especially in light traps. Other Procridinae are also occasionally attracted to light, although they are normally only diurnal. The adults often feed at the flowers of Asteraceae such as Helichrysum spp. or the naturalised Arctotheca calendula (L.) Levyns (originally African) (Pl. 60, Fig. 6). Species of the viridipulverulenta-group prefer open bushland or heath whereas those of the eumetopus-group seem to be restricted to tropical forests.

Remarks There are several Pollanisus specimens in the collections that almost certainly belong to various undescribed species. A formal description is postponed because of insufficient material. These species are briefly described and figured in colour following on from the presumably most closely related named species (pp. 79, 102–104).

Distribution Restricted to the temperate parts of Australia (south-western parts of Western Australia, southernmost South Australia, Tasmania, Victoria, New South Wales and south-eastern Queensland) with the exception of the eumetopus-group that occurs along the coast of Queensland northwards to Cooktown, with a single specimen known from Darwin (Northern Territory).

TAXONOMY

Key to Pollanisus species Pollanisus species are very variable in colour and size and have very uniform genitalic characters with a remarkable variability in details. Determination is therefore extremely difficult. This key is mainly based on external characters as the genitalic characters easily lead to incorrect conclusions. Genitalic characters are only used if constant and significant. The key is based on the examined material and does not include all possible colour variations and aberrations. Moreover, some species are only known from one sex. The key leads to a species or a speciesgroup. In some cases it is necessary to read the differential diagnoses at species level and compare all described characters carefully to come to a satisfactory conclusion. Those species that are supposed to represent undescribed taxa but are not described in this book are not included in the key. They are figured in colour and a short discussion of characters is provided. 1. –

Male (antenna bipectinate, frenulum represented by one single spine) . . . . . . . . . . . . . 2 Female (antenna beserrate, frenulum represented by three small bristles) . . . . . . . . . 22

Male 2. Forewing upperside with shiny scales covering most parts of the wing (Pls 1–10; Pl. 31, Fig. 2; Pl. 32, Fig. 1) . . . . . . . . . . . . . . . . . . . 3 – Forewing upperside without or with only a few shiny scales that are restricted to the proximal part of the costal margin (Pls 11–30) . . . . 11 3. Patagia, tegulae and at least anterior dorsal parts of thorax shiny, unicolorous (Pl. 22) . . . . . 4 – Patagia shinier than rest of thorax, forewing upperside blackish brown, matt, with shiny, metallic, silvery, bluish green or golden green scales, scattered singly (Pl. 6) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. lithopastus (p. 77) 4. From Western Australia . . . . . . . . . . . . . . . . . . 5 – Not from Western Australia . . . . . . . . . . . . . . . 7 5. Forewing upperside dark grey brown, densely covered with metallic, golden, coppery scales proximally, less so more distally, without shiny scales along margin. Large species, forewing length 9.5–11.5 mm (Pl. 4) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. cupreus (p. 72) – Forewing upperside completely covered with metallic, shiny scales except for a small margin. Smaller species, forewing length less than 10.0 mm (Pls 5, 8) . . . . . . . . . . . . . . . . 6

6.

Forewing upperside bright shiny golden yellow and green except for a narrow distal band that is blackish grey; hindwing very dark blackish grey, opaque (Pl. 5) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. nielseni sp. n. (p. 75) – Forewing upperside metallic coppery golden, golden green or bluish green, distal margin and fringe blackish brown; hindwing grey brown, slightly translucent medially (Pl. 8) . . . . . . . . . . . . . . . . . . . . . . P. empyrea (p. 79) 7. Forewing upperside brownish black, densely covered with shiny metallic bluish green scales fading distally, with stronger sheen in basal half. Hindwing opaque. Small species, forewing length 7.5 mm (Pl. 3, Fig. 2) . . . . . . . . . . . . . . . . P. modestus sp. n. (p. 76) – Forewing upperside completely covered with metallic, shiny scales except for a small margin (Pls 1, 2, 9, 32) . . . . . . . . . . . . . . . . 8 8. Frons very broad, more than 2.0× broader than compound eye in frontal view; antenna short, less than 30 segments, with extremely long pectinations; wings rounded. Small species, forewing length 6.5–7.5 mm (Pl. 31, Fig. 2; Pl. 32, Fig. 1) . . . . . . . . . . . . . . . . . . . 9 – Frons less than 2.0× broader than compound eye in frontal view; antenna longer, more than 40 segments, with normal pectinations; wings triangular or elongate triangular. Larger species, forewing length more than 7.5 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 9. Shiny scales on body, forewing upperside and hindwing underside light green (Pl. 31, Fig. 2). Southernmost eastern New South Wales, Victoria, Tasmania . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. calliceros calliceros (p. 106) – Shiny scales on body, forewing upperside and hindwing underside bluish green (Pl. 32, Fig. 1). Eastern and north-eastern New South Wales . . . P. calliceros azurea ssp. n. (p. 107) 10. Forewing broad, strongly triangular, hindwing quadrangular; forewing upperside covered with very shiny metallic yellowish green, dark green, bluish green or bluish scales, hindwing opaque. Large species, forewing length 11.0–13.0 mm (Pls 1, 2) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. viridipulverulenta (p. 67) – Forewing elongate triangular, hindwing subtriangular, rounded distally; forewing upperside covered with shiny metallic blue, bluish green or golden green scales, hindwing slightly translucent medially. Smaller species, forewing length 7.5–9.5 mm (Pl. 9; Pl. 10, Fig. 1) . . . . . . . . . . . . . . . . . P. apicalis (p. 81)

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11. Frons very broad, more than 2.5× broader than compound eye in frontal view; wings short, hindwing broad, rounded, thorax dark dorsally with well separated shiny patagia forming a conspicuous collar that is in strong contrast to the shiny blue or bluish green abdomen . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 – Frons less than 2.5× broader than compound eye in frontal view, hindwing more slender, elongate quadrangular or triangular, thorax without collar or with a collar that is not in strong contrast to the shiny scales on the upperside of the abdomen . . . . . . . . . . . . 13 12. Collar reddish, forewing upperside sprinkled with single white scales (Pl. 30). Aedeagus less than 3× longer than broad P. cyanota (p. 104) – Collar golden, forewing upperside not sprinkled with white scales (Pl. 31, Fig. 1). Aedeagus ca. 5× longer than broad P. isolatus sp. n. (p. 105) 13. Forewing upperside with variable whitish ochreous spots (that may be completely absent); underside of wings always with three streaks of metallic golden green scales along costa of forewing and hindwing (both more or less of equal length) and at anal angle of hindwing (Pl. 11) . . . . . P. trimacula (p. 84) – Forewing upperside without whitish ochreous spots; underside of wings never with three streaks of metallic golden green scales along costa of forewing and hindwing and at anal angle of hindwing . . . . . . . . . . . . . . . . . . . 14 14. Patagia and anterior margin of tegulae shiny greenish blue, abdomen greyish black with shiny, metallic, dark steel-blue scales with a slightly variable green tinge on segments 2 to 8 dorsally; frons very broad, approximately 2.0× broader than compound eyes in frontal view. Small species, forewing length 6.0–7.0 mm (Pl. 20) . . . . . . . . . P. eumetopus (p. 97) – Patagia, tegulae and abdomen of other colour, frons less than 2.0× broader than compound eye in frontal view . . . . . . . . . . . . . . . . . . . 15 15. Frons significantly broader than breadth of compound eye in frontal view . . . . . . . . . 16 – Frons as broad as or narrower than breadth of compound eye in frontal view . . . . . . . . . 19 16. Forewing length more than 9.0 mm; wings broad, forewing broad triangular, hindwing almost rectangular but rounded apically, the only slightly translucent hindwing not contrasting strongly with the opaque forewing (Pl. 12). . . . . . . . . . .P. edwardsi sp. n. (p. 86) – Forewing length less than 9.0 mm; wings more slender . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

17. Patagia, anterior part of tegulae, 20–30% of anterior part of thorax and dorsal part of abdomen shiny metallic copper, the sheen not very intensive; forewing very opaque, hindwing translucent medially, both in strong contrast, (Pl. 17). Forewing length 7.0–7.5 mm. From central-eastern New South Wales to south-eastern Queensland . . . . . . . . . . . . . . . . . P. contrastus sp. n. (p. 91) – Patagia, anterior part of tegulae and dorsal part of abdomen strongly shiny metallic . . . . . 18 18. Patagia and tegulae covered with yellowish green, shiny metallic scales tinged with light blue, abdomen shiny metallic bluish tinged with yellowish green dorsally; forewing narrow, subtriangular, hindwing slightly translucent medially (Pl. 19). Forewing length 6.0 mm (only male type known). Known only from Eungella (central-eastern Queensland) . . . . . P. eungellae sp. n. (p. 94) – Patagia, anterior half of tegulae, anterior margin of dorsal mesothorax and dorsal abominal segments 3 to 8 shiny metallic golden; forewing round elongate, opaque, hindwing very translucent medially (Pl. 21). Forewing length 7.0–7.5 mm. Coastal regions of north-eastern Queensland . . . . . . P. commoni sp. n. (p. 98) 19. Forewing length 7.5 mm or less. Patagia and anterior half of tegulae shiny metallic yellowish green with a bluish tinge, posterior abdominal segments shiny metallic yellowish green dorsally (Pl. 23). Coastal region of north-eastern Queensland .P. angustifrons sp. n. (p. 100) – Forewing length more than 7.5 mm. Patagia and anterior half of tegulae shiny metallic copper, posterior abdominal segments shiny metallic coppery-golden dorsally . . . . . . . . . . . . . . 20 20. Forewing broad, triangular, hindwing subquadrate. Temperate and subtropical rainforests from Victoria to south-eastern Queensland . . . . . . . . . . . . . . . . . . . . . . . . 21 – Wings more slender, elongate (Pl. 22). Coastal tropical forest of north-eastern Queensland . . . . . . . . . . . . . . . . . P. incertus sp. n. (p. 99) 21. Hindwings only slightly translucent, metallic copper sheen on collar and abdomen very dull (Pl. 13). Victoria, New South Wales (southern coastal areas) . . . . . . . . . . . . P. subdolosa subdolosa (p. 87) – Hindwings more translucent, metallic coppery sheen on collar and abdomen stronger (Pl. 14; Pl. 15, Fig. 1). New South Wales (northern coastal parts), south-eastern Queensland . . . . . . . . . . P. subdolosa clara ssp. n. (p. 89)

TAXONOMY

Female 22. Wings reduced (Pl. 32, Fig. 2) . . . . . . . . . . . . . . . . . . . . P. calliceros (p. 106) – Wings normal . . . . . . . . . . . . . . . . . . . . . . . . . 23 23. Abdominal hairtuft greyish-ochreous, collar shiny reddish, forewing upperside sprinkled with single white scales (Pl. 30, Fig. 2) . . . . . . . . . . . . . . . . . . . . . . . . . . P. cyanota (p. 104) – Abdominal hairtuft yellow . . . . . . . . . . . . . . . 24 24. Forewing upperside with shiny scales covering most parts of the wing (Pls 1–5, 8–10) . . 25 – Forewing upperside without or with only a few shiny scales that are resricted to the proximal part of the costal margin (Pls 11–29) . . . 30 25. Patagia, tegulae and at least anterior dorsal parts of thorax shiny, unicolorous . . . . . . . . . . . 26 – Patagia much more shiny than rest of thorax, forewing upperside blackish brown, matt, with shiny, metallic, silvery, bluish green or golden green scales, scattered singly (Pl. 6) . . . . . . . . . . . . . . . . . . . . P. lithopastus (p. 77) 26. From Western Australia . . . . . . . . . . . . . . . . . 27 – Not from Western Australia . . . . . . . . . . . . . . 29 27. Forewing upperside dark grey brown, densly covered with metallic, golden, coppery scales proximally, less so more distally, without shiny scales along margin (Pl. 4) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. cupreus (p. 72) – Forewing upperside completely covered with metallic, shiny scales except for a small margin (Pls 5, 8) . . . . . . . . . . . . . . . . . . . . 28 28. Forewing upperside bright shiny golden yellow and green except for a narrow distal band that is blackish grey; hindwing very dark blackish grey, opaque (Pl. 5) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. nielseni sp. n. (p. 75) – Forewing upperside metallic coppery golden, golden green or bluish green, distal margin and fringe blackish brown; hindwing grey brown, slightly translucent medially (Pl. 8) . . . . . . . . . . . . . . . . . . . . . . P. empyrea (p. 79) 29. Hindwing opaque (Pl. 3, Fig. 1) . . . . . . . . . . . . . . . . . . . . . . . . . . P. viridipulverulenta (p. 67) – Hindwing slightly translucent medially. (Pl. 9, Fig. 2) . . . . . . . . . . . . . . . . . P. apicalis (p. 81) 30. Forewing upperside with variable whitish ochreous spots (that may be completely absent); underside of wings always with three streaks of metallic golden green scales along costa of forewing and hindwing (both more or less of equal length) and at anal angle of hindwing (Pl. 11) . . . . . P. trimacula (p. 84) – Forewing upperside without whitish ochreous spots; underside of wings never with three

streaks of metallic golden green scales along costa of forewing and hindwing and at anal angle of hindwing . . . . . . . . . . . . . . . . . . . 31 31. Patagia and anterior margin of tegulae shiny greenish blue, abdomen greyish black with shiny, metallic, dark steel-blue scales with a slightly variable green tinge on segments 2 to 8 dorsally (Pl. 20) . . . . P. eumetopus (p. 97) – Patagia, tegulae and abdomen not dark steelblue . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 32. Patagia, tegulae and abdomen shiny bluish (Pl. 19) . . . . . . . . P. eungellae sp. n. (p. 94) / P. acharon (p. 95) (The two species are not distinguishable from each other in the female, see p. 96) – Patagia, tegulae and abdomen not shiny bluish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .33 33. Patagia, tegulae and abdomen shiny copper (Pls 12–18) . . . .P. edwardsi sp. n. (p. 86) / P. subdolosa (p. 87) / P. contrastus sp. n. (p. 91) (These three species are not clearly distinguishable from each other in the female, see p. 93.) – Patagia, tegulae and abdomen shiny golden (Pls 21, 22) . . . . . . . . . P. commoni sp. n. (p. 98) / P. incertus sp. n. (p. 99) (The two species are not clearly distinguishable from each other in the female, see p. 99.)

Pollanisus viridipulverulenta (Guérin-Méneville) Pl. 1, Figs 1, 2; Pl. 2, Figs 1, 2; Pl. 3, Fig. 1; Pl. 59, Fig. 1; Pl. 60, Figs 3, 4, 5; Pl. 61, Figs 1–3; Figs 1–13, 99–104, 122–123, 125, 129–130, 138–141, 186–189, 240, 241. Procris viripulverulenta Guérin-Méneville, 1839, Magasin Zool. Paris (2) 1 (Insects): pl. 11, fig. 4. Type not found, [AUSTRALIA] ‘Nova Hollandia’. Procris viridipulverulenta Guérin-Méneville; Meyrick 1886: 794. Pollanisus viridipulverulentus (GuérinMéneville); Walker 1854: 114; Kirby 1892: 87; Aurivillius 1920: 43; Tillyard 1926: 435; Turner 1926b: 444. Pollanisus viridipulverulenta (Guérin-Méneville); Jordan 1907: 10; Hering 1922: 13; Bryk 1936: 119; Turner 1939: 110; Alberti 1954: 298; Fletcher and Nye 1982: 129; Common 1990: 297 (bionomics). Tarmann 1996: 143. Pollanisus adelaidae Turner, 1926, Proc. Linn. Soc. NSW 51: 444. Type not found. South Australia. Pollanisus adelaidae Turner; Tarmann 1996: 143.

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Material examined Queensland: 1 么, Acacia Ridge, 16. ix. 1965 (G. B. Monteith) (UQIC); 1 么, Archerfield, 20. ix. 1969 (V. Stablum) (UQIC); 1 么, Brisbane, viii. 1898 (DPIQ); 7 么, 2 乆, Brisbane, 3. x. 1912, 28. i. 1913, 5. x. 1913 (H. Hacker) (ANIC, QM); 12 么, 6 乆, Brisbane, ix. 1911, ix. 1914 (T. P. Lucas) (QM); 1 么, Brisbane, 20. ix. 1969 (D. L. Hancock) (UQIC); 1 么, 1 乆, Brisbane, May Orchard, ix. 1910 (QM); 4 么, 1 乆, Brisbane, Virginia, 21. ix. 1924 (T. H. Beck) (UQIC); 1 么, Burleigh, 16. ix. 1932 (UQIC); 3 么, Caloundra, 1.–3. ix. 1912, 4. ix. 1915 (ANIC, UQIC); 1 么, Carnarvon Ra., 19. xii. 1938 (W. B. Barnard) (QM); 3 么, 2 乆, Crows Nest, 31. viii. 1920, 6.–9. x. 1921, 14. iii. 1931 (AMS, ANIC, UQIC); 3 么, 1 乆, Duaringa (AMS, QM); 2 么, 1 乆, Goombungee, 31. viii. 1971, 23. ix. 1971 (J. Macqueen) (ANIC); 1 么, Milmerran, 16. ix. 1930 (ANIC); 4 么, Milmerran, 8. x. 1943, 3. ix. 1944, 11. x. 1944, 2. x. 1945 (J. Macqueen) (ANIC, UQIC); 1 么, Redcliffe, Blackford, 10. ix. 1938 (DPIQ); 3 么, 2 乆, Stanthorpe, ii. (ANIC); 1 么, Toowoomba, 6. x. 1921 (QM); 1 么, Warwick, x. (ANIC). New South Wales: 1 乆, Armidale, NE University, 6. xii. 1966 (C. W. Frazier) (ANIC); 1 么, Ashfield (AMS); 1 么, Ash I., (Scott) (AMS); 1 么, Bankstown, 22. ix. 1928 (AMS); 1 么, Barren Grounds, 22. xi. 1959 (C. E. Chadwick) (BCRI); 1 么, Barren Grounds, 13. x. 1962 (V. J. Robinson) (ANIC); 1 么, Barren Grounds Fauna Reserve, 22. x. 1978 (V. J. Robinson) (ANIC); 1 么, Bathurst, 13 mls E, 3200 ft, 27. x. 1964 (I. F. B. Common & M. S. Upton) (ANIC); 2 么, Berowra, 1.–29. xi. 1917 (ANIC); 20 么, 4 乆, Blackheath, 3.–6. xii. 1946, 1. xii. 1947, 3. xii. 1948, 22. xi. 1949, 11. xii. 1952, 17. xi. 1953, 22. xii. 1953 (L. H. MosseRobinson) (ANIC); 1 么, Blackheath, Hat Hill Rd, 11. xi. 1963 (AMS); 1 么, Boyd R., 4100 ft, 26. xi. 1976 (V. J. Robinson) (ANIC); 2 么, 1 乆, Bundanoon, 12.–15. x. 1918 (AMS, ANIC); 1 么, Clyde Mountain, 2400 ft, 17. x. 1962 (I. F. B. Common & M. S. Upton) (ANIC); 1 么, Ebor, 2. i. 1916 (ANIC); 1 乆, Fassifern, 31. x. 1950 (L. H. MosseRobinson) (ANIC); 2 么, Guildford, 30. ix. 1922 (AMS); 2 么, Helensburgh, xi. 1961, iii. 1962 (H. E. Osburne) (BCRI); 1 么, Helensburgh, 27. ix. 1962 (V. J. Robinson) (ANIC); 2 乆, Hoskinstown, 7 km WNW, Whiskers, 19. ii. 1990, 25. ix. 1993 (M. S. Upton) (ANIC); 1 么, Hoskinstown, 9 km E, South Black Ra., 1. x. 1980 (D. J. Ferguson) (ANIC); 2 么, Kanangra-Boyd Nat. Park, 6. xii. 1983 (at light) (B. Hacobian) (ANIC); 4 么, 2 乆, Katoomba, 3.–4. xi. 1909, xi. 1912 (AMS); 2 么, Leura, x. 1913 (AMS, ANIC); 1 么, Lucas Heights, 28. ii. 1970 (G. Daniels) (AMS); 2 么, 1 乆, Manly, 1. x. 1906, 11. x. 1908 (AMS); 2 么, Maroota, ix. 1960 (L. C. Haines) (AMS); 1 乆, Mittagong, 9. xi. 1928 (G. M. Goldfinch) (AMS); 1 么, Mittagong, 25. ii. 1932 (ANIC); 1 么, Mt Canobolas, 4500 ft, 8. xi. 1947 (V. J. Robinson) (ANIC); 1 么, Mt Kaputar, 11. iii. 1969 (V. J. Robinson) (ANIC); 1 么, Mt Irvine, 23. x. 1977 (G. R. Brown) (BCRI); 1 么, Mt Keira, 19. xi. 1966 (V. J. Robinson) (ANIC); 1 么, Mt Vincent, near running stream, 20. xi. 1978 (G. Daniels) (BCRI); 1 乆, Mt Wilson, 20. xi. 1923 (ANIC); 12 么, 2 乆, Mulgoa, 13. iii. 1951,

11.–15. iii. 1954 (L. H. Mosse-Robinson) (ANIC); 3 么, Narara, 26. ix. 1949, 29.–30. ix. 1950 (L. H. MosseRobinson) (ANIC); 2 么, Newcastle Firth, i. 1888 (AMS); 1 么, Oxford Falls, 22. x. 1961 (L. H. Mosse-Robinson) (ANIC); 1 乆, Port Stephens, 29. x. 1940 (AMS); 1 么, Richmond R., xi. 1935 (V. J. R[obinson]) (ANIC); 1 乆, Roseville, 26. ii. 1922 (L. H. Mosse-Robinson) (ANIC); 1 么, Rous, Richmond R., 2. viii. 1925 (V. J. Robinson) (ANIC); 1 么, Royal Nat. Park, Goondera Ridge, 8. x. 1969 (G. Daniels) (AMS); 1 么, 2 乆, Shoalhaven, 13 km NNW of Braidwood, 26. x. 1977 (D. C. F. Rentz & J. Balderson) (ANIC); 3 么, Sydney (W. W. Frogatt) (AMS); 2 么, Sydney, 16. x. 1906 (AMS, ANIC); 2 么, Sydney, 1924 (W. W. Frogatt) (AMS); 1 么, Sydney, Heathcote, 20. ix. 1973 (B. Hacobian) (ANIC); 1 乆, Tahmoor, 28. x. 1981 (B. J. Day) (AMS); 1 么, Tallong, 27. ii. 1935 (Hatter) (ANIC); 1 么, Uralla, 1914 (BCRI); 1 么, Watagan SF, near Wyong, 24. ii. 1979 (B. Hacobian, N. Goodman & V. J. Robinson) (ANIC); 1 么, Warrumbungle, 23. x. 1967 (V. J. Robinson) (ANIC); 3 么, Wirrimbirra, near Picton, 29.–30. xi. 1967 (V. J. Robinson) (ANIC); 8 么, Wilton, CSIRO Experimental Farm, 17.–20. xi. 1966, 22. xi. 1967, 24. ii. 1971, 18. iii. 1976, 2. xi. 1972, 8. iii. 1981, 7. x. 1981 (V. J. Robinson) (ANIC); 2 么, Young, 22. ii. 1935, 6. x. 1935 (Hatter) (ANIC). Australian Capital Territory: 31 么, 6 乆, Aranda Bushland, Canberra, 700 m, 12.–22. xi. 1989 (G. M. Tarmann) (GMT, ANIC); 14 么, 2 乆, Black Mountain, Canberra, light trap, 30. xi. 1955, 18. x. 1957, 29. x. 1959, 8. iii. 1960, 3.–5. xi. 1959, 12. xi. 1962, 20.–27. iii. 1963, 14. iii. 1967, 2. v. 1967 (I. F. B. Common) (ANIC); 2 乆, Black Mountain, N of summit trail, Canberra, 710 m, 17. xi. 2001 (G. M. Tarmann) (ANIC); 2 egg clusters on Hibbertia obtusifolia, Black Mountain, Canberra, 710 m, 25. xi. 2001 (G. M. Tarmann) (ANIC); 1 么, Jervis Bay, 6. x. 1912 (ANIC); 2 么, 1 乆, Jervis Bay, 13. x. 1918 (ANIC); 4 么, 2 乆, Jervis Bay, 6.–12. ix. 1920 (L. H. Mosse-Robinson) (ANIC); 1 么, Jervis Bay, 14. x. 1966 (I. F. B. Common & M. S. Upton) (ANIC). Victoria: 1 么, 4 乆, Beaconsfield, xii. 1903, 12. xi. 1904 (MV); 2 么, Belgrave, 12.–28. xii. 1943 (F. E. Wilson) (ANIC); 3 么, Big Desert, Hermies garden Track, 23. ii. 1992 (E. S. Nielsen & E. D. Edwards) (ANIC); 1 么, Big Desert, Milmed Track, 22. ii. 1992 (E. S. Nielsen & E. D. Edwards) (ANIC); 1 乆, Birchip, 27. xii. 1918 (ANIC); 2 乆, Boronia, i. 1930 (Kershaw) (MV); 3 么, Britton, 9. iii. 1934 (C. Ives) (ANIC); 3 么, 1 乆, Castlemaine, 15. iii. 1907, 8.–13. xi. 1907 (W. E. Drake) (MV); 2 么, 1 乆, Cheltenham (MV); 1 乆, Cheltenham, 25. x. 1919 (L. B. Thorn) (QM); 1 么, Croydon, 22. xi. 1908 (MV); 2 么, Dutson, 7. xi. 1901, ix. 1905 (M. I. Wise) (MV); 2 么, Eltham, 31. x. 1943 (F. E. Wilson) (ANIC, MV); 1 么, Emerald, 28. x. 1907 (G. Lyell) (MV); 1 乆, Fern Tree Gully, 2. xii. 1894 (MV); 1 么, Fern Tree Gully, 24. x. 1927 (AMS); 1 么, Frankston, 28. ix. 1919 (L. B. Thorn) (MV); 4 么, 7 乆, Gisborne, 10. xi. 1892, 18. xi. 1894, 16. xii. 1894, 11.–19. xi. 1895, 18. xii. 1899, 20. xi. 1901, 4.xii. 1904 (MV); 1 么, Gisborne, 5. xi. 1914 (L. H. Mosse-Robinson) (ANIC); 13 么, 2 乆, Gisborne, 14. xi.

TAXONOMY

1909, 8. xii. 1913, 12. xii. 1915, 7. xii. 1917, 3. xi. 1918, 11. xii. 1921, 21.–25. xi. 1926, 11. xii. 1929, 2. xi. 1930, 4. xii. 1932, 23. xi. 1934 (G. Lyell) (MV); 1 么, Halls Gap, 11. xi. 1942 (Le Souef) (ANIC); 2 么, 1 乆, Heathmont, 11.–24. x. 1927 (AMS); 9 么, 2 乆, Little Desert, 27. x. 1946 (C. G. L. Gooding) (ANIC); 4 么, Little Desert, 13 mls S of Kiata, 6.–7. xi. 1966 (I. F. B. Common & M. S. Upton) (ANIC); 11 么, Little Desert Nat. Park, 21. ii. 1992 (E. S. Nielsen & E. D. Edwards) (ANIC); 1 么, 1 乆, Longford, 5. ix. 1900, 30. ix. 1912 (M. L. Wise) (MV); 22 么, 8 乆, Moe, ii. 1917, 1. xi. 1918, xii. 1918, 19. ii. 1920, 17. xi. 1929, 4. xii. 1932, 7. iii. 1935, 12. xii. 1940, 12. iv. 1954, 20. xii. 1954 (C. G. L. Gooding) (ANIC, SAM); 1 么, Moondarra Dam Area, 22. xi. 1972 (C. G. L. Gooding) (ANIC); 1 么, Mt Macedon, 18. xi. 1895 (MV); 1 乆, Nelson, 11. xi. 1966 (I. F. B. Common & M. S, Upton) (ANIC); 2 么, Nhill, x. 1902 (J. R. F[rench]) (BCRI); 1 么, 1 乆, Ocean Grove, 23.–25. x. 1912 (SAW) (AMS); 2 么, Portland, 28 mls NW, 11. xi. 1966 (I. F. B. Common & M. S. Upton) (ANIC); 1 么, 1 乆, Ringwood, 11. xii. 1914 (MV); 1 么, 4 乆, Ringwood, 11. xii. 1913, 15. x. 1918, 25. x. 1920 (F. P. S[pry]) (MV); 1 乆, Ringwood, 13. xii. 1944 (F. E. Wilson) (ANIC); 1 么, Roseberry, 1. xi. 1944 (Le Souef) (ANIC); 2 么, Sandringham, 5. xi. 1905, 24. x. 1908 (MV); 1 乆, Sea Lake, x. 1904 (D. Goudie) (ANIC); 3 么, Shady Creek, 13 mls of Willow Grove, 20. xi. 1963 (C. G. L. Gooding) (ANIC); 2 么, ‘X. Y. Z.’ [Shady Creek, 17 km N Yarragon], 9. xii. 1974 (C. G. L. Gooding) (ANIC); 1 乆, Springvale, 3. xi. 1920 (F. P. Spry) (MV); 11 么, 2 乆, Traralgon, 15. xi. 1972, 4. xi. 1974 (C. Elton, C. G. L. Gooding) (ANIC); 1 么, 1 乆, Valencia, 16. xi. 1953 (Le Souef) (ANIC); 1 么, Willow Grove, 26. xi. 1966 (C. G. L. Gooding) (ANIC); 1 么, Willow Grove, 4 mls W, 17. x. 1972 (C. G. L. Gooding) (ANIC); 1 乆, Wyperfeld Nat. Park, 5. xi. 1966 (I. F. B. Common & M. S. Upton) (ANIC); 1 么, Wyperfeld Nat. Park, 27. ix. 1973 (I. Brooks) (ANIC), 1 么, Yinnar, 22. xi. 1947 (J. H. S) (AMS). South Australia: 7 么, 1 乆, Adelaide (SAM); 1 乆, Adelaide (G. L[yell]) (MV); 1 么, Adelaide, Ashley W, 25. xi. 1899 (SAM); 1 么, Aldgate, 25. xi. 1906 (F. M. Angel) (SAM); 3 么, Aldinga Scrub, 27. ix. 1966 (J. O. Wilson) (ANIC); 1 么, Aldinga Scrub, 28. x. 1976 (at UVL light) (P. B. McQuillan) (SAM); 1 么, Aldinga Dellicks Beach Reserve, 16. ix. 1987 (E. G. Matthews & J. A. Forrest) (SAM); 1 么, Alligator Gorge, 170 m, 8. x. 1962 (N. B. Tindale) (SAM); 2 么, Athelstone, 3.–7. xi. 1967, 4. x. 1973 (at light) (J. J. H. Szent-Ivany) (SAM); 1 么, 1 乆, Balhannah, 7. iii. 1885 (SAM); 1 么, Belair, 24. x. 1885 (J. G. O. Tepper) (SAM); 1 乆, Belair Nat. Park, Adelaide, 9. xi. 1964 (G. F. Gross) (SAM); 2 乆, Belair Nat. Park, Adelaide, 31. x. 1963 (G. F. Gross & H. Oldham) (SAM); 1 么, Belair Nat. Park, Adelaide, 2. xi. 1975 (K. J. Tremelling) (SAM); 4 么, Blackwood, 1899, 24. x. 1906, 31. x. 1939 (F. M. Angel) (ANIC, SAM); 2 么, 4 乆, Blackwood, 12. ix.–17. x. 1926 (J. O. Wilson) (ANIC); 1 么, Blackwood (S. Ashby) (SAM); 1 乆, Blackwood, 18. xi. 1935 (F. M. Angel) (SAM); 1 么, Bridgewater, 1. xii. 1932 (J. O. Wilson) (ANIC); 1 么, 2 乆, Bridgewater, 19. xi. 1939, 16. xi. 1941, 25. xi. 1943 (F. M.

Angel) (ANIC, SAM); 20 么, 8 乆, Burnside (SAM); 3 乆, Burnside, Mt Lofty Ra., 17. x. 1884 (J. G. O. Tepper) (SAM); 3 乆, Burnside, 17. ii. 1905, 18. xi. 1905 (F. M. Angel) (SAM); 1 么, Burnside, 12. x. 1908 (SAM); 6 么, Highbury (N. B. Tindale) (SAM); 1 么, Highbury, 16. ix. 1937 (N. B. Tindale) (SAM); 7 么, 4 乆, Highbury, 18. xi. 1905, ix. 1919, 15.–16. ix. 1927, 15. ix. 1940 (F. M. Angel) (SAM); 1 么, Inglewood, 10. xi. 1953 (J. O. Wilson) (ANIC); 1 么, Kangaroo I., Rocky Ra., xii. 1934 (Museum’s Expedition) (SAM); 1 乆, Marble Ra., Eyre Peninsula, 5. x. 1979 (G. F. Gross) (SAM); 1 么, 1 乆, Meningie, 10 km S, 17. xi. 1967 (J. Szent-Ivany) (SAM) 1 么, Mt Compass, 28. x. 1938 (F. M. Angel) (SAM); 1 么, 1 乆, Mt Compass, 18.–26. ix. 1962 (J. O. Wilson) (ANIC); 2 么, 1 乆, Mt Lofty, 30. x.–5. xi. 1924 (ANIC); 1 么, Mt Remarkable, 28. viii. 1955 (H. M. Cooper) (SAM); 2 么, Munchilla, 22 mls WSW, 8. xi. 1966 (I. F. B. Common & M. S. Upton) (ANIC); 1 乆, Murray Bridge (Lea) (SAM); 1 么, Myponga, 30. iii. 1934 (F. M. Angel) (SAM); 1 么, Port Lincoln (Lea) (SAM); 1 么, 4 乆, Port Victoria, xi. 1896 (E. Mellor, J. W. Mellor) (SAM); 3 么, 2 乆, Reynella, Mt Lofty, 7. xii. 1923 (ANIC); 1 乆, Reynella, Mt Lofty, 5. iv. 1924 (J. O. Wilson) (ANIC); 1 乆, Slapes Gully, 10. i. 1907 (SAM); 1 么, Stansbury, 8. x. 1957 (F. M. Angel) (SAM); 2 么, Stoneyfell, Adelaide, 1. x. 1889 (on Hibbertia stricta) (J. G. O. Tepper); 1 么, Stoneyfell, Adelaide, 1. x. 1906 (F. M. Angel) (SAM); 1 乆, Tea Tree Gully, ix. 1898 (J. M. Mellor) (SAM); 1 么, Tea Tree Gully, 30. i. 1938 (SAM); 2 么, 1 乆, Tea Tree Gully, 13. ix. 1959 (R. V. Southcott) (SAM); 1 么, Tintinara, 29. ix. 1952 (N. B. Tindale) (SAM); 3 么, Upper Sturt, Mt Lofty Ra., 14. xii. 1985 (M. J. Lewis) (ANIC); 1 乆, Wedge, Eyre Peninsula, 30. ix. 1979 (G. F. Gross) (SAM); 1 乆, Willowie E, Refge top Hundred of Gregory, 5. xii. 1984 (E. Colley) (SAM); 1 乆, Woodside, 19. x. 1935 (M. W. Mules) (ANIC); 3 么, Woodside, 17. xi. 1936, 7. xi. 1937 (F. M. Angel) (SAM); 1 么, Yankalilla, 15. x. 1933 (J. O. Wilson) (ANIC). Tasmania: 1 么, Arthur Plains, 6. ii. 1965 (Neboiss) (MV); 1 乆, Bicheno, 6 mls SW, 19. i. 1948 (Key, Carne & Kerr) (ANIC); 1 乆, Bicheno, 1 km S, 10. ii. 1988 (G. & A. Daniels) (UQIC); 1 么, Charlton, 26 km SE of Ross, 28. xi.–3. xii. 1975 (H. D. Barker, R. Barnett et al.) (TMH); 1 么, Coles Bay, 15. ii. 1938 (G. M. Goldfinch) (AMS); 1 么, Cradle Mountain (Carter & Lea) (SAM); 2 么, Cradle Mountain, 15.–21. i. 1917 (G. H. Hardy) (AMS); 1 么, Cradle Mountain, 11. i. 1917 (A. J. Turner) (MV); 2 么, Derwent Bridge, 15. i. 1935 (ANIC, TMH); 1 么, Epping Forest, x. 1940 (J. W. Evans) (TMH); 1 乆, Hobart, 14. xii. 1901 (MV); 4 么, 1 乆, Hobart, 21. x. 1913, 20. x. 1916, 12. xi. 1916 (G. H. Hardy) (AMS, ANIC); 1 么, Hobart, 6. xi. 1916 (C. E. Cole) (SAM); 5 么, 4 乆, Hobart, 2. i. 1934, 12. i. 1935 (D. C. P.) (ANIC, TMH); 1 么, Hobart, Grange, 30. xii. 1945 (F. B. Angel) (ANIC); 2 么, 1 乆, Kingston, 23. x. 1945, 6. iii. 1946 (J. R. Cunningham) (TMH); 1 乆, Lake Loane, 1150 m, 25. i. 1981 (P. B. McQuillan) (ANIC); 1 么, Lake St Clair, 14. i. 1934 (G. M. Davis) (ANIC); 1 么, Lake Tyre, 1240 m, 30. i. 1979 (E. D. Edwards) (ANIC); 1 么, Launceston, 29. xii. 1908 (TL) (MV); 1 乆, Orford, 4 km W, 27. i. 1983 (J. C. Cardale) (ANIC); 1 乆, Orford,

69

70


Thumbs Forest Reservation, 6. i. 1991 (E. S. Nielsen & E. D. Edwards) (ANIC); 4 么, 1 乆, Pelion Hut, 870 m, 8.–11. i. 1991 (E. S. Nielsen & E. D. Edwards) (ANIC); 1 乆, Pelion Plains, 860 m, 11. i. 1991 (E. S. Nielsen & E. D. Edwards) (ANIC); 4 么, Strahan, 12. xi.1899, 7. ii. 1925, 26. i. 1934 (ANIC, MV, QM); 1 么, Wedge Bay, 29. xii. 1913 (G. H. Hardy) (AMS).

Diagnosis The largest Pollanisus-species of eastern and southeastern Australia including Tasmania. The males are unmistakable because of their size, broad wings and the shiny body and forewings; the females are similar to those of P. apicalis and P. lithopastus but the former has narrower wings and the latter a very dark forewing upperside sprinkled with shiny scales. In some parts of New South Wales and southern Queensland P. viridipulverulenta populations containing smaller individuals are known, especially in the late summer generations, and it is difficult to separate them from P. apicalis. However, the more triangular wingshape of the forewing and the broader hindwing in P. viridipulverulenta and the medially slightly translucent hindwings in P. apicalis usually separate the two species without problems.

Redescription Male (Pl. 1, Figs 1, 2; Pl. 2, Figs 1, 2; Pl. 61, Figs 1, 2). Forewing length: 11.0–13.0 mm. Head, thorax, abdomen, forewing upperside and parts of forewing and hindwing underside covered with shiny, metallic, yellowish green to dark green or blue scales, blue specimens sometimes with a purple tinge. The coloration is variable but usually not extremely so in a single population. The shiniest specimens are known from South Australia (Pl. 1, Fig. 1). Frons significantly projecting beyond compound eye in lateral view, of medium breadth, approximately 1.2–1.3× that of the black compound eye in frontal view; labial palps slender, strongly upturned, shiny metallic; proboscis dark brown; ocelli small, white, distance from compound eye longer than diameter of ocelli; chaetosemata brown, the anterior extension not covering the whole space between compound eye and ocellus and, therefore, with a band of metallic scales ventrad of chaetosema. Antenna dark greyish green, shaft sometimes shiny blue, but not metallic; segments 1 to 38–40 bipectinate, 38–40 to 48–51 biserrate; the pectinations very long, 8–9× longer than breadth of the shaft in dorsal view at segment

15, 6–7× longer at segment 25, sensory hairs of medium length. Legs shiny metallic, concolorous with thorax. Abdomen sometimes not concolorous with thorax and forewing upperside but also strongly shiny (Pl. 1, Fig. 1). Forewing upperside with shiny scales except for a narrow, blackish band along distal margin; forewing underside dark greybrown, sometimes with a few shiny scales at costa proximally and costad of distal part of cell; fringe with green tinge, in fresh specimens with a few metallic scales towards hind margin. Hindwing upperside dark grey-brown, only exceptionally with a few metallic scales towards hind margin; hindwing underside as upperside but with golden green or blue metallic scales on anterior half of wing and at anal angle. Female (Pl. 3, Fig. 1; Pl. 61, Fig. 3). Forewing length: 8.0–9.0 mm. Significantly smaller than male with narrower and more rounded wings. The shiny scales on head, thorax and forewing upperside often bluish. Abdomen often not shiny, abdominal hairtuft yellow. Head with significantly broader frons and smaller compound eyes than male; frons 2.2–2.3× broader than compound eye in frontal view. Antenna long and slender, biserrate, the dentations close together, giving the antenna an almost filiform appearance, the dorsoventrally compressed antennal shaft broad, the two rows of dentations forming a deep ventral groove. Male genitalia (Figs 138–141, 186–189). Valva with costa straight proximally and slightly convex distally, apical angle ca. 70°, folded translucent central part triangular, ventral sclerotization broad proximally, strongly tapering distally. Aedeagus ca. 4× longer than broad, cornutus long and slender, almost straight, slightly shorter than length of aedeagus. Female genitalia (Figs 240, 241). Praebursa with large lateral appendix, central sclerotization usually small, with two teeth but variable.

Phenology and bionomics Adults are found from late August and early September (Queensland, South Australia) to late January (Tasmania). Pollanisus viridipulverulenta is an early summer species and about three weeks earlier on the wing than the sympatric P. apicalis. Personal observations of the Black Mountain population in Canberra (ACT) (November 1989 and 2001) follow.

TAXONOMY

The adults have a weak flight. They prefer to be in sunshine but are also frequently found in areas of open bushland in half shadow. When at rest they are difficult to spot. They are shy and fly immediately when disturbed. As soon as they are on the wing they are easily netted. The flight is significantly different from that of the externally very similar European Jordanita and Adscita species (both Procridini). The beat of the wing in the male is slower and the flight is more oscillating, somewhat reminiscent of adelid moths and not as straight as in the European species. In the female the beat of the wing is significantly faster, the flight is not oscillating. The yellow abdominal hairtuft is clearly visible. In good weather, males become active in late morning. They prefer to fly close to the ground but can also be observed between the tops of bushes. The females appear approximately one hour later. They always fly close to the ground and never come out of the vegetation. When landing to rest on stems or leaves they often make quick moves until they have found the right position. During the hottest hours of the day males and females have been observed to visit flowers of Helichrysum spp. to obtain nectar (Pl. 61, Figs 2, 3). In the resting position the males close the pectinations of the antenna against the shaft, the antenna then appearing filiform. Oviposition takes place on the flowers and leaves of Hibbertia spp. in early afternoon. During oviposition the females obtain nectar several times. Egg Ovoid, subquadrate, yellow, with slightly rough surface; length 0.5 mm, breadth 0.4 mm. The eggs are laid in small batches and are covered with the yellow, spiny, deciduous scales from the female abdomen (Pl. 59, Fig. 1; Figs 129, 130). The eggs are always deposited on leaves close to the ground mainly on very small bushes and in the shadow. Duration of egg stage: 5–7 days. Larva First instar (L1). Length 1.1–1.2 mm, yellowish green, head light brown, eyes black, prothoracic shield large, dark brown, with long, light setae; anal comb fully developed, blackish brown, with short dentations that are longer medially, a transversal brown band dorsad of anal comb; thoracic legs long, brown, abdominal prolegs yellowish green. The dorsolateral semi-eversible gland on abdominal segment 2 (see p. 39) is not visible in first instar larvae. Setal combination formula of first abdominal segment (sensu Efetov and Tarmann 1999; Efetov et

al. 2000; Efetov 2001; Efetov, Mollet and Tarmann 2003): D: 1d; SD: 1d, 1l; L: 2l (Fig. 131). [D = dorsal setae, SD = subdorsal setae, L = lateral setae; d = dark, stiff seta (like a spine), l = light, flexible seta] The L1 larvae are very active and walk long distances. Immediately after hatching they first eat the shell of the eggs. Then they feed on the leaves and flowers of Hibbertia, producing small oval grooves by inserting the head into the parenchyma of the plant. These feeding marks turn brown afterwards and are easily visible. One larva usually does not make more than 2–3 holes in one leaf or flower head and continuously changes from one part of the host-plant to another. Duration of L1: ca. 10–12 days. Second instar (L2). Length 2.0–2.2 mm, similar to L1, setal combination D: 5d; SD: 4d, 1l; L: 2l. Habits as in L1. Duration of L2 ca. 12–15 days. Third instar (L3). Length 3.0–3.4 mm, similar to L1 and L2, setal combination D: 8d; SD: 6d, 2l; L: 4l. Habits as in L1 and L2. Duration of L3 ca. 12–20 days. Fourth instar (L4). Length 5.0–5.5 mm, head dark brown, prothoracic shield with two subtriangular plates, separated by a narrow yellowish green dorsal line, densely covered with short and long setae; body with a narrow brown dorsal line and a broader subdorsal and subventral line. Duration of L4 ca. 20 days. Fifth instar (L5). Length 8.5–10.5 mm, head and thoracic shield as in L4, dorsal area of body between subdorsal lines brown, dorsal line narrow, dark brown, subdorsal lines broader, dark brown, subventral line dark brown. Duration of L5 ca. 20 days. Sixth instar (L6), fully grown (Pl. 60, Fig. 3). Length 15.0–15.5 mm; head dark brown, thoracic legs blackish brown, prothoracic shield consisting of two subtriangular plates that are separated by a narrow yellowish green line, anterior 2/3 of shield light brown, posterior 1/3 dark brown, densely covered with long and short setae, the longer pointing forwards; dorsal part of the larva with broad greyish white dorsal band, mesothorax with three pairs of dark brown spots posteriad of dorsal verrucae (DV), metathorax with one pair of dark spots between DV, abdominal segments A1–8 with narrow, partly interrupted dark brown dorsal line, A9 with one dorsal and a pair of subdorsal dark brown spots; beside the greyish white dorsal band larva light brown (along dorsal verrucae), verrucae surrounded by blackish brown rings, subdorsal line blackish brown, space between subdorsal and lateral line white, ventrad of lateral line light brown,

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subventral line light brown, venter yellowish white; anal comb fully developed, with short dentations, two small groups of spiny setae on both sides of anus below anal comb; verrucae very uniform on all segments, setal combination D: ca. 45d; SD: ca. 40d, 6l; L: ca. 20l. According to Tothill, Taylor and Paine (1930) Levuana iridescens Bethune-Baker has five larval instars. However, these observations were based on a tropical species and it is no surprise that in the more temperate regions the larvae have more than five instars. It is known from Palaearctic Procridinae larvae that multiple moulting up to 11–12 instars can take place under tough environmental conditions. Duration of L6 ca. 20–30 days. Pupation takes place in a flat, whitish cocoon on the ground (Pl. 60, Fig. 4).

als than those from Victoria and South Australia. According to external, biological and genitalic characters, the Western Australian species, P. cupreus, is not conspecific with P. viridipulverulenta as indicated by Turner (1926b: 444), although it is very closely related. The South Australian populations usually have a brighter metallic sheen (Pl. 1, Fig 1) and were described as ‘form adelaidae’ by Turner 1926b: 444. They clearly belong to P. viridipulverulenta. However, they can also be separated by the different larval host-plant (see above) and may represent a distinct subspecies.

Distribution From southern Queensland throughout New South Wales, Victoria and South Australia to Tasmania.

Pupa (description based on two female pupae) Length 9.5 mm; light brown, eyes black, appendages very weakly fused to each other and to body (loosening from body if larva is preserved in alcohol); proboscis and antennae not exceeding 5th–6th abdominal segment, abdominal segment 2 with lateral gland ventrad of spiracle (Pl. 60, Fig. 5), segments 3–7 with one row of spines of equal triangular form, cremaster consisting of four sockets bearing one short, distally hooked seta each. Foodplants Confirmed larval foodplants are: Hibbertia obtusifolia DC. (ACT) (I. F. B. Common, G. M. Tarmann, personal observation, M. Horak, personal observation and rearing) (Pl. 62, Fig. 1), H. stricta (DC.) F. v. M. (Pl. 62, Fig. 2) and H. sericea (R. Br. ex DC) Benth. (South Australia) (McFarland, 1979: 62). The main larval host-plant in Queensland, New South Wales, Australian Capital Territory, Victoria and Tasmania seems to be Hibbertia obtusifolia DC., as the distribution of P. viridipulverulenta in those states obviously corresponds with the currently known distribution of that plant. The populations of South Australia must feed on other Hibbertia species, as H. obtusifolia does not occur in that state. McFarland (1979: 62) records H. stricta and H. sericea for South Australia but according to Hnatiuk (1990) only the latter occurs in that state except for Kangaroo Island where there is an isolated colony of H. stricta.

Pollanisus cupreus Walker Pl. 4, Figs 1, 2; Pl. 60, Fig. 6; Figs 142, 143, 190, 191, 242, 243. Pollanisus cupreus Walker, 1854, List Specimens lepid. Insects Colln Br. Mus. 1: 115. Lectotype 么 (here designated), AUSTRALIA: Western Australia (BMNH) [examined]. Pollanisus cupreus Walker; Butler 1877: 15; Aurivillius 1920: 43; Hering 1922: 13; Bryk 1936: 118; Alberti 1954: 298; Tarmann 1996: 143. Pollanisus cuprea Walker; Meyrick 1886: 794. Pollanisus viridipulverulenta cupreus Walker; Jordan 1907: 10.

Material examined Remarks Populations from Queensland and parts of New South Wales contain significantly smaller individu-

Western Australia : 1 乆, Albany, 2. iii. 1926 (W. B. Barnard) (QM); 1 么, Albany, 9 km NNE, 7. x. 1981 (J. C. Cardale) (ANIC); 2 么, 1 乆, Applecross, 12. ix. 1948, 10. ix. 1950 (W. M. O’Donnel) (WADA); 1 么, Arthur R., 27 km

TAXONOMY

SW, 19. x. 1990 (I. F. B. Common) (ANIC); 1 么, Augusta, 5 mls NW, 4. iv. 1968 (I. F. B. Common & M. S. Upton) (ANIC); 1 么, Augusta, 14 mls NE (at Alexandra Bridge), 16. xi. 1968 (N. McFarland) (SAM); 1 乆, Bemier Is., x. (Watson) (MV); 1 么, Bibra Lake, 14. ix. 1912 (WAM); 4 么, Bindoon, 9 km SSW, 25. ix. 1981 (J. C. Cardale) (ANIC); 2 么, 1 乆, Bridgetown, xi. 1904 (H. B.) (MV); 2 乆, Brunswick, 7. x. 1905 (Hamb. S. W.-Austr. Exped. 1905) (ZMB); 1 么, Burma Rd Reserve, 30 km E Walkaway, 14. ix. 1986 (P. M. McMillan) (WAM); 2 么, Burns Beach, 21. ix. 1987 (R. W. Hay) (ANIC); 1 乆, Busselton (ANIC); 1 乆, Busselton, 24. x. 1970 (S. J. Curry) (WADA); 1 么, Cottesloe, 12. ix. 1912 (WAM); 2 乆, Crawley, 31. viii. 1934 (K. R. Norris) (ANIC); 1 么, N Darling Ra., 7. ix. 1963 (V. J. Robinson) (ANIC); 1 么, Deepdene, Karridale, 26. x. 1963 (L. M. O’Halloran) (WADA); 1 乆, Deepdene, 3. xi. 1962 (V. J. Robinson) (ANIC); 1 么, Denmark, 14. xi. 1938 (ANIC); 1 么, Doconing Rd Beechina (Chidlow), 16. ix. 1988 (R. P. McMillan) (WAM); 3 么, Donnybrook, 12 km SW, 3. x. 1981 (J. C. Cardale) (ANIC); 1 么, Dunsborough, 2 mls NW, 15. xi. 1968 (N. McFarland) (SAM), 1 么, Eneabba, 10. ix. 1981 (R. P. McMillan) (WAM); 1 么, Fremantle, 20. viii. 1934 (K. R. Norris) (ANIC); 2 么, Fremantle, Canningsroad, 29. viii. 1905 (Hamb. S. W.Austr. Exped. 1905) (ZMB); 1 乆, Geraldton, 4. ix. 1926 (AMS); 2 么, 1 乆, Gidgiegannup, Napier property, 4. x. 1975 (R. P. McMillan) (WAM); 2 么, Gingin, 17. ix. 1969 (K. T. Richards) (WADA); 1 么, Gingin, 15 mls N, 28. ix. 1973 (M. Trudgen) (WADA); 7 么, 1 乆, Glen Forest, 13. ix. 1975, 24. viii. 1976, 11. ix. 1976 (S. M. Postmus) (WAM); 1 么, 8. x. 1973 (S. M. Wade) (WAM); 2 么, 1 乆, Hamel, 1. x. 1913 (MV, QM); 1 么, Holyoake, 24. viii. 1948 (C. Jenkins) (WADA); 1 乆, Hopetoun, 22. ix. 1986 (L. N. McKenna) (WAM); 3 么, 2 乆, Jandakot (WAM); 1 么, 3 乆, Kalamunda, ix. 1913 (WAM); 1 乆, Kalbarri Reserve, 8 mls N, Northern Coastal Hwy, 11. ix. 1977 (A. M. & M. J. Douglas) (WAM); 1 么, Kelmscott, 21. ix. 1976 (C. Johnstone) (WAM); 3 么, Lennox, 5. xi. 1928, 24. viii. 1929, 22. ix. 1929 (A. S.) (MV); 1 么, Lion Mill, 27. viii. 1905 (Hamb. S. W.-Austr. Exped. 1905) (ZMB); 7 么, 4 乆, Mandagallup, 10. ix. 1947 (C. F. H. Jenkins) (SAM, WADA); 1 么, 1乆, Margaret R., 30. x., 1.–4. xi. 1938 (ANIC, QM); 1 乆, Margaret R. Distr., 1915 (WAM); 1 么, 1 乆, Martin, 15. ix. 1987 (R. W. Hay) (ANIC); 1 么, 1 乆, Muchea, 29. viii. 1969 (K. T. Richards) (WADA); 1 么, Mundaring, 22. ix. 1912 (W. B. Alexander) (WAM); 1 么, Mundaring (J. Clarke) (QM); 1 乆, Nannup, 2. x. 1975 (K. A. Spencer) (WAM); 3 么, Nedlands, 7.–27. x. 1923 (M. L. Wise) (MV); 1 么, 2 乆, New Norcia, 9 mls S, 6. ix. 1964 (P. B. Carne) (ANIC); 1 么, Noongan, 9. x. 1981 (J. C. Cardale) (ANIC); 3 么, 3 乆, Parkerville, 8. ix. 1952, 5. x. 1952, 8. x. 1952 (C. F. A. Jenkins) (WADA); 2 么, Perth (G. H. Hardy) (AMS); 1 么, 1 乆, Perth, xi. (C. Wyatt) (ANIC); 1 么, 1 乆, Perth, viii.–ix. 1913 (G. H. Hardy) (AMS); 5 么, 4 乆, Perth, 4.–30. x. 1920 (AMS); 2 么, Perth, 7. x. 1886, 11. xi. 1902 (SAM); 1 么, 1 乆, Perth, 1913, ix. 1937 (J. Angel) (SAM); 2 么, Perth, 4. ix. 1938 (WAM); 2 么, Perth, 31. viii. 1936, 1. ix. 1946 (AMS); 7 么, 3 乆, Perth, 26. ix. 1913, 3.–6. ix. 1938 (ANIC, QM); 1 么, 1 乆, Perth (J.

Clarke) (QM); 1 么, Perth, Guildford, 28. viii. 1905 (Hamb. S. W.-Austr. Exped. 1905) (ZMB); 16 么, 1 么, Perth, Kings Park, 4. ix. 1940 (A. N. F.) (WADA); 1 乆, Perth, Kings Park, 15. ix. 1965 (L. E. Koch) (WAM); 2 么, Perth, Kings Park, 31. viii. 1995 (G. M. & B. E. Tarmann) (TLMF); 10 么, Perth, Subiaco, 3. ix. 1905 (Hamb. S. W.-Austr. Exped. 1905) (ZMB); 5 么, 1 么, Perth, University, 14. ix. 1940 (P. N. F.) (WADA); 1 乆, Serpentine, 23–25. ix. 1905 (Hamb. S. W.-Austr. Exped. 1905) (ZMB); 1 么, Swan R. (B. A. O’Connor) (WADA); 3 乆, Swan R. (L. J. Newman) (WADA); 1 么, Tallanalla, 8. xi. 1958 (I. F. B. Common) (ANIC); 20 么, 1 么, Walpole-Nornalup Nat. Park, Hilltop Lookout, 8. i. 1988 (G. & A. Daniels) (QM); 12 么, 3 乆, Waroona, 10. xii. 1906, 31. viii.–26. x. 1907, 14. iii. 1908, 20. xii. 1908, 20. ix. 1909, 1. xi. 1909, 24. xi. 1910, 15.–27. x. 1913 (G. F. Berthoud) (BMNH, MV); 3 么, 5 乆, Waroona, 7. x. 1913, 25.–26. x. 1938 (ANIC, QM, SAM, WAM); 2 么, 1 乆, Waroona, Darling Ra., 700 ft., near Waroona Dam, 14. xi. 1968 (N. McFarland) (SAM); 1 么, Warren R., (W. D. Dodd) (SAM); 8 么, 8 乆, Wedge I., 5 km E, N. Lancelin, 90 m, 2. and 7. ix. 1995 (G. M. & B. E. Tarmann) (ANIC, TLMF); 2 么, 3 乆, Wubin, 4. ix. 1963 (V. J. Robinson) (ANIC); 3 么, Wubin, 42 km NE, 26. ix. 1981 (J. C. Cardale) (ANIC); 3 么, 4 乆, Yanchep, 10.–13. ix. 1938 (ANIC).

Diagnosis Unmistakable because of the very characteristic appearance. The largest species of the genus in Western Australia. Pollanisus nielseni sp. n. is smaller than P. cupreus and has a strong metallic sheen on the forewing upperside, the shiny scales extending almost to the outer margin that is distinctive blackish brown. Pollanisus empyrea is much smaller, has more rounded wings and a more translucent hindwing, especially medially.

Redescription Male (Pl. 4, Fig. 1). Forewing length: 9.5–11.5 mm. Head, thorax and abdomen metallic golden, with a coppery sheen on upperside, greenish golden with a coppery sheen also on underside. Frons significantly protruding beyond compound eyes in lateral view, of medium breadth, approximately 1.2–1.3× that of the black compound eye in frontal view; palps slender, upturned, shiny metallic green; proboscis dark brown; ocelli small, white, distance from compound eye longer than diameter of ocellus; chaetosemata brown, the anterior extension not covering the whole space between compound eye and ocellus, therefore, with a band of shiny scales ventrad of chaetosema. Antenna very long, dark greyish green, shaft metallic green; segments 1 to 45–47 bipectinate, 45–47 to 56–58 biserrate; the pectinations very

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long, 10–11× longer than shaft in dorsal view at segments 15 and 25, sensory hairs of medium length. Forewing upperside dark grey-brown, densely covered with metallic, golden, coppery scales proximally, less so more distally, and without shiny scales along margin. Forewing underside uniform dark grey-brown, sometimes with a few shiny scales at costal base (very fresh specimens only). Hindwing upperside uniform dark greybrown, sometimes with a few shiny golden scales basally; hindwing underside dark grey-brown with green-golden shiny scales with a coppery tinge along cell and at anal angle. Female (Pl. 4, Fig. 2; Pl. 60, Fig. 6). Forewing length: 8.0–9.0 mm. Significantly smaller than male with more rounded and narrower wings. Abdomen almost without metallic scales except for first segment, with a bright yellow hairtuft. Head broader and compound eye smaller than in male, frons approximately 2.2–2.3× broader than compound eye in frontal view. Antenna slender, biserrate. Male genitalia (Figs 142, 143, 190, 191). Valva strongly convex dorsally and ventrally, apex bent downwards, ending at right-angles with the translucent, folded, triangular central part of valva; ventral sclerotization broad, lancet-shaped, pointed distally. Aedeagus ca. 3.5× longer than broad, slightly tapering posteriorly. Cornutus equal in length to aedeagus, almost straight, very slender, pointed distally. Female genitalia (Figs 242, 243). Ductus bursae short; praebursa large, translucent, with short and very stout lateral appendix, the central sclerotization somewhat rounded, with mainly two strong, curved spines, but variable.

were weak fliers. They preferred sunshine and were most abundant during the hottest hours of the day but also, if warm enough, were active in dry cloudy weather. The preferred nectar-plant was Arctotheca calendula (L.) Levyns (Pl. 60, Fig. 6), a naturalized, originally African, Asteraceae that was the most common yellow roadside daisy in flower at the time. Phyllanthus calycinus Labill (Euphorbiaceae), a white flowering small bush, was also visited by males and females. No specimens were observed visiting the yellow flowers of Hibbertia. At any time of the day, males and females can be disturbed from plants by simply walking through the vegetation. Then they fly only for a short distance before resting again. Females were observed ovipositing on Hibbertia hypericoides in midday sunshine. The eggs are laid singly although not very far from each other on the sepals, small twigs or on the leaves of the host-plant. The females frequently interrupt ovipositing to feed. It was not possible to obtain eggs from females kept in captivity without providing fresh hostplants, and even then only a few eggs were laid. It seems essential that the female takes nectar from flowers between the oviposition intervals. The eggs are translucent yellowish green. They are covered with the deciduous yellow spiny scales of the female’s abdomen. The larvae hatch after 5–8 days. Foodplant As indicated above it is assumed that Hibbertia hypericoides (DC.) Benth. (Pl. 62, Fig. 3) is most likely the larval host-plant. Rearing attempts have failed so far.

Distribution Phenology and bionomics Two generations occur, in early and late summer. Pollanisus cupreus is an early-flying species in Western Australia and can be found about two or three weeks earlier than the often sympatric P. empyrea. Field observations by the author around Perth, WA, in early September 1995, indicated the preferred habitat to be open forest in native bushland with the main larval foodplant Hibbertia hypericoides (DC.) Benth. (Pl. 62, Fig. 3; Pl. 64, Fig. 1). Some specimens were also found in treeless heath on limestone country near the coast north of Lancelin (Pl. 63, Fig. 1). At this locality, P. cupreus was syntopic with P. nielseni sp. n. (Pl. 60, Fig. 6) but the host-plant of the larva of the latter seems to be Hibbertia spicata F. Muell (see p. 76). The adults

Endemic to the temperate parts of Western Australia. The known distribution corresponds well with that of the assumed larval host-plant H. hypericoides (Hnatiuk 1990).

TAXONOMY

Pollanisus nielseni sp. n. Pl. 5, Figs 1, 2; Pl. 60, Fig. 6; Figs 136, 137, 144, 244, 245.

Material examined Holotype 么: Western Australia: 5 km E Wedge Island, N. Lancelin, GPS: 30°49´04´´S 115°14´16´É, 90 m, 7. ix. 1995 (G. M. & B. E. Tarmann) (ANIC). Paratypes Western Australia: 22 么, 12 乆, Wedge Island, 5 km E, N. Lancelin, GPS: 30°49´04´´S 115°14´16´É, 90 m, 7. ix. 1995 (G. M. & B. E. Tarmann) (ANIC 2 么, 4 乆; WAM 1 么, 1 乆; TLMF); 1 么, Wedge Island, 8 km E, vic. Duggan Well, N. Lancelin, GPS: 115°15´51´É 30°47´44´´S, 80 m, 2. ix. 1995 (G. M. & B. E. Tarmann) (TLMF); 3 么, 1 乆, Drummond Cove, 11 km N of Geraldton, 22. i. 1973 (N. McFarland) (ANIC); 1 乆, Kojarena, 6. ix. 1926 (Waterhouse) (AMS); 1 乆, Moresby Range, Howatharra Road, 25. viii. 1974 (N. McFarland) (WADA); 1 么, 1 乆, Munglinup R., 34 km NW by W Shoal Cape, 21. ix. 1981 (I. D. Naumann & J. C. Cardale) (ANIC); 1 乆, Oldfield R., 52 km WNW Shoal Cape, 21. ix. 1981 (I. D. Naumann & J. C. Cardale) (ANIC); 1 乆, Tombstone Rocks, 4 km W, GPS: 30°41´56´´S 115°12´58´É, 100 m, 2. ix. 1995 (G. M. & B. E. Tarmann) (TLMF).

Diagnosis Very similar to P. cupreus in wingshape, but slightly smaller. Forewing upperside with a very characteristic, extremely bright metallic sheen, the shiny scales extending almost to apex, leaving a broad blackish brown margin. In P. cupreus the shiny scales are less brilliant and concentrated at the basal part of the forewing with only very few single scales apically, a distinct margin is absent. Hindwing very opaque, almost black, in strong contrast to the shiny forewing. Pollanisus empyrea is smaller in the male, and equal in size in the female but has more rounded wings and the forewing upperside is less shiny, the hindwing is more translucent medially.

Description Male (Pl. 5, Fig. 1). Forewing length: 9.0–10.0 mm. Head completely covered with shiny metallic scales, green with a golden highlight; frons projecting dorsally, of medium breadth, approximately 1.2× as broad as breadth of black compound eyes in frontal view; proboscis dark brown; labial palps short, porrect, sharply pointed apically, shiny metallic green; ocelli large, white, distance from dorsal margin of compound eye equal to diameter of ocel-

lus; chaetosemata large, dark greyish brown, with a broad band of shiny metallic golden green scales between chaetosema and dorsal margin of compound eye; antenna greyish black, covered with metallic bluish green scales, segments 1–39 bipectinate, 40–50 biserrate, segment 51 simple and pointed apically, pectinations very long, length of pectinations increasing from segment 1–10, of similar length from 11–20, 8–9× longer than shaft in dorsal view at segment 15, progressively tapering towards apex from 20–34, abrubtly shortening from 35–39, sensory hairs short. Thorax shiny metallic golden green dorsally, more bluish green laterally and ventrally, legs shiny metallic green. Abdomen upper- and underside shiny metallic golden green with slight bluish sheen at abdominal end on underside. Forewing upperside bright shiny golden yellow and green (depending on the angle of the transmitted light) except for a narrow distal band that is blackish grey, under side blackish grey, with a few golden scales at costa basally. Hindwing upper and underside blackish grey, groups of shiny scales along costa, at basal parts of cell and at anal angle; fringe in both wings blackish grey. Female (Pl. 5, Fig. 2; Pl. 60, Fig. 6). Forewing length: 8.0–8.5 mm. Similar but smaller than male with narrower and more rounded wings. Abdominal hairtuft yellow. Head with significantly broader frons and smaller compound eyes; frons 1.7–1.8× broader than compound eye in frontal view. Antenna long and slender, biserrate, the dentations close together, giving the antenna an almost filiform appearance, the dorsoventrally compressed antennal shaft broad, the two rows of dentations forming a deep ventral groove. Male genitalia (Figs 136, 137, 144). Valva convex dorsally almost straight ventrally, apex ending at a slightly acute angle with the translucent, folded, triangular central part of valva; ventral sclerotization broad, pointed distally, sacculus well separated. Aedeagus ca. 4.0× longer than broad, slightly bent upwards, not tapering posteriorly. Cornutus slightly exceeding aedeagus in length, very slender, almost straight and pointed distally, slightly curved upwards proximally. 8th abdominal segment completely covered by sclerotized sternite and tergite. Female genitalia (Figs 244, 245). Ductus bursae short; praebursa large, translucent, with short triangular lateral appendix, a ring-like structure present posteriorly, the central sclerotization a short plate, with two slender, curved spines.

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Phenology and bionomics

Distribution

The presence of this third Pollanisus species in Western Australia was verified on a field trip in early September 1995. The existence of ‘brilliant shiny forms’ of ‘P. cupreus’ had been recognised in collections for some time but unfortunately the genitalic characters are very unsatisfactory and variable in this genus. No attempt had been made to describe this ‘form’ on the basis of only a few more brilliant specimens in series of P. cupreus, which could easily have been aberrations. At the type locality of P. nielseni, both species were syntopic and flying synchronously (i.e. in the same locality at the same time) on 2nd and 7th September 1995, visiting flowers of Arctotheca calendula (L.) Levyns (Asteraceae) (Pl. 60, Fig. 6) and Phyllanthus calycinus Labill (Euphorbiaceae). It was easy to recognise that there were two species involved. The strong sheen on the forewing of P. nielseni was very conspicuous whereas P. cupreus looked almost blackish brown. Pollanisus nielseni had a significantly faster wingbeat than P. cupreus and appeared to be much smaller when flying. Only females of P. nielseni were observed ovipositing at this locality. They deposited their eggs on Hibbertia spicata F. Muell., a small shrub with short, narrow leaves and small drooping flowers. The eggs were laid singly on the sepals, stems or leaves and were pale yellow and darker than those of P. cupreus (that are translucent yellowish green). They were covered with the deciduous dart-like scales of the female’s abdominal hairtuft as is typical for all Pollanisus species. No larvae emerged from the eggs taken (too hot in the car?). Males and females were very abundant at this locality. It was a warm and sunny spring day with almost no wind and hundreds of specimens were on the wing, resting on leaves or twigs and visiting flowers. The type-locality is a heath with almost no large bushes except for a few Banksia ilicifolia R. Br. (Pl. 63, Fig. 1). The rocky soil is calcareous (old reef material?). Although very abundant at the typelocality, no further specimens of P. nielseni were seen southwards and only two single specimens a few kilometres northwards. Near the white sand dunes in the vicinity of the type-locality the host-plant is absent and another Hibbertia, i.e. H. subvaginata (Steudel) F. Muell., was in flower but was absent at the type-locality. That Hibbertia species was obviously not suitable as a foodplant for Pollanisus. Based on information obtained from four specimens collected by N. McFarland in January near Geraldton it can be assumed that this species is bivoltine with one generation in early spring and a second in mid-summer.

Coastal parts of south-western Western Australia.

Etymology Pollanisus nielseni is one of the most beautiful and shiny Australian zygaenids. It is dedicated to the late Ebbe S. Nielsen who was a shining light on the sky of Australian lepidopterology and to whom the author is deeply indebted for much help and support and for years of wonderful friendship.

Pollanisus modestus sp. n. Pl. 3, Fig. 2; Figs 145, 146, 192, 193.

Material examined Holotype 么, New South Wales: Clyde Mountain, 2400 ft, 29. iii. 1960 (I. F. B. Common & M. S. Upton) (ANIC). Genitalia slide Z 3064.

Diagnosis This species is very distinct. It clearly belongs to the group of species including P. viridipulverulenta, P. cupreus, P. nielseni, P. apicalis, P. empyrea and P. lithopastus, with its shiny forewing upperside and the whole body being covered with shiny scales. It is similar in size to P. apicalis, but the shiny scales on the forewing upperside are distributed similarly to those of P. cupreus, although these scales are of different colour and more densely cover the wing proximally. In external appearance as well as in the genitalia it differs from the above-mentioned species and is clearly specifically different. It is amazing that there is only a single male in the rich material of Pollanisus examined. According to I. F. B. Common (personal communication) the type-locality is a slightly damp heath-like clearing with Hibbertia but not an area of ununsual vegetation or climate.

TAXONOMY

Description Male (Pl. 3, Fig. 2). Forewing length: 7.5 mm. Head greyish black dorsally and frontally, with a few shiny metallic green scales, strongly shiny metallic green laterally, frons broad, just slightly projecting dorsally, approximately 1.5× as broad as breadth of the small, black compound eyes in frontal view; proboscis dark brown; labial palps short, porrect, sharply pointed apically, basal segment shiny metallic green, dark grey and not shiny apically; ocelli small, white, distance from dorsal margin of compound eye almost 3× as long as diameter of ocellus; chaetosemata large, greyish brown, lighter posteriorly, with a broad band of shiny metallic green scales between chaetosema and dorsal margin of compound eye; antenna greyish black, with weak bluish sheen but not metallic, segments 1–32 bipectinate, 33–44 biserrate, segment 45 simple and pointed apically, pectinations of medium length, length of pectinations increasing from segments 1–11, of similar length from 12 to 17, 6–7× longer than shaft in dorsal view at segment 15, progressively tapering towards apex from 18–45, sensory hairs very short. Thorax shiny metallic yellowish and bluish green dorsally, bluish green ventrally, legs shiny metallic bluish green proximally, dark grey distally. Abdomen upper- and underside shiny metallic greenish blue. Wings densely scaled. Forewing upperside brownish black, densely covered with shiny metallic bluish green scales fading distally, with stronger sheen in basal half. Forewing underside unicolorous matt dark grey-brown. Hindwing opaque, upperside and underside dark grey-brown, underside with bluish green scales except around margin. Female. Unknown. Male genitalia (Figs 145, 146, 192, 193). Valva slender, rounded apically, slightly convex dorsally and ventrally, the folded translucent central part triangular, ventral sclerotization of valva large, with broad proximal part, lancet-shaped. Aedeagus of medium length, ca. 3.5× longer than broad, slightly upcurved, cornutus strong and long, of same length as aedeagus, pointed apically.

Phenology and bionomics Unknown.

Distribution New South Wales (only male holotype known).

Pollanisus lithopastus Turner Pl. 6, Figs 1, 2; Figs 147–152, 194–197, 246, 247, 426. Pollanisus lithopastus Turner, 1926b, Proc. Linn. Soc. NSW 51: 443. Lectotype 么 (here designated), AUSTRALIA: New South Wales, Ebor, 11.i.1914 (A. J. Turner) (ANIC) [examined]. Pollanisus lithopastus Turner; Turner 1939: 110; Tarmann 1996: 143.

Material examined New South Wales: 1 么, Allyn R. Forestry Park, 31. ii. 1981 (V. J. Robinson & L. Willan) (ANIC); 2 么, Barralier, 7–9. i. 1969 (V. J. Robinson) (ANIC); 1 么, Barren Grounds, 14. i. 1966 (V. J. Robinson) (ANIC); 1 么, Barrington Tops, ii. 1925 (SU Zool. Exp.) (AMS); 1 么, Barrington Tops, 25. i. 1922 (Nicholson) (QM); 2 么, Boonoo Boonoo R., 29. xi. 1981 (G. Daniels & M. A. Schneider) (QM); 1 么, Boyd Creek, Kanangra Walls, 4100 ft., 26. ii. 1956 (K. Fairey, L. Willan & V. J. Robinson) (ANIC); 3 么, Boyd R., Kanangra Walls, 25. i. 1968 (V. J. Robinson) (ANIC); 3 么, Boyd R., 21. ii. 1971 (V. J. Robinson & L. Willan) (ANIC); 3 么, Boyd R. Crossing, 4100 ft., 12. xi. 1973, 6. iii. 1976 (V. J. Robinson) (ANIC); 2 么, Bundanoon, 10. xii. 1971 (V. J. Robinson & N. R. Badans) (ANIC); 1 么, Darkes Forest, 5. xii. 1964 (V. J. Robinson) (ANIC); 13 么, Ebor, 20. ii. 1939, 20. i.–4. ii. 1944, 26. i.–4. ii. 1946, xii. 1948 (AMS); 2 么, Ebor, 9–11. i. 1914 (A. J. Turner) (ANIC); 10 么, Ebor, 15–29. i. 1944, 1. ii. 1944, 5–7. ii. 1946 (L. H. MosseRobinson) (ANIC); 2 么, 4 乆, Ebor, Pt Lookout, 24–25. ii. 1967 (V. J. Robinson) (ANIC); 1 么, Katoomba, 5. i. 1966 (V. J. Robinson) (ANIC); 1 么, Katoomba, e.l. 7. ii. 1978 (B. Hacobian) (ANIC); 1 么, Ku-ring-gai Chase Nat. Park, 26. ii. 1971 (V. J. Robinson) (ANIC); 1 么, Minnamurra Falls, 18. xii. 1968 (V. J. Robinson) (ANIC); 4 么, Mittagong, 3. xii. 1926, 17. xii. 1927 (AMS); 2 么, Mittagong, 25. i. 1936 (ANIC); 1 么, Mt Kaputar, 13. iii. 1969 (V. J. Robinson) (ANIC); 1 么, Mt Keira, 22. xii. 1963 (V. J. Robinson) (ANIC); 3 么, 2 乆, Sawpit Creek, Snowy Mountains, 12.–13. iii. 1972 (V. J. Robinson & R. Badans) (ANIC); 1 么, Saw Pit Creek, Snowy Mountains, 10. ii. 1979 (D. K.

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McAlpine & B. J. Day) (AMS); 2 么, Wilton, CSIRO Exp. Farm, 6. xii. 1971, 14. ii. 1972 (V. J. Robinson) (ANIC). Australian Capital Territory: 1 么, 1 乆, Bendora, 6. ii. 1962 (I. F. B. Common) (ANIC), 1 乆, Condor Creek, Brindabella Ra., 8. ii. 1984 (D. K. McAlpine & B. J. Day) (AMS). Victoria: 11 么, 8 乆, Beaconsfield, xi–xii. 1903, 14. iii.1904 (MV, QM); 2 么, Beaconsfield, 23. xii. 1908 (Drake) (MV); Beenak, 3 km W, 7. i. 1972 (A. Neboiss) (MV); 2 么, Boronia, xii. 1955 (Fleet) (MV); 3 么, Boronia, i. 1930 (Kershaw) (MV); 1 么, Cockatoo, 24. xii. 1929 (MV); 6 乆, Emerald, 20. ii. 1904 (T. K.) (MV); 1 乆, Frankston (MV); 2 么, 1 乆, Gisborne, 13. i. 1895 (MV); 1 么, Grampians, xi. 1892 (MV); 2 么, Latrobe R. Survey St. 17 A, Tanjil Junction, 18. xii. 1972 (MV); 7 么, Wannon Divide, Grampians, 1400 ft., 8–10. ii. 1956 (I. F. B. Common, N. B. Tindale) (ANIC, SAM); 1 么, Wilsons Promontory, 30. xii. 1924 (Cb.) (MV); 1 乆, Windsor, 15. iii. 1904 (T. K.) (MV). Tasmania: 1 么, Cradle Mountain Rd, Moina, 2000 ft., 20. i. 1925 (A. J. Turner) (ANIC); 2 么, Derwent Bridge, 10 mls E, 2500 ft., 21. ii. 1963 (I. F. B. Common & M. S. Upton) (ANIC); 2 么, Don R., Devonport, 22. i. 1991 (E. S. Nielsen, E. D. Edwards & L. Hill) (ANIC); 2 么, Flinders I., 5 km N Ranya, UVL, 21. i. 1982 (P. B. McQuillan) (ANIC); 1 乆, Hobart (BMNH); 1 么, 1 乆, Hobart (A. M. Lea) (ANIC, TMH); 7 么, Huon-Picton R. junction, 18. ii. 1967 (A. Neboiss) (MV); 4 么, Huon Plains, 2–6. ii. 1965 (A. Neboiss) (MV); 3 么, Lake St Clair, 22. i. 1934 (ANIC); 1 么, Lefroy, 21. i. 1897 (MV); 1 么, Marion Bay, 7. i. 1982 (P. B. McQuillan) (ANIC); 60 么, 1 乆, Melaleuca, 12.–16. ii. 1990 (E. S. Nielsen & P. B. McQuillan) (ANIC); 21 么, Melaleuca, 14.–22. i. 1991 (E. S. Nielsen & E. D. Edwards) (ANIC); 9 么, Melaleuca, near Bathurst Harbour, 18.–21 ii. 1991 (A. A. Calder & W. Dressler) (ANIC); 1 么, Mole Creek, 20. ii. 1989 (I. F. B. Common) (ANIC); 1 么, Mt Eliza, 1. iii. 1980 (P. B. McQuillan) (ANIC); 1 么, 2 么, Mt Mueller, 6 km W, 25. i. 1981 (P. B. McQuillan) (ANIC); 5 么, Mt Nelson, 200 m, 8. ii. 1989 (P. B. McQuillan) (ANIC); 1 么, Mt Wellington, 28. i. 1978 (P. B. McQuillan) (ANIC); 1 么, 4 km W Orford, 27. i. 1983 (J. C. Cardale) (ANIC); 1 么, Port Arthur, 30.–31. i. 1992 (I. F. B. Common) (ANIC); 11 么, Port Sorell, 1 km W, 22. i. 1991 (E. S. Nielsen, E. D. Edwards & L. Hill) (ANIC); 1 么, Ridgeway, 8. i. 1978 (P. B. McQuillan) (ANIC); 2 么, Rosebery, 4–9. ii. 1925 (ANIC, QM); 1 么, St Helens, 15. xii. 1979 (T. V. Rangsi) (ANIC); 1 么, St Marys, 23. i. 1938 (ANIC); 1 么, St Patricks Plains, 20. i. 1988 (P. B. McQuillan) (ANIC); 1 么, Scottsdale, 9 km E, 12. i. 1983 (J. C. Cardale) (ANIC); 2 么, Strahan (A. M. Lea & Carter) (SAM); 2 么, Strahan, 5–6. ii. 1925 (ANIC); 4 么, Waratah, 10. ii. 1938 (G. M. Goldfinch) (AMS).

Diagnosis Pollanisus lithopastus is a very characteristic species. This dark moth with its silvery or golden farinose wings is unmistakable. No similar species is known.

Redescription Male (Pl. 6, Fig. 1). Forewing length: 8.5–9.5 mm. Head black, with metallic golden green scales dorsally and frontally, metallic, bluish silvery laterally; frons not projecting beyond compound eye in lateral view, narrow, breadth in frontal view only 0.6× that of large black compound eyes; palps short, dark, basal segment with bluish silvery or golden metallic scales; proboscis dark brown; ocelli large, white, distance from compound eye smaller than diameter of ocellus; chaetosemata dark brown, the anterior extension completely covering the space between compound eye and ocellus. Antenna dark greyish green with weak sheen, antennal shaft slender, dorsoventrally compressed, segments 1 to 34–35 bipectinate, 35–36 to 44–45 biserrate; the pectinations 7–8× longer than breadth of shaft in dorsal view at segment 15, only 3–4× longer at segment 25, sensory hairs on pectinations very short. Thorax dark greenish grey, sometimes almost black ventrally, with or without sheen dorsally, usually with only a few shiny metallic golden green scales; with metallic, silvery, bluish green or golden green, strongly shiny patagia; thorax shiny metallic bluish green ventrally. Legs black, unicolorous. Abdomen shiny metallic blue or bluish green. Forewing upperside blackish brown, matt, with shiny metallic, silvery, bluish green or golden green scales, scattered singly, producing a very characteristic appearance; underside dark grey-brown, with single metallic blue scales variably scattered along anterior half of wing. Hindwing upperside dark brownish grey, matt; underside dark brownish grey, variably scattered with metallic, bluish or bluish green scales, fading towards margin. Fringe of both wings blackish grey with strong sheen. Medial stem absent in forewing, present distally in hindwing. Female (Pl. 6, Fig. 2). Forewing length: 7.0–8.0 mm. Similar to male but smaller, with narrower, more rounded wings. Antenna biserrate. Abdominal hairtuft bright yellow. Male genitalia (Figs 147–152, 194–197). Valva long and slender, much longer dorsally, only slightly convex dorsally, almost straight ventrally, sacculus small; folded translucent central part large, as broad proximally as distally; ventral sclerotization of valva narrow and parallel to ventral margin of valva. Aedeagus ca. 4.5× longer than broad, slightly upcurved, strongly sclerotized, cornutus large and slender, slightly upcurved, pointed distally, its length ca. 80% of aedeagus, vesica seminalis with a plateshaped sclerotization laterally.

TAXONOMY

Female genitalia (Figs 246, 247). Sternite VIII extremely narrow and well sclerotized. Ductus bursae slightly sclerotized (completely translucent in most other species) and broadening into a somewhat bottle-shaped praebursa with short appendix, central sclerotization of praebursa with 2–4 small teeth.

Phenology and bionomics Nothing is known about the habits and biology of this moth. A long series was collected in open heathland at Melaleuca (TAS) by E. S. Nielsen and E. D. Edwards in 1991 (Pl. 63, Fig. 4).

Distribution Found from northern New South Wales to Victoria and in Tasmania. Often collected in mountainous regions at higher elevations, e.g. Barrington Tops, Ebor, Blue Mountains, Snowy Mountains, Grampians, Cradle Mountain, but also at sea level, especially in Tasmania.

very similar to that of P. lithopastus, with only a slightly protruding frons. The eyes are slightly smaller and the frons is slightly broader than in P. lithopastus. The antenna is shorter, segments 1–29 bipectinate, 30–37 weakly biserrate, strongly pointed apically, the pectinations very long, approximately 8–9× longer than breadth of shaft in dorsal view at segment 15. The shiny yellowish green collar on the thorax is more distinct and the abdomen is bright blue. The male genitalia are also similar to those of P. lithopastus, with the aedeagus and the cornutus on the vesica being slightly shorter. The female genitalia have a slightly sclerotized almost bottle-shaped praebursa like those of P. lithopastus but the sclerotization seems different (in the only examined specimen). It is still uncertain whether these two specimens are specifically different or just a very distinct aberration of P. lithopastus, a species that is quite variable. The impression of a specific difference is strong but not sufficient to describe a new species.

Pollanisus empyrea (Meyrick) Pl. 8, Figs 1, 2; Figs 155, 156, 200–205, 250, 251.

Pollanisus sp. 1 Pl. 7, Figs 1, 2; Figs 153, 154, 198, 199, 248, 249.

Material examined Australian Capital Territory: 1 么, 1 乆, Bendora, 19. ii. 1959 (I. F. B. Common) (ANIC).

Procris empyrea Meyrick, 1888, Proc. Linn. Soc. NSW (2) 2: 927. Holotype 么, AUSTRALIA: Western Australia, Albany, 6. xi. 1886 (reared) (BMNH) [examined]. Pollanisus empyrea (Meyrick); Kirby 1892: 87; Jordan 1907: 10; Hering 1922: 13; Bryk 1936: 118; Tarmann 1996: 143. Pollanisus empyreus (Meyrick); Turner 1926b: 443. Procris amethystina Meyrick, 1888, Proc. Linn. Soc. NSW (2) 2: 927. Lectotye 么 (here designated), AUSTRALIA: Western Australia, Perth, 25. xi. 1886 (BMNH) [examined]; Tarmann 1996: 143 (syn. of P. empyrea). Pollanisus amethystina (Meyrick); Kirby 1892: 87; Jordan 1907: 10; Strand 1915: 118; Hering 1922: 13; Bryk 1936: 117. Pollanisus amethystinus (Meyrick); Turner 1926b: 443.

Remarks This species has a very distinctive habitus. It is externally somewhat similar to P. lithopastus, the forewing upperside having scattered shiny scales. It is significantly smaller, the wings are more rounded and there are more shiny scales mixed with the dark scales of the forewing upperside. The head is also

Material examined Western Australia: 2 么, 2 乆, Albany, 5. iii. 1926 (W. B. Barnard) (ANIC); 3 么, 2 乆, Augusta (bred), 27. x. 1963 (V. J. Robinson) (ANIC); 1 乆, Bunbury, 11. ii. 1928 (W. B. Barnard) (QM); 1 乆, Burma Rd Reserve, 30 km E Walkaway, 15. ix. 1986 (R. P. McMillan) (WAM); 4 么,

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Busselton, 8. x. 1920 (AMS); 1 么, Capel, 10. xi. 1940 (D.H.F.) (WADA); 1 乆, Cape Naturaliste, 2. iv. 1968 (I. F. B. Common & M. S. Upton) (ANIC); 1 么, Claremont (WAM); 1 么, Deepdene, Karridale, 17. i. 1967 (M. S. Upton) (ANIC); 2 么, 2 乆, Denmark, 12. iii. 1926, 14.–16. iv. 1926, 14. xi. 1935 (W. B. Barnard) (ANIC); 1 么, Denmark, i. 1967 (R. P. McMillan) (WAM); 1 乆, Dwellingup, 20 km S, Upper Murray Valley, 2. xii. 1985 (O. Mueller) (WAM); 1 么, Ellens Brook, Glenbourne Farm, near Margaret R., 25. xi. 1979 (P. G. Kendrick) (WAM); 1 么, 1 乆, Esperance, 8. i. 1987 (in sand dunes) (G. & A. Daniels) (UQIC); 1 乆, Lancelin, 21 mls S, 10. xi. 1968 (on Hibbertia sp.) (N. McFarland) (SAM); 1 么, Lancelin, 25 mls ESE, Darling Ra. near Moore R., 9. xi. 1968 (N. McFarland) (SAM); 2 么, Margaret R., 10. xi. 1958 (I. F. B. Common) (ANIC); 1 么, Mt Dale, 21. i. 1928 (W. B. Barnard) (QM); 1 么, Perth, 11. xi. 1902 (SAM); 1 么, Scott Nat. Park, 24. iv. 1983 (E. S. Nielsen & E. D. Edwards) (ANIC); 1 么, 1 乆, Tarmela, 1958 (V. J. Robinson) (ANIC); 5 么, 1 乆, Warwick, 4. xi. 1992 (E. D. Edwards, E. S. Nielsen & R. W. Hay) (ANIC); 2 乆, Yanchep, 24. xi. 1938, 27. xi. 1939 (ANIC).

Diagnosis Pollanisus empyrea is endemic to the temperate parts of Western Australia. The sympatric Pollanisus cuprea has much larger males, broader wings with a more triangular wing shape, a darker, more intensely coppery appearance and longer antennae with longer pectinations. The females may be rather similar, those of P. cuprea have broader wings and significantly longer antennae. Pollanisus cuprea is on the wing approximately three weeks earlier than P. empyrea. Pollanisus nielseni has a strongly shiny metallic golden forewing upperside and very dark opaque hindwings.

30–32 to 45–47 biserrate; the pectinations 5–6× longer than breadth of shaft in dorsal view at segment 15, 4–5× longer at segment 25, sensory hairs on pectinations very short. Legs not so intensely shiny as in thorax, but still metallic. Abdomen dark greenish brown, not shiny. Forewing upperside metallic coppery golden, golden green, or rarely bluish green, distal margin and fringe blackish brown; underside dark greybrown, with a shorter subcostal elongate patch of shiny golden scales basally and a longer one between the cell and the costa more distally. Upperside of hindwing uniform dark grey-brown, slightly translucent medially; underside as upperside but with two patches of shiny greenish golden scales, one between the cell and the costa, the other at anal angle. Medial stem absent in forewing, present in hindwing. Female (Pl. 8, Fig. 2). Forewing length: 8.0–8.5 mm. Similar to male and equal in size, wings more rounded, less densely covered with shiny scales, darker, with a yellow abdominal hairtuft. Head much broader and compound eyes smaller than in male, breadth 2.0–2.2× that of compound eyes in frontal view. Antenna biserrated. Male genitalia (Figs 155, 156, 200–205). Valva strongly pointed apically (angle ca. 60°), translucent central part triangular, ventral sclerotization very broad. Aedeagus short, stout, 2.5–3.5× longer than broad, with short cornutus (ca. 80% length of aedeagus). Female genitalia (Figs 250, 251). Praebursa with a strong central sclerotization with 3–4 teeth of very variable shape and length.

Redescription Male (Pl. 8, Fig. 1). Forewing length: 8.0–8.5 mm. Head, thorax, forewing upperside and parts of forewing and hindwing underside covered with shiny, almost metallic, golden, greenish golden or coppery golden scales. Frons projecting beyond compound eyes in lateral view, of medium breadth, approximately 1.2× that of compound eye in frontal view; palps curved proximally, porrect distally, shiny metallic; proboscis dark brown; ocelli of medium size, white, distance from compound eye almost equal to diameter of ocellus; chaetosemata brown, the anterior extension almost covering the whole space between compound eye and ocellus, just allowing a very narrow stripe of metallic green scales (one row) ventrally. Antenna dark grey with weak blue sheen, segments 1 to 30–32 bipectinate

Phenology and bionomics The foodplant of the larva is unknown but is most likely Hibbertia sp. Several specimens have been reared but unfortunately no data of these rearings are available. According to samples from late summer and late autumn it is possible that this species is at least partly bivoltine.

Remarks All golden green or coppery golden shiny metallic parts may be more bluish green. This rare colour variation was described as a distinct species, Procris amethystina Meyrick, 1888, in the same paper and on the same page as Procris empyrea Meyrick, 1888.

TAXONOMY

The type-material and all constant characters of both taxa indicate that P. amethystina is a subjective synonym of P. empyrea Meyrick, 1888 (Tarmann 1996: 143). The genitalia of P. empyrea are so variable that there is strong evidence that this taxon represents an unrecognised species-group. More information, especially on biology, is required to solve this problem.

Distribution Endemic to the temperate parts of Western Australia, particulatly the Jarrah and Karri forests between Perth and Albany (Pl. 64, Fig. 1).

Pollanisus apicalis (Walker) Pl. 9, Figs 1, 2; Pl. 10, Fig. 1; Pl. 58, Fig. 4; Pl. 59, Fig. 6; Pl. 60, Figs 1, 2; Figs 157–160, 206–209, 252, 253. Procris apicalis Walker, 1854, List Specimens lepid. Insects Colln Br. Mus. 1: 111. Lectotype 乆 (here designated), AUSTRALIA: New South Wales, Sydney (BMNH) [examined]. Procris apicalis Walker; Butler 1877: 14; Meyrick 1886: 794. Adscita apicalis (Walker); Kirby 1892: 86. Pollanisus apicalis (Walker); Jordan 1907: 10; Hering 1922: 13; Turner 1926b: 443; Bryk 1936: 117; Alberti 1954: 298; Tarmann 1996: 143. Pollanisus sequens Walker, 1854, List Specimens lepid. Insects Colln Br. Mus. 1: 115. Lectotype 么 (here designated). AUSTRALIA: [Tasmania] ‘Van Diemen’s Land’ [type without locality data] (BMNH) [examined]. Synonymised by Meyrick (1886: 794). Procris novaehollandiae Wallengren, 1860, Wien. ent. Monatschr. 4: 39. Holotype 么, AUSTRALIA: New South Wales, Sydney (NHRM [Stockholm]) [examined].

Material examined Queensland: 1 么, Amiens SF, near Stanthorpe, 28. xi. 1981 (m.v. lamp) (G. Daniels & M. A. Schneider) (UQIC); 1 么, Archerfield, 2. ix. 1969 (V. Stablum) (UQIC); 9 么, 3 乆, Brisbane, 10. i. 1914, ix.–xi. 1914, iii.–iv. 1915 (SAM); 1 乆, Burleigh Heads, 16. ix. 1932 (UQIC); 1 么, Burleigh Heads, 16. ix. 1942 (E. J. Dumigan) (UQIC); 1 么, Carnarvon Ra., 18. xii. 1938 (W. B. Barnard) (QM); 2 么, Carnarvon Ra., xi. 1944 (N. Geary) (AMS); 4 么, Cooloola Forestry Area E Gympie, iv. 1978 (at light) (I. D. Galloway) (DPIQ); 2 么, 1 乆, Crows Nest, 29. x. 1930 (UQIC); 1 么, 2 乆, Dunwich, 22. viii. 1958 (S. S. Sexton) (UQIC); 1 么, Dunwich, 7 km E, north Stradbroke I., 13. iii. 1988 (G. Daniels) (UQIC); 1 么, Girraween Nat. Park, 1. xii. 1981 (mv lamp) (G. Daniels & M. A. Schneider) (UQIC); 1 么, Herberton, ii. 1911 (MV); 11 么, Herberton, 8 km NNW, 820 m, 27. v. 1977 (I. F. B. Common & E. D. Edwards) (ANIC), 1 么, 1 乆, Milmerran, 13./20. x. 1929 (ANIC); 19 么, 8 乆, Milmerran, 24. iii. 1925, 26.–28. ix. 1933, 4.–15. x. 1933, 1. xi. 1933, 8. x. 1934, 13.–19. x. 1935, x. 1944, 7.–8. x. 1944, 28. iv. 1946, x. 1949 (J. Macqueen) (ANIC); 2 么, Millstream Falls near Ravenshoe, 3. xii. 1967 (R. Dobson) (ANIC); 1 乆, Mt Tibberoowuccum, near Beerburrum, 4. xii. 1985 (R. Eastwood) (UQIC); 1 么, Paluma, 15 km W, 27. viii. 1985 (I. F. B. Common) (ANIC); 3 么, Stannary Hills (ANIC); 1 么, 1 乆, Stanthorpe, ii. (WADA); 5 么, 3 乆, Stradbroke I., 30. xi. 1902, 28.–29. ix. 1906, 9. ix. 1911, 28. ix. 1906, 26. xii. 1913 (ANIC, BMNH, WADA); 1 么, 1 乆, Stradbroke I., 2. x. 1911, 2. xii. 1912 (H. Hacker) (DPIQ, QM); 1 么, Stradbroke I., 12. viii. 1942 (I. F. B. Common) (ANIC); 2 乆, Stradbroke I., Brown Lake, 16. ix. 1961 (G. Daniels) (UQIC); 1 么, Stradbroke I., Brown Lake, 26. ii. 1963 (at light) (M. A. Schneider) (UQIC); 1 么, Stradbroke I., Brown Lake, 1. ii. 1980 (T. H. Cribb) (UQIC); 1 乆, Stradbroke I., Brown Lake, 16. ix. 1981 (G. Daniels) (UQIC); 1 乆, Tewah Creek, via Tin Can Bay, 16.–18. x. 1970 (D. L. Hancock) (UQIC); 1 么, Toowoomba, 9. iii. 1929 (UQIC); 2 么, Toowoomba, 14. xi. 1961, 13. i. 1961 (J. Macqueen) (ANIC) . New South Wales: 2 么, Armidale, 11. xi. 1952, 31. x. 1954 (C. W. Frazier) (ANIC); 1 乆, Ash I. (Scott) (AMS); 1 么, Barralier, 7. i. 1969 (V. J. Robinson) (ANIC); 1 么, Bay View, 28. i. 1963 (AMS); 3 么, Berowra, 26./29. x. 1917, 2. xi. 1917 (L. H. Mosse-Robinson) (ANIC); 1 么, Blackheath, 30. xi. 1953 (L. H. Mosse-Robinson) (ANIC); 1 么, Broadwater Nat. Park, 19. iii. 1981 (J. Humphreys & B. J. Laudon) (NSWA); 2 么, Broulee, 24. ii. 1962, 3. iii. 1963 (M. S. Upton) (ANIC); 2 么, 1 乆, Brunswick Heads, 4.–8. i. 1927 (W. B. Barnard) (QM); 1 么, Calga, 40 km N, 6. xi. 1987 (B. Hacobian) (ANIC); 8 么, Canyonleigh, 25. ii. 1968, 28.–29. iii. 1968 (V. J. Robinson) (ANIC); 1 么, Castlereagh SF, 50 km N Sydney, 27. x. 1982 (B. Hacobian) (ANIC); 1 么, Chatswood, 27. i. 1966 (V. J. Robinson) (ANIC); 1 乆, Como, 18. xi. 1897 (MV); 1 么, Como West, 21. xi. 1970 (V. J. Robinson) (ANIC); 3 么, Coonabarabran, 23 km NNE, 488 n, 7. xii. 1974 (I. F. B. Common & E. D. Edwards) (ANIC); 1 么, Crescent Head, 3 mls S, 27. iii. 1965 (I. F. B. Common & M. S. Upton) (ANIC); 1 乆, Dalmorton (Lea) (DPIQ); 2 么, Galston Gorge, Crusader

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Camp, 29.–30. xi. 1985 (at light) (S. G. Hunter) (DPIQ); 1 么, Glenorie, 16. ii. 1969 (at light) (G. Daniels) (AMS); 2 么, Gosford, 23. x. 1897 (MV); 2 乆, Gosford (MV); 2 么, Gosford, 19. xi. 1924 (ANIC); 1 么, Gosford, 25. ii. 1968 (V. J. Robinson) (ANIC); 1 么, Gosford, 4 mls SW, 30. iii. 1965 (I. F. B. Common & M. S. Upton) (ANIC); 1 么, Kuring-gai Chase, 10. i. 1975 (D. McAlpine & R. Sinqham) (AMS); 3 么, Manly, 21.–26. x. 1906, 29. xi. 1908 (AMS); 1 么, Maryland, 6. xi. 1934 (ANIC); 1 么, Megalong Valley, 25. xi. 1978 (B. Hacobian) (ANIC); 1 么, Mosman, 25. xi. 1909 (AMS); 5 么, Mt Kaputar, 10.–13. iii. 1969 (V. J. Robinson) (ANIC); 1 乆, Mt Kosciusko, 3000–3500 ft, 5. ix. 1912 (ANIC); 10 么, 1 乆, Narara, 9. ii. 1919, 1. iv. 1924, 22. xi. 1943, 8.–25. ii. 1951, 26. x. 1951, 29. iii. 1952, 20. ii. 1954, 2. iii. 1954 (L. H. Mosse-Robinson) (ANIC); 1 乆, Narrabeen, 25. xi. 1922 (AMS); 1 么, Newcastle (AMS); 1 么, Pilliga SF, 32 km E of Baradine, 24. x. 1984 (B. Hacobian) (ANIC); 3 么, Port Stephens, 1. xi. 1940 (AMS); 1 么, Port Stephens, 7. ix. 1922 (L. H. Mosse-Robinson) (ANIC); 1 么, Putty, 20 mls S, 14. xi. 1961 (I. F. B. Common & M. S. Upton) (ANIC); 1 么, 1 乆, Roseville, 9. ii. 1920, 20. xi. 1921 (L. H. Mosse-Robinson) (ANIC); 1 么, 1 乆, Rous, Richmond R., x. 1923, v. 1934 (V. J. Robinson) (ANIC); 2 么, Royal Nat. Park, Sydney, 11. iv. 1935 (AMS); 2 么, Singleton, 26 mls S, 7. xi. 1960 (I. F. B. Common & M. S. Upton) (ANIC); 1 乆, Somersby, 30. x. 1953 (L. H. MosseRobinson) (DPIQ); 3 么, 2 乆, Sydney (MV); 1 乆, Sydney, xii. 1918 (AMS); 1 乆, Sydney, 20. x. 1923 (AMS); 1 么, Sydney, Manly, 22. ii. 1935 (ANIC); 1 么, Tabbimoble SF, near Woodburn, e.l. 4. x. 1978 (B. Hacobian) (ANIC); 1 么, 1 乆, Tabulam, 25. xii. 1912 (ANIC); 4 么, Tabrabucca Creek, Barrington Tops, 4500 ft, 7. i. 1967 (V. J. Robinson) (ANIC); 3 么, Toronto, 17. 4. 1917 (MV); 2 么, Wahroonga, 19. xi. 1969 (M. Lovell) (AMS); 1 么, Wahroonga, 7. x. 1970 (A. B. Rose) (ANIC); 1 乆, Warrah, 26. 4. 1921 (MV); 11 么, Wilton, CSIRO Experimental Farm, 17.–25. ii. 1966, 31. x. 1966, 20. i. 1967, 11.–16. iii. 1967, 12. i. 1982 (V. J. Robinson) (AMS, ANIC); 5 么, Wirrimbirra near Picton, 29. iii. 1967, 30. xi. 1967, 1. xii. 1967 (V. J. Robinson) (ANIC); 2 么, 2 乆, Woy Woy, 21. ii. 1952, 12. ii. 1953, 3. x. 1953 (L. H. Mosse-Robinson) (ANIC). Australian Capital Territory: 14 larvae (5th instar), Aranda Bushland, Canberra, 21. xi. 1989 (on Hibbertia obtusifolia) (at night with headlamp) (G. M. Tarmann & E. S. Nielsen) (ANIC, GMT); 1 么, Aranda Bushland, Canberra, 22. xi. 1989 (G. M. Tarmann) (GMT); 27 么, Black Mountain Light Trap, 1. iv. 1961, 21. iii. 1962, 5. xii. 1962, 20.–22. iii. 1963, 18. iii. 1965, 4. iii. 1966, 3.–8. iii. 1967 (I. F. B. Common) (ANIC); 4 么, 2 乆, Black Mountain, e.l. 11.–21. xii. 1959 (larva on Hibbertia obtusifolia) (I. F. B. Common) (ANIC); 2 么, 1 乆, Black Mountain, Canberra, 12. xi. 1962, 21. xii. 1959 (e.l. ex Hibbertia obtusifolia), 4. iii. 1966 (light trap) (I. F. B. Common) (ANIC); 10 么, 6 乆, Black Mountain, CSIRO, 28. xi. 1989 (G. M. Tarmann) (ANIC, GMT); 1 乆, Canberra, 6. xii. 1927 (AMS); 1 么, Canberra, Aranda, 22. xi. 1989 (G. M. Tarmann) (GMT); 7 么, 2 乆, Jervis Bay, 5.–26. ii. 1918, 7. iii. 1918, 29. i. 1920, 9. ix. 1920, 10. x. 1920, 11. iii. 1923 (L. H. Mosse-Robinson) (ANIC).

Victoria: 1 乆, Birchip, 14. x. 1904 (MV); 2 么, 2 乆, Dimboola, 16. xi. 1903, 19. xi. 1907 (MV); 1 么, Kiata, xi. 1933 (MV); 2 么, 2 乆, Sea Lake, 24. xi. 1905, 1. xii. 1905 (MV); 8 么, 2 乆, Wyperfeld Nat. Park, 5. xi. 1966 (I. F. B. Common & M. S. Upton) (ANIC). South Australia: 1 乆, Adelaide, Belair, Sheoak Rd Scrub, 1 ml E, 3. iv. 1967 (SAM); 1 么, Robe, xi. 1960 (J. O. Wilson) (ANIC); 2 乆, Ungarra, 9 mls NW, 29. x. 1969 (Key & Upton) (ANIC). Tasmania: 2 么, Freycinet Nat. Park, 28. ii. 1963 (I. F. B. Common & M. S. Upton) (ANIC); 1 乆, Hobart, 23. ii. 1913 (G. H. Hardy) (AMS).

Diagnosis Smaller than Pollanisus viridipulverulenta, with narrower wings, shorter antenna with shorter pectinations in male and the hindwing with a translucent central part. Males and females almost equal in size (the male much larger in most populations of P. viridipulverulenta). Pollanisus apicalis is two or three weeks later on the wing than the sympatric P. viridipulverulenta.

Redescription Male (Pl. 9, Fig. 1; Pl. 10, Fig. 1). Forewing length: 7.5–9.5 mm. Head, thorax, forewing upperside and parts of forewing and hindwing underside covered with shiny, metallic, bluish, bluish green or golden green scales. Frons projecting beyond compound eyes in lateral view, of medium breadth, approximately 0.9–1.1× that of compound eyes in frontal view; palps short, upturned, shiny metallic; proboscis dark brown; ocelli large, white, distance from compound eye slightly less than diameter of ocellus; chaetosemata brown, the anterior extension covering the whole space between compound eye and ocellus, sometimes just allowing a very narrow strip of metallic bluish green scales (one row) ventrally. Antenna dark grey with weak blue sheen, segments 1 to 28–32 bipectinate, 28–32 to 41–45 biserrate; the pectinations 5–6× longer than breadth of shaft in dorsal view at segment 15, 4–5× longer at segment 25, sensory hairs on pectinations short. Legs not so intensely shiny as thorax. Abdomen dark greenish brown, slightly shiny but not metallic. Forewing upperside metallic blue, bluish green or golden green, distal margin and fringe blackish brown; underside dark grey-brown, with a few shiny bluish or bluish green scales proximally at costa and anterior of cell. Hindwing upperside uniform blackish brown, slightly translucent medially; underside of same colour as upperside, with shiny scales costad of medial stem and at anal angle. Medial

TAXONOMY

stem absent in forewing, present in hindwing. The intensity of the metallic sheen on forewing upperside is very variable and varies from very intense to almost matt. The colour variablility is also remarkable. The forewing upperside of this species very often gives the impression of a creamy yellow tinge in bright daylight. Female (Pl. 9, Fig. 2). Forewing length: 7.5–8.5 mm. Similar to male and equal in size, wings more rounded, with a yellow abdominal hairtuft. Head much broader and compound eyes smaller than in male, breadth 1.9–2.1× that of compound eyes in frontal view. Antenna biserrate. Male genitalia (Figs 157–160, 206–209). Valva with pointed apex (ca. 50–60°), folded, translucent, central part triangular, ventral sclerotization broad. Aedeagus short and stout, 2.5–3.0× longer than broad, cornutus ca. 80% the length of aedeagus, pointed distally. Female genitalia (Figs 252, 253). The sclerotization at centre of praebursa usually with two long, curved teeth.

Phenology and bionomics The adults are found from early September to late March. According to the available label data this species seems to be bivoltine in most of the southern areas but perhaps it is trivoltine in Queensland. Personal observations of the Black Mountain population in Canberra (ACT) (November 1989) follow. The adults are weak fliers. They prefer sunshine and are often found in open bushland and light forest in half-shadow (Pl. 64, Fig. 2). They often feed during the hottest hours of the day, especially at Helichrysum flowers. In the afternoon the males can be observed searching for the females. Mating takes place in late afternoon and lasts till late morning of the next day. At midday the females deposit their eggs in small clusters of 8–15 on the flowers and leaves of Hibbertia bushes, covering them with the deciduous, spiny, yellow scales of their abdominal hairtuft. During oviposition the females obtain nectar several times. Egg Ovoid, subquadrate, yellow, with slightly rough surface; length 0.5 mm, breadth 0.4 mm. The eggs are laid in small layers and are covered with the yellow, spiny, deciduous scales from the female

abdomen (as in P. viridipulverulenta, Figs 129, 130). The eggs are deposited on leaves close to the ground mainly on small bushes and in shadow. Duration of egg stage: 5–7 days. Larva First instar. Length 1.1–1.2 mm, yellowish green, head light brown, eyes black, prothoacic shield large, dark brown, with long, light setae; anal comb fully developed, blackish brown, with short dentations that are longer medially, two brown spots dorsad anal comb; thoracic legs long, brown, abdominal prolegs yellowish green. Setal combination formula of first abdominal segment (see p. 38): D: 1d; SD: 1d, 1l; L: 2l (Fig. 131). The habits of the L1 larvae are similar to those of P. viridipulverulenta (see p. 71). Duration of L1: ca. 10–12 days. Second instar (L2). Length 2.0–2.2 mm, similar to L1, setal combination D: 5d; SD: 4d, 1l; L: 2l. Habits as in L1. Duration of L2 ca. 10–12 days. Third instar (L3). Length 3.0–3.4 mm, simlar to L1 and L2, setal combination D: 8d; SD: 6d, 2l; L: 4l. Habits as in L1 and L2. Duration of L3 ca. 17–19 days. Sixth instar (L6), fully grown (Pl. 59, Fig. 6; Pl. 60, Figs 1, 2; Figs 132, 133). Length 14.5–15.0 mm; head dark brown, thoracic legs blackish brown, prothoracic shield consisting of two dark brown subtriangular plates, densely covered with long and short setae, the longer pointing forwards; ground colour of larva yellow, dorsal line broad, bright white, a broad, reddish brown band along dorsal verrucae, lateral and ventral part of larva yellow, subdorsal verrucae encircled by a reddish brown ring and lateral verrucae by a reddish brown halfring; anal comb fully developed, with short dentations; verrucae very uniform on all segments, setal combination D: ca. 45d; SD ca. 40d, 6l; L: ca. 20l. Pupa (Fig. 134) Length 9.0 mm; light brown, eyes black, appendages very weakly fused to each other and to body, loosening from body if larva is preserved in alcohol; proboscis and antennae not exceeding 6th abdominal segment, abdominal segments 2–7 么 and 3–7 乆 with one row of spines of equal triangular form, cremaster consisting of four sockets bearing each one short, distally hooked seta. Cocoon (Fig. 134) Flat ovoid, silky white, spun between leaves on the ground.

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Parasitoids (Fig. 135) One Tachinidae was reared from a larva from Black Mountain (Canberra, ACT). Identification has not yet been possible. Foodplants Confirmed larval foodplants are: Hibbertia obtusifolia DC. (ACT) (Pl. 62, Fig. 1) (I. F. B. Common, label data, E. S. Nielsen & G. M. Tarmann, personal observation, M. Horak, personal observation and rearing), H. virgata R. Br. ex DC. (South Australia) (McFarland, 1979).

Distribution From northern Queensland throughout New South Wales (including ACT) to Victoria and South Australia, Tasmania.

Pollanisus trimacula (Walker) Pl. 11, Figs 1, 2; Figs 161–164, 210, 211, 254. Procris trimacula Walker, 1854, List Specimens lepid. Insects Colln Br. Mus. 1: 110. Holotype 乆, AUSTRALIA: New South Wales, Sydney (BMNH) [examined]. Procris trimacula Walker; Butler 1877: 13; Meyrick 1886: 794. Pollanisus trimacula (Walker); Jordan 1907: 10; Hering 1922: 13; Bryk 1936: 119; Tarmann 1996: 143. Pollanisus trimaculus (Walker); Turner 1926b: 442; Common 1970: pl. 7N (colour painting).

Material examined Queensland: 1 么, Dandabah, Bunya Mountains Nat. Park, 1000 m, 1. xii. 1988 (I. F. B. Common) (ANIC); 1 乆, Fraser Park, 29. xi. 1946 (A. B.[urns]) (MV). New South Wales: 1 么, Barrington R., 4500 ft, 12. x. 1962 (mv light) (C. W. Frazier) (ANIC); 4 么, Bungwahl, 5 mls

N, 24. iii. 1965 (I. F. B. Common & M. S. Upton) (ANIC); 1 么, Chichester SF, near Dungog, 8. xii. 1979 (B. Hacobian & N. Goodman) (ANIC); 4 (males), Depot Beach, 10 mls NE Bateman’s Bay, 14.–23. iii. 1970, 30. iii. 1975 (I. F. B. Common) (ANIC); 2 乆, Dorrigo (SAM); 6 (males), Dorrigo Nat. Park, 19.–20. iii. 1969 (V. J. Robinson) (ANIC); 2 (males), Ebor, 3. iv. 1938 (B. L. Middleton) (AMS); 1 么, 3 乆, Gosford, 8. xi. 1896, iv. 1943 (AMS, MV); 1 乆, Jamberoo Mountain, e.l. 16. iii. 1963 (V. J. Robinson) (ANIC); 1 乆, Lilyvale, 6. iv. 1907 (MV); 1 乆, Liverpool Ra., 17. iv. 1921 (MV); 2 乆, Manly, 21. xi. 1909 (AMS); 3 (males), 1 乆, Minnamurra Falls, 8. iv. 1969 (V. J. Robinson) (ANIC); 1 么, Mittagong, 10. iv. 1926 (AMS); 1 么, 2 乆, Mooney Mooney Creek, near Gosford, 16.–30. xi. 1978, 3. xii. 1984 (D. K. McAlpine & B. J. Day) (AMS); 1 么, Mt Dromedary, 1000 ft, 24. xi. 1965 (I. F. B. Common & M. S. Upton) (ANIC); 3 么, 1 乆, Mt Keira, 12. iv. 1967, 22.–30. xii. 1967 (V. J. Robinson) (ANIC); 1 乆, Mt Kembla, 2. i. 1968 (V. J. Robinson) (ANIC); 20 么, 3 乆, Narara, 27. iv. 1924, 3. iii. 1935, 2.–8. iv. 1944, 2. iii. 1947, 1. iv. 1947, 1. xi. 1947, 9. iv. 1949, 23. ii. 1950, 29. iii. 1950, 5.–6. iv. 1950, 6.–24. x. 1950, 30. x. 1952, 3. xi. 1952, 23. ii. 1953 (L. H. Mosse-Robinson) (ANIC, MV); 1 么, Newcastle, 15. iv. 1966 (V. J. Robinson) (ANIC); 1 么, Niagara Park, 20. xi. 1948 (W. Brandt) (ANIC); 1 么, 1 乆, Otford, 22. xi. 1962, 23. xii. 1963 (V. J. Robinson) (ANIC); 1 乆, Palm Beach, 22. x. 1917 (MV); 1 么, Pine Creek SF, 6 km NNE Raleigh, 16. xi. 1966 (I. F. B. Common & E. D. Edwards) (ANIC); 1 么, 1 乆, Port Macquarie, 17. iv. 1922 (AMS, MV); 1 么, Port Macquarie, 16. i. 1933 (A. L. Brown) (MV); 2 么, Port Macquarie, 23. ii. 1947 (L. H. Mosse-Robinson) (ANIC); 1 么, Port Macquarie, 2. x. 1955 (C. W. Frazier) (ANIC); 13 (males), Port Macquarie, 3 mls S, 26. iii. 1965 (I. F. B. Common & M. S. Upton) (ANIC); 1 么, Port Macquarie, 7. v. 1966 (D. Reed) (ANIC); 3 (males), Port Macquarie, 3 mls SSE, 15. v. 1966 (M. S. Upton) (ANIC); 1 么, Rosebank, 7 mls W, 1700 ft, 8. xi. 1961 (I. F. B. Common & M. S. Upton) (ANIC); 1 乆, Rose Bay, 1895 (MV); 4 么, 4 乆, Royal Nat. Park, 21. ii. 1925, 12. iv. 1925 (AMS, ANIC, MV); 4 乆, Royal Nat. Park, 5. iv. 1959 (on flowers of Parsonsia straminea (R. Br.) F. Muell.) (C. E. Chadwick) (DPIQ); 1 么, Royal Nat. Park, e.l. 2. x. 1970 (larva on Hibbertia scandens (Willd.) Gilg.) (V. J. Robinson) (ANIC); 1 么, St George’s Basin, 3. xii. 1964 (V. J. Robinson) (ANIC); 2 乆, Sydney (SAM); 2 乆, Stanwell Park, 5. iv. 1898 (W.C.D.C) (MV); 1 么, Stanwell Park, 3. iv. 1898 (SAM); 2 么, 4 乆, Stanwell Park, ii. 1913, 23. iv. 1916, 29. iii. 1924 (AMS, MV); 1 么, Stanwell Park, 16. iii. 1932 (L. H. Mosse-Robinson) (ANIC); 1 乆, Stanwell Tops, iii. 1953 (V. J. Robinson) (ANIC); 2 么, Styx R. SF, E of Armidale, 3. i. 1983 (B. Hacobian & N. Goodman) (ANIC); 1 么, Tuglow, 48 km N Singleton, 3. xii. 1977 (C. N. Smithers) (AMS); 1 么, Tyringham, 10. xii. 1939 (ANIC); 1 乆, Ulladulla, 2 mls N, e.l. xii. 1963 (larva on Hibbertia dentata DC.) (I. F. B. Common) (ANIC); 1 么, Watagan SF, near Wyong, 18. xii. 1984 (B. Hacobian) (ANIC); 1 么, Wollongong, 6. iv. 1964 (V. J. Robinson) (ANIC).

TAXONOMY

Diagnosis Although P. trimacula is supposed to be a very distinct species because of its wing pattern, dark males and especially dark females are very similar to other species of the P. acharon- and the P. eumetopus-groups. The best characters by which to recognise P. trimacula are, except for the ochreous spots on the forewing upperside: dark proboscis, coppery metallic sheen on collar and dorsal parts of abdomen in strong contrast to the bluish green or golden green shiny head-parts, and the distinct bands on the underside of the wings and on the lateral parts of the thorax below the wings. These characters are very constant. Most similar species have a yellow proboscis and lack the significant contrast in colours as described above.

Redescription Male (Pl. 11, Fig. 1). Forewing length: 7.5–9.5 mm. Head dark grey-brown dorsally, frons and lateral areas shiny metallic golden green, ventral margin of frons and labial palps light grey; frons strongly protruding beyond compound eye in lateral view, as broad or slightly narrower than large, black compound eyes in frontal view; labial palps short, porrect distally; proboscis dark brown proximally, yellowish brown distally, if rolled usually appearing dark brown; ocelli of medium size, distance from compound eye as long as diameter of ocellus; chaetosemata brown, with long anterior extension, with a single row of metallic scales ventrally. Antenna long, pectinations of medium length and very often arranged in an acute angle to shaft, giving the antenna a slender appearance; segments 1 to 36–38 bipectinate, 36–38 to 46–50 biserrate, sensory hairs very short. Thorax grey-brown dorsally except for a conspicuous, shiny metallic golden collar with a faint green tinge, grey-brown also ventrally, but strongly shiny metallic bluish green or golden green laterally (concolorous with anal angle of hindwing). Legs dark grey with a few shiny metallic bluish green or golden green scales, especially on distal part of femur (stronger on femur of hindleg). Wings relatively narrow and elongate; forewing upperside grey-brown with variably developed 3–6 whitish ochreous spots, that are sometimes almost completely reduced but a few ochreous scales are always present (visible under high magnification only); hindwing dark grey-brown with translucent central part. Underside of both fore- and hindwing very distinct, dark grey-brown, with a conspicuous, shiny bluish green or golden green metallic band on

each below costa, and with some shiny scales also at anal angle of hindwing. Abdomen upperside shiny metallic coppery, except for anterior segments in some specimens. Female (Pl. 11, Fig. 2). Forewing length: 8.0–9.0 mm. Similar to male but forewing upperside usually darker and with fewer and less well-defined ochreous spots; collar golden coppery; abdominal hairtuft bright yellow. Antenna with slender and dorsoventrally slightly flattened shaft that is slightly thicker at approximately two-thirds of length of antenna, biserrate, the dentations very close together forming a ventral groove proximally, strongly pointed apically. Male genitalia (Figs 161–164, 210, 211). Valva with pointed apex, folded translucent central part triangular, ventral sclerotization broad, ventral margin of valva straight. Aedeagus short, stout, 2.5–3.5× longer than broad, cornutus as long as aedeagus, slender, pointed distally. Female genitalia (Fig. 254). Ductus bursae slender, leading into a large praebursa with strong central sclerotization bearing usually three teeth.

Phenology and bionomics There are no published records available. According to label data the species appears to be bivoltine, the late summer generation being more abundant than the spring generation. According to the label data some specimens in the ANIC have been reared from larvae feeding on Climbing Guinea-Flower (Hibbertia scandens (Willd.) Gilg) (Pl. 62, Fig. 4) and Twining Guinea-Flower (Hibbertia dentata DC.) (Dilleniaceae). These observations indicate that P. trimacula, if monophagous on climbing Hibbertia species, is restricted to the coastal rainforests of New South Wales (Pl. 64, Fig. 3) and southern Queensland, and the known distribution does, in fact, correspond with the range of these plants. Several females have been collected on flowers of Parsonsia straminea (R. Br.) F. Muell. (Apocyanaceae) and the habits therefore seem to be similar to those described earlier for other species (see pp. 71, 74, 76).

Distribution South-eastern Queensland, eastern parts of New South Wales.

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Externally very similar to P. subdolosa subdolosa but differs in the significantly smaller compound eyes and broader frons in the male. P. subdolosa clara ssp. n. has more translucent hindwings and the metallic coppery sheen on the collar and abdomen is more intense. P. contrastus sp. n. is smaller and has narrower wings, the contrast between the dark opaque forewings and the very translucent hindwings is significant. All species from the tropical rainforest of Queensland (i.e. P. acharon, P. eungellae sp. n., P. eumetopus, P. commoni sp. n., P. incertus sp. n., P. angustifrons sp. n.) are smaller and have much narrower wings.

Pollanisus edwardsi sp. n. Pl. 12, Figs 1, 2; Figs 165, 166, 212, 213, 255.

Material examined Holotype 么: Australian Capital Territory: Bendora, 1200 m, 21. i. 1981 (I. F. B. Common) (ANIC). Paratypes Queensland: 1 么, Bunya Mountains, 3500 ft, 4. x. 1919 (ANIC); 1 么, Bunya Mountains, 19. iii. 1926 (W. B. Barnard) (QM); 1 么, Burleigh Heads, 20. ix. 1934 (I. F. B. Common) (ANIC); 1 么, 2 乆, Caloundra, 31. viii. 1912, 2. ix. 1912 (ANIC); 2 么, Stanthorpe, ii. (ANIC); 2 么, 1 乆, Stradbroke Island, 28. ix. 1906, 10. i. 1903 (ANIC). New South Wales: 2 么, 1 乆, Armidale, 3. iii. 1956 (C. W. Frazier) (ANIC); 1 么, Gosford, iv. 1943 (AMS); 1 么, Mount Kosciuszko, 3500 ft, 23. i. 1914 (AMS); 1 么, Mount Kosciuszko, 4500 ft, 23. i. 1914 (ANIC); 1 么, Tenterfield, ii. (ANIC). Victoria: 1 么, Warragul, 15. ii. 1975 (C. G. L. Gooding) (ANIC). There are four male specimens in the material examined that have slightly narrower wings. They are probably not conspecific but they are also not referable to any other described species of Pollanisus. They are mentioned below but excluded from the type series and the distribution map. Queensland: 1 么, Southport, iv. 1937 (J. M[acqueen]) (ANIC); 2 么, Stradbroke I., 10. ix. 1911 (ANIC); 1 么, Toowoomba, 6. iii. 1929 (UQIC); 1 么, Toowoomba, 18. xi. 1933 (E. J. Dumigan) (UQIC).

Diagnosis Pollanisus edwardsi is the largest Pollanisus species without a shiny forewing upperside. It can be recognised by its broad wings with relatively rectangular hindwings, the only slightly translucent hindwings not contrasting strongly with the opaque forewings and by its frons that is significantly broader than the breadth of the small compound eye in frontal view.

Description Male (Pl. 12, Fig. 1). Forewing length: 9.0–11.0 mm. Head dark grey-brown dorsally, sometimes with a few shiny metallic coppery scales, especially medially, frons dark grey dorsally, light grey ventrally, in most specimens covered with shiny metallic yellowish green or bluish green scales, head shiny metallic blue or greenish blue dorsolaterally; frons broad, significantly broader (ca. 1.5×) than small compound eyes in frontal view, protruding beyond compound eye in lateral view, more so dorsally, labial palps dark grey proximally, light grey distally, short, slightly upcurved; proboscis yellow; ocelli small, white, distance from compound eye approximately twice as long as diameter of ocellus; chaetosemata dark grey, lighter posteriorly, a broad stripe of shiny metallic blue, bluish green or yellowish green scales between ocellus and dorsal margin of compound eye; antenna greyish black, segments 1–27 bipectinate, 28–39 biserrate, bluntly pointed distally, the pectinations of medium length, 5–6× longer than breadth of shaft in dorsal view at segment 10, slightly tapering distally, sensory hairs very short. Thorax dark grey-brown dorsally, patagia, anterior part of tegulae and anteriormost part of mesothorax metallic coppery, with weak sheen, metallic blue laterally, dark grey ventrally; legs dark grey, midleg and hindleg metallic blue laterally on coxa and femur. Abdomen metallic coppery dorsally, greyish black ventrally. Wings broad, forewing broad, triangular, hindwing almost rectangular but rounded apically; forewing upperside and underside dark grey-brown, underside slightly paler with a few shiny metallic blue scales below costa proximally; hindwing dark, slightly translucent medially, upperside dark greybrown, underside similar with metallic blue scales in a band between cell and costa and at anal angle, with a green-blue tinge.

TAXONOMY

Female (Pl. 12, Fig. 2). (Description based on four females from localities where males are known and the females are believed to be conspecific). Forewing length: 7.0–8.5 mm. Similar to male but smaller, wings narrower. Frons broader, compound eyes smaller, antenna biserrate. Abdominal hairtuft yellow. Not significantly different from females of closely related species. Male genitalia (Figs 165, 166, 212, 213). Valva pointed distally, slightly convex dorsally, straight ventrally, folded translucent central part triangular, ventral sclerotization of valva broad proximally, lancet-shaped. Aedeagus approximately 3× longer than broad, cornutus as long as aedeagus, slender, pointed apically. Female genitalia (Fig. 255). Ductus bursae short; praebursa large, with short lateral appendix, posterior part with slightly sclerotized wall, a ring-like structure well visible, anterior part of praebursa large and completely translucent, the central sclerotization somewhat triangular, with a doublepointed tooth.

Phenology and bionomics In Victoria and New South Wales probably univoltine, flying in midsummer, in Queensland bivoltine. Bionomics unknown.

Distribution From southern Queensland to Victoria.

Pollanisus subdolosa (Walker) Procris subdolosa Walker, [1865], List Specimens lepid. Insects Colln Br. Mus. 31 (Suppl.): 62. Holotype 么, ‘Austral’ [AUSTRALIA, Melbourne (see remarks below)] (BMNH) [examined]. Procris subdolosa Walker; Meyrick 1886: 793. Pollanisus subdolosa (Walker); Kirby 1892: 87; Jordan 1907: 10; Hering 1922: 13; Bryk 1936: 119; Alberti 1954: 298; Common 1990: 297 (bionomics, ?partim); Tarmann 1994: 122 (male genitalia); Tarmann 1996: 143. Pollanisus subdolosus (Walker); Turner 1926b: 442. Walker’s original description does not correspond very well with the characters of the type-specimen. In particular, the colour of the hindwings is not green but dark grey and the antennae are not cupreous. However, the relatively broad wings, the frons of medium breadth and the large eyes are significant enough to recognise this species clearly in the male. Unfortunately, the females are not clearly referable to P. subdolosa as there are no significant differences to distinguish them from closely related species. Moreover, as there are two subspecies involved (see below) it is necessary to state where the ‘typical P. subdolosa’ has been collected. Walker mentions in the original description ‘Australia. From Mr. Damel’s collection’. According to Musgrave (1932) Edward Damel was a collector for the Godeffroy Museum in Hamburg. From 1853 to 1858 Damel was based in Sydney, but he was travelling and collecting around New South Wales and Victoria during that time (E. D. Edwards, personal communication). The type corresponds best with specimens from Beaconsfield (Victoria) and may originate from the environs of Melbourne.

Pollanisus subdolosa subdolosa (Walker) Pl. 13, Figs 1, 2.

Material examined

Etymology Named in honour of E. D. [Ted] Edwards (CSIRO, Canberra) who is one of the leading lepidopterists of Australia and whose invaluable collaboration has contributed significantly to this volume.

Victoria: 1 么, Bayswater, 12. iii. 1922 (MV); 2 么, 1 乆, Beaconsf[ie]ld, 2.–6. xii. 1904 (MV); 1 么, Beaconsf[ie]ld, 28. xi. 1904 (MV); 1 么, B[ea]consfield, 14. iii. 1904 (MV); 1 么, Belgrave, 9. iii. 1929 (C. [H.] B[orch]) (MV); 1 么, Cann Riv[er], xi. 1928 (J. Clark) (MV); 1 么, Dutson (M. J. Wise) (MV); 1 么, Meeniyan, iv. 1896 (MV); 1 么, Ringwood, 28. ii. 1920 (MV); 3 么, Upwey, 20. xii. 1930, 14. iii. 1931 (C. [H.] B[orch]) (MV).

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Diagnosis Pollanisus subdolosa subdolosa is recognised by its relatively broad wings, the hindwing being somewhat rectangular, almost opaque and not contrasting strongly with the forewings, and by its narrow frons that is almost the same breadth as the size of the compound eye in frontal view. It is very similar to P. edwardsi; it differs only in the significantly narrower frons and larger compound eyes in the male and the slightly more opaque hindwings. Pollanisus subdolosa clara ssp. n. has more translucent hindwings, the metallic coppery sheen on the collar and abdomen is more intensive (in fresh specimens!). Pollanisus contrastus sp. n. is smaller and has narrower wings, the contrast between the dark opaque forewings and the very translucent hindwings is significant and the frons is much broader. All species from the tropical rainforest of Queensland, i.e. P. acharon, P. eungellae sp. n., P. eumetopus, P. commoni sp. n., P. incertus sp. n., P. angustifrons sp. n., are smaller and have narrower wings.

Redescription Male (Pl. 13, Fig. 1). Forewing length: 8.0–10.5 mm. Head blackish brown dorsally, sometimes with a few shiny metallic coppery scales, especially mediodorsally, frons dark grey dorsally, light grey ventrally, covered with a few shiny metallic yellowish green scales, lateral head-parts shiny metallic yellowish green dorsally, sometimes with a bluish tinge; frons slightly narrower or as broad as compound eye in frontal view, protruding beyond compound eye in lateral view, more so dorsally, labial palps dark grey, slightly upcurved; proboscis yellow; ocelli of medium size, white, distance from compound eye approximately 1.5× as long as diameter of ocellus; chaetosemata dark grey, lighter posteriorly, a broad band of shiny metallic blue, bluish green or yellowish green scales between ocellus and dorsal margin of compound eye; antenna greyish black, 1–24(26) bipectinate, 24(26)–35(36) biserrate, bluntly pointed distally, the pectinations relatively short, 4–5× longer than breadth of shaft in dorsal view at segment 10, slightly tapering distally, sensory hairs very short. Thorax greyish black dorsally, patagia, anterior part of tegulae and mesothorax shiny metallic coppery, the sheen weak, in old specimens often absent, shiny metallic yellowish green laterally with bluish tinge, dark grey ventrally; legs dark grey, midleg and hindleg shiny metallic yellowish green or bluish greeen laterally at coxa and femur. Abdomen weakly shiny metallic coppery dorsally, greyish black ventrally.

Wings broad, forewing broad, triangular, hindwing almost rectangular but rounded apically; forewing upperside dark grey-brown, underside slightly paler and in some specimens with a few shiny metallic bluish green scales below costa proximally; hindwing almost opaque, only very weakly translucent, upperside unicolorous greyish black, underside dark grey with a few shiny, metallic bluish green scales anterior of medial stem and at anal angle. Female (Pl. 13, Fig. 2). Forewing length: 7.0–8.0 mm. Similar to male but smaller, wings somewhat narrower. Frons broader, compound eye smaller, antenna biserrate. Abdominal hairtuft yellow. Not significantly different from females of closely related species. Male genitalia (Figs 167, 214, 215). Valva with sharply pointed apex, slightly convex dorsally at half length, mainly straight towards apex, folded translucent central part triangular, ventral sclerotization of valva broad proximally, lancet-shaped. Aedeagus approximately 3× longer than broad, slender, cornutus as long as aedeagus, sharply pointed apically. Female genitalia (Fig. 256). Praebursa with very variable central sclerotization bearing a variable number of teeth. The variation of this sclerotization is so outstanding that it gives the strong impression that the taxon P. subdolosa, as treated in this revision, is still a species-group. However, the variability of the central sclerotization could only be elucidated from examination of a series of material reared from eggs, an interesting task for the future.

Remarks The possibility cannot be excluded that P. subdolosa subdolosa and P. subdolosa clara ssp. n. might be different species. The notable differences in habitus, especially when populations from northern New South Wales and Queensland are compared with those from Victoria, give a strong impression of a specific difference. Nevertheless, the head capsule is very similar. Even the coloration of the frons, i.e. darker grey dorsally and slightly covered with shiny scales, is the same. The large number of specimens that are difficult to identify as one or other taxon makes it almost impossible to recognise two different species at present. Treating P. subdolosa clara ssp. n. as a subspecies enables one to place most of the material. If specific differences between P. s. subdolosa and P. s. clara ssp. n. are eventually recognised, the established names can be used.

TAXONOMY

Phenology and bionomics Bivoltine according to the label data. Bionomics unknown.

Distribution Victoria.

Pollanisus subdolosa subdolosa Pollanisus subdolosa clara

B. Common & M. S. Upton) (ANIC); 1 么, Cordeaux Dam, 10. iii. 1972 (V. J. Robinson) (ANIC); 3 么, Depot Beach, 10 mls NE Bateman’s Bay, 17. ii. 1968, 4. iv. 1970, 17. x. 1970 (I. F. B. Common) (ANIC); 1 么, Fitzroy Falls, 22. iii. 1971 (V. J. Robinson) (ANIC); 3 么, 2 乆, Gosford (MV); 12 么, Gosford, 23. ii. 1968 (V. J. Robinson) (AMS, ANIC); 1 么, Gosford, 4 mls SW, 30. iii. 1965 (I. F. B. Common & M. S. Upton) (ANIC); 2 么, Minnamurra Falls, 9. iii. 1974 (V. J. Robinson) (ANIC); 1 么, Minyon Falls, 6 km W of Rosebank, 19. xi. 1976 (I. F. B. Common & E. D. Edwards) (ANIC); 9 么, 2 乆, Mosman, 20. x. 1909, 8. xi. 1909, x. 1912 (AMS); 1 么, Narara, 21. xi. 1924 (L. H. MosseRobinson) (ANIC); 1 么, 2 乆, Niagara Park, 26. x. 1948, 28. i. 1949 (W. Brandt) (ANIC); 1 么, 1 乆, Roseville, 16. iv. 1904, 20. iv. 1915 (MV); 1 么, Ryan’s Creek Rd, Depot beach, 16 km NE of Bateman’s Bay, 14. iv. 1975 (I. F. B. Common) (ANIC); 2 么, 2 乆, N Sydney, 28. x. 1896, 5. iv. 1903, 24. iv. 1903, 3. iv. 1904 (MV); 1 么, Sydney, 18. x. 1896 (MV); 1 么, Tabbimoble SF, near Woodburn, 4. x. 1978 (at light) (B. Hacobian) (ANIC); 1 么, 1 乆, Watagan SF, near Wyong, 17. iii. 1979 乆, 21. xi. 1982 (B. Hacobian) (ANIC); 1 么, Wilton, CSIRO Experimental Farm, 11. x. 1977 (V. J. Robinson) (ANIC); 1 么, Woolgoolga, 30. iv. 1966 (V. J. Robinson) (ANIC).

Pollanisus subdolosa clara ssp. n. Pl. 14, Figs, 1, 2; Pl. 15, Fig. 1; Pl. 59, Figs 2–5; Figs 167, 214, 215, 257, 436–448.

Material examined Holotype 么, New South Wales: Minyon Falls, 6 km W of Rosebank, 28.37´S 153.23É, 19. xi. 1976 (I. F. B. Common & E. D. Edwards) (ANIC). Genitalia slide Z 3040. Paratypes New South Wales: 6 么, Minyon Falls, 6 km W of Rosebank, 28.37´S 153.23É, 19. xi. 1976 (I. F. B. Common & E. D. Edwards) (ANIC). There is one male in the series of specimens from the type-locality collected together with the types and on the same day that has a slightly broader frons and is slightly larger. This specimen is excluded from the type-series as sympatry of closely related and yet unrecognised species cannot be excluded. While the following specimens also are almost certainly referable to P. subdolosa clara and therefore referred to that subspecies in the distribution map, they are not included in the type series for reasons mentioned above. Queensland: 1 么, Caloundra, 28. viii. 1923 (ANIC); 1 么, Coolangatta, 4. v. 1913 (ANIC); 1 么, C[unning]ham’s Gap, 19.–21. xi. 1949 (ANIC); 1 么, Toowoomba, 19. iii. 1974 (J. Macqueen) (ANIC). New South Wales: 1 么, Bateman’s Bay, 20. iii. 1975 (V. J. Robinson & N. Grant) (ANIC); 1 么, Boonoo Boonoo R., 29. xi. 1981 (at mv light) (G. Daniels & M. A. Schneider) (UQIC); 23 么, Budawang Nat. Park, Western Distributor Rd, Carters Creek, 200 m, 1. xii. 2003 (at mv light) (A. Zwick) (ANIC); 1 么, Bungwahl, 5 mls N, 24. iii. 1965 (I. F.

Diagnosis Pollanisus subdolosa clara differs from P. subdolosa subdolosa by its more translucent hindwings and the metallic coppery sheen on the collar and abdomen is more intensive. The tropical rainforest species of Pollanisus from the coastal parts of northern Queensland, i.e. P. eungellae sp. n., P. angustifrons sp. n., P. eumetopus, P. commoni sp. n., P. incertus sp. n., P. acharon, have significantly narrower wings with a straighter costa on the forewing; the colour of the shiny metallic scales is not so coppery but more shiny. Pollanisus subdolosa clara differs from P. edwardsi in its narrower frons and larger eyes in the male. The female is very similar and once more one has to accept that it might not be possible to separate the females properly, if not collected in a series with males at the same locality.

Description Male (Pl. 14, Fig. 1; Pl. 15, Fig. 1). Forewing length: 7.5–9.0 mm. Head blackish brown dorsally, paler ventrally, with shiny metallic yellowish and/or bluish green scales dorsolaterally; frons as broad or slightly narrower than compound eyes in frontal view, dark grey dorsally light grey ventrally, with a few single shiny metallic yellowish green scales, protruding beyond the large compound eyes in lateral view, more so dorsally; labial palps small and slender, slightly upcurved, dark grey proximally,

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light grey distally, strongly pointed apically; proboscis yellow; ocelli large, white, distance from compound eye as long or slightly shorter than diameter of ocellus; chaetosemata brownish grey, lighter grey posteriorly, with long anterior extension, a single row of shiny metallic blue scales between chaetosema and dorsal margin of compound eye; antenna greyish black with a slighty purplish tinge at shaft basally, segments 1 to 28–30 bipectinate, 28–30 to 38–41 shortly biserrate, pointed apically, the pectinations of medium length, ca. 5× longer than breadth of shaft in dorsal view at segment 10, tapering distally, sensory hairs short. Thorax blackish brown dorsally, patagia and anterior part of tegulae shiny metallic coppery golden, shiny metallic blue laterally, dark grey ventrally; legs dark grey, with a few shiny metallic blue scales on coxa and femur laterally, especially on hindleg. Abdomen shiny coppery golden dorsally, greyish black ventrally. Wings of medium breadth, forewing elongate with a significantly arched costa distally and rounded apex, hindwing broad subtriangular with rounded apex, anterior margin rounded but sometimes slightly concave between CuA1 and CuP; forewing upperside blackish brown, underside sometimes with a few shiny metallic blue scales below costa; hindwing upper and underside dark grey-brown, slightly translucent medially, underside with a band of shiny metallic greenish blue or blue scales along cell below costa and at anal angle. Female (Pl. 14, Fig. 2). Forewing length: 7.5 mm; Similar to male but smaller, with narrower wings, smaller compound eyes and broader frons. The shiny metallic scales on the dorsolateral parts of the head more yellow, and on the lateral areas of thorax more greenish. Antenna biserrate. Abdomen with yellow hairtuft.

Phenology and bionomics Bivoltine according to the label data. On 1 December 2003 a large series of males was collected at Carters Creek, Western Distributor Road in Budawang National Park (NSW) by Andreas Zwick at mv light. The specimens were attracted by light early in the morning before sunrise. In spite of this observation, the main time of activity should be during the day as in all other Australian zygaenid species. However, this observation indicates that there might be early morning activity in the males. A few days later, some more males and females

were observed. One female was enclosed in a container and oviposited after leaves of Hibbertia scandens (Willd.) Gilg., the presumed larval hostplant, were added. Andreas Zwick (CSIRO, Canberra), who collected the specimens and reared the larvae from the eggs, provided excellent cryoSEM figures and colour pictures of the egg and first instar larva and allowed me to publish them in this book as Appendix II (Figs 435–447). Egg Ovoid, subquadrate, yellow, with reticulate surface (Pl. 59, Fig. 2; Figs 437, 438); length 0.5 mm, breadth 0.4 mm. The eggs are laid in small batches and are covered with the yellow, spiny, deciduous scales from the female abdomen (Fig. 436). Duration of egg stage: 5–7 days. Larva First instar (L1) (Pl. 59, Fig. 3; Figs 441–448). Length 1.0–1.1 mm, yellowish green, with irregular brown spots, head light brown, eyes black, prothoracic shield large, brown, with long, light setae; anal comb fully developed, blackish brown, with short dentations that are longer medially; thoracic legs long, brown, abdominal prolegs yellowish green. Setal combination formula of first abdominal segment (sensu Efetov and Tarmann 1999; Efetov et al. 2000; Efetov 2001; Efetov, Mollet and Tarmann 2003): D: 1d; SD: 1d, 1l; L: 2l (Fig. 131, 443) [D = dorsal setae, SD = subdorsal setae, L = lateral setae; d = dark, stiff seta (like a spine), l = light, flexible seta]. The cuticula of the body bears unispined microtubercles (Fig. 447). The L1 larvae feed on the leaves of Hibbertia scandens, producing small oval grooves by inserting the head into the parenchyma of the plant. Third instar (L3). Length 3.0–3.4 mm, similar to L1 but with light brown dorsolateral lines. Habits as in L1 but the feeding marks are now longer (Pl. 59, Figs 4, 5). Final instar (decription from alcohol). Length 10.0–11.0 mm, head and thoracic shield dark brown, colour of body (in alcohol) yellowish white, subdorsal lines dark brown, very variable, sometimes only seen as short streaks on the segments T3 and A1 and A6 and A7, sometimes fully developed on all segments. Foodplants Confirmed larval foodplant: Hibbertia scandens (Willd.) Gilg. (Dilleniaceae) (Pl. 62, Fig. 4).

TAXONOMY

Remarks It should be noted that older specimens in collections are loosing their colour. When this occurs, the shinier collar and abdomen of P. subdolosa clara ssp. n. cannot be recognised easily. However, reference to the localities listed below might be of some help in identifying the two subspecies.

Distribution From south-eastern Queensland to New South Wales.

Male specimens probably conspecific with P. subdolosa The following specimens cannot be clearly assigned to P. subdolosa. Some of them have a slightly broader frons than P. subdolosa, the wing-shape is slightly different and there seems to be variation in the length of the antennal pectinations. These specimens are only listed, not included in the map. Their specific identity is uncertain.

Queensland: 1 么, Brisbane, xi. 1914 (T. P. Lucas) (SAM). New South Wales: 1 么, Allen Beach, 20. x. 1951 (L. H. Mosse-Robinson) (ANIC); 1 么, Barren Grounds Fauna Reserve, 25. iii. 1968 (V. J. Robinson) (ANIC); 1 么, Berowra, 25. x. 1917 (L. H. Mosse-Robinson) (ANIC); 2 么, Como West, 18. iii. 1971, 27. xii. 1976 (L. Willan & V. J. Robinson) (ANIC); 1 么, Cordeaux Dam, 12. iii. 1972 (V. J. Robinson) (ANIC); 3 么, Ebor, 1. xii. 1937, 15. xii. 1939 (AMS); 1 么, Fitzroy Falls, 2. iii. 1973 (V. J. Robinson) (ANIC); 1 么, Geroa, e.p. 9. xi. 1970 (pupa spun on old dead spike of Macrozamia communis L. Johnson [Zamiaceae]) (V. J. Robinson) (ANIC); 1 么, Gosford, ii. 1943 (AMS); 1 么, Mooney Mooney Creek, near Gosford, 16. xi. 1978 (D. K. McAlpine & B. J. Day) (AMS); 3 么, Mt Keira, 19. xi. 1966, 5. iv. 1968, 18. iii. 1971 (V. J. Robinson) (ANIC); 3 么, Narara, 30. xi. 1924, 23. xi. 1950 (L. H. Mosse-Robinson) (ANIC); 2 么, Narrabeen, 10. x. 1914, 8. xi. 1919 (AMS); 1 么, O’Sullivans Gap, 10 km NNE of Buladelah, 15. xi. 1976 (I. F. B. Common & E. D. Edwards) (ANIC); 1 么, Pelican Pt, Bateman’s Bay, xii. 1923 (T. Campbell) (AMS); 1 么, Roseville Chase, 6. xi. 1963 (C. E. Chadwick) (DPIQ); 2 么, Roseville, Sydney, 26. ii. 1926 (L. H. Mosse-Robinson) (ANIC); 2 么, Sydney (AMS); 1 么, Watagan SF, near Wyong, 24. xi. 1978 (B. Hacobian) (ANIC); 1 么, Watagan SF, near Wyong, 24. ii. 1979 (B. Hacobian & V. J. Robinson) (ANIC); 3 么, Wilton, CSIRO Experimental Farm, 25. i. 1971, 5. x. 1972, 18. iii. 1976 (V. J. Robinson) (ANIC); 1 么, Wollongong, 16. xi. 1943 (V. J. Robinson) (ANIC). Australian Capital Territory: 5 么, Jervis Bay, 12. xi. 1918, 26. x. 1919, 18. x. 1920, 7. xi. 1920 (L. H. Mosse-Robinson) (ANIC).

Populations close to, but doubtfully conspecific with, P. subdolosa Population 1. (Slightly smaller, wings narrower, costal margin straighter, frons slightly broader, ca. 1.2× broader than compound eyes in frontal view.) (Pl. 16, Figs 1, 2.) New South Wales: 3 么, 3 乆, Wollongong, 8. i. 1943, iii.–iv. 1943, 29./30. iv. 1966 (ANIC). Population 2. (Slightly smaller, costal margin of forewing strongly arched, frons slightly broader, ca. 1.1–1.2× broader than compound eye in frontal view.) (Pl. 15, Fig. 2.) New South Wales: 3 么, St George’s Basin, 16. iv. 1964 (V. J. Robinson) (ANIC). One single male from a different population but with similar characters: 1 么, Clyde Mountain, 2400 ft, 20. v. 1960 (I. F. B. Common & M. S. Upton) (ANIC). Population 3. (Slightly smaller, wings narrower, frons slightly broader.) Queensland: 2 么, Narangba, 15. ii. 1967 (V. J. Robinson) (ANIC). The material from the above mentioned populations was collected in summer and autumn. The specimens may therefore represent a second or third generation and the differences in size and habitus from typical P. subdolosa could even be the result of different conditions during the development of the larva. Population 4. (Three specimens of different size but narrow frons and large eyes as in P. subdolosa, the proboscis dark brown instead of yellow.) Queensland: 1 么, Dunwich, north Stradbroke I., 22. viii. 1958 (S. S. Sekhon) (UQIC); 1 么, May Orchard, Brisbane, ix. 1910 (SAM). New South Wales: 1 么, Ku-ring-gai Chase Nat. Park, 26. xi. 1971 (V. J. Robinson) (ANIC). Population 5. (Frons slightly broader than compound eyes in frontal view, proboscis dark brown.) New South Wales: 3 么, 1 乆, Canyonleigh, 28. iii. 1968 (V. J. Robinson) (ANIC).

Pollanisus contrastus sp. n. Pl. 17, Figs 1, 2; Figs 171, 172, 216, 217, 260.

Material examined Holotype 么, New South Wales: Narara, 23. xi. 1950 (L. H. Mosse-Robinson) (ANIC). Paratypes Queensland: 1 么, Brisbane (SAM); 1 么, Brisbane, 31. iii. 1907 (ANIC); 1 么, Brisbane, 18. vi. 1898 (R. Illidge) (MV); 1 么, Brisbane, 15. x. 1938 (I. F. B. Common) (ANIC); 1 么, Coolangatta, 10. ix. 1913 (ANIC); 1 么, Stanthorpe, ii. (ANIC). New South Wales: 2 么, 5 乆, Narara, 25. xi. 1924, 12. ii. 1925, 27. xi. 1943, 29. xi. 1943, 19. iii. 1949, 3. xi. 1950 (genitalia slide Z 3199 乆), 23. xi. 1950 (L. H. MosseRobinson) (ANIC); 1 么, 1 乆, Niagara Park, 1948 (W.

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Brandt) (ANIC); 1 么, Sydney, 1861 (SAM); 2 么, Watagan State Forest, near Wyong, 24. ii. 1979 (B. Hacobian & V. J. Robinson) (ANIC); 1 么, Wollongong, 14. ii. 1973 (V. J. Robinson & R. Badans) (genitalia slide Z 3069) (ANIC). One male is smaller and has slightly more rounded wings. It is excluded from the type-series: New South Wales: 1 么, Bendalong, 7. i. 1969 (G. Daniels) (AMS).

Diagnosis Pollanisus contrastus is characterized by its small size, narrow wings and the strong contrast between the very opaque forewing and the hindwing that is translucent medially and has a broad dark margin distally. The frons is broad, the compound eyes are relatively small, and the length of antennal pectination in the male is moderate. Pollanisus edwardsi is larger, has much broader and more rounded wings and there is no strong contrast between the forewing and the hindwing. Pollanisus subdolosa has broader and more rounded wings and a significantly narrower frons and larger compound eyes. Pollanisus angustifrons sp. n. and P. incertus sp. n. have a narrow frons and larger compound eyes. Pollanisus eungellae sp. n. and P. acharon are smaller and more bluish green, P. eumetopus is smaller and darker blue, P. commoni sp. n. is shiny golden and has significantly longer antennal pectinations.

Description Male (Pl. 17, Fig. 1). Forewing length: 6.0–7.5 mm. Head greyish black dorsally, with a narrow row of shiny metallic coppery golden scales medially (sometimes reduced to a few single scales), shiny metallic yellowish golden green frontally and laterally, light grey ventrally; frons broad, approximately 1.4–1.6× broader than compound eyes in frontal view, strongly protruding beyond compound eyes in lateral view, more so dorsally; labial palps light grey, proboscis yellow; ocelli small, white, distance from compound eye twice as large as diameter of ocellus; chaetosemata dark grey, slightly lighter posteriorly, with long anterior extension, clearly separated by a stripe of shiny metallic yellowish green scales from dorsal margin of compound eye; antenna greyish black, segments 1 to 26–27 bipectinate, 26–27 to 36–38 biserrate, pointed distally, pectinations of medium length, ca. 5× longer than shaft in dorsal view at segment 10, only becoming slightly shorter distally, still 4× longer at segment 20, giving the impression that the pectination is for a long distance almost parallel to the shaft, sensory hairs very short. Thorax greyish black dorsally, patagia,

anterior part of tegulae and 20–30% of anterior part of thorax shiny metallic coppery, the sheen not very intensive, shiny metallic bluish green laterally, grey ventrally; legs dark grey, coxa and femur shiny metallic bluish green laterally, especially on hindleg. Abdomen shiny metallic coppery, concolorous with collar, greyish black ventrally. Wings narrow, the very dark and opaque forewing constrasting strongly with the rather translucent hindwing, forewing elongate, anterior margin almost straight, hindwing subtriangular, with rounded apex, posterior margin slightly rounded, almost straight; forewing upperside very densely scaled, dark greyish brownish black, underside dark grey, sometimes with a few shiny metallic bluish green scales at costa proximally; hindwing upperside greyish black, the translucent medial part contrasting with a broad dark distal margin and anal angle, underside with shiny metallic bluish green scales anterior of medial stem and at anal angle. Female (Pl. 17, Fig. 2). Forewing length: 6.5–7.5 mm. Similar to male in size and the narrow wings; compound eyes slightly smaller and frons broader. Abdomen with yellow hairtuft. Antenna strongly biserrate. Male genitalia (Figs 171, 172, 216, 217). Valva pointed apically, slightly convex dorsally, straight ventrally, folded translucent central part triangular, ventral sclerotization of valva broad proximally, lancet-shaped. Aedeagus 3.5–4.0× longer than broad, cornutus straight, slender, as long as aedeagus, pointed apically. Female genitalia (Fig. 260). Ductus bursae short, praebursa large, at the point where the ductus intrabursalis is inserted with a small rounded sclerotization with two small tooth-like structures.

Phenology and bionomics Bivoltine according to the label data. Bionomics unknown.

Remarks There are several specimens that are similar in external appearance to P. contrastus but significantly different in at least one character. They may represent other species and are figured and discussed below as Pollanisus cf. contrastus 1 and 2.

TAXONOMY

Distribution

Material examined

New South Wales and south eastern Queensland.

The included females are based on the character ‘proboscis yellowish brown’. Queensland: 1 乆, Bunya, 10. iv. 1925 (QM); 2 么, 1 乆, Palmwoods, 7. ii. 1930, 5. iii. 1930 (C. [H.] B[orch]) (MV); 1 么, Rainbow beach, Via Tin Can Bay, 21. iii. 1970 (V. Stablum) (UQIC); 4 么, 2 乆, Stradbroke I., 3. xii. 1912 (H. Hacker) (QM). New South Wales: 2 么, 2 乆, Clyde Mountain, 16 mls SE Braidwood, 2400 ft, 15. ii. 1962 (I. F. B. Common) (ANIC); 1 么, Manly, 9. i. 1909 (AMS); 1 乆, Mt Keira, 4. xii. 1970 (V. J. Robinson) (ANIC); 1 么, 1 乆, Narara, 9. iv. 1927, 20. xi. 1943 (L. H. Mosse-Robinson) (ANIC). Australian Capital Territory: 2 乆, Jervis Bay, 25. x. 1919, 1. xii. 1919 (ANIC). There is an additional group of specimens that cannot be placed with certainty, but are also closely related to P. contrastus (all with broad frons and relatively narrow wings). They are excluded from the distribution map. Queensland: 1 么, Brisbane (SAM); 1 么, Coolangatta, 4. v. 1913 (ANIC); 1 么, Eidsvold (ANIC). New South Wales: 1 么, Ebor, 7.-13. ii. 1940 (G. M. Goldfinch) (AMS); 2 么, Geroa, 1. xi. 1970 (V. J. Robinson) (ANIC); 2 么, Murwillumbah, ix. 1969 (G. May) (ANIC); 2 么, Otford, 4. ii. 1963,. 4. ii. 1964 (V. J. Robinson) (ANIC); 1 么, Sydney, 1861 (SAM).

Pollanisus cf. contrastus 1 Pl. 18, Fig. 1.

Material examined The four included females are externally similar to the males and are from the same locality. They are possibly conspecific. Queensland: 1 么, Brisbane, 19. i. 1913 (ANIC); 1 么, Dunwich, 29. iii. 1959 (P. R. Webb) (UQIC); 1 么, 2 乆, M[arooch]ydore, 8. iv. 1920 (J. A. Beck) (UQIC). New South Wales: 1 么, 2 乆, Tweed Heads, 29. xi. 1922, 8. i. 1923 (QM).

Remarks Habitus very similar to that of P. contrastus but male antenna with longer pectinations, especially proximally, at segment 8–10 ca. 6–7× longer than breadth of shaft in dorsal view, tapering distad of segment 11. The hindwing seems to be slightly less translucent. The patagia are less shiny or entirely lack shiny scales. No other differences could be recognised. The length of the antennal pectinations may be slightly variable but variation to this extent is not known in zygaenids elsewhere. It is therefore almost certain that the specimens with longer antennal pectinations belong to a different species. Nevertheless, a description of a new species would have to be based on four males with slightly variable length of antennal pectinations, a character that is considered to be insufficient to warrant distinct specific status.

Pollanisus cf. contrastus 2 Pl. 18, Fig. 2.

Remarks Externally very similar to P. contrastus but with yellowish brown proboscis, the forewing slightly less opaque and the hindwings less translucent. Although there is considerable material with this character combination available, a description of a new species seems risky as it would have to be based on rather old and worn specimens in which all characters cannot be seen clearly. Moreover, the character ‘proboscis yellowish brown’, in contrast to the always bright yellow proboscis of similar species is unreliable, as the colour depends also on whether the proboscis is completely rolled, half rolled or extended. The completely rolled proboscis may even suggest a dark brown colour whereas an unrolled proboscis appears to be almost yellow.

Possible P. edwardsi, P. subdolosa or P. contrastus females These females cannot be assigned to species but belong to P. edwardsi, P. subdolosa, P. contrastus, or possibly a closely allied but undescribed species:

Queensland: 2 乆, Brisbane, xi. 1914 (T. P. Lucas) (SAM); 1 乆, Buderim Mountains, 7. iv. 1912 (H. Hacker) (QM); 1 乆, C[unning]ham’s Gap, 19.-21. xi. 1949 (ANIC); 1 乆, Dulong, 7. iv. 1907 (ANIC); 2 乆, Moreton Beach, Peel I., 29. ix. 1906 (ANIC); 1 乆, Mt Tennyson Woods, near Mt Glorious, 7. i. 1977 (M. A. Schneider)

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(AMS); 1 乆, Sandgate, 6. ix. 1902 (ANIC); 1 乆, Springbrook, 2. xii. 1932 (ANIC); 1 乆, Stanthorpe, ii. (ANIC); 1 乆, Toowoomba, 6. xii. 1937 (I. F. B. Common) (ANIC). New South Wales: 1 乆, Arrawarra, 20. x. 1962 (C. W. Frazier) (ANIC); 1 乆, Bendalong, 9. i. 1969 (G. Daniels) (AMS); 1 乆, Boambee, 31. x. 1939 (C. R. W.) (DPIQ); 1 乆, Brunswick Heads, 27. xii. 1926 (W. B. Barnard) (QM); 1 乆, Bungwahl, 5 mls N, 24. iii. 1965 (I. F. B. Common & M. S. Upton) (ANIC); 1 乆, Congo, 8 km ESE Moruya, 19. iii. 1983 (M. S. Upton) (ANIC); 2 乆, Fassifern, 31. x. 1950 (L. H. Mosse-Robinson) (ANIC); 1 乆, Fingal Bay, 24. x. 1962 (V. J. Robinson & K. English) (ANIC); 1 乆, French’s Forest, 31. x. 1928 (AMS); 1 乆, Geroa, 1. xi. 1970 (V. J. Robinson) (ANIC); 1 乆, Geroa, e.l. 29. xi. 1970 (larva on Hibbertia scandens) (V. J. Robinson) (ANIC); 2 乆, Gosford, 1913 (DPIQ); 1 乆, Kurnell, e.l. 30. xi. 1970 (larva on Hibbertia scandens) (V. J. Robinson) (ANIC); 1 乆, Lismore Area, 10. 1976 (G. R. Brown) (DPIQ); 1 乆, Malua Bay, 20 m, 2. xii. 1989 (G. M. Tarmann) (ANIC); 1 乆, Mt Keira, 19. xi. 1966 (V. J. Robinson) (ANIC); 1 乆, Nambucca, 24. ii. 1936 (AMS); 1 乆, Narara, 24. xi. 1924 (L. H. Mosse-Robinson) (ANIC); 1 乆, Nerriga, 4 km SSW, 6. ii. 1975 (I. F. B. Common, E. D. Edwards & M. Story) (ANIC); 1 乆, Otford, 2. v. 1903 (MV); 1 乆, Picton lakes, 18. iii. 1972 (V. J. Robinson) (ANIC); 1 乆, Roseville, Sydney, 16. x. 1921 (L. H. Mosse-Robinson) (ANIC); 5 乆, Rous, Richmond R., ix. 1933, iii. 1934, x. 1935 (V. J. Robinson) (ANIC); 1 乆, [Royal] Nat. Park, 12. iv. 1925 (MV); 1 乆, Royal Nat. Park, e.l. 31. x. 1976 (larva on Hibbertia scandens) (V. J. Robinson) (ANIC); 4 乆, Stanwell Park, iii. 1912, ii. 1913 (AMS); 1 乆, Stanwell Park, 16. iii. 1952 (L. C. Haines) (AMS); 1 乆, Woy Woy, 19. iv. 1907 (W. W. Frogatt) (DPIQ). Australian Capital Territory: 1 乆, Jervis Bay, 3. xi. 1919 (ANIC); 1 乆, Jervis Bay, 14. x. 1966 (I. F. B. Common & M. S. Upton) (ANIC). Victoria: 1 乆, Bayswater, 12. iii. 1922 (MV); 1 乆, Cann R., xi. 1928 (J. Clark) (MV).

Pollanisus eungellae sp. n. Pl. 19, Figs 1, 2; Figs 173, 174, 218, 261, 262.

Material examined Holotype 么, Queensland: Eungella, 9. x. 1928 (ANIC). Genitalia slide Z 3194. Paratype Queensland: 1 乆, Eungella, 9. x. 1928 (ANIC). Genitalia slide Z 3052.

Diagnosis Pollanisus eungellae shares the broad frons and relatively small compound eye with P. edwardsi, P.

contrastus, P. eumetopus, P. commoni sp. n. and P. acharon. Pollanisus edwardsi differs from P. eungellae in its much larger size and broader wings, and the shiny metallic scales on the collar and dorsal part of the abdomen are coppery. Pollanisus contrastus has darker forewings and more translucent hindwings, the colour of the shiny metallic scales on the collar and dorsal part of the abdomen is coppery. Pollanisus eumetopus is darker, the shiny scales on the collar and abdomen are darker bluish, the male has much smaller compound eyes and a broader frons, the shiny collar of the female is narrower than in P. eungellae. Pollanisus commoni sp. n. has shiny metallic golden scales on collar and abdomen. The female of P. acharon is externally very similar to the female paratype of P. eungellae. Conspecificity is most unlikely (see comment pp. 95, 96). Pollanisus subdolosa, P. angustifrons sp. n. and P. incertus sp. n. have a significantly narrower frons and larger eyes in the male; the shiny scales of P. angustifrons sp. n. are more yellowish green, those of the other two species coppery golden.

Description Male (Pl. 19, Fig. 1). Forewing length: 6.0 mm. Head grey-black dorsally, lighter grey ventrally, partly covered with shiny scales anteriorly and laterally (including eyelashes); frons ca. 1.3× broader than compound eyes in frontal view, significantly protruding beyond compound eyes in lateral view, more so dorsally, light grey ventrally; labial palps short, slender, porrect distally, light grey; proboscis yellow; compound eye of medium size, black, sprinkled with white spots (this character may not be constant as in other species); ocelli large, white, distance from compound eye 2× larger than diameter of ocellus; chaetosemata dark grey, slightly lighter caudally, strongly extended anteriorly; the dorsolateral parts of frons, a small space between chaetosema, dorsal margin of compound eye and the area posterior of compound eye covered with shiny metallic bluish green scales. Antenna greyish black, segments 1–28 bipectinate, 28–38 biserrate, apex pointed, the pectinations of medium length, approximately 4–5× longer than breadth of shaft in dorsal view (at segment 10), tapering distally, sensory hairs very short. Thorax greyish black dorsally, patagia and tegulae covered with yellowish green, shiny metallic scales tinged with light blue; lateral parts of thorax beyond wings shiny metallic blue; legs dark grey, femur and tibia with shiny blue scales laterally, especially on hindleg. Abdomen with segments 1–3 greyish black, with a few shiny scales

TAXONOMY

mediodorsally, segments 4–8 shiny metallic bluish green tinged with yellowish green dorsally, unicolorous greyish black ventrally. Wings narrow, forewing elongate, triangular, apex rounded, hindwing extending 70% of length of forewing, subtriangular, apex rounded, posterior margin almost straight, slightly concave. Forewing upperside blackish brown, underside dark greybrown; hindwing upperside and underside dark greybrown, slightly translucent medially, underside with a broad stripe of shiny metallic greenish bluish scales below costa and at anal angle. The scales on forewing upperside uniform, short, double-lobed apically. Female (Pl. 19, Fig. 2). Forewing length: 6.5 mm. Similar to male, the broad collar and shiny parts of abdomen slightly more bluish, with a bright lemonyellow abdominal hairtuft. Frons broader than in male, compound eyes smaller. Antenna biserrate. Male genitalia (Figs 173, 174, 218). Valva slender, bluntly pointed apically, slightly curved dorsally, almost straight ventrally, folded central part triangular, ventral sclerotization of valva broad proximally, lancet-shaped, sacculus well separated. Aedeagus approximately 4× longer than broad, cornutus relatively broad, ca. 8× longer than broad, slightly longer than aedeagus. Female genitalia (Figs 261, 262). Praebursa large, central sclerotization with two long teeth.


Distribution Only known from the type-locality Eungella, an isolated rainforest area with a rather distinct fauna and flora in north-eastern Queensland.

Pollanisus acharon (Fabricius) Zygaena acharon Fabricius, 1775, Systema Entomologicae: 556. Lectotype 乆 (here designated), [AUSTRALIA] [‘Nova Hollandia’] (type without label data) (BMNH, Banks Coll.) [examined]. Adscita acharon (Fabricius); Kirby 1892: 85. Pollanisus acharon (Fabricius); Jordan 1907: 10 (? as syn. of Pollanisus amethystinus (Meyrick)); Hering 1922: 13; Bryk 1936: 117; Tarmann 1996: 143. Pollanisus acheron [sic] Bryk, 1936, in Strand, E., Lepid. Cat. 71: 117 (misspelling). The true identity of this taxon will never be completely clear. The female type is deposited in the Banks collection in the BMNH in London (Pl. 58, Figs 1–3). It is an unprepared female in fairly poor condition. There is little doubt that it is the genuine type. The type-specimen has no locality label but a label with the BMNH registration number 63.48. It was presented to the BMNH by the Linnean Society in 1863 together with other types of Fabricius. In the right corner of p. 835 of Entomological Accessions Annulosa, Vol. III (1850–1863) ‘New Holland’ is cited as the locality for this specimen. The type has been studied carefully. However, even with the genitalia it is impossible to ascertain its true taxonomic position. There is no doubt that it represents a species of the rainforest group of Pollanisus (small species with narrow wings) and is closely related to P. eumetopus and several other species described in this book. From the female of P. eumetopus it differs in the shape of the wing, the colour of the body and collar and the breadth of the collar. Within the rich material of P. eumetopus, P. subdolosa and similar species studied for this revision, there is only one female of a so far undescribed species from the isolated rainforest area of Eungella in eastern Queensland showing almost all characters that could be recognised in the type of P. acharon. Nevertheless, it can never be proven that they are really conspecific. Most of the early material from Australia was collected during the expedition of Captain Cook. Cook’s ship Endeavour was repaired on the coast of Queensland near the present town of Cooktown in June and July 1770 and many of the species described subsequently had been taken near this locality. Eungella was never visited by the scientists on Cook’s expedition as it is much further south and more inland. Furthermore, its fauna is notably isolated with many endemic species and differs from the coastal fauna around Cooktown, and the two existing specimens from Eungella (1 么,

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1 乆) were collected in October. The possibility of conspecificity with the type of P. acharon is therefore very unlikely. The tropical species of Pollanisus from the rainforests in Queensland represent a group of taxa with only small but mainly very constant external differences. This can be seen in the few species reared from larvae found on different foodplants, e.g. P. eumetopus, P. commoni sp. n., P. incertus sp. n. As the genitalia have no significant characters in these closely related species, and in the genus Pollanisus in general, it is probably not possible to recognise all of the existing species at present. Many of them seem to be restricted to very small isolated areas. It is therefore possible that P. acharon is in fact a species originating from and perhaps even restricted to the coastal areas around Cooktown and is not currently represented in collections. There is no Pollanisus material available from that locality. The rediscovery of P. acharon around Cooktown may therefore be just a matter of more careful collecting in that area and there is no justification for referring the Eungella specimens to P. acharon.

Diagnosis Pollanisus edwardsi differs from P. acharon by its larger size, broader wings and the shiny metallic coppery collar and abdomen. Pollanisus contrastus has darker, more opaque forewings and more translucent hindwings, the colour of the shiny metallic scales on the collar and abdomen is coppery. Pollanisus eumetopus is darker, the shiny scales on the collar and abdomen are darker bluish and the collar is narrower than in P. acharon. Pollanisus commoni sp. n. and P. incertus sp. n. have shiny metallic golden scales on the collar and abdomen, the latter species has a narrower frons and larger eyes. Pollanisus subdolosa and P. angustifrons sp. n. have narrower frons and larger eyes, the shiny metallic scales on the collar and dorsal part of the abdomen are coppery in the former and yellowish green in the latter. Based on one single female of each, no clear differences can be recognised to separate P. acharon from P. eungellae. The frons of P. acharon is slightly broader.

Redescription (of the female type) Female (Pl. 58, Figs 1–3). Forewing length: 7.0 mm. Head greyish black dorsally, paler ventrally, covered with shiny metallic bluish scales frontally; frons very broad, ca. 2.5× broader than compound eyes in

frontal view, slightly protruding beyond compound eye in lateral view; labial palps short, slender, porrect distally, light grey; proboscis brownish; compound eye small, black; ocelli small, white, distance from compound eye 2–3× longer than diameter of ocellus; chaetosemata dark grey, strongly extended anteriorly. Antennae missing. Thorax greyish black dorsally, patagia and tegulae covered with bluish green shiny metallic scales tinged with yellow; lateral parts of thorax below wings shiny metallic blue; legs dark grey, femur and tibia with shiny blue scales laterally. Abdominal segments 1–3 greyish black, with a few shiny scales mediodorsally, segments 4–8 shiny metallic bluish green tinged with yellowish green dorsally, unicolorous greyish black ventrally, with a bright yellow abdominal hairtuft. Wings narrow, forewing elongate triangular, apex rounded, hindwing 70% length of forewing, subtriangular, apex rounded, posterior margin almost straight, slightly concave. Forewing upperside and underside dark grey-brown; hindwing upperside and underside greyish black, slightly translucent medially, underside with a broad band of shiny metallic blue scales below costa and anterior part of cell proximally, and at anal angle. Female genitalia (Fig 263). Praebursa large, ovoid, central sclerotization with two teeth.


Distribution Only the type-specimen is known. Its origin is unknown but it was most likely collected near Cooktown in north-eastern Queensland during the Cook expedition.

TAXONOMY

Pollanisus eumetopus Turner Pl. 20, Figs 1, 2; Figs 175, 219, 220, 264. Pollanisus eumetopus Turner, 1926b, Proc. Linn. Soc. NSW 51: 443. Holotype 么, AUSTRALIA: Queensland, Kuranda, 16. vi. 1911 (A. J. Turner) (ANIC) [examined]. Genitalia slide Z 3053 (ANIC). Pollanisus eumetopus Turner; Bryk 1936: 118; Common 1990: 297 (bionomics); Tarmann 1996: 143.

Material examined Queensland: 2 么, 2 乆, Kuranda, v. 1904 (F. P. Dodd) (MV); 1 乆, Kuranda, 21./23. xi. 1925 (AMS); 4 么, 2 乆, Redlynch, 4 km S, e.l. 3.–16. x. 1985 (larvae on Pipturus argenteus) (I. F. B. Common) (ANIC).

Wings narrow, forewing elongate, triangular, hindwing subtriangular, with rounded apex and slightly concave posterior margin. Forewing upperside greyish black with purple tinge, underside greyish black; hindwing greyish black with translucent medial part, underside with two bands of shiny metallic blue scales below costa and at anal angle, concolorous with lateral parts of thorax. Female (Pl. 20, Fig. 2). Forewing length: 5.5–7.5 mm. Similar to male, with yellow abdominal hairtuft with orange tinge. Antenna shortly biserrate. Head not much broader and compound eyes almost as large as in male.

Differs from all similar species by its broader frons and small compound eyes of almost the same size in both sexes, and its distinctive dark blue colour.

Male genitalia (Figs 175, 219, 220). Valva slender, bluntly pointed apically, slightly curved dorsally, almost straight ventrally, folded central part triangular, ventral sclerotization of valva broad proximally, lancet-shaped. Aedeagus approximately 3× longer than broad, cornutus relatively broad, ca. 8× longer than broad and approximately 80% length of aedeagus.

Redescription

Female genitalia (Figs 264). Praebursa with a central sclerotization bearing two short, triangular teeth.

Diagnosis

Male (Pl. 20, Fig. 1). Forewing length: 6.0–7.0 mm. Head greyish black with a purplish tinge dorsally, shiny metallic greenish blue frontally and laterally, light grey ventrally; frons broad, almost twice as broad in frontal view as the relatively small black compound eyes, strongly protruding beyond compound eyes in lateral view, more so dorsally; labial palps small and slender, porrect distally; proboscis bright yellow; ocelli small, white, distance from compound eye almost twice as large as diameter of ocellus; chaetosemata greyish brown, strongly extended anteriorly. Antenna with slightly shiny purplish black shaft and black pectinations, 1–25 bipectinate, 25–35 biserrate, apex bluntly pointed; the pectinations of medium length, approximately 5–6× longer than breadth of shaft in dorsal view at segment 10, becoming only slightly shorter until segment 17, approximately 2.5× as long as breadth of shaft at segment 20, sensory hairs very short. Thorax greyish black with a purplish tinge dorsally, patagia and anterior margin of tegulae shiny metallic greenish blue; ventral part of thorax shiny metallic blue laterally, dark grey ventrally; legs dark grey, coxa and part of femur shiny metallic blue. Abdomen greyish black, with shiny metallic dark steel-blue scales with a slightly variable weak green tinge on segments 2–8 dorsally.

Phenology and bionomics A series of specimens of this species was reared by I. F. B. Common from larvae obtained from Pipturus argenteus (G. Forster) Wedd. (Urticaceae) (Common, 1990: 297). According to the few available label data presumably multivoltine.

Distribution Coastal parts of north-eastern Queensland.

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Pollanisus commoni sp. n. Pl. 21, Figs 1, 2; Figs 176, 221–224, 265, 266. Pollanisus subdolosa (Walker); sensu Common 1990: 297 (partim).

Material examined Holotype 么, Queensland: Clifton Beach, 16.46´S 145´45 E, e.l. 11. x. 1985 (ex larva on Dillenia alata) (I. F. B. Common) (ANIC). Genitalia slide Z 2860 (ANIC). Paratypes Queensland: 3 么, 8 乆, Clifton Beach, 16.46´S 145.45É, e.l. 5.–22. x. 1985 (ex larva on Dillenia alata) (I. F. B. Common) (ANIC). Genitalia slides Z 2643 么, Z 2861 乆, Z 3049 么, Z 3050 乆 (ANIC). Not included in the type-series but considered to be conspecific: Queensland: 1 么, 1 乆, Cairns, e.l. i. 1989 (ex larva on Dillenia alata) (J. Waite) (DPIQ); 1 么, 1 乆, Port Douglas, 13. v. 1974 (N. R. Badans & V. J. Robinson) (ANIC); 1 么, Port Douglas, 13. v. 1974 (N. R. Badans) (ANIC).

Diagnosis Differs from P. eumetopus, P. angustifrons sp. n., P. eungellae and P. acharon in its broad, shiny metallic golden collar and abdomen. Pollanisus incertus sp. n. is very similar in coloration but the male has a significantly narrower frons and larger eyes. Pollanisus edwardsi is larger, has broader wings and the shiny scales are darker and more coppery. Pollanisus subdolosa has broader wings and a significantly narrower frons and larger eyes. Pollanisus contrastus has more triangular forewings and the shiny metallic scales on the collar and abdomen are coppery.

Description Male (Pl. 21, Fig. 1). Forewing length: 7.0–7.5 mm. Head greyish black dorsally, sometimes with a narrow line of single shiny metallic golden scales medially, frons shiny metallic yellowish green, lateral areas shiny metallic greenish blue, eyelashes light grey; frons as broad or slightly broader than compound eyes in frontal view, strongly protruding beyond compound eyes in lateral view, more so dorsally; labial palps dark grey, short, porrect distally; proboscis yellow; ocelli large, white, distance from compound eye as large as diameter of ocellus; chaetosemata brownish grey, with long anterior extension, a single row of shiny metallic greenish blue scales between chaetosema and dorsal

margin of compound eye; antenna greyish black, segments 1–31 bipectinate, 32–44 biserrate, pointed apically, the pectinations very long, 6–7× longer than shaft in dorsal view at segment 10, only becoming slightly shorter towards the more distal segments, still 4–5× longer at segment 20, 3× longer at segment 25, sensory hairs very short. Thorax greyish black dorsally, patagia, anterior half of tegulae and anterior margin of dorsal mesothorax covered with shiny metallic golden scales with a coppery tinge, shiny metallic blue laterally, dark grey ventrally; legs dark grey, midleg and hindleg shiny metallic blue laterally. Abdominal segments 1 and 2 greyish black, 3–8 increasingly shiny metallic golden dorsally with a coppery tinge, unicolorous greyish black ventrally. Wings narrow, forewing elongate triangular, hindwing broad, subtriangular, with rounded apex, posterior margin straight or slightly concave; forewing upperside greyish black, underside dark brownish grey, with a narrow band of shiny metallic blue scales below costa proximally; hindwing upperside greyish black, strongly translucent medially, underside with shiny metallic blue scales anterior of medial stem and at anal angle. Female (Pl. 21, Fig. 2). Forewing length: 6.5–7.5. Similar to male, with bright yellow abdominal hairtuft. Antenna shortly biserrate. Dorsal row of metallic golden scales on head usually more pronounced. Compound eyes slightly smaller than in male, frons slightly broader. Male genitalia (Figs 176, 221–224). Valva short, triangular, sharply pointed apically, slightly convex dorsally and ventrally, folded central part broad triangular, ventral sclerotization of valva broad proximally, lancet-shaped. Adeagus 3.0–3.5× longer than broad, cornutus straight, relatively broad, ca. 9× longer than broad, rounded apically. Female genitalia (Figs 265, 266). Large praebursa with a comb-like central sclerotization with a row of teeth (three or more).

Phenology and bionomics The type-series was reared from larvae found on Dillenia alata (DC.) Martelli (Dilleniaceae) by I. F. B. Common (1990: 297) who published a short note erroneously referring the specimens to P. subdolosa, a species that is externally similar. Two specimens (one male, one female) have also been reared by J. Waite from larvae found on the same foodplant at Cairns.

TAXONOMY

Distribution North-eastern coast of Queensland.

Etymology Named in honour of I. F. B.Common (Toowoomba), the doyen of Australia’s lepidopterists, who discovered this new species by rearing it from larvae.

Pollanisus incertus sp. n.

nal pectinations and probably also the colour pattern (caused by combinations of the various metallic shiny scales) seem to represent quite constant characters in the eumetopus-group of the genus Pollanisus. Females of P. incertus are almost identical to P. commoni. However, the differences in the males exclude conspecificity and one has to face the fact that the females of both species are very difficult to recognise. Other similar species are P. eumetopus, P. angustifrons sp. n., P. eungellae, P. acharon, P. contrastus, P. edwardsi and P. subdolosa. P. incertus differs from P. eumetopus, P. eungellae and P. acharon in its copperygolden collar and abdomen and the significantly larger eyes and narrower frons in the male. Pollanisus angustifrons sp. n. has similar large eyes and narrow frons in the male but differs in the colour of the collar and abdomen, which are shiny metallic yellowish green with a bluish tinge. Pollanisus edwardsi is larger, has broader wings, a broader frons and smaller compound eyes. Pollanisus contrastus has a significantly broader frons and more triangular forewings. Pollanisus subdolosa has a similar narrow frons but broader wings.

Pl. 22, Figs 1, 2; Figs 225, 267, 268.

Description Pollanisus subdolosa (Walker); sensu Common 1990: 297 (partim).

Material examined Holotype 么, Queensland: 2.5 km N of Kuranda, e.l. 20. x. 1985 (larva on Tetracera nordtiana) (I. F. B. Common) (ANIC). Genitalia slide Z 3195 (ANIC). Paratype Queensland: 乆, 2.5 km N of Kuranda, e.l. 8. xi. 1985 (larva on Tetracera nordtiana) (I. F. B. Common) (ANIC). Genitalia slide Z 3196 (ANIC).

Diagnosis This species is externally very similar to P. commoni and together with it has been referred to P. subdolosa by Common (1990: 297). The significantly larger eyes and narrower frons in the male and the slightly shorter pectinations of the antenna are the only characters separating these two species, except for the different foodplants upon which the larvae have been found. The female of P. incertus differs only slightly from that of P. commoni by the more rectangular form of the frons in frontal view. The form of the frons, the size of the compound eyes, the length of the anten-

Male (Pl. 22, Fig. 1). Forewing length: 8.0 mm. Head greyish black dorsally, frons shiny metallic yellowish green, lateral head-parts shiny metallic yellowish green with a bluish tinge; breadth of frons significantly smaller than that of the very large compound eyes in frontal view (reaching about 80% of their breadth), protruding beyond compound eyes in lateral view, more so dorsally; labial palps dark grey, last segment whitish grey, short, porrect distally; proboscis yellow; ocelli large, white, distance from compound eye slightly less than diameter of ocellus; chaetosemata brownish grey, with long anterior extension, almost touching compound eye ventrally and covering a short part of the shiny metallic scales surrounding the compound eye at that point; antenna greyish black, segments 1–29 bipectinate, 29–40 biserrate, pointed apically, the pectinations long, 5–6× longer than shaft in dorsal view at segment 10, becoming only slightly shorter towards more distal segments, still 4–5× longer at segment 20, 3–4× longer at segment 25; sensory hairs very short. Thorax greyish black dorsally, patagia, anterior half of tegulae and anterior margin of dorsal mesothorax covered with shiny metallic coppery-golden scales, shiny metallic bluish green laterally, dark grey ventrally; legs dark grey, midleg and hindleg shiny metallic bluish green laterally.

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Abdomen greyish black, shiny metallic copperygolden dorsally, the shiny scales covering only a small part of segment 1 dorsomedially, a broader part of segment 2 and almost the whole dorsum of segments 3–8, unicolorous greyish black ventrally. Wings narrow, forewing elongate triangular, hindwing broad subtriangular, with rounded apex, anterior margin straight to slightly concave; forewing upperside greyish black, underside dark grey, with a few shiny metallic bluish green scales at costa proximally; hindwing upperside greyish black, translucent centrally, underside with shiny metallic bluish green scales anteriad of medial stem and at anal angle.

Remarks The female genitalia of the paratype are very distinct and differ from the other species. However, these characters are very variable and more material would be required to confirm constant genitalic differences that can be used for identification.

Distribution Coastal parts of north-eastern Queensland. Only known from the type-locality.

Female (Pl. 22, Fig. 2). Forewing length: 8.5 mm. Similar to male, the metallic coppery-golden scales extending throughout the whole tegulae and the anterior quarter of the mesothorax dorsally; abdomen also shinier than in male, with a bright orange abdominal hairtuft. Antenna biserrate, the shaft slightly broader at about two-thirds of its length, pointed distally. Compound eyes smaller than in male, frons broader, chaetosema not touching dorsal margin of compound eye. Male genitalia (Fig. 225). Valva triangular, bluntly pointed apically, slightly curved dorsally, almost straight ventrally, folded central part triangular, ventral sclerotization of valva broad proximally, lancet-shaped, sacculus only slightly pronounced. Aedeagus approximately 5× longer than broad, cornutus slender, straight distally, curved proximally, c. 12–13× longer than broad and slightly longer than aedeagus. Female genitalia (Figs 267, 268). Ostium broad, ductus bursae short and slender, praebursa large, with a posterior part that bears a weakly sclerotized band-like structure and with a completely translucent, spherical anterior part; the sclerotized structure that ruptures the wall of the spermatophore is a large plate with a row of four short teeth on one side.

Phenology and bionomics Both specimens were reared by I. F. B. Common from larvae found on Tetracera nordtiana F. Muell. (Dilleniaceae). Common (1990: 297) published a short note erroneously referring these specimens to externally similar P. subdolosa.

Pollanisus angustifrons sp. n. Pl. 23, Fig. 1; Figs 226, 227.

Material examined Holotype 么, Queensland: Tully, 25 mls W, 8. iii. 1964 (I. F. B. Common & M. S. Upton) (ANIC). Genitalia slide Z 3051 (ANIC). Paratypes Queensland: 2 么, Babinda, 4 mls N, 10. iii. 1964 (I. F. B. Common & M. S. Upton) (ANIC); 1 么, Mount Lewis, 8 mls NW Mount Molloy, 2700 ft, 15. iii. 1964 (I. F. B. Common & M. S. Upton) (ANIC); 1 么, Paluma, 2 mls W, 2800 ft, 14. iv. 1969 (I. F. B. Common & M. S. Upton), genitalia slide Z 3044 (ANIC); 2 么, Tully, 25 mls W, 8. iii. 1964 (I. F. B. Common & M. S. Upton) (ANIC). There are several more specimens known with the characters of P. angustifrons but with white instead of black ocelli. As it is not yet known whether the pigmentation of the ocellus is a constant character they are not included in the type-material. Queensland: 1 么, Borumba Dam, 19. ii. 1967 (V. J. Robinson) (ANIC); 2 么, Herberton, Baldy SF, 1100 m, 20. iv. 2002 (at mv light) (A. Zwick) (ANIC); 1 么, Jubilee Rd, 4? mls from Innisfail, 4. xi. 1966 (rainforest at light) (E. Britton) (ANIC); 2 么, Mulgrave R., 2. ii. 1980 (V. J. Robinson & N. C. Coleman) (ANIC); 2 么, Mulgrave R., near Gordonvale, 21. iii. 1984 (B. Hacobian) (ANIC); 1 么,

TAXONOMY

Paluma Dam, 12.-13. v. 1980 (I. D. Naumann & J. C. Cardale) (ANIC); 2 么, Tully R. Valley, 12. ii. 1980 (B. Hacobian & N. Goodman) (ANIC). Four males are very similar but significantly larger (forewing length 8.5–9.0 mm) and have slightly larger eyes but also black ocelli. They are here referred to P. angustifrons but may even represent another so far undescribed species (Pl. 23, Fig 2). Queensland: 2 么, Waterview Creek, 22 mls SW Ingham, 2000 ft, 18. iv. 1961 (R. Straatman) (ANIC); 2 么, Windsor Tableland, 8. v. 1984 (A. N. Gillison) (ANIC).

Diagnosis Externally very similar to P. eungellae and P. acharon, with the colour of the shiny scales slightly more yellowish, but with a significantly narrower frons and larger eyes in the male. This, of course, cannot be proven in P. acharon (only female type known). Pollanisus angustifrons is also similar to P. eumetopus, P. commoni, P. incertus, P. contrastus, P. subdolosa and P. edwardsi. Except for P. incertus and P. subdolosa they can be easily separated from P. angustifrons by their smaller eyes and significantly broader frons. Pollanisus incertus has a similar narrow (even slightly narrower) frons and similar large eyes in the male but a shiny metallic coppery golden collar and abdomen and slightly longer pectinations on the antenna, especially distally. Pollanisus subdolosa has broader wings and the shiny scales on the collar and abdomen are coppery.

black dorsally, patagia and anterior half of tegulae covered with shiny metallic yellowish green scales with a bluish tinge; lateral parts of thorax shiny metallic blue; legs dark grey, femur and tibia shiny metallic blue laterally especially on mid- and hindleg. Abdomen greyish black, segments 2–8 shiny metallic yellowish green dorsally. Wings narrow, forewing elongate triangular, apex rounded, hindwing reaching 75% length of forewing, subtriangular, posterior margin almost straight, sometimes slightly concave. Forewing upperside and underside blackish brown, underside with a few shiny metallic blue scales at costa proximally; hindwing upper and underside greyish black, slightly translucent medially, underside with shiny metallic blue scales anteriad of medial stem and at anal angle. Female. Unknown. Male genitalia (Figs 226, 227). Valva short, triangular, sharply pointed apically, convex dorsally, almost straight ventrally, translucent central part triangular, ventral sclerotization broad proximally, lancetshaped. Aedeagus 3× longer than broad, slightly upcurved, cornutus long and broad, as long as aedeagus, dorsoventrally compressed, therefore apex rounded or pointed in slide preparations, depending on the view.

Phenology and bionomics Description Male (Pl. 23, Fig. 1). Forewing length: 7.0–7.5 mm. Head greyish black dorsally, lighter grey ventrally, with shiny metallic bluish green scales dorso-laterally and a few anteriorly especially dorsoanteriorly; frons slightly narrower than breadth of compound eye in frontal view, reaching approximately 90% of its breadth, significantly protruding beyond compound eyes in lateral view, more so dorsally; labial palps short, upcurved, grey; proboscis yellow; compound eyes black, sprinkled with white spots (this character may not be diagnostic; it can be seen in other species); ocelli large, black, distance from compound eye only half diameter of ocellus; chaetosemata dark grey, very narrow, anterior extension touching compound eye ventrally. Antenna greyish black, segments 1 to 29–30 bipectinate, 29–30 to 41–42 biserrate, bluntly pointed apically, the pectinations of medium length, approximately 4–5× longer than shaft in dorsal view (at segment 10), progressively becoming shorter distally, sensory hairs very short. Thorax greyish

All specimens comprising the type material were collected in March and April, but it is most likely that this species is at least bivoltine. One specimen that is not included in the type-series was taken at light in rainforest. Bionomics unknown.

Distribution North-eastern Queensland.

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Pollanisus sp. 2 Pl. 24, Figs 1, 2.

Material examined New South Wales: 2 么, 1 乆, Murramarang Nat. Park, e.l. 1993 (larva on Kennedia rubicunda) (A. Gibbs) (ANIC).

Remarks Probably undescribed species belonging to the Pollanisus subdolosa-eumetopus-group, with insufficient material for a description. A small species (forewing length 6.5 mm in male and female) with rounded wings and very opaque black hindwings. Head shiny yellowish green frontally and dorsolaterally, frons broad (ca. 1.5× broader than compound eyes in frontal view in male, 2.0× in female), proboscis yellowish brown; antenna in male bipectinate from 1–25, biserrate from 25–36, the pectinations relatively short (only about 4× longer than shaft in dorsal view at segment 10), biserrate in female. Thorax black dorsally, patagia and anterior part of tegulae shiny metallic coppery golden, shiny metallic bluish green laterally, dark grey ventrally; legs dark grey, shiny laterally. Abdomen shiny metallic coppery dorsally, blackish ventrally, with a bright yellow abdominal hairtuft in female. Wings very dark and opaque, with shiny metallic bluish green scales on underside at proximal part of costa in forewing and anterior of medial stem and at anal angle in hindwing. These three specimens were reared from larvae found on Kennedia rubicunda Vent. (Fabaceae). One cocoon has been preserved. It is flattened-ovoid, silky white and spun on the surface of the leaf. This is the only Pollanisus species known that feeds on Fabaceae. Unfortunately both male specimens are in very poor condition, the scales partly lost and the antennae or parts of the antennae are missing. The colour painting of the male is a reconstruction compiled from the available characters of both males.

Remarks Probably undescribed species belonging to the Pollanisus subdolosa-eumetopus-group, with insufficient material for a description. A very dark and densely scaled small species (forewing length 8.0–8.5 mm in male, 8.0 mm in female) with a very broad shiny metallic blue frons and small compound eyes. Externally very similar to P. contrastus but hindwings more opaque. Antenna long, 1–31 bipectinate, 32–43 biserrate in male, the pectinations short (only 3–4× longer than shaft in dorsal view), biserrate in female. These two island populations may belong to an undescribed species, could possibly be conspecific with Pollanisus sp. 6 (see below) or represent a distinct subspecies of P. contrastus with more opaque hindwings. The significantly longer antennae and the shiny blue head distinguish it clearly from P. contrastus.

Pollanisus sp. 4 Pl. 26, Figs 1, 2.

Material examined Queensland: 1 么, 1 乆, Waterview Creek, 22 mls SW Ingham, ca. 2000 ft, 21. iii. 1961 乆, 30. v. 1961 么 (R. Straatman) (ANIC).

Remarks Probably undescribed species belonging to the Pollanisus subdolosa-eumetopus-group, with insufficient material for a description. Very close to P. eungellae, similar in all main characters but slightly larger and with very intensive shiny metallic yellowish green scales on dorsal parts of head, collar and dorsal parts of abdomen. The fairly constant coloration of all the rainforest species of Pollanisus in northern Queensland and the isolated habitat at Eungella indicate specific difference.

Pollanisus sp. 3

Pollanisus sp. 5

Pl. 25, Figs 1,2; Pl. 10, Fig. 2.

Pl. 27, Fig. 1, 2.

Material examined

Material examined

Queensland: 2 么, Magnetic I. (A. M. Lea) (SAM); 1 乆, Magnetic I., 17. ix. 1920 (QM); 1 么, Palm I., 18. vii. 1914 (ANIC); 1 乆, Palm I., 1. vi. 1926 (ANIC).

Queensland: 1 么, Kennedy R., 30 km W of ‘Fairview’, 15°35´S 144°03É, 24. xii. 1984 (at m.v. lamp) (G. & A. Daniels) (ANIC); 1 么, Kuranda, 3. v. 1961 (J. M[acqueen]) (ANIC).

TAXONOMY

Remarks Probably undescribed species belonging to the Pollanisus subdolosa-eumetopus-group, with insufficient material for a description. A small dark species with slender elongate forewings and relatively short hindwings (forewing length 7.5–8.0 mm), the forewing opaque, the hindwing slightly translucent, especially medially. Head with very narrow frons and huge eyes, the breadth of frons only ca. 0.6 that of compound eye in frontal view. Head unicolorous greyish black in the Kennedy River specimen (Pl. 27, Fig. 1), with slightly shiny metallic yellowish green frons in the Kuranda specimen. Ocelli large and black; proboscis brownish yellow. Antenna long, 1–26 bipectinate and 27–36 biserrate in the Kennedy River specimen, 1–31 bipectinate (rest broken) in the Kuranda specimen. Thorax without shiny collar but with a few shiny bluish green scales laterally. Abdomen greyish black, the posterior segments weakly shiny metallic coppery dorsally. Forewing upperside greyish black, underside dark grey, with a few shiny metallic greenish blue scales at costa proximally, hindwing black, with a fairly translucent medial part, underside with a shiny metallic greenish blue stripe in anterior part of cell. It is not certain that both specimens are conspecific although they are very similar in habitus. There are three further specimens with the same habitus but some different characters: Queensland: 1 么 (Pl. 27, Fig. 2), Base Cableway, Mt Bellenden-Ker, 80 m, 17º16´S 146º54É, 19. x. 1981 (E. D. Edwards) (ANIC) (head shiny blue dorsolaterally, abdomen more shiny and green); 1 么, Mt Bellenden-Ker, lower slopes, 500 m, 17º16´S 145º53É, 21. x. 1981 (E. D. Edwards) (ANIC) (head shiny blue dorsolaterally, collar with a faint purplish tinge, abdomen upperside slightly shiny coppery, proboscis blackish brown); 1 么, Kuranda, iv. 1907 (F. P. D[odd]) (ANIC) (head shiny yellowish green frontally and dorsolaterally, collar and dorsal abdomen shiny metallic coppery, proboscis blackish brown). Conspecificity of these specimens is uncertain.

Pollanisus sp. 6 Pl. 28, Fig. 1, 2; Figs 177, 178, 228, 229, 269.

Material examined Queensland: 1 么, Evelyn Scrub, i. 1911 (ANIC); 1 乆, Kuranda, iv. 1906 (F. P. Dodd) (MV); 1 乆, Kuranda, iv.

1907 (F. P. D[odd]) (ANIC); 1 乆, Paluma, 15 km W, 19º00´S 146º13É, e.l. 19. ix. 1985 (larva on Hibbertia synandra) (I. F. B. Common) (ANIC); 1 么, 1 乆, Townsville, ix. 1900 (ANIC).

Remarks Probably undescribed species belonging to the Pollanisus subdolosa-eumetopus-group, with insufficient material for a description. A small, very densely scaled species with dark almost varnish-like forewing upperside, broad frons and small eyes in male. Similar to Pollanisus sp. 3 and perhaps even conspecific. Forewing length 7.0 mm in male, 7.5–8.5 mm in female. With a very broad shiny metallic yellowish green frons and small compound eyes. Externally also similar to P. contrastus but hindwings more opaque. Antenna long, 1–30 bipectinate, 31–42 biserrate in male, the pectinations short (approximately 4× longer than shaft in dorsal view), biserrate in female. One female was reared from larvae found on Hibbertia synandra F. Muell. (Dilleniaceae). The two specimens from Kuranda differ in size and wing shape but similar in all other characters except ground coloration that may have changed in these old specimens. More material is required to decide the systematic status of this species.

Pollanisus sp. 7 Pl. 29, Fig. 1.

Material examined Northern Territory: 1 么, P[ort] Darwin (F. P. Dodd) (SAM).

Remarks Probably undescribed species belonging to the Pollanisus subdolosa-eumetopus-group, with insufficient material for a description. This single specimen from Darwin is the only known Pollanisus from the Northern Territory so far. In habitus it is very similar to Pollanisus sp. 6 but has slightly longer antennal pectinations that are more strongly tapering towards the apex and a very strongly shiny metallic yellowish golden frons, collar and abdominal upperside tinged with green. As F. P. Dodd collected in Darwin (= Port Darwin in former times) and many of the species have never been taken since, the label is accepted as

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genuine. More material from Northern Territory would be required to clarify the taxonomic status of this specimen.

Pollanisus sp. 8 Pl. 29, Fig. 2.

Material examined Queensland: 1 么, ‘Jowalbinna’ H. S., 6.7 km W, 15°45´S 144°12É, 10. v. 1989 (G. & A. Daniels) (UQIC).

Remarks Probably undescribed species belonging to the Pollanisus subdolosa-eumetopus-group, with insufficient material for a description. A species of medium size with narrow and very densely scaled wings, very broad frons, small compound eyes and long antennae. Body, antennae and forewing upperside with a very characteristic sheen. The antennal pectinations are short and very characteristic: 1–33 bipectinate, 34–45 biserrate, pectinations 1–9 becoming progressively longer (pectinations at segment 9 ca. 4× longer than breadth of shaft in dorsal view), 10–33 becoming progressively shorter. The entire head (excluding only the eyes, proboscis and chaetosema) intensive, shiny metallic yellowish green with a bluish tinge posteriad of compound eyes. Thorax with shiny metallic coppery collar and shiny bluish green lateral parts. Abdomen dark. This specimen is different from all other species from northern Queensland.

Pollanisus cyanota (Meyrick) Pl. 30, Figs 1, 2; Figs 179, 230–233, 270–272. Procris cyanota Meyrick, 1886, Proc. Linn. Soc. NSW (2) 1: 793. Lectotype 么 (here designated), AUSTRALIA: New South Wales, Sydney, 30. ii. 1879 (G. H. R[aynor]) (BMNH) [examined]. Pollanisus cyanota (Meyrick); Kirby 1892: 87; Jordan 1907: 10; Aurivillius 1920: 43; Hering 1922: 13; Bryk 1936: 118; Tarmann 1996: 143. Pollanisus cyanotus (Meyrick); Turner 1926b: 443.Pollanisus cyanotas [sic] Aurivillius, 1920, Ark. Zool. 13 (2): 43 (misspelling).

Material examined Queensland: 1 么, 1 乆, Howard, 5 km W, 24. viii. 1973 (I. F. B. Common) (ANIC); 1 么, Maryborough, 16 km N, 16. viii. 1973 (I. F. B. Common) (ANIC). New South Wales: 1 乆, Bega, 2. iii. 1960 (V. J. Robinson) (ANIC); 6 么, 1 乆, Berowra, 26. x. 1917, 2. xi. 1917 (AMS, ANIC); 2 乆, Berowra, 22. x. 1950 (L. H. Mosse-Robinson) (ANIC); 1 么, 1 乆, Canyonleigh, 28. iii. 1968 (V. J. Robinson) (ANIC); 1 么, Como, 26. ix. 1914 (AMS); 11 么, 1 乆, Congo, 8 km SE by E of Moruya, 8. xi. 1981, 19. iii. 1983 (M. S. Upton) (ANIC); 1 么, 1 么, Cowan, 12. x. 1953 (L. H. Mosse-Robinson) (ANIC); 1 乆, Depot Beach, 10 mls NE Bateman’s Bay, 25. xi. 1967 (I. F. B. Common) (ANIC); 1 乆, Fingal Bay, 24. x. 1962 (V. J. Robinson & K. English) (ANIC); 1 么, Hornsby, 25. iii. 1910 (G. Lyell) (MV); 1 么, 1 乆, Innaminna, 9. xi. 1897 (ANIC. MV); 1 么, 1 乆, Loddon Falls, 12. xii. 1948 (C. E. Chadwick) (NSWA); 2 乆, Mittagong, 17. xii. 1927 (AMS); 1 么, Oatley, x. 1912 (AMS); 2 么, Penrose SF, 6. v. 1967 (I. F. B. Common) (ANIC); 1 乆, Roseville, 20. ii. 1972 (C. E. Chadwick) (NSWA); 1 么, St George’s Basin, 2. xii. 1964 (V. J. Robinson) (ANIC); 2 么, Wilton, CSIRO Experimental Farm, 19. x. 1973, 2. xi. 1973 (V. J. Robinson) (ANIC); 1 么, Whiskers, 7 km NNW of Hoskingstown, 27. iii. 1993 (M. S. Upton) (ANIC). Australian Capital Territory: 1 么, 1 乆, Jervis Bay, 12.–16. x. 1919 (ANIC). Victoria: 1 么, Cann R., xi. 1928 (J. Clark) (MV); 1 么, Hazelwood, 1. iv. 1907 (MV).

Diagnosis Best recognised by its shiny coppery collar that contrasts with the shiny bluish green abdomen, its small eyes and broad frons, and in the female by its dark grey abdominal hairtuft (yellow in all other species with normally developed wings). These characters seem to be absolutely constant.

Redescription Male (Pl. 30, Fig. 1). Forewing length: 6.0–6.5 mm. Head greyish black except for whitish grey eyelashes below compound eye and some whitish grey scales at margin of chaetosema, especially posteriorly and (only in some specimens) on ventral part of frons; in fresh specimens often with a few coppery metallic scales posteriorly and dorsad of compound eye and sometimes on lateral part of frons; frons more than 2.5× as broad as the relatively small compound eye in frontal view, significantly protruding beyond compound eyes in lateral view; labial palps slender, porrect distally; proboscis dark brown proximally, yellow distally; ocelli small, white, distance from compound eye almost twice as large as diameter of ocellus; chaetosemata greyish,

TAXONOMY

the anterior extension covering the whole space between compound eye and ocellus. Antenna dark grey, segments 1 to 27–28 bipectinate, 27–28 to 37–39 biserrate; the pectinations very long, 7–8× longer than shaft in dorsal view proximally (at segments 8 and 9), becoming progressively shorter distally, only 2.5–3.0× longer than shaft at segment 25, sensory hairs short. Thorax greyish black dorsally, patagia with reddish-coppery metallic scales forming a conspicuous collar; ventral part of thorax covered with shiny bluish green metallic scales; legs dark grey, lighter laterally. Abdomen deep greenish blue. Wings short, forewing elongate triangular, hindwing almost rectangular, reaching almost 90% length of forewing, posterior margin straight, sometimes slightly concave. Forewing upperside black or brownish black, with a few scattered white scales, especially apically (in fresh specimens only); forewing underside greyish black, with a band of scattered bluish green shiny scales below costal margin, especially proximally. Hindwing upperside greyish black, less densely scaled and therefore translucent centrally; hindwing underside greyish black, with some shiny bluish green scales in costal and anal areas. Female (Pl. 30, Fig. 2). Forewing length: 6.0–6.5 mm. Similar to male in size and habitus. Head almost of same breadth as in male. Abdomen with green scales with a greyish-ochreous abdominal hairtuft. Male genitalia (Figs 179, 230–233). Valva slender, much longer dorsally, bluntly pointed apically, convex at two-thirds dorsally but straight proximally and distally, convex ventrally; folded central part almost in the form of a triangle with equally long shorter sides, with the longer third side on top dorsally; ventral sclerotization of valva with broad base, also triangular. Aedeagus very short, only 2.5–3.0× longer than broad, cornutus short, ca. 75% length of aedeagus, pointed apically. Female genitalia (Figs 270–272). Praebursa long and slender, with an ‘octopus-shaped’ central sclerotization with three or more long, partly curved teeth (‘feet of octopus’).

Phenology and bionomics According to the label data bivoltine. Nothing is known about habits and larval foodplants.

Distribution South-eastern Queensland, New South Wales, Victoria.

Pollanisus isolatus sp. n. Pl. 31, Fig. 1; Figs 180, 181, 234, 235.

Material examined Holotype 么, Victoria: Beaconsfield, 23. ii. 1904 (MV). Genitalia slide Z 3041 (MV).

Diagnosis and discussion This specimen is so distinct from all of the Pollanisus specimens examined for this revision, both in habitus and genitalia, that a description based on only one single male is justified. The genitalia are similar to those of P. lithopastus and P. calliceros but clearly different. There is no species known that is similar to P. isolatus.

Description Male. Forewing length: 7.0 mm. Head greyishbrownish black dorsally with a few shiny metallic golden scales, shiny metallic greenish golden dorsolaterally, brownish black frontally with a few shiny metallic golden scales, grey ventrally; frons very broad, almost 3× broader than the very small compound eyes in frontal view, strongly protruding beyond compound eyes in lateral view; labial palps short, grey, upcurved; proboscis blackish brown; ocelli very small, white, distance from compound eye 3.5× longer than diameter of ocellus; chaetosemata large but short, brown, not strongly extended anteriorly; antenna greyish-brownish black, segments 1–26 bipectinate, 27–34 weakly biserrate,

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almost simple, strongly pointed distally, pectinations long, approximately 6–7× longer than shaft in dorsal view. Thorax dark grey-brown dorsally, patagia, tegulae, anterior margin and three spots at posterior end of thorax with shiny metallic golden scales, thorax shiny metallic bluish green laterally, dark greyish brown ventrally; legs greyish black, with a few shiny metallic bluish green scales laterally. Abdomen shiny metallic bright blue dorsally and ventrally. Wings very rounded, especially hindwing, opaque; forewing upperside blackish brown, underside slightly paler, with a few scattered, shiny metallic bluish green scales proximally below costa; hindwing dark brownish grey, very opaque but with a paler central part, underside with scattered, shiny metallic bluish green scales, mostly in anterior half and at anal angle. Female. Unknown. Male genitalia (Figs 180, 181, 234, 235) Valva relatively long, straight proximally for two-thirds of length, dorsal and ventral margin parallel, strongly convex dorsally at two-thirds to apex, bluntly pointed apically, ventral sclerotization narrow, parallel to ventral margin, ending distally at about twothirds length of valva, where the ventral margin is strongly concave for a short distance, then straight distally to apex. Aedeagus relatively long, ca. 5× longer than broad, strongly sclerotized, upcurved; cornutus short and broad, ca. 60% length of aedeagus, strongly curved upwards, pointed apically.

designated), AUSTRALIA: Tasmania, Cradle Mountain Rd, Moina, 2000 ft., 30. i. 1925 (A. J. Turner) (ANIC) [examined]. Genitalia slide Z 3048 (ANIC). Pollanisus calliceros (Turner); Turner 1926b: 443; Bryk 1936: 117; Sattler 1991: 272; Tarmann, 1996: 143; Epstein et al. 1998: 174.

Pollanisus calliceros calliceros (Turner) Pl. 31, Fig. 2; Figs 182–183, 236–237.

Material examined Tasmania: 4 么, Paralectotypes, Cradle Mountain Road, Moina, 2000 ft., 30. i. 1925 (A. J. Turner) (ANIC, MV); 1 么, Cradle Mountain Rd, 2000 ft., 2. ii. 1934 (ANIC); 1 么, idem, 21. i. 1925 (QM). Victoria: 1 么, Native Dog Flat, Buchan R., 4000 ft, 24. i. 1970 (J. Brooks) (ANIC). New South Wales: 3 么, Moonbar, Monaro, 3000–3500 ft., iii. 1889 (Helms) (AMS); 1 么, Tuross, iii. 1936 (A. J. Brandon) (AMS).

Diagnosis Unmistakable owing to the short, rounded wings and the very strongly pectinated antenna in the male. No similar species is known.

Redescription

Pollanisus calliceros (Turner) Pl. 31, Fig. 1, Pl. 32, Figs 1, 2; Figs 14–21, 182–185, 236–239. Pollanisus calliceros Turner, 1926a, Pap. Proc. Roy. Soc. Tasmania 1925: 117 (1926). Lectotype 么 (here

Male (Pl. 31, Fig. 1). Forewing length: 6.5–7.5 mm. Head including antenna, thorax, abdomen, proximal part of legs, forewing upperside, hindwing underside and a distal spot near apex on forewing underside with shiny metallic, light green scales, iridescent, varying from yellowish green to purplish green. Frons very broad, 2.2× broader than compound eyes in frontal view, the latter small, deep black, labial palps short, porrect, strongly scaled, proboscis blackish brown; ocelli very small, white; chaetosema short and broad, dark brown, only slightly extended anteriorly; antenna short, segments 1 to 20–22 bipectinate, 21–23 to 23–25 simple, apex bluntly pointed, the pectinations very long, the length increasing from 1–10, where the pectinations are almost 10× as long as the breadth of shaft in dorsal view, decreasing in length from 10 to 20–22, sensory hairs very short. Wings short and rounded.

TAXONOMY

Female. The female from the nominotypical population is unknown. It is assumed that it is brachypterous as in P. calliceros azurea ssp. n.. Male genitalia (Figs 182–185, 236–239). Valva long and slender, much longer dorsally, strongly convex dorsally, convex also ventrally but only proximally, concave at three-quarters, slightly convex or straight distally, ventral sclerotization of valva strong, parallel to ventral margin. Aedeagus relatively long, ca. 5× longer than broad, strongly sclerotized, upcurved, cornutus short, only ca. 50–60% length of aedeagus, pointed distally.

Phenology and bionomics All specimens were collected in midsummer. Univoltine. Bionomics unknown.

Distribution The nominotypical P. calliceros calliceros occurs in Tasmania and in the mountains of Victoria and the southernmost parts of New South Wales. The four males from southern New South Wales are slightly darker green and one male has a shiny blue abdomen.

么, Ebor, 7. xii. 1940 (G. M. Goldfinch) (AMS); 11 么, Ebor, 14. ii. 1939, 20. ii. 1936, 20. ii. 1942, 4. ii. 1946, 3. iii. 1937, 3. iii. 1940, 6. iii. 1937 (AMS); 2 么, New England Nat. Park, near entrance, 1. i. 1956 (C. W. Frazier) (ANIC); 1 么, New England Nat. Park, iii. 1966 (AMS); 8 么, Tubrabucca Creek, Barrington Tops, 4200 ft., 6. i. 1967 (genitalia slide Z 2796 么) (V. J. Robinson) (ANIC, AMS); 1 么, Tubrabucca Creek, Barrington Tops, 4200 ft., 11. i. 1956 (R. Straatman) (ANIC).

Diagnosis Easy to recognise due to the bluish appearance.

Description Male (Pl. 32, Fig. 1). Forewing length: 7.0–8.0 mm. All shiny parts dark blue tinged with green on head and thorax and posteriorly on abdomen. Female (Pl. 32, Fig. 2). There is only one single female known. It is brachypterous with short rudimentary wings and a huge abdomen. The abdominal hairtuft is large and light brownish green. P. calliceros is the only known zygaenid species in the world that has a short-winged female. Male genitalia. As in nominotypical subspecies. Female genitalia. The only female specimen known has not been dissected so far because it is the only known zygaenid in the world with reduced wings. Moreover, the information that could be gained from the examination of the female genitalia is of minor significance in Pollanisus (see p. 63). I therefore preferred to leave the specimen intact and to wait for additional material.

Pollanisus calliceros azurea Pollanisus calliceros calliceros

Pollanisus calliceros azurea ssp. n. Material examined Holotype 么, New South Wales: Ebor, 11. i. 1914 (ANIC). Paratypes New South Wales: 3 么, Barrington Tops, i. 1925 (SU ZooExp) (SAM, AMS); 3 么, Barrington Tops, 25. i. 1922 (Nicholson) (AMS); 1 乆, Barrington Tops, 29. i. 1921 (G. M. Goldfinch) (AMS); 9 么, Ebor, 4000 ft., 9.-11. i. 1914, 31. i. 1915, 1. ii. 1915 (genitalia silde Z 3059 么) (ANIC); 2

Phenology and bionomics Univoltine. Bionomics unknown.

Distribution Mountainous parts of central and northern New South Wales. All populations north of the Blue Mountains belong to this subspecies.

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Fig. 136. Male genitalia of Pollanisus nielseni, paratype, dorsal view, one valva cut. This type of genitalia represents the primitive artonoid type (without ‘Artona finger’). Arrows: ds, dorsal sclerotization; ft, fan-shaped translucent medial part of valva; pv, pulvinus; sa, saccus; sl, sacculus; tg, tegumen; tt, transtilla; un, uncus; va, valva; vc, vinculum; vs, ventral sclerotization. WA, Wedge Island. Gen. slide no. Z 3191. Fig. 137. Male genitalia of Pollanisus nielseni, paratype, aedeagus, vesica everted by pulling it out. This type of aedeagus represents the primitive artonoid type (with one large cornutus but without additional deciduous cornuti). WA, Wedge Island. Gen. slide no. Z 3191. Figs 138–139. Male genitalia: valva of Pollanisus viridipulverulenta, ACT, Aranda. Gen. slide no. Z 2950. 138: Distal part; 139: proximal part. Figs 140–141. Male genitalia: valva of Pollanisus viridipulverulenta, ACT, Aranda. Gen. slide no. Z 2951. 140: Distal part; 141: proximal part.

TAXONOMY

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Figs 142–143. Male genitalia: valva of Pollanisus cupreus, WA, Perth. Gen. slide no. Z 2954. 142: Distal part; 143: proximal part. Fig. 144. Male genitalia: valva of Pollanisus nielseni, WA, Wedge Island. Gen. slide no. Z 3191.Figs 145–146. Male genitalia: valva of Pollanisus modestus, holotype, NSW, Clyde Mountain. Gen. slide no. Z 3064. 145: Distal part; 146: proximal part, pulvinus missing.

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Figs 147–148. Male genitalia: valva of Pollanisus lithopastus, Tas., Cradle Mountain. Gen. slide no. Z 3007. 147: Distal part; 148: proximal part. Figs 149–150. Male genitalia: valva of Pollanisus lithopastus, Tas., Flinders Island. Gen. slide no. Z 3010. 149: Distal part; 150: proximal part.Figs 151–152. Male genitalia: valva of Pollanisus lithopastus, NSW, Ku-ring-gai-Chase National Park. Gen. slide no. Z 3012. 151: Distal part; 152: proximal part.

TAXONOMY

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Figs 153—154. Male genitalia of Pollanisus sp. 1, ACT, Bendora. Gen. slide no. Z 3046. 153: Overview; 154: valva. Figs 155–156. Male genitalia: valva of Pollanisus empyrea, WA, Perth. Gen. slide no. Z 3020. 155: Distal part; 156: proximal part. Figs 157–158. Male genitalia: valva of Pollanisus apicalis, ACT, Black Mountain. Gen. slide no. Z 3016. 157: Distal part; 158: proximal part.

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Fig. 159–160. Male genitalia: valva of Pollanisus apicalis, Vic., Wyperfeld National Park. Gen. slide no. Z 3017. 159: Distal part; 160: proximal part. Figs 161–162. Male genitalia: valva of Pollanisus trimacula, NSW, Otford. Gen. slide no. Z 3027. 161: Distal part; 162: proximal part. Figs 163–164. Male genitalia: valva of Pollanisus trimacula, NSW, Mt Dromedary. Gen. slide no. Z 3032. 163: Distal part; 164: proximal part.

TAXONOMY

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Figs 165–166. Male genitalia: valva of Pollanisus edwardsi, paratype, NSW, Mt Kosciuszko. Gen. slide no. Z 3068. 165: Distal part; 166: proximal part. Fig. 167. Male genitalia: valva of Pollanisus subdolosa clara, NSW, Gosford. Gen. slide no. Z 3034.Fig. 168. Male genitalia: valva of Pollanisus cf. subdolosa, NSW, Como West. Gen. slide no. Z 3036.Figs 169–170. Male genitalia: valva of Pollanisus cf. subdolosa, NSW, Wollongong. Gen. slide no. Z 3005. 169: Distal part; 170. proximal part.

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Figs 171–172. Male genitalia: valva of Pollanisus contrastus, paratype, NSW, Wollongong. Gen. slide no. Z 3069. 171: Distal part; 172: proximal part.Figs 173–174. Male genitalia of Pollanisus eungellae, holotype, Qld, Eungella. Gen. slide no. Z 3194. 173: Overview; 174: valva.Fig. 175. Male genitalia: valva of Pollanisus eumetopus, Qld, Kuranda. Gen. slide no. Z 3053.Fig. 176. Male genitalia of Pollanisus incertus, holotype, Qld, Kuranda. Gen. slide no. Z 3195.

TAXONOMY

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Figs 177–178. Male genitalia: valva of Pollanisus sp. 6, Qld, Kuranda. Gen. slide no. Z 3043. 177: Distal part; 178: proximal part. Fig. 179. Male genitalia: valva of Pollanisus cyanota, NSW, Wilton. Gen. slide no. Z 3054.Figs 180–181. Male genitalia: valva of Pollanisus isolatus, holotype, Vic., Beaconsfield. Gen. slide no. Z 3041. 180: Distal part; 181: proximal part.

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Figs 182–183. Male genitalia: valva of Pollanisus calliceros calliceros, Tas., Cradle Mountain. Gen. slide no. Z 3058. 182: Distal part; 183: proximal part. Figs 184–185. Male genitalia: valva of Pollanisus calliceros azurea, paratype, NSW, Ebor. Gen. slide no. Z 3059. 184: Distal part; 185: proximal part. Figs 186–187. Male genitalia: aedeagus of Pollanisus viridipulverulenta, ACT, Aranda. Gen. slide no. Z 2950. 186: Overview; 187: detail.

TAXONOMY

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Figs 188–189. Male genitalia: aedeagus of Pollanisus viridipulverulenta, ACT, Aranda. Gen. slide no. Z 2951. 188: Overview; 189: detail. Figs 190–191. Male genitalia: aedeagus of Pollanisus cupreus, WA, Perth. Gen. slide no. Z 2954. 190: Overview; 191: detail. Figs 192–193. Male genitalia: aedeagus of Pollanisus modestus, holotype, NSW, Clyde Mountain. Gen. slide no. Z 3064. 192: Overview; 193: detail.

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Figs 194–195. Male genitalia: aedeagus of Pollanisus lithopastus, NSW, Ebor. Gen. slide no. Z 3011. 194: Overview; 195: detail. Figs 196–197. Male genitalia: aedeagus of Pollanisus lithopastus, Tas., Flinders Island. Gen. slide no. Z 3010. 196: Overview; 197: detail. Figs 198–199. Male genitalia: aedeagus of Pollanisus sp. 1, ACT, Bendora. Gen. slide no. Z 3046. 198: Overview; 199. detail.

TAXONOMY

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Figs 200–201. Male genitalia: aedeagus of Pollanisus empyrea, WA, Perth. Gen. slide no. Z 3020. 200: proximal part, detail. 201: distal part, detail. Figs 202–203. Male genitalia: aedeagus of Pollanisus empyrea, WA, Mt Dale. Gen. slide no. Z 3021. 202: proximal part, detail; 203: distal part, detail. Figs 204–205. Male genitalia: aedeagus of Pollanisus empyrea, WA, Augusta. Gen. slide no. Z 3023. 204: proximal part, detail; 205: distal part; detail.

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Figs 206–207. Male genitalia: aedeagus of Pollanisus apicalis, ACT, Black Mountain. Gen. slide no. Z 3016. 206: Overview; 207: detail. ACT, Black Mountain. Gen. slide no. Z 3016.Figs 208–209. Male genitalia: aedeagus of Pollanisus apicalis, Vic., Wyperfeld National Park. Gen. slide no. Z 3017. 208: Overview; 209: detail. Figs 210–211. Male genitalia: aedeagus of Pollanisus trimacula, NSW, Mt Dromedary. Gen. slide no. Z 3032. 210: Overview; 211: detail.

TAXONOMY

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Figs 212–213. Male genitalia: aedeagus of Pollanisus edwardsi, paratype, NSW, Tenterfield. Gen. slide no. Z 3045. 212: Overview; 213: detail. Figs 214–215. Male genitalia: aedeagus of Pollanisus subdolosa clara, paratype, NSW, Gosford. Gen. slide no. Z 3034. 214: Overview; 215: detail. Figs 216–217. Male genitalia: aedeagus of Pollanisus contrastus, paratype, NSW, Wollongong. Gen. slide no. Z 3069. 216: Overview; 217: detail.

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Fig. 218. Male genitalia: aedeagus of Pollanisus eungellae, holotype, Qld, Eungella. Gen. slide no. Z 3194. Figs 219–220. Male genitalia: aedeagus of Pollanisus eumetopus, Qld, Kuranda. Gen. slide no. Z 3053. 219: Overview; 220: detail.Figs 221–222. Male genitalia: aedeagus of Pollanisus commoni, paratype, Qld, Clifton Beach. Gen. slide no. Z 3049. 221: Overview; 222: detail.

TAXONOMY

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Figure 223. Male genitalia: aedeagus of Pollanisus commoni, paratype, Qld, Clifton Beach. Gen. slide no. Z 2860. 223: Overview; 224: detail. Fig. 225. Male genitalia: aedeagus of Pollanisus incertus, holotype, Qld, Kuranda (twice the size of Figs 223 and 226). Gen. slide no. Z 3195.Figs 226–227. Male genitalia: aedeagus of Pollanisus angustifrons, paratype, Qld, Paluma. Gen. slide no. Z 3044. 226: Overview; 227: detail.

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Figs 228–229. Male genitalia: aedeagus of Pollanisus sp. 6, Qld, Kuranda. Gen. slide no. Z 3043. 228: Overview; 229: detail. Figs 230–231. Male genitalia: aedeagus of Pollanisus cyanota, NSW, Wilton. Gen. slide no. Z 3054. 230: Overview; 231: detail. Figs 232–233. Male genitalia: aedeagus of Pollanisus cyanota, NSW, Congo. Gen. slide no. Z 3055. 232: Overview; 233: detail.

TAXONOMY

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Figs 234–235. Male genitalia: aedeagus of Pollanisus isolatus, holotype, Vic., Beaconsfield. Gen. slide no. Z 3041. 243: Overview; 235: detail. Figs 236–237. Male genitalia: aedeagus of Pollanisus calliceros calliceros, Tas., Cradle Mountain. Gen. slide no. Z 3058. 236: Overview; 237: detail. Figs 238–239. Male genitalia: aedeagus of Pollanisus calliceros azurea, paratype, NSW, Ebor. Gen. slide no. Z 3059. 238: proximal part, detail; 239: distal part, detail.

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Figs 240–241. Female genitalia of Pollanisus viridipulverulenta, ACT, Aranda. Gen. slide no. Z 2952. 240: Overview; 241. detail, sclerotization at insertion of ductus intrabursalis. Figs 242–243. Female genitalia of Pollanisus cupreus, WA, Lancelin. Gen. slide no. Z 3189. 242: Praebursa with spermatophore; 243. detail, sclerotization at insertion of ductus intrabursalis, piercing wall of spermatophore.Figs 244–245. Female genitalia of Pollanisus nielseni, WA, Wedge Island. Gen. slide no. Z 3190. 244: Praebursa with spermatophore; 245: detail, sclerotization at insertion of ductus intrabursalis, piercing wall of spermatophore.

TAXONOMY

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Figs 246–247. Female genitalia of Pollanisus lithopastus, Vic., Yinnar. Gen. slide no. Z 3009. 246: Overview; 247: detail, sclerotizations. Figs 248–249. Female genitalia of Pollanisus sp. 1, ACT, Bendora. Gen. slide no. Z 3197. 248: Overview; 249: detail, sclerotization at insertion of ductus intrabursalis. Figs 250–251. Female genitalia of Pollanisus empyrea, detail, sclerotization at insertion of ductus intrabursalis. WA, Yanchep. 250: Gen. slide no. Z 3022; 251: gen. slide no. Z 3026.

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Figs 252–253. Female genitalia of Pollanisus apicalis, ACT, Black Mountain. Gen. slide no. Z 3018. 252: Overview; 253: detail, sclerotization at insertion of ductus intrabursalis. Fig. 254. Female genitalia of Pollanisus trimacula, NSW, Minnamurra Falls. Gen. slide no. Z 3228. Fig. 255. Female genitalia of Pollanisus edwardsi, paratype, detail, sclerotization at insertion of ductus intrabursalis. Gen. slide no. Z 3192. Fig. 256. Female genitalia of Pollanisus subdolosa subdolosa, detail, sclerotization at insertion of ductus intrabursalis. Gen. slide no. Z 3193. Fig. 257. Female genitalia of Pollanisus subdolosa clara, detail, sclerotization at insertion of ductus intrabursalis. NSW, Niagara Park. Gen. slide no. Z 3037.

TAXONOMY

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Figs 258–259. Female genitalia of Pollanisus cf. subdolosa, NSW, Royal National Park. Gen. slide no. Z 3039. 258: Overview; 259; detail, sclerotization at insertion of ductus intrabursalis. Fig. 260. Female genitalia of Pollanisus contrastus, paratype, detail, sclerotization at insertion of ductus intrabursalis. NSW, Narara. Gen. slide no. Z 3199. Figs 261–262. Female genitalia of Pollanisus eungellae, paratype, Qld, Eungella. Gen. slide no. Z 3052. 261: Overview; 262: detail, sclerotization at insertion of ductus intrabursalis. Fig. 263. Female genitalia of Pollanisus acharon, lectotype, detail, praebursa with sclerotization at insertion of ductus intrabursalis. [AUSTRALIA] [‘Nova Hollandia’]. Gen. slide no. Z 2628: BMNH Zyg. 1626.

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Fig. 264. Female genitalia of Pollanisus eumetopus, detail, sclerotization at insertion of ductus intrabursalis. Qld, Redlynch. Gen. slide no. Z 3048. Figs 265–266. Female genitalia of Pollanisus commoni, paratype, Qld, Clifton Beach. Gen. slide no. Z 3050. 265: aa, apopyses anteriores; ap, apophyses posteriores; apx, appendix of praebursa; db, ductus bursae; cb, corpus bursae; di, ductus interabursalis; ds, ductus seminalis; idi, insertion of ductus intrabursalis; ob, ostium bursae; pa, papillae anales; pb, praebursa; sii, sclerotization at insertion of ductus intrabursalis; 266: detail, sclerotization at insertion of ductus intrabursalis. Figs 267–268. Female genitalia of Pollanisus incertus, paratype, Qld, Kuranda. Gen. slide no. Z 3196. 267: Overview; 268. detail, sclerotization at insertion of ductus intrabursalis.

TAXONOMY

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Fig. 269. Female genitalia of Pollanisus sp. 6, Qld, Paluma. Gen. slide no. Z 3038. Figs 270–271. Female genitalia of Pollanisus cyanota, NSW, Depot Beach. Gen. slide no. Z 3057. 270: Overview; 271: detail, sclerotization at insertion of ductus intrabursalis. Fig. 272. Female genitalia of Pollanisus cyanota, detail, sclerotization at insertion of ductus intrabursalis. NSW, Innaminna. Gen. slide no. Z 3056.

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TAXONOMY

Genus Onceropyga Turner Onceropyga Turner, 1906, Trans. Proc. R. Soc. S. Aust. 30: 137. Type species: Onceropyga anelia Turner, 1906: 137, by monotypy. [Queensland]. Ocneropyga; Bryk 1936: 259. Unavailable; an incorrect subsequent spelling of Onceropyga Turner, 1906. Onceropyga Turner; Jordan 1908: 47; Turner 1926b: 441; Bryk 1936: 259; Common 1970: 830; Fletcher and Nye 1982: 113; Common 1990: 17, fig. 10.8–10.10 (head ultrastructure), 121, fig. 52.10 (larva), 52.11 (cocoon), 296, fig. 97 (wing venation, genitalia, pupa), 298 (bionomics); Tarmann 1994: 118; Tarmann 1996: 143; Tarmann 1998: 75; Epstein et al. 1998: 173.

Diagnosis Small species with narrow wings and brownish ground colour, shining metallic scales absent. Female with abdominal hairtuft as in Pollanisus and Hestiochora. These three genera are considered to be monophyletic (see p. 55) (Pls 33, 34). Head (Figs 22, 23) Frons slightly projected, broad in male, broader in female; labial palps short, curved proximally, porrect distally, maxillary palps short, rounded distally, pilifer short; proboscis brownish yellow; compound eyes small, black, with a small eyelash ventrally; ocelli small, white; chaetosemata strongly projected anteriorly; male antenna strongly bipectinate, the distal segments biserrate or simple, apex rounded, sensory hairs on pectinations short; female antenna biserrate. Thorax Foreleg without tibial epiphysis, tibial spurs 0–2–2, spurs of almost equal length, the outer spur slightly longer; no single medial spur on hind-tibia as in many other genera of Artonini. Wings narrow, forewing triangular in male, elongate rounded in female, hindwing two-thirds length of forewing, rounded. Forewing with all veins free from cell or with R4+R5 (often) and M2+M3 (sometimes) stalked, [R4+R5] sometimes even fused to one vein, medial stem absent; hindwing mainly with M2+M3, rarely with M3+CuA1 stalked. Male frenulum consisting of one strong spine, retinaculum a well-developed hook at base of Sc, female frenulum consisting of three spines in

one row, inserting into a row of anteriorly pointing scales at base of CuP. Pregenital abdomen Abdomen with lateral evaginations only on segment 2. This has been interpreted as a secondary loss of the evaginations on segment 7 (Tarmann 1994: 121). Female with large abdominal hairtuft. Sternum II with a pair of short, slightly curved apodemes, lateral sternal rod weakly developed, short and slender, longitudinal rod weakly developed, slightly curved inwards; sclerotization of segment weak, with broad lateral prolongations anteriorly. Tergum II weakly sclerotized, the sclerotized part covering approximately 30% of segment posteriorly. Sternum VII and VIII in female weakly sclerotized and of minor taxonomic significance. Translucent parts of abdominal end enlarged in female, epidermis with smooth and sticky surface bearing the long, needle-like scales of the abdominal hairtuft. Male genitalia Both known species show clear differences and are easy to recognise in genitalia. We can assume that in this genus the genitalic characters are significant (see also p. 140 for Pollanisus and pp. 63, 65 for Hestiochora). Uncus slender, pointed distally, tegumen and vinculum slender, weakly sclerotized, a distinct saccus is absent (only a slight hook visible). Valva fan-shaped, strongly spreading when pressed, with a rod-shaped sclerotization dorsally and ventrally and a broad, strongly folded translucent central part, without appendages, sacculus very pronounced. Pulvinus transformed into an oval or finger-shaped apically rounded structure bearing setae distally. Juxta broad and rounded. Aedeagus short and stout, everted vesica double-lobed, with one long and slender cornutus. Female genitalia All parts very translucent. Praebursa well developed, with a translucent lateral appendix and a plate-like lateral sclerotization. Ductus intrabursalis inserting laterally approximately at the centre of the praebursa where a small tooth-like sclerotization is developed. The pair of abdominal glands (homologous to Petersen’s gland in Zygaeninae) that are close to ooporus is absent.

Bionomics The bionomics of the type-species has been described by Common (1990: 121) (see below).

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Remarks In Turner’s original generic description the antennae of the male and female are described as ‘in both sexes thickened and moderately pectinated to apex’. This is incorrect. The males of both known species have a strongly bipectinate antenna, the female of O. anelia has a strongly biserrate antenna and that of O. pulchra sp. n. is weakly biserrate. Furthermore, Turner’s description was based on two females. He had no males as he notes in the description of the type-species O. anelia. Later, Turner corrected his error but still describes the female antenna as moderately pectinate (Turner, 1926b: 441). The difference between a ‘moderately pectinate’ antenna and a ‘serrate’ antenna is not dependent upon the length of the pair of appendages on the antennal segment but on their morphological groundplan. A pectination is always an appendage of almost the same breadth basally and apically whereas a sawtooth-like appendage with broad basal and pointed apical part is called a serration. With this definition the female antenna of Onceropyga is clearly biserrate and not bipectinate.

(SAM); 1 么, Mt Tamborine, 28. i. 1912 (ANIC); 1 么, Mt Tamborine, 16. ii. 1960 (E. M. Exley) (UQIC); 1 么, Numinbah, 21. iv. 1935 (DPIQ); 4 么, 1 乆, Prince Henry Heights, Toowoomba, 620 m, e.l. 3. i.–5. ii. 1986 (larva on Cissus antarctica) (I. F. B. Common) (ANIC); 1 么, St Lucia, Brisbane, e.l. 8. viii. 1985 (larva on Cissus antarctica (G. B. Monteith) (ANIC); 2 么, Stradbroke I. N, 23. iii. 1960 (J. Martin) (UQIC); 1 乆, paralectotype, Toowoomba, 12. iv. 1905 ([A. J. Turner]) (ANIC); 2 么, 1 乆, Toowoomba, 13. x. 1916, 16.–17. xii. 1916 ([A. J. Turner]) (ANIC); 2 乆, Toowoomba, 14.–26. ix. 1924 (QM); 1 乆, Toowoomba, 1. iii. 1941 (I. F. B. Common) (ANIC); 4 么, Westwood, 28. x.–4 xi. 1924 (AMS). New South Wales: 1 么, Narrabeen, 9. iv. 1925 (MV); 1 么, Razor Back, near Picton, 28. iv. 1949 (I. F. B. Common) (ANIC); 1 么, Rous, Richmond R., iv. 1936 (V. J. Robinson) (ANIC); 1 么, Saddle Back, 20. x. 1973 (V. J. Robinson) (ANIC); 1 乆, Stanwell Park, i. 1913 (AMS); 1 么, Stanwell Park, 12. iii. 1922 (MV); 1 么, Strickland SF, near Ourimbah, 7. ii. 1986 (B. Hacobian) (ANIC).

Diagnosis Easily recognized by its uniform dark brown appearance. No similar species is known.

Distribution

Redescription

South-eastern Queensland and New South Wales.

Male (Pl. 33, Fig. 1). Forewing length: 6.0–6.5 mm. Head, thorax and abdomen dark grey-brown dorsally, creamy white ventrally; head with broad frons and small black compound eyes, the frons approximately 2.5× broader than compound eyes in frontal view, strongly protruding beyond compound eyes in lateral view; chaetosemata dark brown. Antennal segments 1–28 bipectinate, the pectinations very long, approximately 10–11× longer than shaft in dorsal view at segment 10, segment 29 simple, rounded distally. Forewing upperside blackish brown with some spots and streaks of variable shape and consisting of creamy white scales, underside considerably paler proximally; hindwing opaque, upperside brownish black, underside paler proximally; fresh specimens with a dark spot at distal end of cell in both wings.

Key to Onceropyga species Abdomen brown . . . . . . . . . . . . . . O. anelia (p. 134) Abdomen yellow . . . . . . . . O. pulchra sp. n. (p. 135)

Onceropyga anelia Turner Pl. 33, Figs 1, 2; Figs 22, 23, 273–276, 281. Onceropyga anelia Turner, 1906, Trans. Proc. R. Soc. S. Aust 30: 137. Lectotype 乆 (here designated), AUSTRALIA: Queensland, Toowoomba, 12. iv.1905 ([A. J. Turner]) (ANIC) [examined]. Genitalia slide Z 3066 (ANIC). Onceropyga anelia Turner; Jordan 1908: 47; Bryk 1936: 259; Fletcher and Nye 1982: 113; Common 1990: 121, fig. 52.10 (larva), 52.11 (cocoon), 296, fig. 97 (wing venation, genitalia, pupa), 298 (bionomics); Tarmann 1996: 143; Tarmann 1998: 75, Epstein et al. 1998: 175 (wing venation).

Material examined Queensland: 1 么, Binna Burra, Lamington Nat. Park, 700 m, 15. xi. 1990 (I. F. B. Common) (ANIC); 1 么, Cairns,

Female (Pl. 33, Fig. 2). Forewing length: 7.0 mm. Similar to male but slightly larger, forewings more rounded, general appearance paler, more brownish. With a large, grey-brown abdominal hairtuft. Antenna strongly biserrate. Male genitalia (Figs 273–276). Uncus approximately half length of valva dorsally; the ventral apically rounded sclerotization of valva broad, strongly extending beyond distal margin. Pulvinus-

TAXONOMY

like structure finger-shaped, situated anteriorly and dorsally at base of valva. Aedeagus approximately 4× longer than broad, cornutus as long as aedeagus, slender, curved basally. Female genitalia (Fig. 281). Praebursa with a short, stout lateral appendix.

Phenology and bionomics This species has been reared by I. F. B. Common. Larvae are pale yellowish green with a reddish dorsal pattern (Common 1990: 121, fig. 52.10, 298). It is sometimes common on its host-plant Cissus antarctica Vent. (Vitaceae). The larva pupates in a very flat, oval, white cocoon (Common 1990: 121, fig. 52.11). Pupa stout, flattened dorsoventrally, with moderately sclerotized cuticle, appendages only weakly fused to one another and to body, proboscis extending beyond wing-tips; maxillary palps absent, labial palps, fore coxae and fore femora exposed; antennae broad, two-thirds length of wings; abdominal segments 2–8 with a single row of dorsal spines, spiracles of abdominal segment 1 fully exposed, abdominal segments 2–7 movable in male, 2–6 moveable in female, cremaster absent but a few hooked setae at abdominal end of pupa (Common 1990: 296, fig. 97.4, 297). Some of the described larvae are preserved in alcohol in ANIC (Canberra). From this material that has lost its colour it must be concluded that there is no major difference between the larva of Pollanisus and Onceropyga. The same can be said for the pupa. It should be noted that in alcohol the wings of the pupa become completely loose and it can be seen that fore- and hindwing are accommodated in two different independent compartments.

Distribution Eastern Queensland and northerly parts of eastern New South Wales.

Onceropyga pulchra sp. n. Pl. 34, Figs 1, 2; Figs 277–280, 282.

Material examined Holotype 么, New South Wales, Watagan State Forest, near Wyong, 18. xii. 1984 (B. Hacobian) (ANIC). Genitalia slide Z 3060 (ANIC). Paratypes Queensland: 1 么, Saddletree Creek, via Maidenwell, 12.–14. iii. 1976 (A. Postle) (genitalia slide Z 3061) (UQIC). New South Wales: 3 乆, Mt Keira, 1. ii. 1963 (V. J. Robinson) (genitalia slides Z 3062, Z 3063) (ANIC).

Diagnosis A very characteristic and beautiful species that can be easily recognised. There are only two other Australian species with a coloured abdomen, viz. Myrtartona rufiventris and M. mariannae. Their abdomina are more reddish or orange, not yellow, and the males have long antennae with short pectinations, the females lack the abdominal hairtuft.

Description Male (Pl. 34, Fig. 1). Forewing length: 6.5 mm. Dorsal parts of head with antenna, thorax and first abdominal segment dark grey-brown, ventral half of scapus, frons, eyelashes, palps, lateral parts of legs and ventral part of abdomen ochreous yellow, thorax light brown ventrally, abdominal segments 2–8 bright yellow dorsally. Frons broad, 2.3× broader than black compound eyes in frontal view, rounded and only slightly projecting beyond compound eyes in lateral view; chaetosemata brownish yellow. Antennal segments 1–22 bipectinate, 23–31 biserrate, 32 simple, the pectinations very long, approximately 8–9× longer than shaft in dorsal view at segment 10. Forewing upperside dark brown with a slightly lighter spot at end of cell, distally bordered by a suffusion of blackish scales, underside dark grey-brown, slightly lighter medially; hindwing on both sides blackish, slightly lighter basally, with a light brown costal margin. Female (Pl. 34, Fig. 2). Forewing length: 8.0 mm. Similar to male but larger, wings slightly more rounded, lighter greyish brown with a dark spot at distal margin of cell. Hindwing opaque, dark greyish brown, only very slightly lighter basally. Abdominal hairtuft yellow anteriorly, greyish poste-

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riorly. Antenna weakly biserrate. Male genitalia (Figs 277–280). Uncus long, only slightly shorter than length of valva dorsally; tegumen long and slender, vinculum short and rounded; ventral sclerotization of valva slender, only shortly extending beyond distal edge, pulvinus-like structure oval, short, with short setae; aedeagus approximately 6× longer than broad, curved upwards, with a very long and slender cornutus of almost same length as aedeagus, cornutus slightly curved upwards distally but straight basally.

Distribution South-eastern Queensland and eastern New South Wales.

Female genitalia (Figs 282). Lateral appendix on praebursa long and slender.


273

274

275

276

Figs 273–276. Male genitalia of Onceropyga anelia, Qld, Brisbane. Gen. slide no. Z 3065. 273: One valva cut off; 274: valva; 275: aedeagus; 276: aedeagus, detail.

TAXONOMY

277

278

280 279

281

282

Fig. 277. Male genitalia of Onceropyga pulchra, paratype, Qld, Saddletree Creek. Gen. slide no. Z 3061. Figs 278–280. Male genitalia of Onceropyga pulchra, holotype, NSW, Watagan State Forest. Gen. slide no. Z 3060. 278: Valva; 279: aedeagus; 280: aedeagus, detail. Fig. 281. Female genitalia of Onceropyga anelia, lectotype, Qld, Toowoomba. Gen. slide no. Z 3066. Fig. 282. Female genitalia of Onceropyga pulchra, paratype, NSW, Mt Keira. Gen. slide no. Z 3062.

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TAXONOMY

Genus Hestiochora Meyrick Hestiochora Meyrick, 1886, Proc. Linn. Soc. NSW (2)1: 788. Type species: Procris tricolor Walker, 1854, List Specimens lepid. Insects Colln Br. Mus. 1: 111, by subsequent designation by Turner (1926b: 441). [Tasmania]. Histiochora; Pagenstecher 1909: 44. Unavailable; an incorrect subsequent spelling of Hestiochora Meyrick, 1886. Hestiochora Meyrick; Jordan 1907: 9; Hering 1922: 13; Simmonds, 1924: no page; Turner 1926b: 441; Bryk 1936: 116; Alberti 1954: 218; Common 1970: 830, pl. 6ZC; Fletcher and Nye 1982: 76; Common 1990: 297; Tarmann 1994: 118; Tarmann 1996: 143; Epstein et al. 1998: 172; Efetov and Tarmann 1999: 8. Hestiochora was originally described to accommodate the four species Procris rufiventris Walker, 1854, Hestiochora xanthocoma Meyrick, 1886, Hestiochora erythrota Meyrick, 1886 and Procris tricolor Walker, 1854. Jordan (1907: 9) believed that there is a close relationship between this genus and the Southeast Asian genus Callizygaena Felder, 1874, mainly because of the bee- or wasp-like habitus of the adults of both groups. This opinion was followed by Alberti (1954: 218) who correctly excluded Procris rufiventris from the genus Hestiochora. Alberti (1954: 209) divided the subfamily Procridinae into two tribes, Callizygaenini and Procridini. Hestiochora was included in Callizygaenini. Tarmann (1994: 120) raised Callizygaenini to subfamily level based on genitalia characters but excluded Hestiochora as this genus has the autapomorphies of the tribe Artonini of the Procridinae (Tarmann 1994: 120) (for characters see p. 69). Within the Artonini, Hestiochora shares a striking synapomorphy with the genera Pollanisus and Onceropyga. The female has an abdominal hair-tuft consisting of long, spiny scales that are attached to the surface of the eggs after oviposition. These hairlike scales give the eggs a pincushion-like appearance. The tips of the scales seem to be poisonous as aphids die after touching them (Tarmann 1994: 118). Corresponding to this character is the reduction of a pair of abdominal glands close to the ooporus, present in all Procridinae, known as Petersen’s gland in Zygaeninae, which produce a poisonous liquid presumably to protect the eggs from egg parasitoids or predators (Naumann 1988). The monophyly of the above three Australian genera is supported by the characters listed above.

Diagnosis Externally, species of Hestiochora are colourful and most of them resemble wasps. The main autapomorphies of Hestiochora are: 1. wasp-like habitus (Pls 35–43); 2. complete reduction of lateral abdominal evaginations; 3. reduction of praebursa. Head (Figs 24–28) Frons only slightly projected dorsally, twice as broad as compound eye in frontal view in male, slightly broader in female; proboscis yellow or brownish yellow; labial palps short, orange, red or yellow; pilifer very short; compound eyes small, deep black; ocelli small, white; chaetosemata well developed, artonoid (see Fig. 26); antenna bipectinate in male, tapering towards apex, the last segments biserrate, antenna biserrate in female. Thorax Foreleg without tibial epiphysis, tibial spurs 0–2–2, spurs of equal length on midleg, the outer spur slightly longer on hindleg; no single medial spur on hindtibia as in many other genera of Artonini. Wings narrow, rounded apically, length of hindwing approximately three-quarters that of forewing. Forewing with medial stem mainly reduced, all veins free from cell or R4+R5 shortly stalked; hindwing with medial stem present, Sc fused with anterior margin of cell, at least for a short distance, [Rs+M1] fused; frenulum strongly developed, a long, single spine in male with a retinaculum consisting of a very pronounced cuticular fold, two or three smaller spines in female with a retinaculum consisting of a row of scales at basal part of CuP, pointing anteriorly. Pregenital abdomen Lateral evaginations on segments II and VII absent, a character that has been interpreted as a secondary loss (Tarmann 1994: 121). Sternum II with a pair of short, almost straight apodemes, lateral sternal rod strongly developed, longitudinal rod slender, slightly curved inwards, longer in male than in female; sclerotization of segment with broad lateral elongations anteriorly, projecting beyond lateral sternal rod that crosses posteriorly. Tergum II with a sclerotized part covering approximately two-thirds of the segment posteriorly; the small connecting sclerite to metathorax of characteristic shape. Sternum VIII in male covering only the anterior part of segment. Sterna VII and VIII in female transformed into sclerotized plates of taxonomic significance (Figs 328, 329).

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Translucent parts of abdominal end enlarged in female, epidermis with smooth and sticky surface bearing the long, needle-like scales of the abdominal hair-tuft (Pl. 58, Figs 5, 6). Male genitalia (Figs 283–327) Uncus slender, slightly constricted and pointed distally. Tegumen and vinculum fused into a slender ring, more strongly sclerotized ventrally, saccus small, sometimes almost absent. Valva with a large proximal sclerotization ventrally that may be interpreted as a swollen sacculus with a group of setae at its ventral margin; in its natural position this sclerotization is hemispherical with the opening anteriorly; when the male genitalia are spread, flattened and prepared on a slide this sclerotization often collapses and its form changes significantly; distal part of valva with a dorsal and a ventral sclerotization and a translucent central part; the ventral sclerotization sometimes bears tooth-like dorsal or finger-like ventral processes and is often curved upwards distally and pointed apically. Pulvinus (sensu Alberti 1954: 157) present but small, with short setae. Dorsal diaphragm translucent, ventral diaphragm sclerotized, forming a band, with a central, ring-like structure in some species; these structures often differ from the typical juxta form. Aedeagus long and slender, strongly sclerotized, straight or curved, pointed distally; insertion of ductus ejaculatorius dorsally at 25–30% length of aedeagus, the latter therefore with well-developed coecum; a strong, needle-like sclerotization fused to inner wall of aedeagus dorsally or ventrally, resembling a cornutus; vesica not eversible. Female genitalia (Figs 328–335) Sternum VII forming a sclerotized plate and partly or completely fused to sternum VIII. Sclerotized tergum VIII connected with sternum VIII by a wellsclerotized bridge that not only forms a lateral ring but is sclerotized medially, forming an abdominal end-plate ventrad of the papillae anales. Praebursa absent. Papillae anales small, apophyses posteriores long and slender. Apophyses anteriores absent. Ductus bursae mainly small, slender, translucent, folded, sometimes with small lateral appendix or with a large lateral praebursa-like structure (in H. furcata sp. n. only). Antrum short, slightly sclerotized or larger and well sclerotized, large and funnel-shaped in one species (H. furcata sp. n.). Corpus bursae small and translucent. Glandula sebacea without a bulla at the junction of the glandular part (present in most Procridinae).

Phenology and bionomics Adult Hestiochora species have been observed by various lepidopterists. They fly in bright sunshine, are weak flyers and visit flowers to take nectar (McFarland 1979: 62). According to the available label data, most species seem to have two generations per year: one in spring and another in late summer. This might be different for the more northern H. xanthocoma. Meyrick (1886: 790) mentions Angophora sp. (Myrtaceae) as a preferred nectar-plant for species of the H. erythrota-tricolorgroup. McFarland (unpublished field notes) observed males and females of H. continentalis sp. n. visiting the white flowers of Eucalyptus fasciculosa F. v. M. in southern Australia. He also observed that both males and females rest on leaves and stems of their larval foodplants and make only short flights between frequent landings. Eggs were obtained from a freshly captured female that oviposited on the wall of the jar without any foodplant present. There was no sun required for oviposition. The yellow eggs were laid in flat batches of 15–20 eggs. The larval foodplants are Myrtaceae, especially small mallee eucalypts (e.g. Eucalyptus fasciculosa F. v. M., E. odorata Behr, E. gittinsii Brooker & Blaxell) but also Syncarpia sp. (see also Common 1970: 830) and Buckinghamia celsissima F. Muell. (Proteaceae). The larvae feed freely on the leaves making unmistakeable feeding-tracks on their surface. These are groove-like and turn dark brown after a certain amount of time. McFarland observed dormant larvae freely resting on the leaves of E. fasciculosa in winter. Pupation takes place in a silky, white cocoon that is hidden under the bark of the foodplants. The pupa is stout, slightly compressed dorsoventrally, light brown, with the appendages only slightly attached to the body.

Remarks In preparing the male genitalia it is recommended that at first a cut is made at the connection between the sacculus and the vinculum, then a second cut to the strong medial connection of the ventral part of the valva. After this the connection between the sclerotized band of the ventral diaphragm (juxta) and the valva should be cut. At least one valva should be completely separated. The rest of the genitalia can be easily mounted on a slide without deformation of the vinculum and sacculus. All characters can be seen perfectly on the separated valva.

TAXONOMY

Distribution

8.

Hestiochora occurs in all Australian states. –

Key to Hestiochora species 1. – 2. –

With white scales on wings and/or body . . . . 2 Without white scales . . . . . . . . . . . . . . . . . . . . . 8 Male (antenna bipectinate, frenulum represented by one single spine) . . . . . . . . . . . . . 3 Female (antenna biserrate, frenulum represented by three small bristles). . . . . . . . . . . 9

Male 3. Valva divided into two long processes ventrally, the dorsal one forming a long, slender, downwards curved finger; aedeagus strongly curved upwards distally (Figs 297, 298, 299, 317) . . . . . . . . . . . . H. furcata sp. n. (p. 148) – Valva not strongly divided . . . . . . . . . . . . . . . . 4 4. Valva with a short process ventrally, aedeagus with tooth-like sclerotization distally . . . . 5 – Valva without ventral process (Figs 300–309) 6 5. Ventral sclerotization of valva slender, more than 5× longer than broad, curved upwards distally, with broad, triangular, pointed ending and with a very short process ventrally; aedeagus with a short, tooth-like lateral sclerotization distally pointing sideways or downwards (Figs 284, 285–287, 311, 312). From Tasmania or Victoria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. tricolor (p. 144) – Ventral sclerotization of valva broader, less than 4× longer than broad, with short, slender, upwards pointing tooth distally and with a long process ventrally; aedeagus with a slender, tooth-like, ventral, upwards pointing sclerotization distally (Figs 288, 289–296, 313–316) . . . H. continentalis sp. n. (p. 145) 6. From Western Australia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. occidentalis sp. n. (p. 151) – From Queensland or New South Wales . . . . . 7 7. Frons yellow, vertex orange. Valva with short, curved dorsal tooth on ventro-distal sclerotization; aedeagus straight, with very long, straight, slender coecum (Figs 309, 326, 327) . . . . . . . . . . . . . H. intermixta sp. n. (p. 152) – Frons and vertex shining unicolorous black (almost bluish black). Valva without dorsal tooth on ventro-distal sclerotization of valva, the latter narrow, long and bent upwards distally; aedeagus straight, with slightly upwards curved coecum (Figs 300, 301, 318–320) . H. queenslandensis sp. n. (p. 149)

Forewing with yellow stripes (Pl. 41) . . . . . . . . . . . . . . . . . . . . . . . . . . H. xanthocoma (p. 150) Forewing with yellow distal spot (Pl. 35) . . . . . . . . . . . . . . . . . . . . . . . . H. erythrota (p. 143)

Female 9. Sterna VII+VIII fused to a large, funnel-shaped antrum, ductus bursae with large, translucent, strongly folded, lateral, praebursa-like enlargement (Fig. 331) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. furcata sp. n. (p. 148) – Without large, funnel-shaped antrum and lateral praebursa-like enlargement of ductus bursae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 10. Antrum very pronounced, slightly asymmetrical, ductus bursae curved sidewards proximally (Figs 329, 330) . . . . . . . . . . . . . . . . . 11 – Antrum not very pronounced, ductus bursae almost straight . . . . . . . . . . . . . . . . . . . . . . 12 11. Sternum VII plate-like but not fused with sternum VIII to form a drop-shaped plate (Fig. 329). From Tasmania or Victoria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. tricolor (p. 144) – Sterna VII+VIII fused to form a strongly sclerotized, drop-shaped plate (Fig. 330) . . . . . . . . . . . . . . . . . . . H. continentalis sp. n. (p. 145) 12. From Western Australia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .H. occidentalis sp. n. (p. 151) – From Queensland or New South Wales . . . . 13 13. Ostium and antrum unsclerotized (Fig. 335) . . . . . . . . . . . . . . . H. intermixta sp. n. (p. 152) – Ostium and antrum slightly sclerotized (Fig. 332) . . . . H. queenslandensis sp. n. (p. 149)

‘Hestiochora tricolor auctt.’ – a complicated species-complex Hestiochora tricolor auctt. represents a hitherto unrecognised complex of species. Of the eight species recognised at present in the genus Hestiochora, six are included in this complex, having reliable and constant genitalic differences. Although distinctly different in colour and wing-pattern from other Australian zygaenids, these six species are externally very similar, with a striking, black and white wasp-like habitus (Pls 36–40, 42, 43). It is impossible to recognise any of them with certainty, based only on external characters. One species, Hestiochora tricolor, can be recognised by its distribution since it is the only species of the group that occurs in Tasmania. On the Australian mainland, the species of H. tricolor auctt. are partly sympatric and can only be recognised by genitalic characters.

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It is an interesting question as to why the complicated wasp-like pattern of these adults is so constant. Based on genitalic characters, some of these black and white wasp-like species seem to be more closely related to the two yellow species of the genus (i.e. H. erythrota and H. xanthocoma, that do not show this peculiar pattern), than to each other. This means that the wasp-like Hestiochora intermixta sp. n. and H. occidentalis sp. n., although significantly different in pattern and colour, are closer to the yellow H. xanthocoma than to the externally almost identical taxa H. tricolor, H. continentalis sp. n. and H. furcata sp. n. If a species in Western Australia has remarkably different genitalic characters compared with a species from Tasmania or Queensland, but has the same conspicuous external appearance, we have to assume either a functional correlation creating a striking convergence, or that this pattern has evolved early and is a synapomorphy for all the species with this black and white wasp-like habitus. In this case, we have to assume that the two yellow species, H. xanthocoma and H. erythrota, are derived from species with tricolorhabitus. But this does not concur with genitalic characters. According to genitalia morphology we have to assume that the burnet-like yellow Hestiochora erythrota is the most primitive existing Hestiochora. It has female genitalia with a simple but well developed ductus bursae, a large corpus bursae, and ostium and antrum that lack complicated structures (Fig. 328). The male genitalia are of typical Hestiochora type (see p. 140), the ventro-distal part of the valva is strongly sclerotized to the distal end, the dorsal and ventral margins of this sclerotization are almost parallel with a short, upwardly pointing tooth-like structure distally and ventral process are lacking; the aedeagus is straight, with a very short, tooth-like dorsal sclerotization distally and with a straight coecum (Figs 283, 310). This is supposed to represent a more ancestral character combination. The black and white wasp-like H. tricolor is very similar in male genitalia. The ventro-distal part of the valva has become slightly narrower, the dorsal and ventral magins are still parallel, the distal end is slightly extended and bent upwards to a greater extent with the tooth-like sclerotization at its tip more pronounced; furthermore, a short ventral process is developed. The aedeagus is no longer straight but slightly bent upwards distally, with the distal tooth larger than in H. erythrota (Figs 285–287, 311, 312). The female genitalia of the two species, although remarkably different, have at least the following characters in common: the ostium

and antrum are of medium breadth and of similar shape (i.e. slightly twisted), and sternite VII is not fused with the sternite VIII (as in H. continentalis sp. n. and H. furcata sp. n.) (Figs 328, 329). It can be easily surmised that H. continentalis sp. n. is a further evolutionary step to a more complicated genitalic structure (i.e. shorter and broader ventro-distal part of valva with a longer tooth distally and with a prominent ventral process and the aedeagus more curved distally with longer distal tooth) (Figs 288, 289–296, 313–316). The homologies are easily recognizable. Finally, H. furcata sp. n. is a last step to a very complicated structure combination. But even here the homologies are clear, at least in the male (Figs 297, 298, 299, 317). If we accept the arguments presented above, we still have the problem that there is a second group of black and white wasp-like species with exactly the same habitus as H. tricolor, H. continentals and H. furcata sp. n. but with significantly different genitalic structures. The black and white wasp-like H. intermixta sp. n. and H. occidentalis sp. n. have very similar male and female genitalia and are clearly closely related to the yellow H. xanthocoma that also has very similar genitalia. Once again it seems that the yellow species is the most primitive. Hestiochora xanthocoma has a somewhat triangular ventro-distal sclerotization on the valva with an inwardly bent, short, dorsal tooth and distally tapering aedeagus with a strong, straight cornutus that is fused to the dorsal wall of the distal part of the aedeagus (Figs 302–304, 321–323). The female has a characteristic end-plate at its abdominal end, with the ostium, ductus bursae and corpus bursae being very small and unsclerotized (Fig. 333). The conclusions are therefore: 1. Hestiochora is divided into three speciesgroups, the H. erythrota-tricolor-group (H. erythrota, H. tricolor, H. continentalis sp. n., H. furcata sp. n.), the H. xanthocomaoccidentalis-group (H. xanthocoma, H. occidentalis sp. n., H. intermixta sp. n.) and the isolated H. queenslandensis sp. n., with characters of both groups (i.e. valva like H. erythrota-tricolor-group, aedeagus like H. xanthocoma-occidentalis-group and wasp-like habitus but the white colour tinged with yellow). 2. The yellow species (i.e. H. erythrota and H. xanthocoma) are more primitive than the wasp-like black and white species. 3. The wasp-like black and white habitus must have been derived more than once from yellow ancestors.

TAXONOMY

4. There must be a strong selection pressure in favour of the characteristic black and white wasp-like ‘Hestiochora tricolor habitus’. Several braconid wasps (Common 1970) or even horse flies (e.g. Scapha auriflua (Donovan)) show this typical black and white abdomen and have similar red markings on the collar and abdominal tip. It is a fascinating task for the future to elucidate these relationships.

The ‘Hestiochora tricolor habitus’ – external appearance wasp-like. Male. Head black except labial palps, scales caudad of compound eye and posterior part of chaetosemata that are bright red and concolorous with the red prothoracic collar (the red is of variable shades); frons black except for a small white margin ventrally, sometimes with a few red scales (except for H. intermixta sp. n. that always has a yellow frons and orange or red vertex); proboscis brownish yellow; labial palpi short; compound eye small, black, breadth of frons 2.2× that of compound eye in frontal view; antenna shining brownish black to blackish, antennal shaft sometimes with a narrow white or yellow stripe caudo-laterally, segments 1–35(–38) bipectinate, 36(–39)–48(–50) biserrate, pointed distally. Thorax black with a blue sheen, with two white spots ventro-laterally below the wings; tegulae with a touch of blue and a white posterior margin in fresh specimens; legs shining brownish black, coxa of foreleg white, femur and tibia ochreous laterally, hind legs unicolorous. Forewing dark grey-brown to blackish with three variable translucent stripes between Sc and proximal part of cell, in proximal part of cell and in proximal part between cell and CuA2; in these translucent areas white or yellow scales are present in fresh specimens; underside of forewing with white scales also between CuA2 and anal vein. Hindwing brownish black distally, translucent with white or yellow scales proximally, anal fold white, densely scaled. Abdomen black, with dense white or yellow scales on segments 1–3 and 7 dorsally, 1–6 laterally and 1–6(–7) ventrally. Female. Similar to male but slightly larger and wings darker, with blackish brown abdominal hairtuft. Head often deep yellow to orange-red dorsally. Antenna biserrate.

Remarks All specimens mentioned below under ‘Material examined’ have been determined by examination of

the genitalia, based mainly on dissected material. Some species can be recognised by external genitalic characters (e.g. H. furcata sp. n., H. continentalis sp. n. 乆) and some of these have not been dissected.

Hestiochora erythrota-tricolor-group Four species, H. erythrota, H. tricolor, H. continentalis sp. n. and H. furcata sp. n. form this group that is distinguished on genitalic characters.

Hestiochora erythrota Meyrick Pl. 35, Figs 1, 2; Figs 24–28, 283, 310, 328. Hestiochora erythrota Meyrick, 1886, Proc. Linn. Soc. NSW (2)1: 789. Lectotype 么 (here designated), AUSTRALIA: New South Wales, Goulburn (ANIC, Macleay Museum) [examined]. Hestiochora erythrota Meyrick; Jordan 1907: 10; Hering 1922: 13; Turner 1926b: 441; Bryk 1936: 116; Tarmann 1996: 143.

Material examined Queensland: 9 么, 4 乆, Brisbane (SAM); 2 么, Brisbane (R. Illidge) (ANIC, SAM); 1 乆, Brisbane, v.1912 (T. P. Lucas) (SAM); 1 么, Brisbane, Acacia Ridge, 14.iii.1964 (E. C. Dahms) (QM); 1 么, Cedar Creek, 30.iii.1964 (J. A. Woods) (DEUQ); 1 乆, Killarney, 11.i.1933 (E. J. Dumigan) (DEUQ); 1 么, Lake Broadwater, 19.xii.1985 (QDPI); 19 么, 8 乆, Milmerran, 18.xii.–6.i.1927–49 (J. Macqueen) (ANIC, DEUQ); 1 么, 1 乆, Mt Lindsay Gate, 3.iii.1947 (E. J. Dumigan) (DEUQ); 1 乆, Samford, 2.x.1920 (ANIC); 1 么, Stanthorpe, 26.iii.1929 (W. B. Barnard) (QM); 1 么, 1 乆, Toowoomba, 23.xii.1961, 25.iii.1971 (J. Macqueen) (ANIC). New South Wales: 1 么 Arrawarra, 9.xii.1961 (C. W. Frazier) (ANIC); 6 么, Barralier, 7.i.1964 (V. J. Robinson) (ANIC, AMS); 2 么, Berowra, 27.–30.x.1917 (ANIC, MV); 1 乆, Dangar’s Falls, Armidale, 5.xii.1940 (C. W. Frazier) (ANIC); 1 乆, Dorrigo, 24.xi.1911 (R. J. T[illyard]) (MV); 1 么, Goulburn (ANIC); 1 乆, Herberton, i.1911 (F. P. D[odd]) (MV); 1 么, Mt Kunnagai, 13.x.1935 (A. Musgrave) (AMS); 2 么, 1 乆, Mulgoa, 2.i.1933 (L. H. Mosse-Robinson) (ANIC); 1 乆, Narara, 20.ix.13.x.1947–48, ii.1947 (L. H. Mosse-Robinson) (ANIC); 1 么, Richmond R., 7.v.1935 (V. J. Robinson) (ANIC); 1 乆, Rooty Hill, 7.v.1978 (J. L. Lawson) (ANIC); 1 么, 1 乆, Sydney, 12.x.1924 (AMS); 1 么, Tahmoor, 19.x.1979 (D. S. Kent) (AMS); 1 么, Wilton, CSIRO Experimental Farm., 1.xii.1973 (V. J. Robinson) (ANIC); 1 么, Woolgoolga, 24.iv.1966 (V. J. Robinson) (ANIC).

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Diagnosis


This species is unmistakeable since it is the only Australian zygaenid with an almost burnet-like appearance (i.e. pattern with stripes and spots).

According to the label data this species seems to have two generations. Nothing is known about the early instars or the foodplant of the larva.

Redescription

Distribution

Male. (Pl. 35, Fig. 1). Forewing length 9.0–10.0 mm. Frons densely covered with blackish scales; proboscis yellow, labial palps short and yellow; chaetosemata dark brown; head covered with black scales except frons and a small lateroventral part behind the eye that varies from black to orange. Antenna black. Thorax entirely black with weak bluish sheen except for a yellow-orange collar. Abdomen with segments 1–3 deep yellow with orange tinge, segments 4–8 black, with bluish sheen, and segment 8 with a narrow orange ring anteriorly, end of abdomen black, with blue sheen. Forewing blackish, with a short yellow subcostal band proximally, a longer yellow stripe caudad of cell extending to middle of wing with a spot-like yellow-orange extension distally and a round yellow-orange mark covering end of cell. Hindwing yellow or orange proximally, with a broad blackish band distally. Fringe dark grey.

Hestiochora erythrota occurs in southern Queensland and the subtropical parts of New South Wales.

Female (Pl. 35, Fig. 2). Forewing length 9.0–10.0 mm. Similar to male in habitus and size. Antenna biserrate. Abdomen paler, abdominal hairtuft dark grey-brown. Male genitalia (Figs 283, 310). Uncus short and stout. Tegumen slender, vinculum broad, strongly sclerotized. Juxta lip-shaped with lateral elongations. Valva with very strongly sclerotized, large sacculus; ventral sclerotization extended, curved upwards and sharply pointed distally, without ventral process. Aedeagus straight and slender with a strong, distally sharply pointed and downwardcurved dorsal sclerotization. Sternum VIII covering approximately 15–20% of segment. Female genitalia (Fig. 328). Sternum VII a large sclerotized plate. Sternum VIII forming a small, strongly sclerotized plate around ostium. Tergum VIII small, helmet-shaped, connected to sternum VIII by a large, strongly sclerotized abdominal endplate. Ostium with a strongly sclerotized ring; antrum well developed, slightly asymmetrical; ductus bursae almost straight, relatively long (compared with other Hestiochora-species); corpus bursae large.

Hestiochora tricolor (Walker) Pl. 36, Figs 1, 2; Pl. 58, Fig. 5; Figs 284, 285–287, 311, 312, 329. Procris tricolor Walker, 1854, List Specimens lepid. Insects Colln Br. Mus. 1: 111. Lectotype 么 (here designated), AUSTRALIA: Tasmania, “Van Diemen’s Land” (BMNH) [examined]. Procris tricolor Walker; Butler 1877: 13. Hestiochora tricolor (Walker); Meyrick 1886: 789; Kirby 1892: 86; Jordan 1907: 10; Hering 1922: 13; Tillyard 1926: 435; Turner 1926b: 441; Bryk 1936: 116; Alberti 1954: 218; Common 1970: 830, pl. 6ZC; McFarland 1979: 62; Fletcher and Nye 1982: 76; Common 1990: 297; Tarmann 1996: 143.

Material examined [Australian Capital Territory: 1 么, Canberra, 25. xii. 1937 (G. M. Goldfinch), genitalia slide Z 2939 (AMS).] Victoria: 1 乆, Beech Forest, 11.-19. i. 1932 (J. Clark) (MV); 1 么, Gisborne (G. L[yell]), genitalia slide Z 2926 (SAM); 1 乆, Healesville, genitalia slide Z 2935 (MV). [South Australia: 1 么, Prospect, 17. xii. 1904, genitalia slide 2921 (SAM); 1 么, Russell, 3. i. 1916 (C. Cole), genitalia slide Z 2927 (SAM).] Tasmania: 1 么, Hobart (Lea) (ANIC); 1 么, Hobart, xii. 1890 (J. J. W[alker].) (BMNH); 1 乆, Hobart, 4. i. 1902 (MV); 3 么, 1 乆, Hobart, i. 1924 (G. H. Hardy) (ANIC, UQIC); 1 乆, Hobart, 29. ii. 1941 (ANIC); 1 乆, Kingston, 19. x. 1950 (J. R. Cunningham) (TMH); 1 么, 1 乆,

TAXONOMY

Launceston, 20. i. 1903 (AMS, MV); 1 么, Launceston 13. i. 1904 (G. H. Hardy) (AMS); 1 么, Mt Nelson, 200 m, 1. xi. 1979 (P. B. McQuillan) (ANIC); 1 么, Mangalore, 29. xi. 1914 (G. H. Hardy) (AMS); 1 乆, Moorina, 17. xi. 1975 (K. R. Norris) (ANIC);1 乆, Triabunna, 28. xii. 1915 (G. H. Hardy) (AMS).

cent appendix proximally (hardly visible); corpus bursae of medium size. Region between ostium and papillae anales transformed into a strongly sclerotized abdominal end-plate.

Phenology and bionomics Diagnosis Hestiochora tricolor is the only species of Hestiochora in Tasmania and is the largest species of the group. It is not possible to identify this species except by the locality (if from Tasmania) and the genitalia that are characteristic. Most specimens have veins R4+R5 in the forewing stalked. The scales are slightly narrower than in related species, giving the specimens a ‘rough’ appearance.

Redescription Male (Pl. 36, Fig. 1). Black and white wasp-like habitus (description see p. 143). Forewing length: 11–12 mm. Forewing with all veins free from cell except R4+R5 that are mainly shortly stalked or originating from one point. The scales of the wings are very slender. Female (Pl. 36, Fig. 2). Forewing length: 12–13 mm. Similar to male, slightly larger and darker. Male genitalia (Figs 284, 285–287, 311, 312). Uncus of medium length (approximately half length of tegumen). Tegumen long and slender. Vinculum slender, saccus small, almost absent. Ventral part of diaphragma with a tongue-shaped sclerotization with narrow lateral elongations that may be interpreted as a juxta. Ventral part of valva with very large sacculus proximally and a slender sclerotization distally, the latter with parallel dorsal and ventral margins, tapering towards apex, pointed and curved upwards apically, with a very short, distally rounded process ventrally. Aedeagus almost straight, only very slightly curved upwards, with a lateral, often downward-curved tooth-like sclerotization distally. Sternum VIII of abdomen narrow, covering only 20% of segment. Female genitalia (Fig. 329). Sterna VII and VIII separate, not fused. Sternum VII short and broad, band-like, broader posteriorly, sternum VIII a strongly sclerotized trapezoid plate with a central ring. Ostium relatively broad (compared with allied species); antrum pocket-like, asymmetrical; ductus bursae strongly folded, with a small lateral translu-

Nothing is known about the biology of H. tricolor as all published and unpublished records about habits, larva and foodplants come from the Australian mainland and may refer to other species.

Remarks Almost all material of this species originates from Tasmania. Three old records from continental Australia (southern Victoria) may be reliable but it is strange that no recent material from the Australian mainland is known. Two further records may be referable to localities in South Australia but they cannot be located clearly (many places with the same name are known in South Australia, Victoria and New South Wales). A single record from Canberra seems very doubtful. The occurrence of H. tricolor from continental Australia needs confirmation.

Distribution Tasmania, Victoria, [South Australia?, Australian Capital Territory?].

Hestiochora continentalis sp. n. Pl. 37, Figs 1, 2; Figs 288, 289–296, 313–316, 330.

Material examined Holotype 么. South Australia, Aldinga Scrub, e.l. 2. xii. 1970 (J. O. Wilson), genitalia slide Z 2872 (ANIC).

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Paratypes Queensland: 2 乆, Glen Aplin, 30. iii. 1940 (L. H. Mosse Robinson) (ANIC); 3 么, 1 乆, Milmerran, 24. iii. 1928, 3. xii. 1928, 26. xii. 1936, 17. x. 1945 (J. Macqueen) (ANIC, AMS); 1 乆 Stanthorpe, 8. i. 1929 (W. B. Barnard) (QM). New South Wales: 1 乆, Church Point, e.l. 17. xii. 1966, larva on Syncarpia sp. (I. F. B. Common) (ANIC); 1 么, 2 乆, Manly, 14. iv. 1906 (AMS); 1 么, Maryland, 21. ii. 1932 (W. B. Barnard), genitalia slide Z 2909 (ANIC); 2 么, 1 乆, Narara, 9. xii. 1928, 1. iv. 1934, 29. iii. 1939 (I. M. Mosse Robinson) (AMS, ANIC), Genitalia slide Z 2937 (ANIC); 1 么, Newcastle Firth, i. 1888 (AMS); 4 么, 1 乆, Niagara Park, 20. xi. 1948, xii. 1948, 1. ii. 1949 (W. Brandt) (ANIC); 1 乆, Stanwell Park, 15. iii. 1924 (AMS); 1 么, St George’s Basin, 24. xi. 1965 (V. J. Robinson) (ANIC); 2么, Sydney (Helms) (AMS, Bishop Museum, Honolulu); 2 乆, Sydney (W. W. Froggatt) (AMS); 1 乆, Sydney, Manly Reservoir, 3. iv. 1961 (D. H. Colless) (ANIC); 1 么, Sydney, 14. iii. 1918 (MV); 1 么, Sydney, Strathfield (BMNH); 1 么, Sydney, 29. xii. 1906 (W. B. G.) (DASS). Victoria: 1 么, 2 乆, ‘Victoria’ (ANIC, MV, QLMB); 2 么, Beaconsfield, 27. xii. 1904, 3. xii. 1908, genitalia slide Z 2936 (MV); 1 乆, Sale, 4. xii. 1919 (M. I. Wise) (MV). South Australia: 3 么, 2 乆, Adelaide (MNHU, SAM), genitalia slides Z 2924, Z 2931 (SAM); 1 么, Aldinga, 27. iii. 1971 (K. Sandery) (SAM); 3 么, 7 乆, Aldinga Scrub, 17. iii. 1970, e.l. 2.– 10. xii. 1970 (J. O. Wilson), genitalia slides Z 2878, Z 2920 (ANIC); 1 么, Halbury, xi. 1892, genitalia slide Z 2928 (SAM); 1 乆, Hoyleton, xi. 1892, genitalia slide Z 2963 (SAM); 1 么, Mt Hall, Eyre Peninsula, 6. xii. 1968 (N. McFarland) (SAM). Western Australia: 1 乆, Dryandra Forest (32.45 S / 115.58 E), 5. iv. 1984 (R. P. McMillan), genitalia slide Z 2834 (WAM); 1 乆, Eneabba, 15. x. 1985 (R. P. McMillan) (WADA); feeding marks of larvae on Eucalyptus gittinsii, Hawkshead Lookout, Kalbarri Nat. Park, 180 m, 6. ix. 1995 (B. E. & G. M. Tarmann) (TLMF); 1 么, Leederville, 14. ii. 1963 (J. Berg) (WADA); 1 乆, Nedlands, 24. iii. 1941 (K. R. Norris), genitalia slide Z 2930 (ANIC); 1 么, 1 乆, Perth, 1933 (G. H. Hardy) (AMS, ANIC); 1 乆, Swan R. (L. J. Newman), genitalia slide Z 2916 (ANIC).

Diagnosis Male genitalia similar to those of H. tricolor with which it is sympatric in Victoria. The ventral part of the valva is broader and shorter distally and the ventral process is longer than in H. tricolor. The sclerotization of the ventral diaphragma (‘juxta’) is band-like in H. continentalis and tongue-shaped in H. tricolor. The aedeagus in H. continentalis has a ventral sclerotization distally, while H. tricolor has a lateral one. There are also clear differences in the female genitalia: the drop-shaped plate of the fused sterna VII and VIII in H. continentalis can be easily distinguished from the band-like, separated, unfused sterna VII and VIII of H. tricolor, even externally by

brushing away the ventral abdominal scales. Males of H. furcata sp. n. (southern Queensland to South Australia) are also sympatric with H. continentalis but can be easily recognised by their furcate valva, which is visible by brushing off the ventral scales of the abdomen. Females of H. furcata sp. n. are darker, with almost unicolorous forewings, the large funnel-shaped antrum is visible externally or when the scales are removed by brushing the abdomen. Hestiochora continentalis is sympatric with H. queenslandensis sp. n. in southern Queensland and with H. intermixta sp. n. in southern Queensland and northern New South Wales. Both are more yellowish and slightly smaller. The genitalia are characteristic. In Western Australia H. continentalis is sympatric with H. occidentalis sp. n., but the latter is smaller. The genitalia are characteristic.

Description Male (Pl. 37, Fig. 1). Black and white wasp-like habitus (description see p. 143). Forewing length: 7.5–8.5 mm. Externally similar to H. tricolor and H. furcata sp. n., slightly smaller in size than the former but within the range of variability of the latter. Medial stem in forewing present or absent, all veins free from cell. Female (Pl. 37, Fig. 2). Forewing length: 8.5–10.0 mm. Similar to male and slightly larger; coloration similar or slightly darker. Male genitalia (Figs 288, 289–296, 313–316). Uncus of medium length (approximately half length of tegumen). Tegumen long and slender. Vinculum slender, saccus small, almost absent. A transverse, band-like structure replaces the juxta and/or anellus. Ventral part of valva with very large sacculus proximally and a slender sclerotization distally, the latter tapering towards apex, pointed and curved upwards apically, with a finger-like, distally rounded process ventrally. Aedeagus straight anteriorly, slightly curved upwards posteriorly, with a toothlike upward-pointing ventral sclerotization distally. Sternum VIII covering 20–40% of segment. Female genitalia (Fig. 330). Sterna VII and VIII fused to a strongly sclerotized drop-shaped plate, with the pointed end posteriorly. This plate can easily be recognised on the abdomen externally. Tergum VIII lip-shaped. Sternum VIII and tergum VIII connected by a very pronounced sclerotized plate with longitu-

TAXONOMY

dinal ribs. Ostium situated at posterior end of the drop-shaped ventral plate; antrum short, heartshaped, slightly asymmetrical; ductus bursae with a small lateral translucent appendix proximally, strongly folded; corpus bursae of medium size.

Phenology and bionomics According to the label data, H. continentalis is at least partly bivoltine. The data given by McFarland (1979: 62) for H. tricolor from Aldinga Scrub in South Australia are referable to this species (original material examined). In the original hand-written fieldnotes of Noel McFarland’s visit (in SAM) to this area he records exact details on the habits of the adults as well as oviposition, eggs, larvae, pupae, cocoons and foodplants. The following information is provided with the generous permission of Mr McFarland. The adults were locally abundant on 14 March 1970 at a locality three miles south of Port Willunga, at the northern edge of ‘Aldinga Scrub’, South Australia. They were on the wing in the early afternoon between 1.00 and 3.00 p.m., or resting on leaves and stems of the mallee eucalypts of the area, mainly on Eucalyptus fasciculosa F. v. M. It was a sunny, dry day. Some males and females were observed visiting the white flowers of that eucalypt species. They look like braconid or ichneumonid wasps when on the wing. One of the females began ovipositing on the side of a small jar within minutes after capture and confinement. After a few hours it had deposited 60 eggs in groups of 15–20, evenly spaced, not touching each other. The eggs were lightly ‘fur-coated’ by the hairs of the female abdomen, the hairs bristling out in all directions, mostly around the edge of the egg, with the top more or less bare (see also p. 36 and Figs 129, 130). The colour of the eggs was at first yellow, later dull yellowish cream and pale grey before hatching. The larvae hatched after 10 days. The small larvae began feeding on E. fasciculosa almost immediately. The growth of the larvae was rapid. They made very charactersistic ‘feedingtracks’ on the surface of the leaf. These were groovelike and turned deep brown after a few days. The locomotion of the larva was rapidly lumbering. They were easily disturbed into activity. All larvae were fully grown by late May and pupated in a typical white zygaenid cocoon. In winter, on 28 July 1970 the locality was revisited. Dormant larvae (fifth instar) were found sitting exposed on the surface of the leaves. The old feedinggrooves had turned brown and were clearly visible. As the rearing of the larvae under artificial (warmer)

conditions led to a generation in May, and according to McFarland’s opinion the dormant larvae were clearly from the same females (observed in March), we have to assume that under natural conditions they pupate in September and that there is a spring generation in October and a summer generation in March. The species is therefore bivoltine in that locality. There is another short note by McFarland (1979: 62) that may refer to this species. He observed a single female on the wing at midday on 23 September 1974 at Hawkshead Lookout in the Kalbarri National Park, 23 miles [37 km] east of Kalbarri. Inspection of the vegetation showed characteristic groove-like feeding marks caused by larvae earlier in the year on leaves of small mallee eucalypts that are presumed to be the foodplant of this species. The two mentioned localities (Hawkshead Lookout in the Kalbarri National Park (Western Australia) and Aldinga Scrub (South Australia)) were revisited by G. M. and B. E. Tarmann on 6 September 1995 and 15 September 1995. At Hawkshead Lookout the above-mentioned feeding tracks could be found on leaves of Eucalyptus gittinsii Brooker & Blaxell, a medium-sized mallee eucalypt endemic to the western coast of Australia between Cataby and Kalbarri. The Aldinga Scrub locality has been almost entirely transformed into farmland or used for housing during the last 25 years and no evidence of the presence of larvae could be found. One single female was reared by I. F. B. Common from larvae that he had found at Church Point (New South Wales) on Syncarpia sp. (Myrtaceae) (see ‘Material examined’). One larva and several pupae from the same locality are preserved in alcohol in ANIC (Canberra). From that material it must be concluded that there is no major difference between the larvae of Pollanisus, Onceropyga and Hestiochora. Perhaps they are different in colour and pattern, but in the alcohol material all colours have been lost. All other structures are as described under Pollanisus (see p. 71) except for the lack of the second abdominal semi-eversible evagination in Hestiochora. The same can be said for the pupa. It should be noted that in alcohol the wings of the pupa become completely loose and it can be seen that the fore- and hindwing are accommodated in two different independent compartments.

Distribution From southern Queensland throughout New South Wales to Victoria and South Australia, also in Western Australia.

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(SAM); 1 乆, Moe, xii. 1945 (C. G. L. Gooding) (ANIC); 1 乆, Red Hill, 22. xii. 1951 (Holmes) (ANIC). South Australia: 2 么, ‘S. Australia’ (BMNH); 1 乆, Aldgate, 5. v. 1969 (A. Smith), genitalia slide Z 2918 (ANIC).

Diagnosis

Hestiochora furcata sp. n. Pl. 38, Figs 1, 2; Pl. 39, Figs 1, 2; Figs 297, 298, 299, 317, 331.

Material examined Holotype 么, New South Wales, Wilton, CSIRO Experimental Farm, 23. ii. 1974 (V. J. Robinson), genitalia slide Z 2493 (ANIC). Paratypes Queensland: 1 乆, Mooloolaba, xi. 1890 (C. J. W[ilde]) (QLMB). New South Wales: 1 么, 3 乆, Armidale, 14. i. 1957, 21. i. 1960 (C. W. Frazier) (ANIC); 2 乆, Bald Rock National Park, xii. 1982 (G. Daniels) (UQIC); 1 乆, Bendalong, 8. i. 1969 (G. Daniels) (AMS); 2 么, 2 乆, Ebor, 9. xii. 1936, 1. xii. 1937, 14. xii. 1939, 1. xii. 1951 (AMS); 1 么, French’s Forest, 22. ii. 1936 (D. F. Waterhouse) (ANIC); 1 乆, Mosman, 13. iii. 1910 (MV); 1 么, Narara, 10. ii. 1948 (L. H. Mosse Robinson), genitalia slide Z 2905 (ANIC); 1 么, North Durras, 8 mls W Bateman’s Bay, 9. iii. 1969 (R. J. Bartell) (ANIC); 1 乆, Saw Pit Creek, 2.5 km ESE of Kosciuszko National Park Headquarters, 990 m, 6. i. 1988 (E. D. Edwards) (ANIC); 1 么 Stanwell Park, 10. iii. 1952 (L. H. Mosse Robinson) (ANIC); 1 么, Watagan St. Fo. [State Forest], nr Wyong, 20. ii. 1984 (B. Hacobian) (ANIC); 2 么, 1 乆, Wilton, CSIRO Experimental Farm, 25. ii. 1969, 30. x. 1972, 19. xii. 1972 (V. J. Robinson), genitalia slide Z 2929 (ANIC). Australian Capital Territory: 3 么, 1 乆, Black Mountain Light Trap, 16. xii. 1955, 27. xi. 1959, 12. xii. 1960, 24. i. 1962 (I. F. B. Common) (ANIC); 1 乆, Canberra, 25. xii. 1937 (G. M. Goldfinch) (AMS). Victoria: 1 么, ‘Victoria’, i. 1931 (A. Musgrave) (AMS); 3 么, Buln Buln, 16. xii. 1962 (C. G. L. Gooding) (ANIC); 1 么, Drouin, 15. xi. 1958 (C. G. L. Gooding) (ANIC); 1 么, Emerald, 28. x. 1907 (MV); 1 么, 1 乆, Gisborne (SAM); 4 么, 2 乆, Gisborne, 18. xii. 1892, 12. xii. 1915, 1. i. 1926, 25. xii. 1932 (G. Lyell) (MV); 2 么, 4 乆, Gisborne, 9. ii. 1896, 24. xi. 1901, 1. xii. 1903, 20. xi. 1904, 7.–12. i. 1906 (MV); 3 么, Healesville (MV); 1 么, 1 乆, Kiata, 14.–15. xi. 1945 (AB), genitalia slide Z 2934 (MV); 1 么, Melbourne

Hestiochora furcata is sympatric with H. continentalis in southern Queensland, New South Wales, Victoria and South Australia and with H. queenslandensis sp. n. in southern Queensland and with H. intermixta sp. n. in southern Queensland and northern New South Wales. The males of H. furcata and H. continentalis are externally similar, those of H. queenslandensis sp. n. and H. intermixta sp. n. are usually more yellowish, especially on the forewings. The female of H. furcata is very dark, almost without translucent parts, whereas those of the other three species have stripes on the forewings and a significantly larger translucent basal part on the hindwing. Hestiochora furcata is the only wasp-like species of the H. erythrota-tricolor-group that can be easily determined by the genitalia in both sexes by brushing off the scales at the end of the abdomen; then the long processes of the valva in the male and the large funnel-shaped antrum in the female are clearly visible.

Description Male (Pl. 38, Figs 1, 2; Pl. 39, Fig. 1). Black and white wasp-like habitus (description see p. 143). Forewing length: 7.5–11.0 mm. Externally similar to H. tricolor and H. continentalis. Very variable in size. Forewing with all veins free from cell. Female (Pl. 39, Fig. 2). Forewing length: 8.0–12.5 mm. Slightly larger and significantly darker than male. Specimens with almost unicolorous dark brown or blackish brown forewings and blackish brown, almost opaque, only slightly translucent hindwings with a white spot basally are common. Male genitalia (Figs 297, 298, 299, 317). Uncus short and slender (approximately one-third length of tegumen). Tegumen long and slender. Vinculum also slender, without saccus. Juxta ring-like, somewhat triangular, with broad sclerotization ventrally. Ventral part of valva with large sacculus proximally, with a very long characteristic ventral process and a strongly sclerotized, long, hook-like, downwardcurved dorsal process distally; translucent medial part of valva narrow, dorsal sclerotization broader

TAXONOMY

proximally but very narrow distally. Aedeagus slightly curved downwards anteriorly but strongly curved upwards posteriorly, with a strong, distally pointed, dorsal sclerotization that protrudes beyond the distal end of the aedeagus. Sclerotized part of sternum VIII reduced to a small band at anterior margin of segment, covering only 5–10% of segment. Female genitalia (Fig. 331). Sterna VII and VIII fused to a very characteristic, strongly sclerotized, broad, funnel-shaped structure that is fused with the lip-shaped tergum VIII by a slender sclerotization; the latter is significantly constricted laterally ventrad of the papillae anales. This funnel-shaped structure may be interpreted as an enlarged antrum. Ductus bursae narrow proximally for a very short distance but transformed into a very large, translucent, strongly folded, praebursa-like enlargement distally, which is located slightly laterally and has two small, plate-like sclerotizations like signa; the most distal part of ductus bursae again narrow; corpus bursae relatively large, very translucent and fragile.

Phenology and bionomics Nothing is known about the biology. The label data indicate that this species is possibly univoltine. The moths are on the wing in midsummer between November and February.

Hestiochora queenslandensis-group Currently the queenslandensis group consists of only one species.

Hestiochora queenslandensis sp. n. Pl. 40, Figs 1, 2; Figs 300, 301, 318–320, 332.

Material examined Holotype 么, Queensland: Milmerran, 12. ii. 1944 (J. Macqueen), Genitalia slide Z 2844 (ANIC).

The praebursa-like structure of the central part of the ductus bursae is, in my opinion, not homologous with the real praebursa in other genera (e.g. Pollanisus). It seems to be only a lateral enlargement of the ductus bursae. Hestiochora continentalis and also (almost invisible) H. tricolor have a characteristic lateral appendix of the ductus bursae in the same position.

Paratypes Queensland: 1 乆, Brisbane (R. Midge) (ANIC); 1 乆, Brisbane, Moggill, 27. iii. 1937 (C. Burwell) (UQIC); 1 乆, Kenmore, 16. i. 1958 (R. Kleinschmidt), genitalia slide Z 2941 (DPIQ); 1 乆, Miles, iii. 1907 (ANIC); 4 么, 3 乆, Milmerran, 2. i. 1944, 15. xii. 1945, 16. xii. 1945, 14. xii. 1947 (J. Macqueen), genitalia slide Z 2889 (AMS, ANIC, UQIC); 1 么, Scottsville, 16. ix. 1950 (E. F. Riek), genitalia slide Z 2848 (ANIC); 1 么, Tansey, Planted Ck, 12. xii. 1976 (S. R. Monteith) (ANIC); 1 么, Thruston Park, Mulga, 17–18 x. 1985 (R. Harmsen) (AMS). New South Wales: 1 么, Round Hill Fauna Reserve, 17. ii. 1968 (V. J. Robinson) (ANIC).

Distribution

Diagnosis

Remarks

From southern Queensland throughout New South Wales to Victoria and South Australia.

Together with H. intermixta sp. n. it is the most ‘yellow’ of the six species with black and white wasp-like habitus. Sympatric with H. intermixta sp. n., H. continentalis and H. furcata in southern Queensland and northern New South Wales. Hestiochora intermixta sp. n. is very similar externally but the male has a yellow frons and an orange vertex. The females are externally similar. The two other species are more whitish, mainly larger, and the female of H. furcata is almost uniform dark blackish brown on the forewing upperside. The genitalia are different from those of all sympatric species.

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Description Male (Pl. 40, Fig. 1). Black and white wasp-like habitus (description see p. 143). Forewing length: 7.5–9.0 mm. The translucent stripes on the wings often covered with yellow scales, sometimes mixed with white, rarely completely translucent. R4 and R5 of forewing free from cell. Female (Pl. 40, Fig. 2). Forewing length: 8.5–10.0 mm. Similar to male but darker and slightly larger. Male genitalia (Figs 300, 301, 318–320). Uncus long (almost of same length as tegumen). Tegumen and vinculum slender, saccus small, rounded, well pronounced. Valva without process ventrally but with a deep emargination between sacculus and ventro-distal sclerotization, the latter narrow, strongly curved upwards distally and pointed apically; translucent medial part of valva broad; dorsal sclerotization very narrow. Aedeagus straight, tapering distally, slightly curved upwards proximally, with long, apically pointed dorsal sclerotization. Sternum VIII covering 20% of segment. Female genitalia (Fig. 332). Sternum VII only covering half of segment. Sternum VIII transformed into a plate that extends towards papillae anales. Tergum VIII characteristic, somewhat V-shaped, with two narrow, parallel sclerotizations medially. Ostium small, sclerotized, antrum funnel-shaped, ductus bursae S-shaped proximally, almost straight distally.

Phenology and bionomics Unknown. According to the label data the species is at least bivoltine.

Distribution

Hestiochora xanthocoma-occidentalis-group

Hestiochora xanthocoma Meyrick Pl. 41, Figs 1, 2; Pl. 58, Fig. 6; Figs 302–304, 321–323, 333. Hestiochora xanthocoma Meyrick, 1886, Proc. Linn. Soc. NSW (2) 1: 788. Holotype 么, AUSTRALIA: Queensland, Duaringa 1886 (G. Barnard) (BMNH) [examined]. Hestiochora xanthosoma Kirby, 1892, Synonymic Cat. Lepid. Heterocera 1: 86 [misspelling]. Hestiochora xanthocoma Meyrick; Jordan 1907: 10; Hering 1922: 13; Turner 1926b: 441; Bryk 1936: 116; Alberti 1954: 218; Tarmann 1996: 143.

Material examined It is possible that not all specimens that are mentioned here under H. xanthocoma really are conspecific and that the taxon H. xanthocoma consists in fact of a group of closely related species. At the moment there is not sufficient material available to solve this problem. Therefore all specimens with ‘H. xanthocoma’ habitus are treated as one species in this book. Northern Territory: 1 么, Darwin (H. Wesselman) (SAM); 1 么, Groote Eylandt (N. B. Tindale) (SAM); 1 么, Howard Springs, 5. vi. 1973 (E. D. Edwards) (ANIC); 1 么, Jarrnarm, 8 km NE, Keep R. Nat. Park, 9. vi. 2001 (M. Horak) (ANIC); 1 乆, Katherine, 17. iv. 1962 (I. F. B. Common) (ANIC); 1 么, Port Darwin, 4. x. 1908 (F. P. Dodd) (ANIC). Queensland: 1 乆, ‘Queensland’ (MV); 1 乆, Carnarvon Ra., 12. i. 1940 (W. B. Barnard) (QM); 1 么, Biloela, 2. xii. 1946 (I. F. B. Common) (ANIC); 1 乆, Heifer Creek Rd, 2. i. 1965 (A. Macqueen) (UQIC); 1 乆, Goondiwindi, 24 km NE, 28. xi. 1976, on Atalaya hemiglauca (E. M. Exley & T. Low) (UQIC); 2 么, Jowalbinna Homestead, 6.7 km W, 10. v. 1989 (G. & A. Daniels) (UQIC); 3 么, Rosedale, 27. x. 1974 (H. Frauca) (ANIC); 1 么, Wacol, 16. ii. 1950 (UQIC); 1 么, Westwood, 1. ii. 1925 (A. N. B[urns]) (MV).

South-eastern Queensland and northern New South Wales.

Diagnosis Hestiochora xanthocoma has an unmistakeable colour and pattern. It is remarkable that the genitalia are very similar in the male, and almost identical in the female, to those of H. occidentalis sp. n. and H. intermixta sp. n., the former being endemic to Western Australia and the latter only known from a few specimens collected in southern Queensland and northern New South Wales, both with black and white wasp-like habitus.

TAXONOMY

Redescription Male (Pl. 41, Fig. 1). Forewing length: 7.0–8.0 mm. Head with frons partly or completely covered with yellow scales, vertex usually bright yellow to orange but also blackish in some specimens; labial palps yellow; compound eye black, ocelli white, the scales caudad of compound eye yellow; chaetosemata dark brown anteriorly, orange posteriorly; antenna blackish brown, sometimes with orange scales on distal end of pectinations, 49–50 segments. Thorax black, with a blue sheen and an orange collar; legs blackish brown, femur with orange scales dorsally, tibia and tarsi with yellow scales ventrally. Forewing blackish brown with yellow longitudinal stripes, hindwing yellow, blackish around margin, except at anal angle, with all veins free from cell. Abdomen dorsally with segments 1–3 dull yellow, 4–6 black with blue sheen, and segment 7 black, with a narrow yellow band, completely yellow ventrally, end of abdomen black. Female (Pl. 41, Fig. 2). Forewing length: 8.0–8.5 mm. Similar to male but slightly larger, with collar deeply orange. Forewing and sometimes hindwing upperside with a deep yellow spot of variable size at anal angle (sometimes absent). Abdomen and abdominal hair-tuft deep dark yellow dorsally, margins of last few segments and tip of abdominal hairtuft edged with blackish scales, abdomen lighter yellow ventrally. Male genitalia (Figs 302–304, 321–323). Uncus long (almost as long as tegumen). Tegumen of medium length and breadth. Vinculum slender, saccus small, rounded. Ventral part of diaphragma translucent, slightly folded, the fold holding the aedeagus, juxta absent. Ventral part of valva with very pronounced sacculus that is smaller and less sclerotized than in most other species of the genus; the ventral sclerotization of valva broad with a slightly pointed apex distally and a tooth-like dorsal process; distad of sacculus with deep emargination; unsclerotized medial part of valva triangular, dorsal sclerotization narrow. Aedeagus curved anteriorly, straight posteriorly, strongly tapering towards its posterior end and pointed apically, with a short, strongly sclerotized dorsal spine. Sternum VIII covering 20–30% of segment. Female genitalia (Fig. 333). Sternum VII translucent. Sternum VIII forming a strongly sclerotized half-moon-shaped plate around the unsclerotized ostium. Tergum VIII large and strongly sclerotized,

connected with sternum VIII by two slender, transverse sclerotizations. Ostium small, antrum absent, ductus bursae slightly curved.

Phenology and bionomics Unknown. A tropical species with possibly several generations a year.

Distribution Only known from the tropical parts of Australia: Northern Territory and Queensland.

Hestiochora occidentalis sp. n. Pl. 42, Figs 1, 2; Figs 305–307, 324, 325, 334.

Material examined Holotype 么. Western Australia, Mt Dale, 21. I. 1926 (W. B. Barnard), Genitalia slide Z 2792 (ANIC). Paratypes Western Australia: 1 乆, Katanning, 10 km N, 22. xii. 1977 (R. P. McMillan), genitalia slide Z 2902 (WAM); 1 乆, Margaret R., 10 mls S, 15. i. 1971 (G. A. Holloway & H. Hughes), genitalia slide Z 2919 (AMS); 1 么, Moir’s Rock, Salmon Gums, 42 km NNW, 2. i. 1986, at mv lamp (G. & A. Daniels), genitalia slide Z 2946 (UQIC); 4 么, 4 乆, Mt Dale, 21–23. i. 1926 (W. B. Barnard), genitalia slides Z 2881, Z 2932 (QM, WADA); 1 乆, Mt Dale, 23. i. 1926 (WADA); 1 么, Nedlands, CSIRO Light Trap, 15. i. 1963 (M. M. H. Wallace) (SAM); 3 么, Perth, Boya, 12. iii. 1977 (B. G. Muir) (WAM).

Diagnosis Hestiochora occidentalis is sympatric with H. continentalis but is smaller. The genitalia are characteristic.

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Description Male (Pl. 42, Fig. 1). Black and white wasp-like habitus (description see p. 143). Forewing length: 7.5–9.0 mm. Smaller than most of the other species with wasp-like habitus but externally similar. Forewing with all veins free from cell. Female (Pl. 42, Fig. 2). Forewing length: 8.5–9.0 mm. Darker than male but almost equal in size. Male genitalia (Figs 305–307, 324, 325). Uncus long (almost as long as tegumen). Tegumen of medium length and breadth. Vinculum slender, saccus small, rounded. Ventral part of diaphragma with small, double-lobed juxta. Ventral part of valva with very pronounced sacculus proximally that is smaller and less sclerotized than in most related species; the ventral sclerotization of valva short and broad distally with a rounded apex and a tooth-like dorsal process; between sacculus and ventro-distal sclerotization there is a deep emargination. Aedeagus slightly curved upwards proximally, straight distally, strongly tapering towards its posterior end and pointed apically, with a short, strongly sclerotized dorsal spine. Sternum VIII covering 30–50% of segment.

Hestiochora intermixta sp. n. Pl. 43, Figs 1, 2; Figs 308, 309, 326, 327, 335.

Material examined Holotype 么. New South Wales, Pilliga State Forest, 40 km N of Baradine, 21. iii. 1982 (B. Hacobian). Genitalia slide Z 2885 (ANIC).

Female genitalia (Fig. 334). Sternum VII translucent. Sternum VIII forming a sclerotized almost rectangular plate around the unsclerotized ostium. Tergum VIII large and strongly sclerotized, connected to sternum VIII by two slender, transverse sclerotizations. Ostium small, antrum absent, ductus bursae almost straight.

Paratypes Queensland: 1 么, Goondiwindi, 24 km NE, 28. xi. 1976 (on Atalaya hemiglauca) (E. M. Exley & T. Low), genitalia slide Z 2852 (UQIC); 2 么, Milmerran, 26. xii. 1946, 21. xii. 1947 (J. Macqueen), genitalia slides Z 2869, Z 2890 (ANIC, UQIC). New South Wales: 1 么, 1 乆, Pilliga State Forest, 40 km N of Baradine, 21. iii. 1982 (B. Hacobian) (Coll. B. Hacobian). South Australia: 1 么, Hoyleton, xi. 1892, genitalia slide Z 2922 (SAM).


Diagnosis

Unknown. Some of the localities, especially the type locality Mt Dale west of Perth, were visited by G. M. and B. E. Tarmann in early September 1995. All of them are woodland dominated by tall eucalypts. At Mt Dale the following species were identified around the summit: Eucalyptus calophylla Lindley, E. marginata Smith and E. rudis Endl.

Hestiochora intermixta is very closely related to H. occidentalis and H. xanthocoma. Although it has the typical black and white wasp-like habitus like the former, it shares the yellowish orange frons and vertex of the male with the latter and is somewhat intermediate in genitalia. It is partly sympatric with, and externally very similar to, H. queenslandensis, sharing with it the yellowish scales around and on the translucent parts of the wing. The genitalic differences from H. queenslandensis are striking. Hestiochora intermixta is also sympatric with H. continentalis and H. furcata. Both are larger, lack the yellowish scales on the wings and their genitalia are significantly different.

Distribution Endemic to the temperate parts of Western Australia.

TAXONOMY

Description Male (Pl. 43, Fig. 1). Black and white wasp-like habitus (description see p. 143). Forewing length: 7.5–8.0 mm. Externally similar to H. queenslandensis. Translucent stripes on wings covered with whitish yellow scales except for anal angle of hindwing that is white. Head with yellow scales on frons and orange scales on vertex. R4 and R5 in forewing free from cell. Female (Pl. 43, Fig. 2). Forewing length: 9.5 mm (only one female known). Darker than the males and larger. Male genitalia (Figs 308, 309, 326, 327). Uncus long (almost of same length as tegumen). Tegumen and vinculum slender, saccus small, rounded. Ventral part of diaphragma with small, translucent, double-lobed juxta that has a stronger sclerotization medially. Ventral part of valva with very pronounced, distally rounded sacculus that is smaller than in most other species of the genus (except H. occidentalis and H. xanthocoma); the ventro-distal sclerotization of valva broad with a slightly triangular edge proximally, sharply pointed distally, with a small tooth-like process dorsally; between sacculus and ventral sclerotization there is a deep emargination. Aedeagus curved upwards proximally, straight posteriorly, strongly tapering towards its posterior end and pointed apically, with a strongly sclerotized dorsal spine. Sternum VIII covering 10–20% of segment.

Female genitalia (Fig. 335). Sternum VII slightly sclerotized anteriorly, translucent posteriorly. Sternum VIII forming a sclerotized, almost rectangular plate around the unsclerotized ostium. Tergum VIII large and strongly sclerotized, connected to sternum VIII by two slender, transverse sclerotizations. Ostium small, antrum absent, ductus bursae only slightly twisted, almost straight.

Phenology and bionomics Unknown. This species is only known from four localities, all situated in the dry sclerophyll Eucalyptus forest. One specimen was taken from a flower of Atalaya hemiglauca (F. Muell.) F. Muell. (Sapindaceae).

Distribution Southern Queensland, New South Wales, eastern South Australia.

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Fig. 283. Male genitalia: valva of Hestiochora erythrota, detail, Qld, Toowoomba. Gen. slide no. Z 2894. Fig. 284. Male genitalia: valva of Hestiochora tricolor, overview, Vic., Gisborne. Gen. slide no. Z 2926. Figs 285–287. Male genitalia: valva of Hestiochora tricolor, detail. 285: Vic., Gisborne. Gen. slide no. Z 2926; 286: SA, Russell. Gen. slide no. Z 2927; 287: SA, Prospect. Gen. slide no. Z 2921. Fig. 288. Male genitalia: valva of Hestiochora continentalis, paratype, overview, SA, Adelaide. Gen. slide no. Z 2931.

TAXONOMY

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Figs 289–296. Male genitalia: valva of Hestiochora continentalis, detail. 289: Vic., Beaconsfield. Gen. slide no. Z 2936; 290: NSW, Maryland. Gen. slide no. Z 2909. 291: SA, Adelaide. Gen. slide no. Z 2924; 292: SA, Adelaide. Gen. slide no. Z 2931; 293: SA, Aldinga. Enlargement. Gen. slide no. Z 2920; 294: SA, Halbury. Enlargement. Gen. slide no. Z 2928; 295: NSW, Sydney. Gen. slide no. Z 2943; 296: NSW, Narara. Gen. slide no. Z 2937.

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Fig. 297. Male genitalia: valva of Hestiochora furcata, paratype, overview, NSW, Wilton. Gen. slide no. Z 2929. Figs 298–299. Male genitalia: valva of Hestiochora furcata, paratype, detail. 298: NSW, Wilton. Gen. slide no. Z 2929; 299: NSW, Narara. Gen. slide no. Z 2905. Fig. 300. Male genitalia of Hestiochora queenslandensis, paratype, overview, Qld, Scottsville. Gen. slide no. Z 2848. Fig. 301. Male genitalia: valva of Hestiochora queenslandensis, paratype, detail, Qld, Milmerran. Gen. slide no. Z 2889.

TAXONOMY

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Figs 302–304. Male genitalia: valva of Hestiochora xanthocoma, detail. 302: Qld, Wacol. Gen. slide no. Z 2944; 303: Qld, Rosedale. Gen. slide no. Z 2892; 304: Qld, Jowalbinna. Gen. slide no. Z 2891. Figs 305–307. Male genitalia: valva of Hestiochora occidentalis, paratype, detail. 305: WA, Mt Dale. Gen. slide no. Z 2932; 306: WA, Mt Dale. Gen. slide no. Z 2881; 307: WA, Moir’s Rock. Gen. slide no. Z 2946.

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Fig. 308. Male genitalia of Hestiochora intermixta, paratype, overview, Qld, Goondiwindi. Gen. slide no. Z 2852. Fig. 309. Male genitalia: valva of Hestiochora intermixta, paratype, detail, Qld, Milmerran. Gen. slide no. Z 2890. Fig. 310. Male genitalia: aedeagus of Hestiochora erythrota, detail, Qld, Toowoomba. Gen. slide no. Z 2894. Figs 311–312. Male genitalia: aedeagus of Hestiochora tricolor, detail. 311: Vic., Gisborne. Gen. slide no. Z 2926; 312: SA, Prospect. Gen. slide no. Z 2921.

TAXONOMY

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Figs 313–316. Male genitalia: aedeagus of Hestiochora continentalis, paratype, detail. 313: SA, Adelaide. Gen. slide no. Z 2931; 314: SA, Adelaide. Gen. slide no. Z 2924; 315: NSW, Narara. Gen. slide no. Z 2937; 316: NSW, Sydney. Gen. slide no. Z 2938. Fig. 317. Male genitalia: aedeagus of Hestiochora furcata, paratype, detail, NSW, Wilton. Gen. slide no. Z 2931. Figs 318–319. Male genitalia: aedeagus of Hestiochora queenslandensis, paratype, Qld, Scottsville. Gen. slide no. Z 2848. 318: Detail, proximal part; 319: detail, distal part.

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Figure 320. Male genitalia: aedeagus of Hestiochora queenslandensis, paratype, detail, Qld, Milmerran. Gen. slide no. Z 2889. Figs 321–323. Male genitalia: aedeagus of Hestiochora xanthocoma, detail. 321: Qld, Rosedale. Gen. slide no. Z 2892; 322: Qld, Wacol. Gen. slide no. Z 2944; 323: Qld, Jowalbinna. Gen. slide no. Z 2891. Figs 324–325. Male genitalia: aedeagus of Hestiochora occidentalis, paratype, detail. 324: WA, Mt Dale. Gen. slide no. Z 2881; 325: WA, Moir’s Rock. Gen. slide no. Z 2946.

TAXONOMY

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Figs 326–327. Male genitalia: aedeagus of Hestiochora intermixta, paratype, detail. 326: Qld, Goodiwindi. Gen. slide no. Z 2852; 327: SA, Hoyleton. Gen. slide no. Z 2922. Fig. 328. Female genitalia of Hestiochora erythrota: aep, abdominal end plate; an, antrum; db, ductus bursae; cb, corpus bursae; ds, ductus seminalis; ob, ostium bursae; smVIII, sticky membrane of abdominal segment VIII; stVII, 7th sternite. Qld, Milmerran. Gen. slide no. Z 2896. Fig. 329. Female genitalia of Hestiochora tricolor: aep, abdominal end plate; an, antrum; cb, corpus bursae; db, ductus bursae; ds, ductus seminalis; ob, ostium bursae; pa, papillae anales; smVIII, sticky membrane of abdominal segment VIII; stVII, 7th sternite. Tas., Kingston. Gen. slide no. Z 2876. Fig. 330. Female genitalia of Hestiochora continentalis, SA, Aldinga Scrub. Gen. slide no. Z 2871. Fig. 331. Female genitalia of Hestiochora furcata, Qld, Mooloolaba. Gen. slide no. Z 2900.

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Fig. 332. Female genitalia of Hestiochora queenslandensis: aep, abdominal end plate; ob, ostium bursae; smVIII, sticky membrane of abdominal segment VIII; spVII, spiracle of 7th abdominal segment; stVIII, 8th sternite. Qld, Kenmore. Gen. slide no. Z 2941. Fig. 333. Female genitalia of Hestiochora xanthocoma: aep, abdominal end plate; cb, corpus bursae; db, ductus bursae; ob, ostium bursae; pa, papillae anales; rs, receptaculum seminis; smVIII, sticky membrane of abdominal segment VIII; stVII, 7th sternite; ut, utriculus. Qld, Carnavon Range. Gen. slide no. Z 2917. Fig. 334. Female genitalia of Hestiochora occidentalis, paratype: aep, abdominal end plate; cb, corpus bursae; db, ductus bursae; ob, ostium bursae; smVIII, sticky membrane of abdominal segment VIII; spVII, spiracle of 7th abdominal segment; stVII, 7th sternite. WA, Katanning. Gen. slide no. Z 2902. Fig. 335. Female genitalia of Hestiochora intermixta, paratype, NSW, Pilliga State Forest. Gen. slide no. Z 2886. This slide was made at a stage of investigation before the peculiar characters of the female genitalia of Hestiochora were understood by the author. Unfortunately many important characters were destroyed; even worse, this is the only known female of the species.

TAXONOMY

Genus Turneriprocris Bryk Turneriprocris Bryk, 1936, in Strand, Lepid. Cat. 4 (71): 304. Type species: Procris dolens Walker, 1854: 112, by original designation (for Neoprocris Turner, 1926a). [Tasmania]. Turneriprocris is an objective replacement name for Neoprocris Turner, 1926a. Neoprocris Turner, 1926a, Proc. Linn. Soc. NSW 51: 445. Type species: Procris dolens Walker, 1854: 112, by original designation. A junior homonym of Neoprocris Jordan, 1915: 300 (Lepidoptera, Zygaenidae). Neoprocris Turner; Bryk 1936: 141; Alberti 1954: 300; Tarmann 1996: 143. Turneriptocris; Tarmann 1994: 121, fig. 3. Unavailable; an incorrect subsequent spelling of Turneriprocris Bryk, 1936. The type-species of Turneriprocris Bryk, Procris dolens Walker, 1854, is rather unusual in many of its characters. The three other species that are similar to it in biology, form of the antenna and male genitalia are characterised by a striking synapomorphy in the female genitalia and clearly form a monophyletic unit. One of them, Procris rufiventris Walker, was transferred to Turneriprocris by Alberti (1954: 300), based on similarities in the male genitalia. Turneriprocris is here treated as a monotypic genus with the single included species T. dolens. Procris rufiventris is excluded and transferred to the new genus Myrtartona.

Diagnosis Small species with rounded wings, dark, unicolorous, without shining metallic scales (Pl. 44). Antenna dark with broad shaft and pointed apex, densely scaled dorsally, unscaled ventrally, leaving an unscaled ventral groove from base to apex, short pectinations in male, strong serrations in female. Abdomen dorsoventrally compressed, the lateral evaginations only on segment II. Female without abdominal hairtuft. Female with normal tube-like ductus, without praebursa. Larva on Myrtaceae. Head (Figs 29–36) Frons slightly projecting, rounded, very broad; labrum tongue-shaped medially, pilifer with about 12 long setae; proboscis with spiny surface uniform also at base; maxillary palps short; labial palps

extending towards anterior margin of head, slightly curved proximally, porrect distally; compound eyes small, with broad eyelash medioventrally consisting of several layers of scales; ocelli small; chaetosemata small, of typical artonoid form. Antennal shaft broad, shortly bipectinate in male, weakly biserrate in female, apex rounded, densely scaled dorsally and laterally but without scales ventrally, leaving an unscaled ventral groove from base to apex between pectinations that are very close together. Thorax and wings (Figs 105–111) Foretibia without epiphysis, tibial spurs short, stout, 0–2–2, the outer spurs slightly longer, no single medial spur on hind-tibia. Wings rounded, narrow, forewing more densely scaled than hindwing, the latter slightly translucent. Both wings with medial stem present, forewing with all veins free from cell, hindwing with all veins present, Sc touching costal margin of cell and at one point connected to it by very short cross-vein. M3+CuA1 either stalked with the length of the stalk very variable or both veins may be free but originating from the same point. Frenulum one long spine in male, three short spines in female; retinaculum a well-developed hook at base of Sc in male, and a number of scales on base of CuP in female. Pregenital abdomen (Fig. 428) Abdomen short, dorsoventrally flattened in dry specimens, especially in female; the artonoid lateral evaginations present only on segment 2, very small. Sternum II with pair of small, proximally straight and distally curved apodemes, lateral sternal rod very weak, almost absent; sclerotization of segment without lateral prolongations anteriorly. Tergum II with a sclerotized part covering approximately onethird of segment posteriorly in male and at onequarter in female. Sternum VIII in male reduced to a small anterior margin, rest of segment translucent. Sternum VII in female covering the whole segment, sternum VIII reduced. No abdominal hair-tuft. Male genitalia (Figs 336–341) Uncus triangular basally, slender distally. Tegumen consisting of two broad lobes, vinculum broad, Ushaped, not strongly sclerotized, saccus very small and slender, vestigial. Valva with strong ventral sclerotization, sacculus not pronounced; dorsomedial part of valva fused with dorsolateral part of diaphragma forming a characteristic, finger-like process that is forked basally and densely covered with setae distally. Normally, this process points backwards parallel to the valva; one part of the fork

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points sideways and forms a joint with the valva, the other points upwards and is probably an apodeme for a muscle that moves the process. Pulvinus absent. Transtilla and juxta present. Aedeagus very short and stout with strong sclerotization ventrodistally; vesica not eversible. Female genitalia (Figs 342, 343) Sternum VIII reduced. Tergum VIII slender, free, not connected to sternum VIII, with short, slender, lateral elongations. Papillae anales with short setae, apophyses posteriores slender, as long as diameter of papillae. Ostium unsclerotized; ductus bursae simple, slender, translucent; corpus bursae simple. Glandula sebacea bearing distally a pair of spherical bulbs with long and very slender ducti inserting into a heart-shaped ‘bulla’, and with a broad common ductus proximally.

Phenology and bionomics See T. dolens (p. 165). The larva feeds on Leptospermum spp. (Myrtaceae).

Remarks The principal autapomorphy of Turneriprocris is the complete loss of the praebursa (see further discussion on p. 170).

Distribution Turneriprocris occurs from New South Wales to Victoria and South Australia and Tasmania.

Turneriprocris dolens (Walker) Pl. 44, Figs 1, 2; Figs 29–36, 105–111, 126–127, 336–343. Procris dolens Walker, 1854, List Specimens lepid. Insects Colln Br. Mus. 1: 112. Lectotype 么 (here designated), AUSTRALIA: Tasmania (‘Van Diemen’s Land’) (BMNH) [examined]. Procris dolens Walker; Meyrick 1886: 791; Jordan 1907: 10. Neoprocris dolens (Walker); Turner 1926a: 445; Bryk 1936: 141; Alberti 1954: 300. Pollanisus dolens (Walker); McFarland 1979: 62; Yen 2003: 300, 333. Turneriprocris dolens (Walker); Tarmann 1996: 143.

Material examined Queensland: 1 乆, Stanthorpe, 3. iv. 1930 (W. B. Barnard) (QM). New South Wales: 1 么, Barrington Tops, i. 1925 (SU Zoo Exp.) (AMS); 1 么, Barrington Tops, 5. xi. 1970 (V.J. Robinson) (ANIC); 1 乆, Brown Mountain, 1200 m , 9. i. 1968 (I. F. B. Common, A. E. May & M. S. Upton) (ANIC); 1 么, Shoalhaven R., 13 km NNW Braidwood, 26. x. 1977 (D. C. F. Rentz & J.Balderson) (ANIC); 1 么, Tinderry Mts., 1300m , 8. i. 1963 (I. F. B. Common & M. S. Upton) (ANIC); 1 么, Tubrabucca, Barrington Tops, 1450 m , 10. i. 1967 (V. J. Robinson) (ANIC). Australian Capital Territory: 1 乆, Bendora, 6. ii. 1962 (I. F. B. Common) (ANIC); 3 (males), 3 (females), Camels Hump, ca. 2 km S Tidbinbilla, 1200 m , 9. xii. 1989 (G. M. Tarmann & E. S. Nielsen) (ANIC, TLMF); 1 么, Canberra, 25. xii. 1937 (G. M. Goldfinch) (AMS); 1 么, Canberra (Farrer), 7. xii. 1980 (D. C. F. Rentz) (ANIC); 1 么, Mt Franklin, 1500 m , 5. ii. 1969 (I. F. B. Common & A. E. May) (ANIC). Victoria: 4 么, Beaconsfield, xi. 1903 (MV); 1 乆, Belgrave, 16. xii. 1943 (J. C. Le Souef) (ANIC); 2 么, Cheltenham (MV); 1 乆, Emerald, 20. xii. 1904 (J. A. Kershaw) (MV); 1 乆, Fern Tree Gully, 30. xi. 1907 (J. A. Kershaw) (MV); 1 么, Forrest, 25. xii. 1892 (MV); 1 么, Gisborne, 15. xii. 1895 (MV); 1 乆, Kallista, 6. xii. 1944 (J. C. Le Souef) (ANIC), 1 么, Mordialloc, 27. xi. 1931 (A. L. Brown) (MV); 5 么, 2 乆, Mt Macedon, 27. xii. 1902, 29. xii. 1905, 24.–28. xi. 1907 (ANIC, MV); 1 么, nr. Mt Pinnibar, 24. xi. 1961 (K. L. Taylor) (ANIC); 3 乆, Upwey, 24. xii. 1930, 5. i. 1931 (C. H. B[orch]) (MV); 1 么, Wandin, 26. xii. 1892 (MV); 1 么, Woodend, 15. xii. 1919 (MV). South Australia: Larvae (L4, L5), Aldgate (Heather Rd.), Mt Lofty Ra., 13. ix. 1969 (on Leptospermum myrsinoides) (N. McFarland & D. Bakker) (SAM); Larvae (L5), Belair Railway Station, 1 ml E, Adelaide, 14. ix. 1969 (on Leptospermum myrsinoides) (N. McFarland & D. Bakker) (SAM); 1 么, Balhannah, 1883 (E. Guest) (SAM); 4 么, Balhannah, 21. xi. 1886, 7. xi. 1892 (SAM); Larvae, Longwood, 7 ml N, Mt Lofty Ra., 3. ix. 1966 (half grown), 17. ix. 1966 (fully grown) (on Leptospermum myrsinoides and rarer on L. juniperinum) (N. McFarland) (SAM). Tasmania: 5 么, 13 乆, Cradle Mt, 3000 ft, 20.–25. i. 1925, 21. i. 1934 (ANIC, QM); 2 么, 1 乆, Cradle Mt, Moina, 2000 ft, 30. i. 1925 (ANIC); 1 么, Derby, 28. i. 1938 (TMH); 1 么, Derwent Br., 13. i. 1938 (ANIC); 1 乆, Kingston, 12. ii. 1953 (J. A. Cunningham) (TMH); 1 么, Hobart (Lea) (SAM); 1 乆, Hobart, 14. xii. 1904 (MV); 1 么, 2 乆, Hobart, 4. xii. 1913, 16. xii. 1913, (G. H. Hardy) (AMS); 1 乆 Huon R., 1907 (Lea) (ANIC); 1 么, 1 乆, Lake St Clair, 21.-24. i. 1934 (ANIC, QM); 1 乆, Mt Nelson, 200 m , iv. 1980 (P. B. McQuillan) (ANIC); 4 么, 2 乆, Pelion Plains, 860 m , 8. ii. 1991 (E. S. Nielsen & P. B. McQuillan) (ANIC); 1 么, 3 乆, Pelion Plains, 880 m , 8. ii. 1990 (E. S. Nielsen & P. B. McQuillan) (ANIC); 1 乆, Pelion Hut, 900 m, 8. i. 1989 (P. B. McQuillan & E. S. Nielsen) (ANIC); 1 乆, Pelion Hut, 870 m , 7. ii. 1990 (E. S. Nielsen & P. B. McQuillan) (ANIC); 5 么, 4 乆, Pelion Hut, 870 m , 8. i. 1991, 11. i. 1991 (E. S. Nielsen & E. D. Edwards) (ANIC); 1

TAXONOMY

么, Ridgeway, 15. xii. 1935 (MV); 1 乆, Ridgeway, 26. xi. 1977 (P. B. McQuillan) (ANIC); 1 乆 St Marys, 13 mls ESE, 19. i. 1984 (Key, Carne & Kerr) (ANIC); 1 乆 Stone Hut, 30. i. 1979 (P.B. McQuillan) (ANIC); 1 乆, Thumbs Forest Reserve, near Orford, 6. i. 1991 (E. S. Nielsen & E. D. Edwards) (ANIC).

Diagnosis Turneriprocris dolens is one of the smallest zygaenids in Australia. The dark overall appearance without metallic sheen, rounded wings, broad frons, small compound eyes, very short antennal pectinations in the male and the lack of an abdominal hair-tuft in the female render this species unmistakable. The blackish brown proboscis is a good diagnostic character for distinguishing large specimens of T. dolens from small specimens of Myrtartona coronias that have a yellow proboscis.

Redescription Male (Pl. 44, Fig. 1). Forewing length: 5.5–6.5 mm. Head, thorax, abdomen and forewing upperside blackish brown with bluish green sheen in fresh specimens, densely covered with long and slender scales, hindwing black, without sheen, slightly translucent, especially basally. Frons slightly projecting, rounded, very broad, 2.5× broader than compound eye in frontal view; proboscis dark brown; compound eyes small, deep black; ocelli small, white; chaetosema dark. Antenna with 29–31 segments, bipectinate from segments 1–22, weakly biserrate from segments 22 to apex; pectinations short, only 1.5× longer than shaft at segment 5 and approximately as long as shaft at segment 15, apex rounded. Female (Pl. 44, Fig. 2). Similar to male but slightly larger. The abdomen is remarkably flattened dorsoventrally, especially in dried specimens. Frons 3.0× broader than compound eye in frontal view. Antenna weakly biserrate. Male genitalia (Figs 336–341). Uncus of medium length. Valva with a strong, finger-like, ventral sclerotization with a distal point, a slender, translucent medial and a very slender, weakly sclerotized dorsal part, sacculus not pronounced. Mediodorsal process of valva almost equal in length to ventral process, slender, slightly pointed apically. Transtilla forming a narrow bridge. Juxta half-moon-shaped. Aedeagus very short and stout, more or less pear-shaped, broad basally, slightly upcurved and tapering distally, completely translucent proximally at inser-

tion of ductus ejaculatorius, but strongly sclerotized ventrodistally, broad at first, then tapering distally into a quite slender tube. Female genitalia (Figs 342, 343). Sternum VIII reduced to two round setae-bearing plates laterad of ostium. Ostium small, translucent; ductus bursae simple, straight, slender, translucent; corpus bursae spherical, broadly connected to slender ductus seminalis laterally.

Phenology and bionomics Turneriprocris dolens is a univoltine early summer species inhabiting the temperate parts of south-eastern Australia and Tasmania. In the more northern parts of its range it seems to be restricted to higher altitudes. A single April record indicates that there may be a partial second generation in the autumn. The adults are weak flyers and are known to be abundant in many localities, flying just above or amongst vegetation, close to the larval foodplants. They seem to be most active in the late afternoon and it is possible that the females secrete pheromones at that time, as in many other Procridinae. McFarland (unpublished field notes) collected final instar larvae on Leptospermum myrsinoides Schl. (Pl. 62, Fig. 5) and L. juniperinum Smith (Myrtaceae) at Longwood, Mt Lofty Range, South Australia, on 3 September 1966. The half grown larvae were very abundant and could be collected by beating twigs with old leaves. Leptospermum myrsinoides was preferred as a host-plant. On 17 September 1966 final instar larvae were obtained at the same locality.

Distribution Eastern and southern Australia (south-eastern Queensland, New South Wales, Australian Capital Territory, Victoria, South Australia) and Tasmania.

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Figs 336–337. Male genitalia of Turneriprocris dolens, Tas. Lake St Clair. Gen. slide no. Z 2957. 336: Overview; 337: detail. Figs 338–339. Male genitalia of Turneriprocris dolens, detail, Tas., Cradle Mountain. Gen. slide no. Z 2956. 338: Overview; 339: detail. Figs 340–341. Male genitalia: aedeagus of Turneriprocris dolens. 340: Tas., Cradle Mountain. Gen. slide no. Z 2956; 341: Tas., Lake St Claire. Gen. slide no. Z 2957.

TAXONOMY

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343

Fig. 342. Female genitalia of Turneriprocris dolens: gs, glandula sebacea; gsbl, glandula sebacea, bulla; gscd, glandula sebacea, common duct; gsg, glandula sebacea, glands; rs, receptaculum seminis. Tas., Cradle Mountain. Gen. slide no. Z 2959. Fig. 343. Female genitalia of Turneriprocris dolens: cb, corpus bursae; db, ductus bursae; ds, ductus seminalis; ob, ostium bursae; pa, papillae anales; rs, receptaculum seminis; sppl, spiny plate; stVII, 7th sternite. Tas., Cradle Mountain. Gen. slide no. Z 2958.

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Plate 1: Figs 1–2. Pollanisus viridipulverulenta, 6.8× original size. 1: (male), SA, Aldinga Scrub; 2: (male), Qld, Brisbane.

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Plate 2: Figs 1–2. Pollanisus viridipulverulenta, 6.8× original size. 1: (male), Vic., Gisborne; 2: (male), blue morph, Vic., Gisborne.


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Plate 3: Fig. 1. Pollanisus viridipulverulenta, (female), Vic., Gisborne, 6.8× original size. Fig. 2: P. modestus, holotype (male), NSW, Clyde Mountain, 6.8× original size.

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Plate 4: Figs 1–2. Pollanisus cupreus, 6.8× original size. 1: (male), WA, Perth; 2: (female), same data.


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Plate 5: Figs 1–2. Pollanisus nielseni, 6.8× original size. 1: (male), WA, Wedge Island N of Lancelin; 2: (female), same data.

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Plate 6: Figs 1–2. Pollanisus lithopastus, 6.8× original size. 1: (male), Vic., Beaconsfield; 2: female), same data.


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Plate 7: Figs 1–2. Pollanisus sp. 1, 6.8× original size. 1: (male), ACT, Bendora; 2: (female), same data.

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Plate 8: Figs 1–2. Pollanisus empyrea, 8× original size. 1: (male), WA, Albany; 2: (female), same data.


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Plate 9: Figs 1–2. Pollanisus apicalis, 8× original size. 1: (male), ACT, Black Mountain; 2: (female), same data.

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Plate 10: Fig. 1. Pollanisus apicalis, (male), Qld, Stradbroke Island, 8× original size. Fig. 2. Pollanisus sp. 3, (male), Palm Island, 8× original size.


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Plate 11: Figs 1–2. Pollanisus trimacula, 8× original size. 1: (male), NSW, Narara; 2: (female), same data.

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Plate 12: Figs 1–2. Pollanisus edwardsi, 8× original size. 1: (male), NSW, Tenterfield; 2: (female), Qld, Stradbroke Island.


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Plate 13: Figs 1–2. Pollanisus subdolosa subdolosa, 8× original size. 1: (male), Vic., Beaconsfield; 2: (female), same data.

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Plate 14: Figs 1–2. Pollanisus subdolosa clara, 8× original size. 1: (male), NSW, Niagara Park; 2: (female), same data.


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Plate 15: Fig 1: Pollanisus subdolosa clara, (male), NSW, Minyon Falls, 8× original size. Fig 2: Pollanisus cf. subdolosa, (male), NSW, St. George’s Basin, 8× original size.

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Plate 16: Figs 1–2. Pollanisus cf. subdolosa, 8× original size. 1: (male), NSW, Wollongong; 2: (female), same data.


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Plate 17: Figs 1–2. Pollanisus contrastus, 8× original size. 1: (male), NSW, Narara; 2: (female), same data.

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Plate 18: Fig. 1. Pollanisus cf. contrastus 1, (male), NSW, Tweed Heads, 8× original size. Fig. 2. P. cf. contrastus 2, (male), Qld, Stradbroke Island, 8× original size.


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Plate 19: Figs 1–2. Pollanisus eungellae, 8× original size.. 1: holotype (male), Qld, Eungella; 2: paratype (female), same data.

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Plate 20: Figs 1–2. Pollanisus eumetopus, 8× original size. 1: (male), Qld, Redlynch; 2: (female), same data.


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Plate 21: Figs 1–2. Pollanisus commoni, 8× original size. 1: (male), Qld, Clifton Beach; 2: (female), same data.

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Plate 22: Figs 1–2. Pollanisus incertus, 8× original size. 1: holotype (male), Qld, Kuranda; 2: paratype (female), same data.


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Plate 23: Figs 1–2. Pollanisus angustifrons, 8× original size. 1: holotype (male), Qld, Tully; 2: (male), Qld, Waterview Creek.

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Plate 24: Figs 1–2. Pollanisus sp. 2, 8× original size. 1: (male), NSW, Murramarang National Park; 2: (female), same data.


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Plate 25: Figs 1–2. Pollanisus sp. 3, 8× original size. 1: (male), Qld, Magnetic Island; 2: (female), same data.

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Plate 26: Figs 1–2. Pollanisus sp. 4, 8× original size. 1: (male), Qld, Waterview Creek; 2: (female), same data.


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Plate 27: Figs 1–2. Pollanisus sp. 5, 8× original size. 1: (male), Qld, Kennedy River; 2: (male), Qld, Mt Bellenden-Ker.

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Plate 28: Figs 1–2. Pollanisus sp. 6, 8× original size. 1: (male), Qld, Townsville; 2: (female), same data.


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Plate 29: Fig. 1. Pollanisus sp. 7, (male), NT, Darwin, 8× original size. Fig 2: Pollanisus sp. 8, (male), Qld, Jowalbinna, 8× original size.

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Plate 30: Figs 1–2. Pollanisus cyanota, 10× original size. 1: (male), Qld, Howard; 2: (female), same data.


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Plate 31: Fig. 1. Pollanisus isolatus, holotype (male), Vic., Beaconsfield, 10× original size. Fig. 2. Pollanisus calliceros calliceros, (male), TAS, Cradle Mountain, 10× original size.

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Plate 32: Figs 1–2. Pollanisus calliceros azurea, 10× original size. 1: (male), NSW, Ebor; 2: (female), NSW, Barrington Tops.


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Plate 33: Figs 1–2. Onceropyga anelia, 8.5× original size. 1: (male), Qld, Brisbane; 2: (female), Toowoomba.

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Plate 34: Figs 1–2. Onceropyga pulchra, 8.5× original size. 1: holotype (male), NSW, Watagan State Forest; 2: (female), NSW, Mt Keira.


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Plate 35: Figs 1–2. Hestiochora erythrota, 8.5× original size. 1: (male), Qld, Milmerran; 2: (female), same data.

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Plate 36: Figs 1–2. Hestiochora tricolor, 8.5× original size. 1: (male), TAS, Mt Nelson; 2: (female), TAS, Moorina.


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Plate 37: Figs 1–2. Hestiochora continentalis, 8.5× original size. 1: (male), NSW, Niagara Park; 2: (female), WA, Nedlands.

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Plate 38: Figs 1–2. Hestiochora furcata, 8.5× original size. 1: (male), ACT, Black Mountain; 2: (male), NSW, Wilton.


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Plate 39: Figs 1–2. Hestiochora furcata, 8.5× original size. 1: (male), NSW, Barrington Tops; 2: (female), NSW, Sawpit Creek, Mt Kosciuszko National Park.

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Plate 40: Figs 1–2. Hestiochora queenslandensis, 8.5× original size. 1: (male), Qld, Planted Creek; 2: (female), Milmerran.


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Plate 41: Figs 1–2. Hestiochora xanthocoma, 8.5× original size. 1: (male), Qld, Biloela; 2: (female), NT, Katherine.

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Plate 42: Figs 1–2. Hestiochora occidentalis, 8.5× original size. 1: (male), WA, Moir’s Rock; 2: (female), WA, Mt Dale.


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Plate 43: Figs 1–2. Hestiochora intermixta, 8.5× original size. 1: (male), NSW, Pilliga State Forest; 2: (female), same data.

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Plate 44: Figs 1–2. Turneriprocris dolens, 10× original size. 1: (male), TAS, Cradle Mountain; 2: (female), same data.


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Plate 45: Figs 1–2. Myrtartona coronias, 7.5× original size. 1: (male), NSW, Blackheath; 2: (female), same data.

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Plate 46: Figs 1–2. Myrtartona leucopleura, 7.5× original size. 1: (male), Sydney; 2: (female), same data.


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Plate 47: Figs 1–2. Myrtartona rufiventris, 7.5× original size. 1: (male), WA, Salmon Gums; 2: (female), WA, Mandurah.

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Plate 48: Fig. 1. Myrtartona mariannae, holotype (male), Qld, Milmerran, 7.5× original size. Fig. 2. Australartona mirabilis, (male), Qld, Lamington National Park, 7.5× original size.


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Plate 49: Figs 1–2. Australartona mirabilis, 7.5× original size. 1: (male), NSW, Barrington Tops; 2: (female), NSW, Mt Dromedary.

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Plate 50: Fig. 1. Australartona mirabilis, (male), NSW, Brown Mountain, 7.5× original size. Fig. 2. Pseudoamuria neglecta, holotype (female), Qld, Taylor Range, 7.5× original size.


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Plate 51: Figs 1–2. Homophylotis thyridota, 9× original size. 1: (male), Qld, Moses Creek; 2: (female), Qld, Kuranda.

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Plate 52: Figs 1–2. Homophylotis pseudothyridota, 9× original size. 1: (male), Qld, Kuranda; 2: (male), Qld, Mt Tozer.


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Plate 53: Figs 1–2. Homophylotis pseudothyridota, 9× original size. 1: (male), Qld, Kuranda; 2: (female), same data.

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Plate 54: Figs 1–2. Homophylotis artonoides, 9× original size. 1: holotype (male), Qld, Iron Range; 2: paratype (female), same data.


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Plate 55: Figs 1–2. Pseudoamuria uptoni, 9× original size. 1: holotype (male), Qld, Iron Range; 2: paratype (female), Banks Island.

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Plate 56: Figs 1–2. Palmartona catoxantha, 9× original size. 1: (male), Malaysia, Malakka; 2: (female), Qld, Taylor Range.


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Plate 57: Figs 1–2. Thyrassia inconcinna, 6.5× original size. 1: (male), Qld, Townsville; 2: (female), same data.

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Plate 58: Figs 1–3. Zygaena acharon, lectotype (female) (BMNH, Banks Coll.). 1: Upperside; 2: underside; 3: labels. Fig. 4. Pollanisus apicalis, (female), abdominal hairtuft. Fig. 5. Hestiochora tricolor , (female), abdominal hairtuft. Fig. 6. Hestiochora xanthocoma, (female), abdominal hairtuft.


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Plate 59: Biology. Fig. 1. Eggs of Pollanisus viridipulverulenta on Hibbertia obtusifolia DC. (Dilleniaceae), ACT, Black Mountain. Fig. 2. Eggs of Pollanisus subdolosa clara, NSW, Budawang National Park (photo V. Rangsi, CSIRO). Fig. 3. First instar larva of Pollanisus subdolosa clara, NSW, Budawang National Park (photo A. Zwick, CSIRO). Fig. 4. Third instar larva of Pollanisus subdolosa clara, dorsal view, NSW, Budawang National Park (photo A. Zwick, CSIRO). Fig. 5. Third instar larva of Pollanisus subdolosa clara, lateral view, NSW, Budawang National Park (photo A. Zwick, CSIRO). Fig. 6. Adult larva of Pollanisus apicalis on Hibbertia obtusifolia DC., ACT, Aranda.

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Plate 60: Biology: Figs 1–2. Pollanisus apicalis, ACT, Aranda. 1: Adult larva (dorsal); 2: adult larva (lateral) (photos J. Green, CSIRO). Figs 3–5. Pollanisus viridipulverulenta, ACT, Black Mountain. 3: Adult larva; 4: cocoon; 5 : detail of pupa with abdominal gland on second segment visible (photos V. Rangsi, CSIRO). Fig. 6. Pollanisus cupreus, (female) (right specimen) and Pollanisus nielseni, (female) (left specimen) on flower of Arctotheca calendula (L.) Levyns (Asteraceae), WA, Wedge Island E.


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Plate 61: Biology. Fig. 1. Pollanisus viridipulverulenta, male, blue morph, Vic., Anglesea (photo J. Landy). Fig. 2. P. viridipulverulenta, male, green morph, on flower of Helichrysum viscosum Sprengel, ACT, Black Mountain. Fig. 3. P. viridipulverulenta, female, ACT, Black Mountain. Fig. 4. Myrtartona leucopleura, female, Qld, Seventeen Seventy.

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Plate 62: Larval host-plants. Fig. 1. Hibbertia obtusifolia DC. (Dilleniaceae), ACT, Black Mountain. Fig. 2. Hibbertia stricta (DC.) F. v. M. (Dilleniaceae), ACT, Black Mountain. Fig. 3. Hibbertia hypericoides (DC.) Benth. (Dilleniaceae), WA, Perth. Fig. 4. Hibbertia scandens (Willd.)Gilg. (Dilleniaceae), NSW, Mt. Dromedary. Fig. 5. Leptospermum myrsinoides Schl. (Myrtaceae), ACT, Black Mountain. Fig. 6. Tetrarrhena juncea R. Br. (Poaceae), NSW, Mt. Dromedary.

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Plate 63: Biotopes. Fig. 1. WA, Wedge Island, 5 km E, heathland on sand dunes, type-locality of Pollanisus nielseni and habitat for Pollanisus cupreus. Fig. 2. Qld, Seventeen Seventy, coastal forest, habitat of Myrtartona leucopleura. Fig. 3. Qld, Seventeen Seventy, dense coastal scrub with Hibbertia scandens (Willd.)Gilg., the larval host-plant of various Pollanisus species. Fig. 4. TAS, Melaleuca, treeless heathland, habitat of Pollanisus lithopastus (photo E. S. Nielsen, CSIRO).

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Plate 64: Biotopes. Fig. 1. WA, Perth, western Australian sclerophyll forest, habitat of Pollanisus cupreus and P. empyrea. Fig. 2. ACT, Aranda, eastern Australian sclerophyll forest, habitat of P. viridipulverulenta and P. apicalis. Fig. 3. NSW, Mt Dromedary, eastern temperate rain forest, potential biotope of P. subdolosa and P. trimacula. Fig. 4. NSW, Mt Dromedary, summit area, clearing in misty open temperate rain forest with Tetrarrhena juncea R. Br. (Poaceae), type-locality of Australartona mirabilis.

TAXONOMY

Genus Myrtartona gen. n. Type-species: Procris rufiventris Walker, 1854: 110.

Diagnosis Species of medium size with variable wing shape (Pls 45–47; Pl. 48, Fig. 1) and colour, without shining metallic scales. Antenna dark with broad shaft and pointed apex, densely scaled dorsally, unscaled ventrally, leaving an unscaled ventral groove from base to apex, short pectinations in male, strong serrations in female. Abdomen with the lateral evaginations only on segment II. Female without abdominal hairtuft. Female with a prominent praebursa that bears sclerotized ring-shaped structures and a large dagger-like cornutus. Larva on Myrtaceae. Head (Figs 37–40) Frons slightly projecting, dorsally more than ventrally, rounded, broad; labrum small; proboscis yellow, maxillary palps short; labial palps short and slender, not reaching as far as anterior margin of head, upcurved, pointed distally; compound eyes large, with a large eyelash medioventrally consisting of several layers of scales; ocelli large (M. rufiventris) to medium-sized (M. leucopleura, M. coronias); chaetosema typically artonoid. Antenna with broad shaft, bipectinate in male, biserrate in female, apex pointed, densely scaled dorsally, unscaled ventrally, leaving an unscaled ventral groove from base to apex, which is especially evident in females. Thorax and wings (Figs 112–118) Fore-tibia without epiphysis, tibial spurs 0–2–2, no single medial spur. Wings of different shapes, in the male broader and more rounded in one species (M. coronias), more slender and slightly pointed at apex in the three others, slender and apically rounded in the female, forewing more densely scaled than hindwing, the latter with a slightly translucent part mediobasally. Medial stem in forewing absent, present in hindwing; forewing with all veins free from cell, sometimes R4+R5 and/or M2+M3 originating from one point; hindwing with Sc touching the costal margin of cell at one point, venation in hindwing extremely variable: RR and M1 free, stalked or fused into a single vein, M2 and M3 free, stalked or fused, sometimes even M2, M3 and CuA1 stalked. Asymmetries in the venation of the left and right wings are frequent. Frenulum a single strong spine in male, three smaller bristles in female; retinaculum a strong hook at base of Sc in male, a number of scales at base of CuP in female.

Pregenital abdomen (Fig. 429) Abdomen longer in female; artonoid lateral evaginations only on segment II, small. Sternum II with a pair of small, curved apodemes, lateral sternal rod variable, usually not strongly developed. Tergum II sclerotized posteriorly, translucent anteriorly. Sternum VIII in male covering only proximal part of segment. Sternum VII in female large and broad, covering whole segment. No abdominal hair-tuft. Male genitalia (Figs 344–372). Uncus short, with an almost triangular base, slender and down-curved distally. Tegumen long and slender, vinculum broad U-shaped, not strongly sclerotized, with a small rounded or slightly triangular saccus or without saccus. Valva with a strong ventral sclerotization that is pointed distally and extends beyond the distal margin of the valva, sacculus not well developed, bearing a field of long setae; dorsomedial part of valva fused with dorsolateral part of diaphragma, forming a characteristic finger-like process that is forked basally and densely covered with setae distally (‘Myrtartona finger’). Normally, this process points backwards parallel to the valva, one part of the forked base points sidewards and forms a joint with the valva, the other points upwards (see also p. 165 T. dolens). Pulvinus absent. Transtilla present but very small. Juxta forming a large plate. Aedeagus very large, broad and stout, the vesica is very difficult to evert; when everted it is large and bulb-shaped, with one prominent cornutus and a large sclerotized plate. Female genitalia (Figs 373–377). Sternum VIII reduced to a small, narrow band connected to the broader tergum VIII forming a narrow ring. Apophyses anteriores present but very short. Papillae anales large, apophyses posteriores narrow, equal in length to diameter of papillae anales. The main autapomorphy of Myrtartona is a large, translucent praebursa with a very characteristic structure for rupturing the wall of the spermatophore. If a spermatophore is present in the praebursa, the way in which this functions is often visible, with the structure actually piercing the bulb of the spermatophore. This structure consists of a very strongly sclerotized thorn in the lateral wall of the praebursa with a comma-shaped, sharply pointed inner part and a knob-like, rounded outer part giving the whole a dagger-like appearance. There is a large sclerotized plate around the thorn although not connected to it. The short, broad ductus intrabursalis inserts at exactly the point

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where the thorn is situated. Corpus bursae small. The glandula sebacea consists of a pair of pearshaped bulbs distally with very slender ducti leading into a ‘bulla’, and a slender common duct proximally.

Phenology and bionomics The early instars are only known for the typespecies M. rufiventris (see p. 175), the larva of which feeds on Melaleuca spp. (Myrtaceae). M. coronias larvae feed on Melaleuca spp. and Leptospermum juniperinum (Myrtaceae) (according to labels on specimens, see p. 172, M. leucopleura presumably on Eucalyptus spp. (see p. 174).

Remarks Myrtartona gen. n. is closely related to Turneriprocris. The dorsomedial finger-like process in the male genitalia (Myrtartona-finger) and the strong, distally pointed ventral sclerotization on the valva are important synapomorphies for these two genera, indicating their monophyly. The larvae of both genera feed on Myrtaceae, mainly on Leptospermum spp., Melaleuca spp. and Kunzea spp. One species of Myrtartona, M. coronias, is also similar to Turneriprocris dolens in habitus, sharing with it the distinctive weak bluish green sheen on the upperside of the forewing and body, and the uniform dark greyish black ground colour. Nevertheless, differences in other characters are too great for the two groups to be considered congeneric. Turneriprocris seems to be a relict genus with a number of ancestral characters (e.g. fully developed medial stem in both wings). The female genitalia of the type-species, T. dolens, have no praebursa, but a simple translucent ductus bursae. This character was interpreted by Alberti (1954: 300, 338) as the secondary loss of a former praebursa. Alberti’s conclusion, i.e. that a translucent enlargement of the ductus bursae, as in Pollanisus and Myrtartona, can easily be reduced to a single tube, may be correct. Even if T. dolens is a relict species, it is most unlikely that the simple ductus bursae is a primitive character state. If it were, the praebursa in Artonini would have developed independently several times and this possibility cannot be completely excluded. In some groups, e.g. Pollanisus, Onceropyga, the praebursa is very translucent and is clearly just an expanded posterior part of the ductus bursae. However, in most cases the praebursa has a specialized structure to accommodate the cornuti of the male vesica and/or specialized structures to rupture the spermatophore.

These structures can be seen even in very translucent praebursae and, although they are very different in detail in the various groups, the locations of the functional parts are the same and homologous structures can usually be recognized clearly. This makes independent development of a praebursa in Artonini most unlikely. Myrtartona is characterised by a striking autapomorphy in the female genitalia, i.e. the thorn-like structure on the praebursa wall to rupture the spermatophore, and shows several other derived characters, e.g. the loss of the medial stem in the forewing and very variable wing venation in the hindwing with frequent reductions.

Distribution Temperate and subtropical Australia from the south-west (Perth to Albany) throughout the dry south (Nullarbor Plain) to the east (South Australia, Victoria to New South Wales and southern Queensland) and Tasmania.

Key to Myrtartona species 1. – 2. – 3. –

Body and wings unicolorous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. coronias (p. 170) Body with white or orange markings . . . . . . . 2 Abdomen dark dorsally, partly white ventrally and laterally . . . . . . .M. leucopleura (p. 172) Abdomen orange dorsally . . . . . . . . . . . . . . . . 3 Abdomen dark ventrally M. rufiventris (p. 174) Abdomen yellowish white ventrally . . . . . . . . . . . . . . . . . . . . . . . M. mariannae sp. n. (p. 176)

Myrtartona coronias (Meyrick) comb. n. Pl. 45, Figs 1, 2; Figs 344–346, 356–358, 373, 374. Procris coronias Meyrick, 1886, Proc. Linn. Soc. NSW (2) 1: 792. Lectotype 么, AUSTRALIA: Victoria, Mt Macedon, xii. 1877 (G. H. Raynor) (BMNH), here designated [examined]. Adscita coronias (Meyrick); Kirby 1892: 85. Pollanisus coronias (Meyrick); Jordan 1907: 10; Hering 1922: 13; Turner 1926b: 444; Bryk 1936: 118. Turneriptocris [sic] coronias (Meyrick); Tarmann 1994: 121 (misspelling). ‘No available genus’ [sic] coronias (Meyrick); Tarmann 1996: 143.

TAXONOMY

Material examined Queensland: 1 乆, Stanthorpe, 31. x. 1944 (J. M[ac]q[ueen]) (ANIC). New South Wales: 1 乆, Amosfield, 5. xii. 1970 (J. F. Donaldson) (DPIQ); 1 么, Barren Grounds Fauna Reserve, 30. i. 1973 (V. J. Robinson) (ANIC); 3 么, 1 乆, Barrington Tops, i.-ii. 1925 (Sydney University Zoo. Exped.) (AMS); 15 么, 1 乆, Blackheath, 7. xii. 1946, 20. xii. 1946, 14.–17. xii. 1952 (L. H. Mosse-Robinson) (ANIC); 1 么, Boyd River Crossing, 4100 ft, 12. xii. 1978 (V. J. Robinson) (ANIC); 1 么, Clyde Mt, 2400 ft., 23. i. 1961 (I. F. B. Common & M. S. Upton) (ANIC); 3 么, Ebor, 3. i. 1914, 12. iii. 1940, 17. xi. 1946 (AMS, QM); 1 么, Ebor, 20. i. 1946 (L. H. MosseRobinson) (ANIC); 8 么, 1 乆, Glen Innes, 26. iii. 1913 (ANIC); 1 乆, Gosford, 7. xi. 1896 (MV); 1 么, Kiandra, 10 mls E, 9. i. 1964 (M. S. Upton) (ANIC); 3 么, 1 乆, Manly (AMS); 1 乆, Mt Kosciusko, 3500 ft, 23. i. 1914 (AMS); 1 乆, Mt Kosciusko, 4000 ft, 23. i. 1914 (ANIC); 1 乆, New England Nat. Park, 19. xi. 1954 (C. W. Frazier) (ANIC); 2 乆, Petersham (AMS); 3 么, Point Lookout, New England Nat. Park, 17.–19. iii. 1969 (V. J. Robinson) (ANIC); 1 乆, Stanwell Tops, iii. 1953 (V. J. Robinson) (ANIC); 1 么, Tinderry Mtns, 4000 ft, 5. i. 1963 (I. F. B. Common & M. S. Upton) (ANIC); 2 乆, Sydney (AMS); 3 么, Sydney, ii. (ANIC, WADA); 1 么, Sydney (Helms) (AMS); 1 么, Sydney (A. & F. R. Zietz) (SAM); 1 么, Wahroonga, e.p. 11. x. 1971 (pupated on Kunzea ambigua) (A. B. Rose) (ANIC). Australian Capital Territory: 1 乆, Canberra, 25. xii. 1937 (G. M. Goldfinch) (AMS). Victoria: 6 么, 2 乆, Beaconsfield, xi. 1903, 30. i. 1904, 9. iv. 1904, 28. ii. 1905, 28. xi. 1908 (MV); 1 么, Belgrave, 16. xii. 1943 (Le Souef) (ANIC); 2 么, Blackwood (SAM); 3 么, 5 乆, Clayton, xii. 1915 (W. J. R.) (MV); 2 乆, Dandenong, 21. ii. 1910, 25. xi. 1911 (QM, SAM); 2 么, Fern Tree Gully, 30. xii. 1895 (MV); 2 乆, Frankston, 10. iii. 1930 (A. L. B[rown]) (MV); 7 么, 5 乆, Gisborne, 16. xii.1894, 20. i. 1895, 3. ii. 1895, 5. ii. 1903, , 11. i. 1904, e.l. 2. i. 1906 (2 males, larva on Melaleuca), 29. xi. 1914, 17. xii. 1922 (MV, NSWA, WADA); 1 么, 1 乆, Grampians, xi. 1892 (MV); 2 么, 2 乆, Macedon, i- 1894, 13. i. 1901, 20. i. 1906 (MV); 1 么, Macedon, 6. i. 1913 (G. Lyell) (MV); 1 么, Mitcham, 24. xii. 1946 (A. L. Brown) (MV); 5 么, 6 乆, Moe, xii. 1918, xii. 1945 (C. G. L. Gooding) (ANIC); 4 么, 1 乆, Mordialloc, 15.-18. xi. 1931 (A. L. Brown) (MV); 1 乆, Mt Evelyn, 30. xi. 1933 (A. L. Brown) (MV); 3 么, Mt Buffalo, 1. ii. 1933 (ANIC); 1 乆, Ringwood, 8. xii. 1923 (MV); 2 乆, Wandin, 28. xii. 1891 (MV); 1 么, Wannon Divide, Grampians, 1400 ft, 8. ii. 1950 (at mv light) (N. B. Tindale) (SAM); 2 么, Warburton, i. (MV); 1 么, Wilsons Promontory, 18. i. 1983 (on buds and flowers of Leptospermum juniperinum) (A. N. Andersen) (ANIC); 4 么, 6 乆, Yinnar, 18.–28. xii. 1949, 5.–10. i. 1950 (A. L. Brown) (MV). South Australia: 2 么, 2 乆, Aldgate, 28. xii. 1905, 17. xii. 1939 (F. M. Angel) (SAM); 6 么, Burnside (SAM); 1 么, Inglewood, 17. x. 1926 (J. O. Wilson) (ANIC); 1 么, Lucindale, 15. xi. 1964 (T. Saunders) (SAM); 1 么, Mt Lofty, 1. i. 1906 (F. M. Angel) (SAM). Tasmania: 1 么, Arthur Plains S, 4. ii. 1965 (Neboiss) (MV);

1 么, Cradle Mt, 2000 ft, 30. i. 1925 (QM); 3 么, Freycinet Nat. Park, 28. ii. 1963 (I. F. B. Common & M. S. Upton) (ANIC); 1 么, Gladstone, 1 km SSE, 6. ii. 1983, at light (J. C. Cardale) (ANIC); 1 乆, Hobart (ANIC); 1 乆, Hobart, 18. ii. 1917 (G. H. Hardy) (ANIC); 1 么, Karoola (SAM); 4 么, 1 乆, Kelso (SAM); 4 么, Melaleuca, 12. ii. 1990 (E. S. Nielsen & P. B. McQuillan) (ANIC); 1 么, Strahan, 7. ii. 1925 (QM); 1 么, Zeehan, i. 1924 (G. H. Hardy) (UQIC); 1 么, 1 乆, Zeehan, 4. ii. 1925 (ANIC, QM).

Diagnosis Myrtartona coronias is externally similar to Turneriprocris dolens and Australartona mirabilis sp. n. The former is smaller, has more rounded wings and a blackish brown proboscis (M. coronias yellow). The latter has an epiphysis on the foretibia, a more projected frons, strongly pectinate antennae in the male, the chaetosema is surrounded by white scales and the palpi are at least partly white. The genitalia are also characteristic.

Redescription Male (Pl. 45, Fig. 1). Forewing length: 8.0–9.0 mm. Head, thorax, abdomen and upperside of forewing dark greyish black with distinct bluish green sheen that can be pronounced and almost varnish-like in fresh specimens, densely covered with long and slender scales, hindwing dark grey-brown, almost matt but sometimes with weak blue-green sheen near anal angle; underside of both wings dark greybrown with weak blue-green sheen on forewing apically and on hindwing along costa apically, along vein CuP and at anal angle. Frons slightly projecting, especially dorsally, rounded, broad, 2.0× broader than compound eye in frontal view; proboscis bright yellow; compound eyes black; ocelli of medium size, white; chaetosema dark brown. Antenna with +/- 35 segments, bipectinate from segments 1–25, biserrate from segments 26 to apex; pectinations 2.5× longer than breadth of antennal shaft at segment 5, 2.0× longer at segment 15, apex pointed. Female (Pl. 45, Fig. 2). Similar to male with narrower and more rounded wings. Frons 2.5× broader than compound eye in frontal view. Abdomen long, olive green, not shiny. Antenna strongly biserrate with dentations almost as long as breadth of shaft at segment 15. Male genitalia (Figs 344–346, 356–358). Uncus slender distally, longer than in related species. Tegumen and vinculum slender, saccus small, rounded. Valva

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broad, rectangular, the ventral sclerotization only slightly projecting beyond distal margin of valva, sharply or bluntly pointed. Mediodorsal process strongly developed, longer than tegumen. Transtilla with two small lateral sclerotizations and an extremly narrow bridge-like connection that can be absent in some specimens. Juxta large and rounded, slightly emarginate dorsally. Aedeagus large and stout, approximately 4× longer than broad, slightly tapering caudally. Everted vesica with a long, slender cornutus, pointed apically and with a complicated proximal structure consisting of an invaginated plate. At this invagination a narrow, band-like sclerotization is inserted, forming a joint with the base of the cornutus. If the vesica is fully everted the cornutus can be moved by pulling at this band. In addition to the cornutus and the sclerotized band, the vesica bears a plate-like sclerotization and a small field of minute spines. Female genitalia (Figs 373, 374). Sternum VIII very narrow, connected to the broader tergum VIII by an almost translucent bridge. Ostium sclerotized, ductus bursae tube-like, sclerotized proximally, translucent distally, with a band-like lateral sclerotization running from ostium to insertion of ductus into praebursa; at this insertion there is a group of cornuti (one long and two short); praebursa spherical, translucent, thorn relatively short and surrounded by a ring-like sclerotization; ductus intrabursalis broader with spiraled ribs proximally, more slender and longitudinally folded distally; corpus bursae small, very translucent; ductus seminalis inserting laterally at approximately one-fifth of corpus bursae.

that this was also the foodplant of the larva. As both plants belong to the Myrtaceae, they may be genuine foodplants for larvae of M. coronias.

Remarks It is rather difficult to distinguish males from females simply on habitus and the antenna alone. The male can completely flatten the pectinations along the shaft and, when like this, the antenna looks like that of a female. The more rounded wings and abdomen in the female may not be sufficient as a distinguishing character, especially if the material is in poor condition. The easiest and most certain way to separate males from females without dissection is to examine the frenulum and retinaculum.

Distribution Eastern and south-eastern Australia, from southern Queensland to Tasmania. Only one record from Queensland. It is probable that most of the records from southern Queensland are referable to the externally similar Australartona mirabilis sp. n.

Phenology and bionomics There is one reliable label record of the larval foodplant. Two male specimens from Gisborne (Victoria) deposited in MV were reared from larvae found on Melaleuca (species not mentioned) (Myrtaceae). The larvae were collected in December 1905, the adults emerged on 2 January 1906. One male in the ANIC collection bears a handwritten label ‘Leptospermum juniperinum buds & fls’. As it is a fresh specimen and the label refers to buds as well as flowers it is possible that this male was reared from a larva found feeding on buds and flowers of Leptospermum juniperinum Smith (Myrtaceae) although confirmation is required. There is another male specimen in the ANIC collection reared from a pupa spun on Kunzea ambigua (Smith) Druce (Myrtaceae) but there is no proof

Myrtartona leucopleura (Meyrick) comb. n. Pl. 46, Figs 1, 2; Pl. 61, Fig. 4; Figs 37–40, 112–118, 347–349, 359–364, 375. Procris leucopleura Meyrick, 1886, Proc. Linn. Soc. NSW (2) 1: 792. Lectotype 么 (here designated), AUSTRALIA: New South Wales, Sydney (Macleay) (ANIC) [examined]. Adscita leucopleura (Meyrick); Kirby 1892: 85. Pollanisus leucopleura (Meyrick); Jordan 1907: 10; Hering 1922: 13; Bryk 1936: 118.

TAXONOMY

Pollanisus leucopleurus (Meyrick); Turner 1926b: 442. ‘No available genus’ [sic] leucopleura (Meyrick); Tarmann 1996: 143.

Material examined Queensland: 1 么, 4 乆, Brisbane, 13. v. 1912, 7. iv. 1914 (H. Hacker) (ANIC, QM); 5 么, 1 乆, Glen Aplin, 27. iii. 1940 (L. H. Mosse-Robinson) (ANIC); 1 乆, Round Hill Head W, Seventeen Seventy, 1. xi. 2001 (G. M. Tarmann) (ANIC); 1 么, 7 乆, Stanthorpe, 24. xii. 1929, 27.-28. xii. 1929, 7.-12. i. 1930 (W. B. Barnard) (QM); 1 乆, Stradbroke I., 28. ix. 1906 (ANIC); 1 么, 1 乆, Stradbroke I., 2. x. 1911 (H. Hacker) (QM); 1 么, Toowoomba, 2. xii. 1924 (W. B. Barnard) (QM). New South Wales: 1 么, Berowra, 23. x. 1950 (L. H. MosseRobinson) (ANIC); 1 乆, Bradfield Drive [central Sydney], 22. iii. 1941 (C. E. Chadwick) (NSWA); 1 乆, Brunswick Heads, 28. xii. 1926 (W. B. Barnard) (QM); 1 么, Como, 22. xi. 1931 (MV); 2 么, Como West, 31. iii.–4. iv. 1970 (L. Willan & V. J. Robinson) (ANIC); 1 么, 1 乆, Field of Mars Reserve, 9 km W of Sydney, 11.- 17. xi. 1984 (B. Hacobian) (ANIC); 1 么, Gosford, 7. xi. 1896 (MV); 1 么, 1 乆, Gosford, e.l. 17. xi. 1903 (larva on teatree) (W. W. Fogatt) (NSWA); 1 乆, Hornsby, 25. iii. 1910 (G. Lyell) (MV); 1 么, 1 乆, Hornsby, xi. 1916, 25. xii. 1916 (ANIC, MV); 1 乆, Lane Cove R., 10 km W of Sydney, 17. xii. 1983 (B. Hacobian) (ANIC); 1 么, 1 乆, Manly, 6. xii. 1905, 20. iii. 1907 (AMS); 5 乆, Narara, 28. xi. 1943, 25. iv. 1944, 24. iv. 1949, 7. xi. 1950 (L. H. Mosse-Robinson) (ANIC); 1 么, Nelson Bay, 12. xi. 1960 (I. F. B. Common & M. S. Upton) (ANIC); 2 乆, Nerriga, 12 km NNE, 6. ii. 1975 (I. F. B. Common, E. D. Edwards & M. Story) (ANIC); 1 么, Peats Ridge, 80 km N Sydney, 11. xii. 1983 (B. Hacobian) (ANIC); 1 乆, Pisga Ridge, near Glenbrook, Blue Mtns Nat. Park, 210 m , 18. x.–6. xi. 1987 (malaise trap) (G. R. Brown) (NSWA); 2 么, Roseville, 9. iv. 1904 (MV); 1 么, Sydney N, 6. xii. 1896 (MV); 15 么, Wilton, CSIRO Experimental Farm, viii. 1925, v. 1934, 16. xi. 1972, 17. ii. 1972, 19. x. 1973, 14.-19. xi. 1973, 1. xii. 1973, 14. i. 1974, 24. ii. 1974, 24. iii. 1974, 8. i. 1975, 24. i. 1976, 24. xii. 1976, 9. iv. 1981, 12. i. 1982 (V. J. Robinson) (AMS, ANIC). Australian Capital Territory: 2 么, Jervis Bay, 15. xi. 1918, 16. xi. 1919 (ANIC).

Diagnosis Myrtartona leucopleura is unmistakable: it is the only Australian zygaenid with a bright white lateral line along the abdomen.

Redescription Male (Pl. 46, Fig. 1). Forewing length: 8.0–8.5 mm.

Head, thorax, abdomen ground colour dark brownish grey. Caudal part of chaetosema, coxa of foreleg, lateral part of thorax, lateral parts of abdominal segments 2–7 and ventral part of segment 7 covered with pure white scales. Very often frons, parts of compound eyes, labial palps and legs also white. Frons slightly projected, especially dorsally, approximately 2.0× broader than compound eye in frontal view; proboscis yellow; ocelli of medium size, white. Antenna with 38–44 segments, shortly bipectinate almost to apex, the pectination approximately 2.0× longer than breadth of antennal shaft at segment 5 and 15, slightly tapering towards apex and only the last 6–7 segments biserrate. Forewing upperside dark grey-brown, underside slightly paler; hindwing upperside blackish grey, paler proximally, underside similar, slightly paler. Female (Pl. 46, Fig. 2). Similar to male, of same size but with narrower and more rounded wings. Frons only slightly broader than in male, of same colour but with a distinct bluish green tinge. Abdomen upperside with yellowish green sheen in fresh specimens, underside with segments 2–6 laterally and ventral part of segment 6 white. Antenna strongly biserrate. Male genitalia (Figs 347–349, 359–364). Uncus short, triangular, only the distal end strongly sclerotized. Tegumen and vinculum very slender, saccus broader, rounded. Valva slender, the ventral sclerotization very strong, down-curved, significantly longer than distal margin of valva, rounded distally. Mediodorsal process slender, longer than tegumen, the distal part of same length as the sideways-pointing base (sideways in natural position), the upward pointing part of the forked base short. Transtilla weakly sclerotized, slender. Juxta large, weakly sclerotized, rounded. Aedeagus large, stout, approximately 4× longer than broad, slightly broader proximally. Everted vesica with a long, slender cornutus that has the same kind of base as described in M. coronias (p. 172) and is movable; the slender band that inserts at the base of the cornutus is very long; in addition to this band there is a large sclerotized plate of characteristic shape. Female genitalia (Fig. 375). Sternum VIII narrow, connected to the broader tergum VIII by an almost translucent band. Ostium sclerotized, ductus bursae tube-like, completely sclerotized; there is a group of thorn-like spines at insertion of ductus bursae into the translucent praebursa, one longer and several shorter ones; the dagger-like structure large, larger

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proximally than distally, surrounded by a sclerotized ring; ductus intrabursalis broader posteriorly than anteriorly; corpus bursae small, very translucent; ductus seminalis inserted laterally at orifice of corpus bursae opposite insertion of ductus intrabursalis into corpus bursae.

Phenology and bionomics According to label data this species is bivoltine. Two old specimens from Gosford (New South Wales) were reared from larvae found on ‘teatree’ (Leptospermum sp.) (Myrtaceae). On 1 November 2001 at 11.45 a.m. one female was observed by the author resting on a leaf of a Eucalyptus sp. (Myrtaceae) (Pl. 61, Fig. 4). The locality is a coastal rainforest at the rocky seashore west of Round Hill Head near the town of Seventeen Seventy on the Capricorn Coast of Queensland (Pl. 63, Fig. 2). It was a fine, hot and sunny day, almost without wind, the temperature was 33.5°C. The female was resting on the leaf in the shade about 1.5 m above the sandy ground, slowly moving its antennae. Some ring-flash photos were taken but then the female dropped to the ground, moved about 5 cm, turned on its back and remained in that position pretending to be dead (exhibiting thanatosis). The dark and white pattern of the abdominal underside made the specimen almost disappear in the strong contrast of light and shadow on the ground. The female was taken for eggs and put into a container with leaves of the Eucalyptus sp. on which it had been found, but died on the same evening without having oviposited. It had been offered several possible nectar plants and leaves of two other Eucalyptus spp. and a Melaleuca sp. from a garden. The locality was carefully examined but no other specimen could be found. Moreover, neither a Melaleuca nor a Leptospermum species could be found. It is therefore assumed that the larval hostplant at that locality is Eucalyptus. Although the leaves of the eucalypts were also carefully examined no eggs could be found. However, only the lower leaves of the trees could be reached. The habit to drop to the ground when the species is disturbed was also observed in Myrtartona rufiventris (see p. 176), which is closely related to M. leucopleura.

Distribution Queensland, New South Wales, Australian Capital Territory (Jervis Bay only).

Myrtartona rufiventris (Walker) comb. n. Pl. 47, Figs 1, 2; Figs 350–352, 365–370, 376, 377. Procris rufiventris Walker, 1854, List Specimens lepid. Insects Colln Br. Mus. 1: 110. Lectotype 么 (here designated), AUSTRALIA: Western Australia, Swan R. [Perth] (BMNH) [examined]. Hestiochora rufiventris (Walker); Meyrick 1886: 790; Jordan 1907: 10; Turner 1926b: 442; Bryk 1936: 116; McFarland 1970: 348 (egg); McFarland 1973: 219; McFarland 1979: 62; Common 1990: 298. Neoprocris rufiventris (Walker); Alberti 1954: 300. ‘No available genus’ [sic] rufiventris (Walker); Tarmann 1996: 143.

Material examined Victoria: 1 么, ‘Mallee District, Victoria’ (MV). South Australia: 1 么, Belair (SAM); 1 么, Brookfield Conservation Park, Blanchetown, 11. xii. 1990 (E. S. Nielsen) (ANIC); 1 么, Ceduna, 35 mls E, 28. xi. 1958 (I. F. B. Common) (ANIC); 2 么, Fowler Bay, 25. x. 1965 (N. B. Tindale) (SAM); 1 乆, Fulham, Holmfirth, v. 1907 (J. W. Melor) (SAM); 1 乆, Macclesfield, xii. 1890 (SAM); 1 乆, Mt Hall, Eyre Peninsula, 6. xii. 1968 (N. McFarland) (SAM); 2 乆, Woodside (SAM). Western Australia: 1 么, Syntype, without locality label (BMNH); 5 么, 7 乆, Albany, 14. ii. 1926, 21. ii. 1926, 2.–6. iii. 1926 (W. B. Barnard) (AMS, ANIC, MV, QM); 3 乆, Booanya, xi. 1932 (Mrs Baesjou) (SAM); 1 乆, Bunbury (L. J. Newman) (WADA); 2 (males), Caiguna, 1. xii. 1968 (N. McFarland) (SAM); 7 么, Dundas Rock, S of Norseman, 12. i. 1993 (E. D. Edwards & E. S. Nielsen) (ANIC); 2 么, 1 乆, Eucla W (N. McFarland) (ANIC); 27 么, 21 乆, Eucla, 43 mls W, 2. xii. 1968, ([imagines] on Melaleuca) (N. McFarland & N. B. Tindale) (AMS, ANIC, BMNH, SAM); 57 么, 17 乆, Eucla, 24 mls W, 3. xii. 1968 ([imagines] on

TAXONOMY

Melaleuca) (N. McFarland) (AMS, ANIC, SAM); 2 么, Jimberlana Hill, 1 km W Norseman, 11. i. 1993 (E. D. Edwards & E. S. Nielsen) (ANIC); 1 么, Kalbarri, 4 km S, 12. xi. 1971 (N. McFarland) (ANIC); 1 乆, Mandurah, 74 km S, 18. i. 1987 (G. & A. Daniels) (UQIC); 2 (males), Moir’s Rock, 42 km NNW Salmon Gums, 2. i. 1986 (G. & A. Daniels) (UQIC); 2 么, 1 乆, Salmon Gums, 8 km E, 2. i. 1986, 5. i. 1987 (G. & A. Daniels) (UQIC); 1 乆, Salmon Gums, 3 km NE, 10. i. 1993 (E. D. Edwards & E. S. Nielsen) (ANIC); 3 么, Yule Brook Reserve, Kenwick, 7. i. 1986 (M. J. Lewis) (ANIC).

Diagnosis Myrtartona rufiventris is well characterised by its remarkable habitus and genitalic characters. Only two other zygaenids in Australia, Myrtartona mariannae sp. n. and Onceropyga pulchra sp. n., also have an orange or yellow abdomen. Both are smaller and their wings are more rounded. Myrtartona mariannae sp. n. has a whitish yellow abdominal underside whereas the underside of the abdomen of M. rufiventris is dark. The female of Onceropyga pulchra sp. n. has the typical abdominal hair-tuft of the Pollanisus-Hestiochora-group, a character that is absent in Myrtartona. There are also differences in the genitalia.

Redescription Male (Pl. 47, Fig. 1). Forewing length: 9.0–10.5 mm. Head densely covered with dark grey scales; breadth of frons 1.5× broader than compound eye in frontal view; labial palps short, white; proboscis yellowish brown; compound eye black, surrounded by white scales; ocelli large (larger than in related species); chaetosema lighter in colour posteriorly. Antenna bipectinate with broad and very densely scaled antennal shaft, pectinations short, approximately 3× longer than breadth of antennal shaft at segments 5 and 15, short distally, apex pointed; number of segments 37–39. Thorax dark brownish grey dorsally, with white scales laterally and ventrally. Forewing triangular, costa almost straight, blackish brown with scattered whitish scales in fresh specimens, giving the wing a farinose appearance. Hindwing elliptical, dark grey-brown, paler proximally. Dorsal abdominal segments 2–8 bright orange, segments 1 and 9 dark, all segments dark brown, with scattered white scales ventrally. Female (Pl. 47, Fig. 2). Forewing length: 7.5–8.5 mm. Very similar to male but smaller. Anterior part of head slightly broader. Antenna shortly bipectinate.

Male genitalia (Figs 350–352, 365–370). Uncus short, strongly sclerotized and pointed distally. Tegumen hood-like dorsally with a triangular elongation surrounding the tuba analis lateroventrally, the tips of the latter nearly touching each other ventromedially, forming a transtilla; mediodorsal process strongly developed, slightly longer than tegumen, pointed distally, bearing a group of spines at its tip and with some backward pointing setae just below this. Vinculum V-shaped, strongly sclerotized. Juxta double-lobed laterally with a proximal groove and a distal cone medially. Valva weakly sclerotized dorsally but with very strong sharks-tooth like sclerotization ventrally, the pointed apex of this ventral part longer than translucent outer margin of valva. Aedeagus large, very stout; on the everted vesica a broad, sharp cornutus of typical shape with a spiny surface and a large sclerotized plate. Sternum VIII unsclerotized except for proximal margin. Female genitalia (Figs 376, 377). Sternum VIII reduced to a very narrow clasp that is connected to the broader tergum VIII, forming a narrow ring; apophyses anteriores very short. Papillae anales large, apophyses posteriores as long as diameter of papillae. Ostium without any sclerotizations; ductus with a very short tube-like translucent proximal part leading into a large translucent praebursa; thorn strongly developed, surrounded by a large sclerotized plate; ductus intrabursalis short, translucent; corpus bursa small, spherical, translucent.

Phenology and bionomics Adults of M. rufiventris were collected by N. McFarland and N. B. Tindale near Eucla in Western Australia. The following field observations were made on 2 December 1968 by Noel McFarland at that locality, and the original hand-written notes were kindly sent by him to the author for publication in this book. The habitat is a rocky limestone area with fine powdery soil in some parts. The dominant shrubs were Eucalyptus spp., Melaleuca lanceolata Otto and Atriplex paludosum R. Br., a ‘saltbush’ with whitish leaves. Freshly emerged adults were observed in the late afternoon on Atriplex bushes, quite well camouflaged by their greyish white-brown colour. A long series of adults, mostly males, was collected between 5.00–6.30 p.m. by searching these bushes. The females were mostly resting on the larval foodplant, Melaleuca lanceolata Otto, where there were also numerous males. The adults were very cryptic on the foodplant (though much more so on Atriplex); the orange abdomen was not visible

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in resting specimens. Three specimens had been caught by a spider of the genus Oxyopes (Oxyopidae) (det. D. Lee, 1970) other individuals of which were later observed hunting the zygaenids, even when the latter were on the wing, by jumping while running. The females were not observed flying at all but only resting on the larval foodplant, in copula, or ovipositing. Even when disturbed the females did not take to the wing; when severely disturbed they dropped to the ground. The males were active in bright sunshine during the hottest part of the day (1–2 p.m.), flying weakly over short distances. When netted or struck at they feigned death. After landing, the males were seen to run a few steps with a rather short, quick motion before resting. The scent given off by an aggregation of females was noted as similar to that of ladybirds (Coleoptera, Coccinellidae) and can be detected by man.

Remarks According to McFarland, this scent is an attractant for the males. However, it is not certain that it is really produced by the females since it is also possible that it comes from the arriving males. It could also be produced by the abdominal lateral glands; the function of which is still unknown. The females move forward while ovipositing, usually laying their eggs on the narrow leaves of the host-plant in short rows of 5–10 eggs. In some cases these rows are longer and may be very long (up to 107 eggs in one row, according to an observation by McFarland). Oviposition takes place in the early afternoon. Generally, the eggs do not touch each other, although sometimes they touch or overlap slightly (see McFarland 1970: 349, 1972: 219). The eggs are opaque, smooth, pale yellow-cream. They are not covered by ‘spines’ or scales (see p. 36; Figs 129, 130).

Distribution Western Australia, South Australia, Victoria

Myrtartona mariannae sp. n. Pl. 48, Fig. 1; Figs 353–355, 371, 372.

Material examined Holotype 么, Queensland: Milmerran, 4. iv. 1946 (J. Macqueen) (ANIC). Genitalia slide Z 3198.

Diagnosis Based on the characters of the male genitalia P. mariannae sp. n. is closely related to M. rufiventris. However, externally as well as in the genitalic characters there are significant differences. Myrtartona rufiventris is larger, has a more rounded hindwing and a dark abdominal underside (yellowish white in M. mariannae). In the male genitalia M. rufiventris has a shorter and more strongly sclerotized uncus, the mediodorsal process is shorter, the ventral sclerotization of the valva is longer distally, the tegumen is more V-shaped and more strongly sclerotized, the aedeagus is larger and shorter and the cornutus on the vesica has a completely different size and shape. Onceropyga pulchra sp. n. is also similar in external appearance but has a different wing-shape, a bright yellow abdomen and completely different genitalia.

Description Male (Pl. 48, Fig. 1). Forewing length: 8.5 mm. Head greyish brown dorsally, frons dark grey dorsally, light grey ventrally; frons of medium breadth, ca. 1.2× broader than the black compound eyes in frontal view, protruding beyond compound eye in lateral view, more so dorsally, labial palps in type missing; proboscis yellow; ocelli small, white, distance from compound eye ca. 1.5× diameter of ocellus; chaetosemata grey; both antennae broken distally in type (reconstructed in painting Pl. 48, Fig. 1), greyish, segments 1–25 bipectinate, rest missing, the pectinations of medium length, 3–4× longer than breadth of shaft in dorsal view at segment 10, slightly tapering distally (as far as visible), sensory hairs very short. Thorax uniform greyish brown (as far as can be seen because ventral part almost without scales); legs dark grey. Abdominal segments 1 and 2 greyish brown, 3–8 yellowish ochreous dorsally, whitish ventrally, abdominal end with a few dark brown scales. Wings elongate, forewing slender, more triangular, hindwing almost rectangular but rounded apically; forewing upperside and underside greyish brown; hindwing dark grey with a brownish tinge,

TAXONOMY

slightly paler medially, fringe of forewing unicolorous with wing, mixed with whitish scales in hindwing. Male genitalia (Figs 353–355, 371, 372). Uncus short, not very strongly sclerotized, pointed distally. Tegumen hood-like, tuba analis surrounded by a translucent transtilla; mediodorsal process strongly developed, longer than tegumen, pointed distally, bearing a group of spines at its tip and with some backward pointing setae just below this. Vinculum broad, U-shaped, weakly sclerotized. Juxta doublelobed laterally with a slender distal prolongation medially. Valva weakly sclerotized dorsally but with strong shark’s-tooth like sclerotization ventrally, the pointed apex of this ventral part slightly longer than translucent outer margin of valva. Aedeagus short and stout, 4× longer than broad; on the everted vesica a short, slender, crest-like cornutus with a spiny surface. Sternum VIII unsclerotized except for proximal margin. Female. Unknown.

Phenology and bionomics Unknown. All related species live on Myrtaceae.

Distribution Only known from the type-locality, Milmerran, in south-eastern Queensland.

Etymology This species is dedicated to Marianne Horak, ANIC, CSIRO Canberra, one of the leading Australian lepidopterists, with thanks for supporting this work, for providing important information and discussions and for her friendship.

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345

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347

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349

Figs 344–346. Male genitalia of Myrtartona coronias, NSW, Blackheath. Gen. slide no. Z 2960. 344: Overview; 345: valva; 346: dorsal part with ‘Myrtartona finger’. Figs 347–349. Male genitalia of Myrtartona leucopleura, NSW, Nelson Bay. Gen. slide no. Z 2977. 347: Overview; 348. valva and ‘Myrtartona finger’; 349: dorsal part with ‘Myrtartona finger’.

TAXONOMY

350

351

352

353

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355

Figs 350–352. Male genitalia of Myrtartona rufiventris, WA, Eucla. Gen. slide no. Z 2975. 350: Overview; 351: valva; 352: dorsal part with ‘Myrtartona finger’. Figs 353–355. Male genitalia of Myrtartona mariannae, holotype, Qld, Milmerran. Gen. slide no. Z 3198. 353: Overview, (twice size of Fig. 350); 354: valva and ‘Myrtartona finger’; 355: dorsal part with ‘Myrtartona finger’.

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356

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359

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361

Figs 356–358. Male genitalia: aedeagus of Myrtartona coronias, NSW, Blackheath. Gen. slide no. Z 2960. 356: Overview; 357: detail, cornutus; 358: detail, field of spines on vesica. Figs 359–361. Male genitalia: aedeagus of Myrtartona leucopleura, NSW, Nelson Bay. Gen. slide no. Z 2977. 359: Overview; 360: detail, movable base of cornutus; 361: detail, cornutus.

TAXONOMY

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367

Figs 362–364. Male genitalia: aedeagus of Myrtartona leucopleura, Qld; Glen Aplin. Gen. slide no. Z 2978. 362: Everted vesica; 363: detail, sclerotization on everted vesica; 364: detail, movable base of cornutus. Figs 365–376 Male genitalia: aedeagus of Myrtartona rufiventris, WA, Eucla. Gen. slide no. Z 2975. 365: Overview; 366: detail, cornutus; 367: detail, field of spines on vesica.

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372

Figs 368–370. Male genitalia: aedeagus of Myrtartona rufiventris, WA, Yule Brook Reserve. Gen. slide no. Z 2973. 368: Overview; 369: detail, cornutus; 370: detail, field of spines on vesica. Figs 371–372. Male genitalia: aedeagus of Myrtartona mariannae, holotype, Qld, Milmerran. Gen. slide no. Z 3198. 371: Overview; 372: detail, cornutus.

TAXONOMY

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377

Fig. 373. Female genitalia of Myrtartona coronias: ag, accessory gland; an, antrum; cb, corpus bursae; di, ductus intrabursalis; ds, ductus seminalis; dsp, ductus spiralis; gsbl, glandula sebacea, bulla; gscd, glandula sebacea, common duct; gsg, glandula sebacea, gland (one gland cut off); ob, ostium bursae; oc, oviductus communis; pb, praebursa; pbth, praebursa-thorn; sp, spiracle; tVII, 7th tergite; tVIII, 8th tergite; ut, utriculus. Vic., Dandenong. Gen. slide no. Z 2962. Fig. 374. Female genitalia of Myrtartona coronias, detail, praebursa-thorn, NSW, Blackheath. Gen. slide no. Z 2963. Fig. 375. Female genitalia of Myrtartona leucopleura, detail, praebursa-thorn, NSW, Narara. Gen. slide no. Z 2979. Fig. 376. Female genitalia of Myrtartona rufiventris, WA, Eucla. Gen. slide no. Z 2962. Fig. 377. Female genitalia of Myrtartona rufiventris, detail, praebursa-thorn, WA, Mandurah. Gen. slide no. Z 2974.

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TAXONOMY

Genus Australartona gen. n. Type-species: Australartona mirabilis sp. n.

Diagnosis Species of medium size, body and wings unicolorous, without shining metallic scales (Pl 48, Fig. 2; Pl. 49, Figs 1, 2; Pl. 50, Fig. 1). Male antenna with long curved pectinations with very long setae. Fore tibia with large epiphysis. Lateral evaginations on abdominal segment II and VII, those on segment II enlarged. Male genitalia with ‘Artona finger’. Female without abdominal hairtuft. Larva on Poaceae. Head (Figs 41–48) Frons strongly projecting, rounded; labrum triangular medially, pilifer enlarged, pocket-like, with long and slender scales directed medially; maxillary palps small; proboscis with significantly longer spines laterally at base than on the rest of its surface; labial palps porrect distally, slightly extending beyond anterior margin of head; compound eyes large, with a narrow eyelash ventrally consisting of only one layer of scales; chaetosemata of typical artonoid form. Antennal shaft compressed dorsoventrally, with very long pectinations in male, the pectinations curved, with very long setae giving the antenna a cobweb-like appearance; antenna biserrate in female, dentations also curved inwards; the antenna scaled dorsally but completely unscaled ventrally, the ventral surface consisting of a large number of upright, strongly sclerotized crests and ribs, the distal segments of the antenna covered with setae ventrally (Figs 45–48). Thorax and wings Fore-tibia with large epiphysis equal in length to tibia. Tibial spurs 0–2–2, no single medial spur on hind tibia. In the male, wings broader and forewing triangular, in the female more rounded and slender; medial stem absent in forewing, present in hindwing but only distally, forewing with the crossvein at distal end of cell reduced or often absent, all veins free from cell, R2+R3 connate or shortly stalked; hindwing with very variable venation, often asymmetrical between the two wings, Sc touching anterior margin of cell at one point, RR+M1 fused, M3+CuA1 stalked or connate. Frenulum in male a strong, single spine, three bristles in female; retinaculum a strong hook at base of Sc in male, a number of upright scales at base of CuP in female.

Pregenital abdomen (Figs 430, 431) Artonoid lateral evaginations on segment 2 and segment 7, the anterior evagination on segment 2 very large, more than twice as large as the posterior one that is of normal size. Sternum II with a pair of long, very slender apodemes, slightly curved distally, lateral sternal rod almost invisible; sclerotization of segment with lateral elongations anteriorly. Tergum II with triangular anteriorly pointed sclerotization on caudal half of segment. Sternum VIII in male reduced to a narrow rectangular distal plate that is approximately 5× longer than broad. Tergum VIII in male tapering distally, covering almost the whole segment. Sternum VII in female covering the whole segment. Female without abdominal hair-tuft. Male genitalia (Figs 378–384) Uncus slender, of normal length, approximately as long as tegumen. Tegumen slender, weakly sclerotized. Vinculum very slender, weakly sclerotized, saccus small. Valva with a strongly developed, long ‘Artona’-finger (see p. 52) that inserts ventrodistally; sacculus not strongly pronounced; an oval, disc-like sclerotization bearing a group of very long scales and setae, probably representing an androconial organ, dorsal to sacculus; where a pulvinus is developed in other genera, e.g. Pollanisus, there is an oval sclerotized plate similar to a pulvinus but without setae, slightly dorsad of this a finger-like upcurved process with pulvinus-like setae distally (this may replace the pulvinus functionally); transtilla absent; juxta a large plate. Aedeagus small, long, slender, ductus ejaculatorius inserting at proximal end dorsally at about one-fifth of length; a strongly sclerotized spine fused to wall of aedeagus (as in Hestiochora). Female genitalia (Fig. 385) Sternum VIII sclerotized laterally, almost translucent medially; tergum VIII band-like; apophyses anteriores absent except for a translucent edge; papillae anales large with long setae, apophyses posteriores short and slender. Ductus bursae and corpus bursae translucent, no praebursa, the posterior end of the ductus bursae forms a sclerotized antrum, the anterior part has a distinctive crumpled appearance and can be described as approximately two turns of a spiral; ductus seminalis inserting into ductus bursae slightly proximal to its insertion into the corpus bursae; glandula sebacea with a long and slender, proximal common duct leading into a tubelike enlargment, with two very long and slender ducts leading into spherical bulbs distally.

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Material examined

See Australartona mirabilis sp. n.

Holotype 么, New South Wales: Mt Dromedary, 30 m below top, 770 m , 36°18.377´ S 150°01.701É (GPS), 3. xii. 1989 (G. M. Tarmann & E. S. Nielsen) (ANIC).

Remarks Australartona gen. n. forms the sister group to Homophylotis and Pseudoamuria, adapted to temperate habitats. All three genera form a sister group to the Asian artonoid genera with a single medial tibial spur on the hindleg (Artona, Amuria, Palmartona and related genera). They all feed on monocotyledons. The posterior end of the ductus bursae, which forms a sclerotized antrum, and the anterior part, which has a distinctive crumpled appearance and can be described as approximately two turns of a spiral, are good autapomorphies for Australartona. The development of an ‘Artonafinger’ in the male genitalia is a clear synapomorphy between Australartona and the other genera of the Artona-group. Australartona also shares with these genera the well-developed lateral abdominal bulbs on segments 2 and 7 and the large fore-tibial epiphysis. The single medial spur on the hind-tibia that is present in most Artonini is absent in Australartona and the wing venation is reduced; these characters can be interpreted as secondary losses. The aedeagus bears a long, sclerotized spine that is fused to its wall in exactly the same way as in Hestiochora. The absence of a praebursa is more likely to be a reduction than a plesiomorphy because in almost all Artonini a well-developed praebursa is present (except in Hestiochora, Turneriprocris and a few other genera) (see also p. 51). The male antenna is very similar to that of Pseudoamuria and Homophylotis, especially in the long pectinations that bear very long setae. Finally, the external appearance of the type-species, Australartona mirabilis sp. n., is very similar to that of Myrtartona coronias and all the material examined in collections was misidentified as the latter.

Distribution New South Wales, southern Queensland.

Australartona mirabilis sp. n. Pl. 48, Fig. 1; pl. 49, Figs 1, 2; Pl. 50, Fig. 1; Figs 41–48, 378–384, 385.

Paratypes New South Wales: 2 么, Barrington Tops, 30. xii. 1978 (B. Hacobian & N. Goodman) (ANIC); 1 么, Brown Mt, 27. xi. 1994 (E. S. Nielsen, E. D. Edwards & M. Kozlov) (ANIC); 6 么, 2 乆, Mittagong, 30. xii. 1927 (G. M. Goldfinch) (AMS); 6 (males), 5 (females), Mittagong, Mt Gibraltar, summit plateau, 15. xii. 2003 (G. M. Tarmann) (TLMF); 8 么, 1 乆, Mt Dromedary, 28. xii. 1918 (AMS, ANIC); 1 乆, Mt Dromedary, 1000 ft., 24. xi. 1965 (I. F. B. Common & M. S. Upton) (ANIC); 19 么, 1 乆, Mt Dromedary, 30 m below top, 770 m , 3. xii. 1989 (G. M. Tarmann & E. S. Nielsen) (ANIC, TLMF); 6 么, 1 乆, Mt Wilson, 9.-10. xii. 1958 (H. Thirkell) (ANIC); 1 么, 2 乆, Robertson, 8. i. 1943 (V. J. Robinson) (ANIC). Queensland: 1 么, Brisbane, 4. iv. 1960 (E. A. Bernays) (UQIC); 1 么, ‘N. Park’ [Lamington Nat. Park, O’Reillys Guest House], 3000 ft, 4. ii. 1925 (QM).

Diagnosis Externally very similar to Myrtartona coronias but with an epiphysis on the foretibia, white scales around the eyes and much longer antennal pectinations in the male. There is no other similar species known from Australia.

Description Male (Pl. 48, Fig. 1; Pl. 49, Fig. 1, Pl. 50, Fig. 1). Forewing length: 8.0–8.5 mm. Head, thorax, abdomen and upperside of forewing densely covered with deeply shiny scales, giving the body and forewing a varnish-like, non-metallic appearance, from olive-green to blue-green, with a purple tinge, the forewing often strongly suffused with brown; upperside of hindwing and underside of both fore- and hindwings dark greyish brown with variable bluish sheen. Frons olive brown, with a white ventral edge, 2.0× broader than compound eye in frontal view; proboscis yellow; labial palps white (at least basal segment), pointed distally; compound eyes surrounded by white scales; ocelli small, white; chaetosema dark brown. Antenna shining bluish green, with 35–39 segments, length of pectinations almost 10× longer than breadth of shaft at segment 10, shorter distally, the last 8–9 segments strongly biserrate, apex rounded. Forewing without medial stem, all veins free from cell. Hindwing with medial stem distally, at one point Sc touches the anterior margin of cell, RR+M1 fused, M3+CuA1 stalked.

TAXONOMY

Female (Pl. 49, Fig. 2). Similar to male with slightly narrower and more rounded wings. Frons 2.5× broader than compound eye in frontal view. Antenna strongly biserrate. Male genitalia (Figs 378–384). Uncus slender, of normal length, approximately as long as tegumen, triangular basally, slender distally and strongly constricted, hook-like pointed apically. Tegumen and vinculum slender, weakly sclerotized, saccus a round disc connected to vinculum by an almost translucent connection (this detaches very easily when spreading open the valvae or when separating the valva from the tegumen and vinculum). Valva bearing complex structures; its basic form is rectangular, slightly longer dorsally, ventral sclerotization broad, central part translucent and strongly folded, dorsal sclerotization narrow; a strongly developed, long ‘Artona-finger’ (see p. 52) that inserts ventrodistally and is only slightly shorter than length of valva is present, this finger is broad and covered with setae basally, is very slender distally and without an apical hairbrush (present in most Artona and Amuria species); sacculus weakly developed, with a group of setae; disc-like sclerotization dorsad of sacculus with very long hairs; pulvinus-like process well developed; juxta large, tongue-shaped, with a spiny distal part. Aedeagus small, long, slender, approximately 10× longer than broad; a long, strongly sclerotized, distally down-curved, apically sharply pointed spine is fused with dorsal part of aedeagus (as in Hestiochora); vesica seminalis not eversible, covered with minute triangular spines. Female genitalia (Fig. 385). Sternum VIII strongly sclerotized laterally but almost translucent medially, connected to the broader, band-like tergum VIII by a slender connection; apophyses anteriores absent except for a translucent edge; papillae anales large with long setae, apophyses posteriores short and slender. Ostium sclerotized ventrally, translucent dorsally, antrum short, pipe-bowl shaped, weakly sclerotized; ductus bursae long, slender, translucent, S-shaped, with strong transverse folds; corpus bursae translucent, bag-like.

Phenology and bionomics Australartona mirabilis is a weak flyer. Males and females were observed on Mt Dromedary (New South Wales) between 770 and 790 m, just below and on the summit on 3 December 1989 (G. M. Tarmann, E. S. Nielsen & M. Shaffer) (Pl. 64, Fig. 4). During the middle of the day (11.45 a.m.–14.00

p.m.) numerous males and two females were observed on the wing and resting on grasses in a small clearing in the mountain forest where Pomaderis apetala Labill. (Rhamnaceae), Hakea salicifolia Meissner (Proteaceae), Doryphora sassafras Endl. (Monimiaceae), Senecio sp. (Asteraceae), Coprosoma sp. (Rubiaceae) and several unidentified bushes and herbs were growing. One of the two females taken from a grass stem began to oviposit on a piece of paper in a glass tube shortly after capture. The plants offered to the female for ovipositing were Hibbertia obtusifolia, Leptospermum myrsinoides, Eucalyptus sp. and Melaleuca sp. However, the female ignored all of these plants. On 5 November 2001 the locality was revisited (G. M. Tarmann & E. D. Edwards). It was a cloudy misty day and no specimens were found. Nevertheless, a comprehensive survey was undertaken to find a possible larval host-plant. We came to the conclusion that the most likely larval hostplant is Tetrarrhena juncea R. Br. (Poaceae) (Pl. 62, Fig. 6) as none of the other known host-plants of Australian Artonini was found except Eucalyptus spp. Poaceae (especially bamboo) are known as host-plants for larvae of many Asian Artonini and the appearance of the climbing Tetrarrhenia with its short and hard lanceolate leaves and the strongly branched stems in fact somehow reminds one of mountainous tropical bamboo-like grasses. Moreover, all Artonini known so far that feed on Poaceae have a foretibial epiphysis, two pairs of lateral abdominal glands and a well-developed ‘Artona-finger’ in the male genitalia as in Australartona mirabilis sp. n. The biotope can be described as a southern temperate mountain rainforest. That means that the place is very humid and most of the humidity is coming from the drizzling mist. The habitat is rocky, the rocks are of dark volcanic type and heat up very strongly in sunshine. On 15 December 2003 the author visited Mt Gibraltar at Mittagong (New South Wales). On the summit plateau Tetrarrhenia or a similar grass could be found. Several specimens of A. mirabilis sp. n. were on the wing in bright sunshine nectaring on the yellow flowers of Helichrysum spp. (Asteraceae). Although several females were followed in the hope to observe oviposition no eggs were laid by these females during the two hours of observation (12.00 noon–14.00 p.m.). One female was put into a box with some grass. It laid some eggs on the wall of the box on the next day. The larvae emerged after 25 days that is extremely long for a Procridinae (5–8 days in Pollanisus and 8–15 days in most Palaearctic species).

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Egg Elliptoid, subquadrate, whitish green, with slightly rough surface; length 0.5 mm, breadth 0.4 mm. The eggs are deposited separately in small groups (Fig. 128). Duration of egg stage: 25 days. Larva First instar (L1). Length 1.0–1.1 mm, white, head dark brown, eyes black, prothoracic shield large, light brown, with long, light setae; anal comb fully developed, blackish brown, with short dentations that are longer medially; thoracic legs long, light brown, abdominal prolegs white. Setal combination formula of first abdominal segment (sensu Efetov and Tarmann 1999; Efetov et al. 2000; Efetov 2001; Efetov, Mollet and Tarmann 2003): D: 1d; SD 1d; L: 2l (Fig. 131) [D = dorsal setae, SD = subdorsal setae, L = lateral setae; d = dark, stiff seta (like a spine), l = light, flexible seta]. The setal combination of the third abdominal segment of the first instar larva of A. mirabilis is the same as in the first abdominal segment and corre-

sponds with that figured by Tothill, Taylor & Paine (1930: 85, fig. 20) for Levuana iridescens BethuneBaker, 1906, the coconut moth of Fiji. The L1 larvae are very active and walk long distances. Immediately after hatching they first eat parts of the shell of the eggs. As the original laval host-plant or any allied species was not available in Austria a small decorative bamboo grass from a greenhouse was offered to the larvae. They immediately started to feed producing small rectangular grooves by scratching on the parenchyma of the plant. One larva usually does not make more than 2–3 holes in one leaf and continuously changes from one part of the host-plant to another. Unfortunately the plant had been treated by a systemic insecticide and the larvae died after a few days.

Distribution New South Wales, southern Queensland. Apart from a few records, this species has always been observed on mountains.

378

379

380

Fig. 378. Male genitalia of Australartona mirabilis, paratype, NSW, Barrington Tops. Gen. slide no. Z 2964. Figs 379–380. Male genitalia: valva of Australartona mirabilis, paratype, NSW, Mt Dromedary. Gen. slide no. Z 2965. 379: Detail, distal part; 380: detail, proximal part.

TAXONOMY

381

382

383

384

385

Figs 381–382. Male genitalia: valva of Australartona mirabilis, paratype, NSW, Mt Wilson. Gen. slide no. Z 2949. 381: Detail, distal part, with coremata and pulvinus-like process; 382: detail, central part. Figs 383–384. Male genitalia: aedeagus of Australartona mirabilis, paratype, NSW, Robertson. Gen. slide no. Z 2947. 383: Overview; 384: detail. Fig. 385. Female genitalia of Australartona mirabilis, paratype, NSW, Robertson. Gen. slide no. Z 2948.

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TAXONOMY

Genus Homophylotis Turner Homophylotis Turner, 1904, Trans. Proc. R. Soc. S. Aust. 28: 243. Type species: Homophylotis thyridota Turner, 1904: 243, by monotypy. [Australia, Queensland.] Homophylotis Turner; Jordan 1908: 46; Turner 1926b: 440; Bryk 1936: 257; Alberti 1954: 281; Common 1970: 830; Tarmann 1996: 143.

Diagnosis Small species with short, elongate, triangular forewings and rounded hindwings (Pls 51–54). Hindwings with a translucent central part covered only with a few upright scales. Head, thorax and abdomen dark dorsally, white or yellow ventrally. Antenna with a conspicuous white spot subapically. Male antenna with asymmetrical pectinations. Head (Figs 49–56) Frons strongly projecting, rounded; labrum small, rounded; pilifer with long, slender scales pointing medially; maxillary palps small; proboscis of normal length, yellow; labial palps long, curved upwards, reaching anterior edge or slightly extending beyond anterior margin of head; compound eye of medium size, with a narrow eyelash ventrally; ocelli small, chaetosemata very slender. Male antenna bipectinate, with long, asymmetrical pectinations, the distal segments weakly biserrate, the pectinations with very long setae; female antenna biserrate; antenna of male and female dark with a conspicuous white dorsal spot close to apex. Thorax and wings Legs very long, fore-tibia with epiphysis, tibial spurs 0–2–2, no single medial spur on hindtibia (as in Artona Walker, 1854, Amuria Staudinger, 1887, and Palmartona gen. n.). Frenulum a very strong single spine in male, two or three smaller bristles in female, retinaculum a strong hook in male at base of Sc, a long row of hook-like scales in female at base of CuP. Forewing with R4+R5 stalked, hindwing with a short crossvein between Sc and cell, RR+M1 and M2+M3 fused, medial stem reduced in forewing, mainly well developed in hindwing. Pregenital abdomen (Fig. 432–433) Artonoid lateral evaginations on segments 2 and 7, the anterior evaginations on segment 2 in male very large, forming a balloon-like globe, evaginations on segment 7 in male and on segments 2 and 7 in

female of normal size. Sternum II with a pair of small, slightly curved, proximal apodemes, lateral sternal rod almost invisible. Tergum II very translucent. Last segments of abdomen very translucent in male and female. Male genitalia (Figs 386–396) Very small. Uncus long and slender; tegumen and vinculum weakly sclerotized, slender, saccus absent. Valva simple, without processes ventrally, dorsal sclerotization with a finger-shaped prolongation at basal edge pointing upwards (in natural situation pointing posteriorly) and covered with setae distally. Dorsal diaphragma translucent, ventral diaphragma with broad juxta with slightly spiny surface. Aedeagus stout, vesica without cornutus but with spiny surface, spines varying from small to very large. Female genitalia (Figs 397–401) Sternum VIII translucent, tergum VIII forming a strongly sclerozized narrow band, apophyses anteriores short. Papillae anales boad, with short setae, apophyses posteriores long and slender. Without praebursa, ostium broad, antrum moderately sclerotized, with spiny surface, corpus bursae long, translucent, strongly folded; ductus seminalis inserting into corpus bursae anteriorly.

Phenology and bionomics Unknown. Totthill, Taylor and Paine (1930: 228) reared some species of the ‘Artona-Levuana-type’ from New Guinea, the Bismarck Archipelago and the Solomon Islands during an extended search for palm-feeding zygaenids in that region. Larvae of five different species were found, but only one species was reared to imago. This species from the Solomon Islands was named ‘Zygaenid X’ but has never been described. It is possible that some of the species belong to Homophylotis. All larvae were found on monocotyledons, one species from New Britain on a Zingiberaceae of the ‘Nicolaia-type’, one from New Guinea on a palm of the genus Licuala (Arecaceae), another from New Guinea on a palm of the genus Areca (Arecaceae) and one from the Solomon Islands (not species X but ‘Zygaenid Y’!) on Musa (Musaceae). As stated under Pseudoamuria gen. n. (p. 201), one species of that genus from New Guinea was reared from bananas (Musa sp., Musaceae). It cannot be ascertained from that information whether the above mentioned ‘Zygaenid Y’ belongs to Pseudoamuria, gen. n., Homophylotis, or another genus.

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Remarks Of the 15 species referred to this genus by Bryk (1936: 258 ff.), only one (Homophylotis doloides (Pagenstecher 1900)) is congeneric with Homophylotis thyridota Turner, 1904, the typespecies of Homophylotis. The balloon-like enlarged lateral evaginations on abdominal segment II in the male, the reduction of the ‘Artona-finger’ on the male valva, the lack of a praebursa with rows of dentations and the insertion of the ductus seminalis into the anterior part of the corpus bursae in the female are supposed to represent autapomorphic characters of Homophylotis. The fact that the ductus seminalis inserts very far anteriorly into the corpus bursae indicates that the current form of the latter was derived from a fusion of a praebursa and a formerly separated corpus bursae. Normally, if a praebursa is present in Procridinae, the corpus bursae is connected with the praebursa by a short ductus intrabursalis (Alberti 1954: 155). The ductus seminalis is usually inserted close to the opening of the ductus intrabursalis into the corpus bursae. The conspicuous white spot close to the antennal apex and the reduction of the cornutus on the vesica seminalis are synapomorphies of Homophylotis and Pseudoamuria gen. n. These two genera are considered to represent sister-groups. Only four species are known in this genus so far: H. thyridota Turner, 1904, H. pseudothyridota sp. n., H. artonoides sp. n. (all occurring in northern Queensland) and H. doloides (Pagenstecher 1900) (Bismarck Archipelago). It is almost certain that the genus is more widely distributed in the New Guinea area but this cannot be confirmed at present.

–

extending beyond posterior margin of tegumen when valva is spread (Figs 386, 387). Aedeagus with vesica that is covered with small triangular and large almost rectangular spines (Fig. 393) . . . . . H. thyridota (p. 192) Proximal finger like prolongation on valva not extending beyond posterior margin of tegumen when valva is spread (Figs 388, 389). Aedeagus with vesica that is covered with small triangular spines only (Fig. 394) . . . . . . . . . . . . . . H. pseudothyridota sp. n. (p. 194)

Female 4. Corpus bursae simple, not divided into a tubelike posterior and a spherical anterior part (Figs 397, 399) . . . . . . . . . . . . . . . . . . . . . . . 5 – Corpus bursae divided into a long, tube-like posterior and a spherical anterior part (Fig. 398) . . . . H. pseudothyridota sp. n. (p. 194) 5. Antrum with broad, spiny, ring-like structure, not funnel-shaped (Fig. 397) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .H. thyridota (p. 192) – Antrum strongly funnel-shaped (Fig. 399) . . . . . . . . . . . . . . . . . . H. artonoides sp. n. (p. 195)

Homophylotis thyridota Turner Pl. 51, Figs 1, 2; Figs 49–56, 386, 387, 393, 397, 432. Homophylotis thyridota Turner, 1904, Trans. Proc. R. Soc. S. Aust. 28: 243. Lectotype 么 (here designated), AUSTRALIA: Queensland, Kuranda, ii. 1904 (F. P. Dodd), genitalia slide Z 2994 (ANIC) [examined].

Queensland, Bismarck Archipelago, ?New Guinea.

Homophylotis thyridota Turner; Jordan 1908: 47; T. B. Fletcher 1925: 10; Alberti 1954: 281; Tarmann 1996: 143.

Key to Homophylotis species

Material examined

The Australian species of Homophylotis can only be distinguished by genitalic characters. 1. Male (antenna bipectinate, frenulum represented by one single spine) . . . . . . . . . . . . . 2 – Female (antenna biserrate, frenulum represented by two or three small bristles) . . . . 4

Queensland: 2 乆, (paralectotypes), Kuranda, iii. 1904 (F. P. Dodd), genitalia slides Z 2991, Z 2993 (ANIC); 1 乆, Kuranda, xi. 1906 (F. P. Dodd), genitalia slide Z 2999 (ANIC); 1 么, Moses Creek, 4 km NE Mt Finnigan, 360 m, 15. x. 1980 (E. D. Edwards), genitalia slide Z 3002 (ANIC).

Male 2. Valva without any process ventrally (Figs 386–389) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 – Valva with a slender ventral process (Figs 390, 391) . . . . . . . . . H. artonoides sp. n. (p. 195) 3. Proximal finger-like prolongation on valva

Diagnosis

Distribution

Homophylotis thyridota is externally indistinguishable from H. pseudothyridota sp. n. Homophylotis artonoides sp. n. has a slightly larger translucent area in the hindwing and a creamy white forehead in the male; the female is also indistinguishable externally

TAXONOMY

from that of H. thyridota. The genitalia are distinctive. The male of H. pseudothyridota sp. n. has a narrower valva that is significantly longer dorsally than ventrally and not rectangular distally, the proximal finger-like prolongation is shorter and does not extend beyond the tegumen posteriorly when the valva is spread, the large rectangular spines on the vesica are absent; in the female of H. pseudothyridota sp. n. the spiny ring-like sclerotized antrum is narrower and leads directly into the large tube-like posterior part of the corpus bursae, which has a spiny surface, the anterior part of the corpus bursae is spherical, without spines, the ductus seminalis inserts at its centre. Homophylotis artonoides sp. n. has a small ‘Artona-finger’ on the valva ventrally but the female is similar to that of H. thyridota, although the corpus bursae is larger. Externally H. thyridota can also be confused with Pseudoamuria uptoni sp. n., but the latter is slightly larger, has a distinctive bluish and purple sheen, a smaller, not translucent, but white streak on the hindwing, and symmetrical antennal pectinations in male.

Redescription Male (Pl. 51, Fig. 1). Forewing length: 5.0–5.5 mm. Head densely scaled, flat, dark grey-brown dorsally; frons heart-shaped, silky fuscous with a few white scales dorsally, whitish yellow ventrally; breadth of frons 2.0× that of compound eye seen from frontal view; pilifer large; labial palps white, long, reaching anterior edge of head, slightly curved upwards; proboscis yellow; compound eyes black, surrounded by whitish scales; ocelli black; chaetosemata white. Antenna dark fuscous except segments 15–18, which are white dorsally, number of antennal segments 28, segments 1–14 long, with long asymmetrical pectinations, the anterior row almost twice as long as the posterior, segments 15–28 very short, without pectinations, weakly dentated. Thorax and abdomen grey-brown dorsally, whitish ventrally. Legs very long, fuscous dorsally, whitish ventrally, completely white at edge of segments, giving the legs a typical dark and white ringed pattern. Forewing with costa straight proximally (threequarters length of forewing), rather strongly arched towards apex, upperside dark grey-brown, sometimes with a faint greenish tinge, underside paler. Hindwing with upper- and underside dark greybrown, with a translucent central area of slightly variable size and shape crossed by densely scaled fuscous veins. Fringe variable, from whitish grey to grey-brown, with very short scales.

Female (Pl. 51, Fig. 2). Forewing length: 5.0 mm. Very similar to male. Antenna biserrate with very short dentations. Male genitalia (Figs 386, 387, 393). Uncus long, slender, bluntly pointed distally. Valva rectangular distally, the proximal finger-like prolongation long and slender, extending beyond posterior margin of tegumen when valva is spread. Aedeagus stout, approximately 4× longer than broad, vesica covered with small triangular and large almost rectangular spines. Female genitalia (Fig. 397). Papillae anales small, with short setae distally and minute spines proximally, apophyses posteriores long and slender, pointed distally. Tergum VIII covered with minute spines, with short, distally pointed apophyses anteriores. Ostium bursae broad, antrum very broad, spiny, ring-like; ductus bursae broad, very short, inserting laterally into the long, broad, curved, bean-shaped corpus bursae; corpus bursae covered with short triangular spines posteriorly, with smooth surface anteriorly. Ductus seminalis inserted laterally into anterior part of corpus bursae.


Remarks Turner’s original description is based on four specimens collected by F. P. Dodd at Kuranda in February and March. This original material consists of one male collected in February 1904, two females collected in March 1904 and another male collected in October 1904. However, the male collected in February and the two females collected in March belong in fact to H. thyridota, whereas the male collected in October is referable to H. pseudothyridota sp. n. There is an additional female specimen in the collection of MV taken at the same locality by the same collector in November 1904. It also belongs to H. pseudothyridota sp. n. As Turner clearly mentions February and March in his original description there is no doubt that the male specimen collected in February 1904 is one of the syntypes upon which Turner based his description. This specimen is therefore designated as lectotype because the complete series of syntypes consists of two species.

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According to genitalic characters, H. thyridota is most closely related to H. doloides from the Bismarck Archipelago [type examined]. Although the latter is much darker and opaque in coloration the genitalia are very similar, more so than those of the three Australian species one to each other (Figs 392, 396, 400, 401).


Homophylotis pseudothyridota sp. n. Pl. 52, Figs 1, 2, Pl. 53, Figs 1, 2; Figs 388, 389, 394, 398.

Material examined Holotype 么, Queensland: Kuranda, 1 ml E, 11. iii. 1964 (I. F. B. Common & M. S. Upton), genitalia slide Z 3001 (ANIC). Paratypes Queensland: 2 么, 1 乆, Kuranda, x. 1904, genitalia slide Z 2990 么, Z 3000 乆, xi. 1906 (F. P. Dodd), genitalia slide Z 2992 么 (ANIC, MV); 1 么, 1 乆, Kuranda, 9 mls N, 12. iii. 1964 (I. F. B. Common & M. S. Upton) (ANIC); 1 么, Mt Tozer, 9 km ENE, 10. vii. 1986 (J. C. Cardale), genitalia slide Z 2995 (ANIC).

Diagnosis Externally not significantly different from H. thyridota or H. artonoides sp. n. (see pp. 192, 195). The genitalia are distinctive (see pp. 197–200).

grey-brown dorsally; frons heart-shaped, silky fuscous with a few white scales dorsally, whitish yellow ventrally; breadth of frons 2.0× that of compound eye seen from frontal view; pilifer large; labial palps white, long, reaching anterior edge of head, slightly curved upwards; proboscis yellow; compound eyes black, surrounded by whitish scales; ocelli black; chaetosemata white. Antenna dark fuscous except segments 15–18, which are white dorsally, number of antennal segments 28–29, segments 1–14(–15) with long asymmetrical pectinations, the anterior row almost twice as long as the posterior, segments (14–)15–28(–29) very short, without pectinations, weakly dentate. Thorax and abdomen grey-brown dorsally, sometimes shiny blue, whitish ventrally. Legs very long, fuscous dorsally, whitish ventrally, completely white at edges of segments, giving the legs a typical dark and white ringed pattern. Forewing upperside dark greyish brown, sometimes with a faint greenish tinge, underside paler. Hindwing with upper- and underside dark greyish brown, with a translucent central area of variable size and shape crossed by densely scaled fuscous veins. Fringe variable, from whitish grey to greybrown, with very short scales. Female (Pl. 53, Fig. 2). Forewing length: 5.0 mm. Very similar to male. Antenna biserrate with very short dentations. Male genitalia (Figs 388, 389, 394). Uncus long, slender, bluntly pointed distally. Valva rounded distally, longer dorsally, the proximal finger-like prolongation not extending beyond posterior margin of tegumen when valva is spread. Aedeagus stout, approximately 4× longer than broad, vesica covered with small triangular spines but large rectangular spines absent. Female genitalia (Fig. 398). Papillae anales small, with short setae distally and minute spines proximally, apophyses posteriores long and slender, pointed distally. Ostium bursae broad, antrum a broad and spiny, slightly V-shaped, ring-like structure; ductus bursae fused with the broad, tube-like posterior part of corpus bursae that bears short triangular spines, anterior part of corpus bursae spherical with smooth surface. Ductus seminalis inserted at centre of anterior part of corpus bursae.

Description Male (Pl. 52, Figs 1, 2; Pl. 53, Fig. 1). Forewing length: 5.0–5.5 mm. Head densely scaled, flat, dark


TAXONOMY


Homophylotis artonoides sp. n. Pl. 54, Figs 1, 2; Figs 390, 391, 395, 399, 433.

Material examined Holotype 么, Queensland: Iron Ra., 11. iv. 1964 (I. F. B. Common & M. S. Upton), genitalia slide Z 2483 (ANIC). Paratype 乆, Queensland: Iron Ra., 14. iv. 1964 (I. F. B. Common & M. S. Upton), genitalia slide Z 2996 (ANIC).

Diagnosis Externally, H. artonoides is not significantly different from H. thyridota or H. pseudothyridota. It has a slightly broader translucent streak on the hindwing and a creamy white frons in the male. The females of all three species do not differ externally. The genitalia are distinctive. Homophylotis thyridota and H. pseudothyridota lack the Artona-finger on the valva and have the antrum of the female ring-like and not funnel-shaped. Homophylotis artonoides is also similar to Pseudoamuria uptoni sp. n. but the latter is slightly larger, has a distinctive purple and bluish sheen, a smaller white, not translucent streak on the hindwing and symmetrical antennal pectinations in the male.

Description Male (Pl. 54, Fig. 1). Forewing length: 5.0 mm. Head densely scaled, flat, dark greyish brown dorsally; frons heart-shaped, silky creamy white, slightly light brown laterally; breadth of frons 1.3× that of compound eye when seen from frontal view; pilifer large; labial palps white, long, slightly extending beyond anterior edge of head, porrect; proboscis

yellow; compound eyes black; ocelli white; chaetosemata white ventrally, fuscous dorsally. Antenna dark fuscous except segments 15–18, which are white dorsally, number of antennal segments 28, segments 1–14 long, with long asymmetrical pectinations, the anterior row almost twice as long as the posterior, segments 15–28 very short, without pectinations, weakly dentate. Thorax and abdomen greyish brown dorsally, whitish-ochreous ventrally. Legs very long, greyish brown dorsally, whitish ventrally, completely white at edges of segments, giving the legs a typical dark and white ringed pattern. Forewing with costa straight proximally (fourfifths length of forewing), arched towards apex, upperside dark greyish brown with a green tinge, underside yellowish grey proximally, darker distally, with a pale streak below costa. Hindwing upperside greyish brown around margin, an extended translucent central part with densely scaled dark veins, costal margin paler, underside similar but more yellowish grey. Fringe dark grey proximally, whitishochreous distally, with very short scales. Female (Pl. 54, Fig. 2). Forewing length: 5.0 mm. Very similar to male. Antenna biserrate with very short dentations. Male genitalia (Figs 390, 391, 395). Uncus approximately half as long as costal part of valva, broader basally, strongly pointed apically. Valva rounded distally, with a slender ‘Artona-finger’ with broad double-lobed base but without a distal brush; the proximal finger-like prolongation short and rounded, strongly resembling a pulvinus; with an additional oval field of short setae more ventrally where in many Artona species a group of very long setae is present, probably representing an androconial organ. Aedeagus approximately 6× longer than broad, slightly broader proximally, vesica without spines (perhaps incomplete in male holotype). Female genitalia (Fig. 399). Papillae anales small, with short setae distally and minute spines proximally, apophyses posteriores long and slender, pointed distally. Tergum VIII very slender, with short, distally pointed apophyses anteriores. Ostium bursae broad, antrum funnel-shaped, spiny; ductus bursae broad, very short, inserting laterally into a large, oval corpus bursae with smooth surface. Ductus seminalis inserted laterally into corpus bursae anteriorly.

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Distribution Queensland (Cape York).

TAXONOMY

386

387

388

389

390

391

Figs 386–387. Male genitalia of Homophylotis thyridota. 386: Lectotype, Qld, Kuranda. Gen. slide no. Z 2994; 387: Qld, Moses Creek. Gen. slide no. Z 3002. Figs 388–389. Male genitalia of Homophylotis pseudothyridota. 388: Paratype, Qld, Kuranda. Gen. slide no. Z 2990; 389: holotype, Qld, Kuranda. Gen. slide no. Z 3001. Figs 390–391. Male genitalia of Homophylotis artonoides, holotype, Qld, Iron Range. Gen. slide no. Z 2483. 390: Overview; 391: detail. Magnification as in figs 386–389.

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392

393

394

395

396

Fig. 392. Male genitalia of Homophylotis doloides, syntype, Bismark Archipelago. Gen. slide no. Z 3071. Fig. 393. Male genitalia: aedeagus of Homophylotis thyridota, lectotype, Qld, Kuranda. Gen. slide no. Z 2994. Fig. 394. Male genitalia: aedeagus of Homophylotis pseudothyridota, holotype, Qld, Kuranda. Gen. slide no. Z 3001. Fig. 395. Male genitalia: aedeagus of Homophylotis artonoides, holotype, Qld, Iron Range. Gen. slide no. Z 2483. Vesica lost distally. Fig. 396. Male genitalia: aedeagus of Homophylotis doloides, syntype, Bismark Archipelago. Gen. slide no. Z 3071.

TAXONOMY

397

398

399

Fig. 397. Female genitalia of Homophylotis thyridota, Qld, Kuranda. Gen. slide no. Z 2991. Fig. 398. Female genitalia of Homophylotis pseudothyridota, paratype, Qld, Kuranda. Gen. slide no. Z 3000. Fig. 399. Female genitalia of Homophylotis artonoides, paratype, Qld, Iron Range. Gen. slide no. Z 2996.

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200


400

401

Fig. 400. Female genitalia of Homophylotis doloides, syntype, Bismarck Archipelago. Gen. slide no. Z 3072. Fig. 401. Female genitalia of Homophylotis doloides, syntype, detail, Bismarck Archipelago. Gen. slide no. Z 3072.

TAXONOMY

Genus Pseudoamuria gen. n. Type-species: Pseudoamuria uptoni sp. n.

Diagnosis Small species with short, triangular forewings and rounded hindwings (Pl. 50, Fig. 2; Pl. 55). Hindwings with a light but not translucent central part, densely covered with scales. Head, thorax and abdomen dark dorsally, white or yellow ventrally. Antenna with a conspicuous white spot subapically. Male antenna with symmetrical pectinations. Head (Figs 57–64) Frons strongly projecting, rounded; labrum small, rounded, pilifer large, pocket-like, with long slender scales; maxillary palps small; proboscis of normal length, yellow; labial palps large, porrect or slightly curved upwards, not extending beyond anterior margin of head; compound eye of medium size, black, with a narrow eyelash ventrally; ocelli small; anterior extension of chaetosemata very slender. Male antenna bipectinate proximally, slightly biserrate distally, pectinations symmetrical; female antenna biserrate; antenna of male and female dark, white spot on distal part of antenna present but variable. Thorax and wings Legs very long; foretibia with long slender epiphysis slightly exceeding length of tibia distally, tibial spurs 0–2–2, no single medial spur on hindtibia (as in Artona Walker, 1854, Amuria Staudinger, 1887, and Palmartona gen. n.). Frenulum in male one strong single spine, two smaller spines in female; retinaculum a strong hook in male at base of Sc, only a few small scales at base of CuP in female. Wings with complete number of veins or with reductions, variable, forewing with all veins free from cell, medial stem absent, hindwing with a long transverse vein between Sc and anterior margin of cell, RR+M1 stalked or fused, M2+M3 stalked, medial stem reduced in forewing, present in hindwing. Pregenital abdomen (Fig. 434) Artonoid lateral evaginations on segments 2 and 7 of equal size in male and female. Sternum II with a pair of short almost straight proximal apodemes, lateral sternal rod very narrow, concave, longitudinal rod absent or just visible as a weak fold; sclerotization with broad lateral prolongations anteriorly. Sclerotized part of tergum II covering almost threequarters of segment posteriorly. Sternum VIII in male long, with a significant medial fold. Sternum VII in female broad, sternum VIII reduced.

Male genitalia (Figs 402–409) Genitalia large compared with the size of the specimens. Uncus short and slender, triangular basally, pointed distally; tegumen small and narrow, concave posteriorly, vinculum weakly developed, saccus not well-pronounced but with a small hook medially. Valva broad, weakly sclerotized distally, with a large well-sclerotized sacculus; Artona-finger (see p. 152) well developed, very long and slender, with a few setae distally; a field of short setae at half length of valva and a more proximal oval spot with very long setae ventrally, the latter probably representing an androconial organ; dorsal base of valva fused with part of the dorsal diaphragma, forming a finger-like structure pointing upwards with setae of medium length, probably of same function as the pulvinus in other groups, although not homologous. Dorsal diaphragma translucent, ventral diaphragma with broad juxta with spiny surface. Aedeagus stout, everted vesica with spiny surface; vesica bearing a bundle of loose, eversible, distally pointed, rod-shaped cornuti. Female genitalia (Figs 410, 411) Sternum VIII translucent, tergum VIII a broad, sclerotized band-like structure, apophyses anteriores absent. Papillae anales broad, with very short setae, apophyses posteriores long and very slender. With well-developed praebursa consisting of a more strongly sclerotized central part with a crest that bears tooth-like structures and a lateral translucent part that serves as a pocket to store the deciduous, eversible cornuti of the male vesica; corpus bursae small, ductus seminales inserted posteriorly close to the very short ductus intrabursalis.

Phenology and bionomics One species, Pseudoamuria melaleuca (Jordan, 1908) comb. n. from New Guinea (Figs 403, 409), was reared from bananas (Musa sp., Musaceae) (reference material in BMNH, London).

Remarks Pseudoamuria is the sister-group of Homophylotis based on the following synapomorphic characters: 1, reduction of cornutus on male vesica; 2, development of a conspicuous white spot close to antennal apex and of unknown function. A convincing autapomorphic character for Pseudoamuria is not known at this moment. For autapomorphies of Homophylotis see p. 192. Although externally very similar to Homophylotis, this genus has retained many charac-

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ters of the Asian species-groups related to Amuria Staudinger, 1887, and Artona Walker, 1854. Several genitalic characters such as the long Artona-finger, an oval spot on the valva medioventrally that bears very long setae and probably represents an androconial organ, eversible deciduous slender cornuti, a praebursa with a pocket in which to store them and a strongly sclerotized crest with tooth-like structures can be found in Pseudoamuria, Amuria and Artona. Pseudoamuria is considered to represent a link between the Southeast Asian species-groups and the Australian Homophylotis, the latter showing several reductions of characters.

Distribution Queensland, New Guinea. It is not yet clear how many species occurring outside Australia and currently included in Artona and Homophylotis belong to this genus but there is no doubt that at least some of the species from New Guinea have to be transferred to Pseudoamuria (see Figs 403, 409).

Key to Pseudoamuria species Forewing upperside uniformly dark greenish grey, with purple sheen, hindwing with a white streak (Pl. 55, Figs 1, 2) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. uptoni sp. n. (p. 202) Forewing upperside dark brownish grey, with a small, divided yellow spot, hindwing with two indistinct converging yellow streaks and a small spot (Pl. 50, Fig. 2) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. neglecta sp. n. (p. 203)

Pseudoamuria uptoni sp. n. Pl. 55, Figs 1, 2; Figs 57–64, 402, 404–408, 410, 434.

Material examined Holotype 么, Queensland: Iron Ra., 11. iv. 1964 (I. F. B. Common & M. S. Upton), genitalia slide Z 2997 (ANIC). Paratypes Queensland: 1 乆, Banks Island, 24. v. 1928 (W. B. Barnard), genitalia slide Z 2998 (QM); 1 么, Cape York, 12. vi. 1928 (W. B. Barnard) (abdomen missing) (QM); 1 么, Lockerbie, Cape York, 1. iv. 1964 (I. F. B. Common & M. S. Upton), genitalia slide Z 2484 (ANIC). There is one further female in the collection of SAM without a label or abdomen.

Diagnosis Similar to Homophylotis thyridota, H. pseudothyridota and H. artonoides, but without translucent hindwings and with symmetrical antennal pectinations in the male. The genitalia are much larger with a long Artona-finger on the valva and with eversible deciduous cornuti on the vesica in the male and a praebursa in the female. Pseudoamuria neglecta sp. n. has narrower wings and lighter brown coloration. The female genitalia are significantly different.

Description Male (Pl. 55, Fig. 1). Forewing length: 6.5 mm. Head dark grey dorsally and anterolaterally, with weak bluish sheen, creamy white anteromedially, shiny white ventrally. Frons strongly projecting, rounded; labial palps long, porrect, shiny white; compound eye of medium size, black, with a shiny white eyelash ventrally; ocelli small, black or white; chaetosemata creamy white. Antenna brownish greyish black, with a bluish shaft, segments 1–17 bipectinate, 18–33 slightly biserrate, pectinations very long, 6–7× longer than breadth of shaft in dorsal view at segment 7, often strongly curved, the distal ends crossing into each other, forming a tube with the shaft, brush-like apically, with very long setae, giving the antenna a cobweb-like appearance, white spot at distal part of antenna covering segments 18–23(–24), but sometimes reduced to a few white scales. Thorax dark grey dorsally with bluish sheen, dark grey ventrally, with a broad white streak laterally; coxa and femur white, tibia and tarsi dark grey with white rings. Abdomen upperside dark grey with strong blue sheen dorsally, white ventrally. Wings short, upperside of forewing dark greenish grey with purple sheen, underside whitish grey, darker towards apex, fringe dark grey basally, creamy white distally; hindwing opaque, dark greenish grey, with a very distinct, densely scaled, white streak behind cell from base to centre of wing, underside similar but whitish grey proximally, darker distally, fringe dark grey basally (one layer of scales), shiny white distally. Female (Pl. 55, Fig. 2). Forewing length: 7.0 mm. Similar to male. Forewing slightly more bluish distally. Frons slightly broader and eyes smaller. Antenna biserrate, the dentations very close together, forming a broad ventral groove. Male genitalia (Figs 402, 404–408). For main characters see generic description. Aedeagus 4× longer

TAXONOMY

than broad; the bundle of eversible deciduous cornuti consisting of 15–18 slender, distally sharply pointed cornuti, their length reaching approximately one-third length of aedeagus. Female genitalia (Fig. 410). Ostium broad, antrum broad, with spiny surface, a short ductus bursae leading into a large, double-lobed praebursa with a strongly sclerotized part bearing a row of long, slender, strongly pointed teeth and a lateral coneshaped, translucent pocket-like part that often contains a bundle of deciduous cornuti from the male. Ductus intrabursalis short and broad; corpus bursae spherical, translucent; ductus seminalis inserted into ductus intrabursalis close to opening of corpus bursae.

Diagnosis Pseudoamuria neglecta is similar to Homophylotis thyridota, H. pseudothyridota and H. artonoides but with more elongate wings and without translucent hindwings (more brownish). It has a praebursa in the female. Pseudoamuria uptoni has shorter wings, a purplish sheen and the streak on the hindwing is short and white. The genitalia are significantly different. Externally, P. neglecta is very similar to Palmartona catoxantha, a species not native to Australia. The underside of P. catoxantha is yellow and it has yellow patagia, the antenna is significantly shorter, the hind-tibia has a single medial-spur and the genitalia are different.

Description Phenology and bionomics Unknown

Distribution Queensland (Cape York).

Etymology Named in honour of the Australian lepidopterist Murray S. Upton who, together with I. F. B. Common, collected this important new species in Queensland.

Pseudoamuria neglecta sp. n. Pl. 50, Fig. 2; Fig. 411.

Material examined Holotype 乆, Queensland: Taylor Ra., Brisbane (F. P. Dodd) (Joicey Bequest 1934–120.) (BMNH). Genitalia slide no. Z 2298.

Male unknown. Female (Pl. 50, Fig. 2). Forewing length: 7.0 mm. Head brownish grey dorsally and anterolaterally, with a whitish spot dorsoanteriorly, shiny white ventrally. Frons strongly projecting, rounded; labial palps long, slightly curved upwards, shiny white basally, brown apically; compound eye of medium size, black, with a shiny white eyelash ventrally; ocelli small, white; chaetosemata creamy white, with a narrow brown stripe dorsally. Antenna brownish black, biserrate, 35 segments, with white spot on segments 25–28 ventrally. Thorax dark brownish grey dorsally, brown laterally, white ventrally; legs with coxa and femur white, tibia and tarsus brownish with white rings. Abdomen upperside brownish grey, underside white. Wings elongate, upperside of forewing dark brownish grey, with a small divided yellow spot on distal part of cell, underside greyish yellow proximally, darker and brownish distally, fringe greyish; hindwing opaque, densely scaled, upperside greybrown, with two indistinct, converging yellow streaks in basal half of wing, and a small spot across distal end of cell, underside similar but slightly paler, with an additional yellow streak along costa, fringe greyish basally (one layer of scales), shiny whitish distally. Female genitalia (Fig. 411). Ostium broad, antrum broad, with smooth surface, a short ductus bursae leading into a large, very strongly sclerotized praebursa with a long row of strongly pointed teeth, a lateral pocket to accommodate the deciduous cornuti of the male is not visible in the holotype but is probably lost as the specimen had a strongly laterally pressed abdomen (papered specimen?) and even the strongly sclerotized spherical praebursa was broken. Ductus intrabursalis long, of medium

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breadth, inserted laterally into praebursa; corpus bursae spherical, translucent; ductus seminalis inserted into ductus intrabursalis close to opening of corpus bursae.

Phenology and bionomics Unknown

Distribution Queensland. Only the holotype is known.

TAXONOMY

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407

Fig. 402. Male genitalia of Pseudoamuria uptoni, holotype, Qld, Iron Range. Gen. slide no. Z 2997. Fig. 403. Male genitalia of Pseudoamuria melaleuca, Papua New Guinea, Mt Hagen. Gen. slide no. Z 2989. Figs 404–407. Male genitalia of Pseudoamuria uptoni, holotype, Qld, Iron Range. Gen. slide no. Z 2997. 404: valva; 405: valva, detail, ‘Artona finger’; 406: dorsal part; 407: aedeagus.

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411

Fig. 408. Male genitalia of Pseudoamuria uptoni, holotype. Qld, Iron Range. Gen. slide no. Z 2997. Aedeagus, detail, vesica, deciduous cornuti lost (male must already have copulated). Fig. 409. Male genitalia: aedeagus of Pseudoamuria melaleuca, detail, vesica with bundles of deciduous cornuti still present. Papua New Guinea, Mt Hagen. Gen. slide no. Z 2989. Fig. 410. Female genitalia of Pseudoamuria uptoni, paratype, Qld, Banks Island. Gen. slide no. Z 2998. The deciduous cornuti from the male are visible in the lateral pocket of the praebursa. Fig. 411. Female genitalia of Pseudoamuria neglecta, holotype, detail, Qld, Taylor Range. Gen. slide no. Z 2298. The abdomen of the type was damaged, only parts of the praebursa could be saved.

TAXONOMY

Genus Palmartona gen. n. Type-species: Brachartona catoxantha Hampson, [1893]: 233

Diagnosis Small species with elongate triangular forewing and rounded or subtriangular hindwing (Pl. 56). Antenna very short, male antenna with strongly asymmetrical pectinations. Hindleg with three spurs (single medial spur present). Head Frons projecting, rounded; pilifer of normal size; maxillary palps small; proboscis of normal length, yellow; labial palps long, porrect, pointed distally; compound eye large, black, with a narrow eyelash ventrally; ocelli of medium size; chaetosemata small. Antenna very short, bipectinate proximally, slightly biserrate distally, pectinations strongly asymmetrical, the anterior row of pectinations significantly shorter, sensory hairs very long. Thorax and wings Foretibia with long slender epiphysis slightly exceeding length of tibia distally. Legs long. Tibial spurs 0–2–3, with single medial spur on hindtibia (as in Artona Walker, 1854, and Amuria Staudinger, 1887). Wings elongate, narrow, forewing with costal margin, arched distally, hindwing subtriangular, posterior margin rounded; forewing with all veins free from cell, medial stem present distally, hindwing with short transverse vein between Sc and anterior margin of cell, Rs+M1 fused, medial stem present. Frenulum in male one strong single spine, two smaller spines in female; retinaculum a strong hook in male at base of Sc, only a few small scales at base of CuP in female. Pregenital abdomen Artonoid lateral evaginations on segments 2 and 7 of equal size in male and female. Sternum II with a pair of short curved proximal apodemes, lateral sternal rod very narrow, concave, longitudinal rod absent or just visible as a weak fold; sclerotization with broad lateral prolongations anteriorly. Tergum II with a sclerotized area covering half of segment posteriorly. Sternum VII in female broad, sternum VIII reduced. Male genitalia (Figs 412–418) Uncus short and slender, triangular basally, pointed distally. Tegumen small and narrow, vinculum well

developed, V-shaped, with short saccus. Valva slender proximally, broad distally, weakly sclerotized, with a peculiar sacculus that has a hook-like appendix with a large oval field of very long setae and that probably represents an androconial organ; Artonafinger well developed, very long and slender, without distal setae and without a real articulation to distal edge of ventral sclerotization of valva. Ventral sclerotization of valva with a field of short dorsally pointing setae at two-thirds length of valva. Dorsal base of valva fused with part of dorsal diaphragma, forming a short finger-like structure pointing posteriorly, with setae of medium length and probably of same function as the pulvinus in other groups although not homologous. Dorsal diaphragma translucent, ventral diaphragma with triangular, posteriorly strongly concave juxta. Aedeagus long and slender, everted vesica with a short tube-like and a large spherical part, with several strong and fixed, but without deciduous, eversible cornuti. Female genitalia (Figs 419–422) Sternum VIII translucent, tergum VIII a strongly sclerotized, broad, band-like structure, apophyses anteriores very short. Papillae anales broad, with very short setae, apophyses posteriores long, very slender. Ostium and ductus bursae broad, without pronounced antrum; praebursa large, translucent, with a large spherical part with several strongly sclerotized crests bearing series of tooth-like structures and a triangular, pocket-like appendix that probably accommodates the cornuti of the male vesica (that has exactly the same spherical form as the larger part of the praebursa); ductus intrabursalis long, corpus bursae very small; ductus seminalis inserting close to insertion of ductus bursae into corpus bursae.

Phenology and bionomics Tropical species with several generations a year. Larvae on palms (Arecaceae), especially coconut palm. The larva has a significantly inflated second thoracic segment.

Remarks Brachartona catoxantha, the type-species of this genus, is a rather isolated species. Hence it could not be placed in any described genus. The type-species of Brachartona Hampson, 1891, is Artona quadrimaculata Moore, 1879, from northern India. It is considered to be congeneric with Amuria cyclops Staudinger, 1887, from the Amur region, the typespecies of Amuria Staudinger, 1887. Amuria was

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treated as a synonym of Artona Walker, 1854, by subsequent authors such as Bryk (1936: 246) or as a subgenus of Balataea Walker, [1865] by Alberti (1954: 272). Efetov and Tarmann (1995: 86), after examination of the types, considered Amuria to be a distinct genus, significantly different from Artona, with Brachartona as a synonym of Amuria. They considered Balataea to be a synonym of Artona. Brachartona catoxantha is not congeneric with either of them as it lacks the autapomorphies of Artona and Amuria (e.g. the characteristic blackyellow pattern of Artona and the distinctive form and sclerotization of the praebursa, the movable joint at half length of the split ventral sclerotization of the valva and the brush-like distal field of setae on the Artona-finger of Amuria). The following characters are considered to represent autapomorphies of Palmartona gen. n.: 1. an inflated second thoracic segment in the larva; 2. the very short, strongly asymmetrical male antenna; 3. a specialized sacculus with a very large characteristic field of long setae; 4. a series of sclerotized bands of characteristic shape with teeth-like structures in the praebursa. The host-plant relationship to palms (Arecaceae) may represent another autapomorphic character for this genus as all true Artona species, as far as is known, feed on Poaceae (mainly bamboo) and Amuria species on Zingiberaceae and related genera. Levuana iridescens Bethune-Baker, 1906, the coconut moth of Fiji, is not closely related. Palmartona gen. n. is therefore erected here as a monotypic genus to accommodate Brachartona catoxantha Hampson, 1892.

Distribution Southeast Asia, from Malaysia to New Guinea. In Australia probably introduced.

Palmartona catoxantha (Hampson) comb. n. Pl. 56, Figs 1, 2; Figs 412–422. Brachartona catoxantha Hampson, [1893], Fauna Br. India, Moths 1 (1892): 233. Holotype 么, BURMA: Tenasserim Valley (Doherty) (BMNH) [examined]. The style of biobliographic notes is slightly modified for this economically important species. The papers mentioned below are compiled from original literature (seen) and also from literature mentioned

in Review of applied Entomology (RAE) (not seen). As Palmartona catoxantha is a serious pest of coconut palms in Southeast Asia and may eventually invade Australia a more or less complete bibliography is given with, in brackets, comments on the content of the mentioned papers. The references in Review of applied Entomology (RAE) are placed in square brackets. For those papers taken from RAE the complete number of pages is given. Brachartona catoxantha (Hampson); Piepers and Snellen 1903: 237 (bionomics); Pratt 1909: 6 pp. (bionomics, pest control); Jepson 1915: 17–27 (bionomics) [RAE 1916: 91]; Dammermann 1919: 110 (bionomics); Knowles 1924: 1–14 (bionomics, pest report) [RAE 1924: 565]; Gater 1924: 374 (pest report) [RAE 1926: 90]; Gater 1925a: 92 (pest report); Gater 1925b: 92 (pest report) [RAE 1926: 549]; Aldrich 1925: 13 (parasitoid) [RAE 1926: 171]; Van Hall 1925: 53 pp. (pest report) [RAE 1926: 401]; Eecke 1930: 166. Artona catoxantha (Hampson); Jordan 1907–1908: 44–45 (taxonomy); Fletcher 1925: 2 (catalogue); Tothill 1925: 3 pp. (pest control, parasitoids) [RAE 1926: 559]; Bezzi 1925: 113–123 (parasitoids) [RAE 1926: 605]; Gater 1926: 304 (bionomics, pest control, pest report Malaysia); Van Hall 1926: 51 pp. (pest report) [RAE 1927: 520]; Corbett 1926: 171–174 (pest report) [RAE 1927: 557]; Gahan 1927: 39 pp. (parasitoids) [RAE 1928: 395]; Leefmans 1927: 260–270 (pest control) [RAE 1929: 190]; Corbett and Dover 1928: 1–7 (climatic influence on outbreaks) [RAE 1929: 475]; Leefmans 1928: 19 pp. (pest report, pest control, parasitoids) [RAE 1929: 576]; Corbett 1928: 136–140 (pest report) [RAE 1929: 628]; Corbett and Miller 1928: 404–424 [RAE 1929/1930: 413]; Wilkinson 1928: 79–105, 109–146 (parasitoids) [RAE 1929: 153]; Simmonds 1930: 215–219 (pest report, pest control, parasitoids); Corbett 1930: 511–513 (pest control) [RAE 1931: 139]; Tothill, Taylor and Paine 1930: 210 (bionomics, pest report Malaysia and Java, pest control); Corbett 1932: 370 (pest control) [RAE 1932: 557]; Corbett 1933a: 41–47 (pest control) [RAE 1933: 260]; Corbett 1933b: 39–52 (pest control) [RAE 1933: 676]; Leefmans and Awibowo 1933: 441 (pest controlt); Leefmans 1934: 1–92 (pest report, pest control) [RAE 1934: 547]; Corbett 1934: 43.56 (pest report) [RAE 1936: 103]; Van der Goot 1935: 1–79 (pest report) [RAE 1935: 560]; Leefmans and Awibovo 1935: 1–20 (pest control) [RAE 1937: 452]; Bryk 1936: 247 (catalogue, systematics); Corbett 1936: 41–53 (pest report)

TAXONOMY

[RAE 1937: 195]; Vecht 1936: 401–465 (pest control) [RAE 1936: 765]; Corbett 1937: 29–48 (pest report) [RAE 1937: 792]; Merino 1938: 31 (bionomics, parasitoids, pest report Philippines); Vecht 1938: 1055–1062 (pest control) [RAE 1938: 674]; Silayan 1938: 182 pp. (pest report) [RAE 1939: 451]; Vecht 1939: 389–428 (pest report, pest control) [RAE 1940: 231]; Betrem 1941: 45–101 (parasitoids) [RAE 1942: 319]; Corbett and Pagden 1941: 347–375 (pest control by rain and wind) [RAE 1942: 396]; Gadd & Fonseka 1945: 9–18 (parasitoids) [RAE 1946: 325]; Vecht 1947: 427 (pest report, pest control, parasitoids); Vecht 1950a: 1 (bionomics, pest control, pest reports Java); Vecht 1950b: 702 (bionomics, pest control, pest reports Java); Blunck 1953: 204 (pest report, pest control, parasitoids); Lever 1953: 20 (bionomics, pest report Malaysia, Singapore); Vecht 1954: 122–123 (parasitoids) [RAE 1956: 424]; Crosskey 1963: 1–16 (parasitoids) [RAE 1964: 45]; Lever 1964: 42–43 (parasitoids) [RAE 1964: 506]; Lever 1969: 190 pp. (pest report Papua New Guinea); Wirjosuhardjo 1969: 149 (pest control); Ho 1972: 329 (pest report, pest control); Ooi et al. 1975: 159 (pest control); Ooi 1977: 128 (pest report); Ooi, Soon and Ghee 1979: 1 (pest report, pest control); Kalshoven 1981: 297 (bionomics, summary of pest reports, pest control); Barlow 1982: 35 (taxonomy, bionomics, pest report); Davis and Sudasrip 1982: 8 (pest report); Peter, Nagarkatti and Jayanth 1987: 63 (parasitoids); Cock and Perera 1987: 283 (pest control); Godfray and Hassell 1989: 153 (population dynamics); Tarmann 1992a: 149 (list of larval host-plants of Zygaenidae); Sastrodihardjo et. al. 1993: 134 (bionomics, pest control, parasitoids); Zhang 1994: 91 (list of pest species); Tarmann 2003: 157 (bibliography, distribution, first report for Australia). Zeuxippa catoxantha (Hampson); Barlow 1982: 35 (pest report); Ginting and Desmier de Chenon 1987: 63 (pest control).

Material examined (Only examined Australian material mentioned.) Queensland: 1 乆, Taylor Ra., Brisbane (F. P. Dodd) (BMNH).

Diagnosis Externally similar to species of Homophylotis and Amuria. Homophylotis lacks the single medial spur on the hind-tibia but this character is present in

Amuria. The very short antenna of P. catoxantha is very characteristic and differs from those of both the above-mentioned genera. The pectinations in the male are asymmetrical in Homophylotis but symmetrical in Amuria. The genitalia are also characteristic.

Redescription Male (Pl. 56, Fig. 1). Forewing length: 6.0–7.5 mm. Head dark brown dorsally, yellow anteriorly, laterally and ventrally, with a brown stripe on frons medially. Compound eyes large, black; ocelli white; chaetosemata brown. Antenna very short, dark brown dorsally, yellow ventrally, with very thick shaft, 1–13 bipectinate, 14–28 very shortly biserrate, almost simple, apex bluntly pointed, anterior pectinations short, approximately 2× longer than breadth of shaft at segment 7, posterior pectinations longer, approximately 4× longer than breadth of shaft at segment 7 in dorsal view. Thorax dark brown dorsally with yellow patagia, yellow ventrally. Legs yellow, fore tibia with epiphysis, tibial spurs 0–2–3. Abdomen brown dorsally, yellow ventrally. Forewing and hindwing upperside dark brown, hindwing with a yellow costal margin, fringe of forewing ochreous grey, fringe of hindwing greyish white or light ochreous; underside of both wings light grey-brown with yellow scales along costa and a yellow mark before apex on forewing and two parallel yellow stripes on hindwing anteriorly. Female (Pl. 56, Fig. 2). Forewing length: 6.5–8.0 mm. Similar to male but slightly larger. Antenna very short, almost filiform, only slightly dentate, pointed distally. Male genitalia (Figs 412–418). For main characters see generic description. Vesica seminalis with one very slender large cornutus of same length as aedeagus, one smaller slender cornutus slightly shorter than half length of the larger cornutus, and with a third small, drop-shaped broader cornutus. Female genitalia (Figs 419–422). For main characters see generic description. Ductus bursae approximately 3× longer than broad, spherical part of praebursa at right angles to ductus bursae, with three long, band-like sclerotizations bearing short, tooth-like spines, appendix of praebursa in axis of ductus bursae, large, cone-shaped. Ductus intrabursalis very slender. Corpus bursae very small, translucent (easily lost during preparation!).

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Phenology and bionomics Palmartona catoxantha has been recorded as a pest on coconut palms by various authors (for summaries see e.g. Piepers and Snellen 1903; Tothill, Taylor and Paine 1930; Vecht 1950a, 1950b; Kalshoven 1981). Outbreaks have been known since 1892 and serious damage has been reported, especially from Malaysia (Malayan Peninsula) and Indonesia (Java). Outbreaks are comparatively rare and never begin during the wet season. There are always periods of several years between notable infestations. Moreover, they are always controlled by natural enemies and usually before serious damage is done to the leaves of the palms. The lower leaves of the palms are attacked first and are killed in most cases. Rarely the whole palm is seriously damaged, but it may be weakened and the growth of the coconuts diminished although the palms are rarely killed. There is no clear regular periodicity in the occurrence of the outbreaks. Normally, P. catoxantha is a rare species restricted to tropical climates and hardly ever observed in the field. Presumably the populations survive in small aggregations in the dense rainforests. Both sexes are diurnal. Copulation and oviposition take place in bright daylight. Egg Oval, transparent, shiny, yellow. The eggs are laid in small irregular batches (3–13) on the underside of the leaves of the host-plant, usually on their distal part, and hatch after 3–5 days. Larva The larva is slug-like. The early instars especially have the second thoracic segment markedly swollen. The ground colour is pale yellow or almost white in the early instars; the mature larva has a yellow head with dark brown mandibles, first and second thoracic segments yellow with a few dark stripes on segment 2, segments 3–10 greenish yellow with a blackish violet dorsal stripe and dark lateral stripes. The early instar larvae feed on the underside of the leaves of the host-plant and skeletonize the leaves. The last instar larva feeds freely on the edges of the leaves of the host-plant, making characteristic holes. The preferred host-plants are Arecaceae such as coconut palm (Cocos nucifera L.) and sago palms (Metroxylon spp.), but ornamental palms (Livistona spp.), are also known to suffer severe damage. Musaceae (Musa spp.) and even Poaceae such as sugar cane (Saccharum spp.) may serve as temporary hosts for later instar larvae.

Pupa Yellowish brown. Pupation takes place in an oval, greyish yellow, dense cocoon on the underside of the leaves of the host-plant, mainly close to the veins of the leaves, sometimes together in clusters. The pupa in the cocoon is always facing the surface of the leaf. Under tropical conditions P. catoxantha is able to produce 8–9 generations a year. Imago The moth rests at an angle of approximately 60° to the substrate, the head high up, touching the leaf with the tip of the abdomen, gripping the surface with the hind legs only. Vertically hanging leaves are preferred for resting. The adults take nectar from the inflorescences of the palms. The females normally do not leave the host-plants and rest close to the place where they have emerged where they wait for males. The males have been observed flying in great numbers around the crowns of the palms and searching for females. Copulation takes place on the leaves but observations of mating during flight have also been recorded, a phenomenon that is very unusual for Procridinae (Kalshoven 1981: 300). Oviposition starts in the early afternoon and lasts till early evening. Palmartona catoxantha moths have been observed several times on short-distance migration flights between coconut plantations, flying over distances of 1–1.5 km across open country (Vecht 1950a: 21).

Natural enemies and pest control Extensive studies have shown that the population density of P. catoxantha is controlled by a number of parasitoids and hyperparasitoids (Gater 1925, 1926; Vecht 1950a, 1950b). Two tachinid flies (Ptychomyia remota Aldr., Cadurcia leefmansi Bar.), two braconid wasps (Cotesia artonae Rohw., Microgaster indicus Wilk.), eight ichneumonids (Photoptera erythronota Vier., Goryphus infernus Szepl., G. rufibasalis Betr., G. javanicus Roman, G. fasciatipennis Szepl., G. bituberculatus Szepl., and two Hemiteles spp.), four chalcids (Brachymeria punctiventris (Cam.), B. lugubris Walk., B. apicornis Cam., Dirhinus banksi Rohw.), one eurytomid (Eurytoma albotibialis Ashm.), two eupelmids (Anastatoidea brachartonae Gah., Eupelmus catoxanthae Ferr.), 13 eulophids (Euplectromorpha viridiceps Ferr., E. artonae Ferr., Neoplectrus bicarinatus Ferr., N. maculatus Ferr., Pleurotropis detrimentosus Gah., P. ptychomyiae Ferr., Trichospilus pupivora Ferr., Syntomosphyrum obscuriceps Ferr., S. zygaenarum Ferr., S. nigricoxis

TAXONOMY

Ferr., two unidentified Syntomosphyrum spp., Mellitobia hawaiiensis Perk.) and one scelionid (Ceraphron sp.) are recorded by Vecht (1950a). The tachinid fly, Ptychomyia remota, and the braconid wasp, Cotesia artonae, are by far the most important parasitoids controlling outbreaks of P. catoxantha. The former lays its eggs on third and fourth instar larvae, the larva dying after it has constructed its cocoon. The latter parasitoid lays its eggs on second instar larvae, the larva dying before the cocoon is constructed. In Malaysia, Ptychomyia remota has been recorded as a more important parasitoid; in Indonesia (Java) Cotesia artonae has greater importance as a parasitoid. There is also a beetle (Callimera arcufer Chap.) recorded as a predator of the early instars of both the moth and their parasitoids (Lever 1953: 23). Biological control of P. catoxantha is also frequently exercised by the fungus Beauveria bassiana (Bals.) Vuill. (Lever 1953: 24).

Remarks Palmartona catoxantha is not native to Australia; as only one female is known it seems possible it was introduced but is now extinct. However, small colonies may still exist somewhere on the coast of Queensland. One has to take into account that P. catoxantha is a rare species and difficult to observe in normal years but that infestations of palms can start suddenly; moreover, there is no clear periodicity between outbreaks.

Of course the possibility that this single known female from Australia may be mislabelled must also be considered. As the label is a genuine original label of F. P. Dodd and as the introduction of a tropical species to Queensland can never be completely excluded, I take the view that this female actually originated from Australia. The species is therefore included in this revision.

Distribution Palmartona catoxantha is distributed from Myanmar [Burma] (Tenasserim) eastwards throughout Malaysia, Singapore, Indonesia (Sumatra, Nias, Java, Bangka, Kalimantan, Sulawesi), the Philippines (Palawan) to Papua New Guinea. Only one female is known from Australia (Queensland).

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Figs 412–418. Male genitalia of Palmartona catoxantha, Malaysia, Genting Highlands. Gen. slide no. Z 2525. 412: Overview; 413: dorsal part; 414: ventral, detail; 415: valva, detail; 416: aedeagus; 417: aedeagus, detail, vesica; 418: aedeagus, detail, vesica with cornuti.

TAXONOMY

419

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422

Figs 419–422. Female genitalia of Palmartona catoxantha, Qld, Taylor Range. Gen. slide no. Z 2202. 419: Overview; 420:detail, papillae anales, antrum. Qld, Taylor Range; 421: detail, praebursa with spiny rings and ductus seminalis (right upper corner); 422: detail, praebursa with spiny rings and corpus bursae.

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TAXONOMY

Genus Thyrassia Butler Thyrassia Butler, 1876, J. Linn. Soc. (Zool.) 12: 355. Type species: Syntomis subcordata Walker, 1854, List Specimens lepid. Insects Colln Br. Mus. 1: 132, by original designation. [N. India.] Thyrassia Butler; Jordan 1907: 51; Turner 1926b: 440; Tillyard 1926: 435; Bryk 1936: 270; Alberti 1954: 218; Common 1970: 830, 831, fig. 36.36.B (habitus); Common 1990: 297; Heppner 1991: 21; Tarmann, 1992a: 151; Tarmann 1996: 143; Tarmann 1998: 69; Xue & Han 2003: 266; Yen 2003: 311. Atucia Watson, 1980, The Generic Names of Moths of the World. 2: 20. Type species: Acutia bidens Kaye, 1919, Ann. Mag. Nat. Hist. (9) 4: 90, by original designation (for Acutia Kaye, 1919). Borneo: Sarawak,Bidi. Objective replacement name for Acutia Kaye, 1919. Atucia Watson; Holloway 1988: 6. Acutia Kaye, 1919, Ann. Mag. Nat. Hist. (9) 4: 89. Type species: Acutia bidens Kaye, 1919, Ann. Mag. Nat. Hist. (9) 4: 90, by original designation. A junior homonym of Acutia Ragonot, 1891, Annls Soc. Ent. Fr. (6) 10: 539 – Lepidoptera, Pyralidae.

Diagnosis Zygaenids of medium size with ctenuchid habitus, large, triangular, spotted forewings, very small, oval hindwings, abdomen with yellow and dark rings (Pl. 57). Head (Figs 65–74) Frons only slightly projecting, rounded; mouthparts very broad. Pilifer very small, maxillary palps small, slender, pointed distally, proboscis well developed, yellowish brown, labial palps small. Compound eyes large, ocelli large; chaetosemata large, not very strongly extending anteriorly but clearly artonoid, also extended dorsomedially, the two ends of each side almost touching each other medially; the free space between ocellus, base of antenna, compound eye and chaetosema not smoothly scaled, but covered with a series of tiny hair-like scales. Antennae shortly bipectinate proximally, strongly biserrate distally in both sexes. Thorax and wings Legs short and robust. Foretibia with epiphysis, tibial spurs absent (0–0–0). Frenulum a single spine

in male and female, retinaculum a long fold at Sc of forewing in male; no retinaculum visible in female. Wings with medial stem almost absent in forewing and hindwing, sometimes a rudimentary short vein visible distally; R1 in forewing arising from cell very distally, R2+R3 stalked, CuA2 arising very proximally at half length of cell; hindwing with Sc fused with anterior margin of cell, RR+M1 fused. Wings dark with characteristic pattern consisting of window-like spots that can be weakly scaled and translucent or scaled and opaque, coloured mainly yellow or orange. Pregenital abdomen (Fig. 435) Abdomen with characteristic pattern of yellow and dark rings. Artonoid lateral evaginations absent. Sternum II with a pair of strong, straight, proximal apodemes with broad base, lateral sternal rod very narrow, almost invisible, longitudinal rod absent; sclerotization with broad lateral prolongations anteriorly. Tergum II with a sclerotized area covering half of segment posteriorly. Sternum VII in female broad, sternum VIII reduced. Sternum and tergum VIII in male with characteristic sclerotizations. Male genitalia (Figs 423, 424) Uncus short and stout, strongly triangular, rounded distally, completely fused with the large tegumen, vinculum strongly sclerotized, with short saccus. Valva almost rectangular distally, slightly bent downwards, longer dorsally, strongly sclerotized, the fan-shaped central part small, sacculus not strongly pronounced, without appendages. A long finger-like process inserted at dorsal base of valva, this process forked basally, the short lateral process obviously representing an apodeme for a muscle to move the finger, the latter bent distally and bearing a brush of strong, short, backward-pointing spines. Dorsal diaphragma with a large, spiny transtilla, ventral diaphragma with well-developed juxta. Aedeagus short and stout, strongly sclerotized; everted vesica without cornuti. Female genitalia (Fig. 425) Sternum VIII translucent, tergum VIII sclerotized, broad, apophyses anteriores well developed, forked basally. Papillae anales fused, forming a short ovipositor, with very short setae, apophyses posteriores very long and very slender. Ostium and ductus bursae translucent, without antrum; praebursa absent. The insertion of the ductus seminalis into the bag-like translucent corpus bursae at its very centre indicates that the praebursa is fused with the corpus bursae. Otherwise the ductus seminalis

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would be inserted close to the point where the ductus bursae widens into the corpus.

Phenology and bionomics Piepers and Snellen (1903: 228, pl. 13, fig. 12) described the larva of Thyrassia procumbens Snellen, 1895 from Java. It feeds on Cayratia trifolia L. (Domin.) (Vitaceae).

Remarks Thyrassia is a tropical south and Southeast Asian genus. Its systematic position is not yet clear. Alberti (1954: 218) included Thyrassia in the tribe Callizygaenini (sensu Alberti) of Procridinae. Callizygaeninae were excluded from Procridinae and given subfamily rank by Tarmann (1994: 120) as they lack the autapomorphies of Procridinae. Thyrassia, although clearly belonging to Procridinae (e.g. according to the reduced lagena in the female), shows several reductions of characters (e.g. absence of tibial spurs, single female frenulum). The chaetosema is artonoid but not strongly extended anteriorly. The valva is artonoid due to the fanshaped translucent central part but is more strongly sclerotized than in all other Artonini. The lateral evaginations on abdominal segments 2 and 7 are absent. The genus is here included in Artonini but more information, especially on bionomics, is required to confirm its correct placement. The main autapomorphies of this genus are the ctenuchid-like habitus, the unusual form of the chaetosema, the complete reduction of tibial spurs, the broad, triangular uncus that is completely fused with the tegumen, the long, movable, finger-like pair of processes that are inserted at the proximal edge of the dorsal valva in the male, the short ovipositor and the reduced praebursa in the female.

Distribution From Sri Lanka and India to Australia (Queensland).

Thyrassia inconcinna Swinhoe Pl. 57, Figs 1, 2; Figs 65–74, 423–425, 435. Thyrassia inconcinna Swinhoe, 1892, Cat. Aust. Lep. Het. Colln Oxford Univ. Mus. 1: 55. Holotype 么, AUSTRALIA: Queensland, Lizard I. (OUM) [examined].

Monoschalis mimetica Turner, 1902, Trans. R. Soc. S. Aust. 26: 200. Lectotype 么 (here designated), AUSTRALIA: Queensland, Townsville, 22. ii. 1900 (F. P. Dodd) (ANIC) [examined]. Thyrassia inconcinna Swinhoe; Jordan 1907: 51; Turner 1926b: 440; Bryk 1936: 270; Alberti 1954: 218; Common 1970: 830, 831, fig. 36.36.B (habitus); Common 1990: 297; Tarmann 1992a: 151; Tarmann 1996: 143; Tarmann 1998: 69.

Material examined Queensland: 8 么, 11 乆, Townsville, 22. ii .1900, 10. iii. 1900, 23. vii. 1900 (F. P. Dodd) (ANIC, MV, QDPI, QM); 2 么, Meringa, i. 1925, e.l. 30.–31. iii. 1925 (F. H. Taylor) (ANIC, MV). There is additional material in ANIC and BMNH with printed labels ‘Townsville’ without date and collector.

Diagnosis Strongly resembling ctenuchids and unmistakable. There are no similar species in Australia.

Redescription Male (Pl. 57, Fig. 1). Forewing length: 10.0–11.0 mm. Head dark brown mixed with yellow dorsally, frons brown dorsally and around compound eye, yellow ventrally. Frons broad, 1.5× broader than compound eyes in frontal view, labial palps dark brown, proboscis yellowish brown; compound eyes black, ocelli white; chaetosemata brown mixed with yellow. Antenna blackish brown, shaft broad from base to apex, segments 1–30 shortly bipectinate, 31–36 strongly biserrate, bluntly pointed apically, pectinations tapering towards the strongly pointed apex, sensory hairs long. Thorax dark brown, with orange patagia, legs dark brown. Abdomen with blackish brown and yellow rings, posterior end black. Forewing large, triangular, grey-brown with a short orange-yellow basal streak, three large, whitish yellow spots at two-thirds and two small spots of same colour close to apex. Hindwing small and rounded, only 0.4 × length of forewing, base of wing with three partially confluent yellow spots, dark brown distally. Female (Pl. 57, Fig. 2). Forewing length: 12.0–12.5 mm. Similar to but significantly larger than male, with longer abdomen. Head and metathorax yellow dorsally. Head and antennae as in male. Male genitalia (Figs 423, 424). See generic diagnosis for main characters.

TAXONOMY

Female genitalia (Fig. 425). See generic diagnosis for main characters.

Phenology and bionomics According to label data the species is at least bivoltine. The larva is on Vitaceae (Vitis sp.) (according to a label on a specimen from ‘Townsville’ in BMNH). The early instars are unknown.

Distribution Northern Queensland.

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Figs 423–424. Male genitalia of Thyrassia inconcinna, Qld, Meringa, Gen. slide no. Z 3003. 423: Overview; 424: aedeagus. Fig. 425. Female genitalia of Thyrassia inconcinna, Qld, Townsville, Gen. slide no. Z 3004.

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Fig. 426. Male abdomen: segments I–V of Pollanisus lithopastus, Tas., Cradle Mountain. Gen. slide no. Z 3007. Lateral evaginations on segment II can be seen as holes. They are easily brushed away when the abdomen is cleaned of scales. Fig. 427. Male abdomen of Onceropyga anelia, Qld, Brisbane. Gen. slide no. Z 3065. Fig. 428. Female abdomen: segments I–VI of Turneriprocris dolens, Tas., Cradle Mountain. Gen. slide no. Z 2958. Fig. 429. Female abdomen: segments I–IV of Myrtartona coronias, Vic., Dandenong. Gen. slide no. Z 2962. Figs 430–431. Female abdomen of Australartona mirabilis, NSW, Robertson. Gen. slide no. Z 2948. 430: segments I–VI; 431: segment II, detail, second spiracle and strongly enlarged lateral gland.

TAXONOMY

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Fig. 432. Female abdomen of Homophylotis thyridota, Qld, Kuranda. Gen. slide no. Z 2991. Fig. 433. Female abdomen of Homophylotis artonoides, paratype, Qld, Iron Range. Gen. slide no. Z 2996. Fig. 434. Female abdomen of Pseudoamuria uptoni, paratype, Qld, Banks Island. Gen. slide no. Z 2998. Fig. 435. Male abdomen: segments I–IV of Thyrassia inconcinna, Qld, Meringa. Gen. slide no. Z 3003.

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Appendix I List of localities

The distribution maps are based on the latitudes and longitudes given below. Most of the examined material was collected before the use of GPS systems and the label data mainly refer to a larger place and not to an exact habitat. Therefore most co-ordinates are taken from the Australian Gazetter CD (eGAZ). The few more recent data based on GPS readings are given with full details and marked with ‘GPS:’. Acacia Ridge, Qld (27°35'S 153°01'E) Adelaide, SA (34°56'S 138°36'E) Albany, WA (35°00'S 117°52'E) Aldgate, SA (35°01'S 138°44'E) Aldinga Dellicks Beach Reserve, SA (35°17'S 138°27'E) Aldinga Scrub, SA (35°16'S 138°29'E) Alligator Gorge, SA (32°47'S 138°04'E) Allyn River Forestry Park, NSW (32°31'S 151°33'E) Amiens State Forest, Qld (28°35'S 151°48'E) Amosfield, NSW (28°40'S 152°04'E) Applecross, WA (32°01'S 115°50'E) Aranda, ACT (35°15'S 149°05'E) Aranda Bushland, ACT (35°15'S 149°05'E) Archerfield, Qld (27°34'S 153°00'E) Armidale, NSW (30°29'S 151°40'E) Arrawarra, NSW (30°04'S 153°12'E) Arthur Plains, Tas. (43°07'S 146°23'E) Arthur River, WA (33°53'S 116°54'E) Ashfield, NSW (33°53'S 151°07'E) Ash Island, NSW (32°52'S 151°42'E) Athelstone, SA (35°52'S 138°42'E) Augusta, WA (34°19'S 115°09'E) Babinda, Qld (17°21'S 145°56'E) Bald Rock National Park, NSW (31°32'S 152°18'E) Balhannah, SA (35°00'S 138°50'E) Banks Island, Qld (10°12'S 142°16'E) Bankstown, NSW (33°56'S 151°01'E) Barralier, NSW (34°16'S 150°04'E) Barren Grounds, NSW (34°40'S 150°42'E) Barren Grounds Fauna Reserve, NSW (34°42'S 150°43') Barrington River, NSW (31°59'S 151°37'E) Barrington Tops, NSW (31°59'S 151°27'E) Bateman’s Bay, NSW (35°44'S 150°15'E) Bathurst, NSW (33°25'S 149°35'E) Bayswater, Vic. (37°51'S 145°16'E) Bay View, NSW (34°45'S 146°00'E) Beaconsfield, Vic. (38°03'S 145°15'E) Beech Forest, Vic. (38°38'S 143°34'E) Beenak [Mountain], Vic. (37°53'S 145°42'E) Bendalong, NSW (35°15'S 150°32'E)

Bega, NSW (36°41'S 149°51'E) Belair Railway Station, SA (35°00'S 138°38'E) Belgrave, Vic. (37°/55'S 145°21'E) Bendalong, NSW (35°15'S 150°32'E) Bendora [Dam], ACT (35°27'S 148°50'E) Berowra, NSW (33°38'S 151°09'E) Bibra Lake, WA (32°06'S 115°49'E) Bicheno, Tas. (41°52'S 148°18'E) Biloela, Qld (24°24'S 150°31'E) Bindoon [Creek], WA (29°57'S 115°12'E) Binna Burra, Lamington National Park, Qld (28°12'S 153°11'E) Birchip, Vic. (35°59'S 142°55'E) Blackheath, NSW (33°38'S 150°17'E) Black Mountain, ACT (35°16'S 149°06'E) Black Mountain, N of summit trail, ACT (GPS: 35°16.184'S 149°06.156'E) Blackwood, SA (35°01'S 138°37'E) Boambee, NSW (30°21'S 153°06'E) Booanya [Rock], WA (32°46'S 123°36'E) Boonoo Boonoo River, NSW (28°43'S 152°11'E) Boronia, Vic. (37°52'S 145°17'E) Borumba Dam, Qld (26°33'S 152°35'E) Boyd Creek, NSW (33°46'S 148°05'E) Boyd River, NSW (33°46'S 148°05'E) Boyd River Crossing, NSW (33°46'S 148°05'E) Bridgetown, WA (33°58'S 116°08'E) Bridgewater, SA (35°00'S 138°46'E) Brisbane, Qld (27°28'S 153°02'E) Broadwater National Park, NSW (27°21'S 151°06'E) Brookfield Conservation Park, Blanchetown, SA (34°21’S 139°36'E) Broulee, NSW (35°51'S 150°11'E) Brown Lake, Stradbroke Island, Qld (27°29'S 153°26'E) Brown Mountain, NSW (36°36'S 149°23'E) Brunswick Heads, NSW (28°33'S 153°33'E) Brunswick, WA (33°15'S 115°50'E) Buderim Mountains, Qld (26°41'S 153°03'E) Buln Buln, Vic. (38°06'S 145°56'E) Bunbury, WA (33°20'S 115°38'E) Bundanoon, NSW (34°39'S 150°18'E) Bungwahl, NSW (32°03'S 152°27'E) Bunya, Qld (27°22'S 152°55'E) Bunya Mountains, Qld (26°51'S 151°34'E) Burleigh, Qld (28°05'S 153°27'E) Burleigh Heads, Qld (28°05'S 153°27'E) Burma Road Reserve, WA (31°46'S 116°20'E) Burns Beach, WA (31°44'S 115°43'E) Burnside, SA (33°17'S 138°27'E) Busselton, WA (33°39'S 115°20'E)

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Caiguna, WA (32°16'S 125°28'E) Cairns, Qld (16°55'S 145°46'E) Calga, NSW (33°25'S 151°13'E) Caloundra, Qld (26°48'S 153°08'E) Cann River, Vic. (37°34'S 149°09') Canyonleigh, NSW (34°35'S 150°07'E) Capel, WA (33°33'S 115°33'E) Cape Naturaliste, WA (33°31'S 115°00'E) Cape York, Qld (10°41'S 142°32'E) Carnarvon Range, Qld (25°20'S 148°40'E) Castlemaine, Vic. (37°04'S 144°13'E) Castlereagh State Forest, NSW (33°40'S 150°41'E) Cedar Creek, Qld (27°20'S 152°50'E) Ceduna, SA (32°08'S 133°41'E) Charlton, Tas. (42°06'S 147°31'E) Chatswood, NSW (31°52'S 147°30'E) Cheltenham, Vic. (37°58'S 145°03'E) Chichester State Forest, NSW (32°14'S 151°41'E) Chidlow, WA (31°52'S 116°15'E) Church Point, NSW (33°39'S 151°17'E) Claremont, WA (31°59'S 115°47'E) Clayton, Vic. (37°56'S 145°07'E) Clifton Beach, Qld (16°46'S 145°40'E) Clyde Mountain, NSW (35°33'S 149°57'E) Cockatoo, Vic. (37°57'S 145°29'E) Coles Bay, Tas. (42°08'S 148°17'E) Como, NSW (34°01'S 151°04'E) Como West, NSW (34°01'S 151°04'E) Condor Creek, Brindabella Range, ACT (35°19'S 148°50'E) Congo, near Moruya, NSW (35°37'S 150°09'E) Cooktown, Qld (15°28'S 145°15'E) Coolangatta, Qld (28°10'S 153°32'E) Cooloola Forestry Area, Qld 25°58'S 153°09'E) Coonabarabran, NSW (31°16'S 149°17'E) Cordeaux Dam, NSW (34°20'S 150°45'E) Cottesloe, WA (31°59'S 115°15'E) Cowan, NSW (33°11'S 149°21'E) Cradle Mountain, Tas. (41°41'S 145°57'E) Crawley, WA (31°59'S 115°48'E) Crescent Head, NSW (31°12'S 152°59'E) Crows Nest, Qld (27°16'S 152°03'E) Croydon, Vic. (37°48'S 145°17'E) Crusader Camp, Galston Gorge, NSW (33°40'S 151°05'E) Cunningham's Gap, Qld (28°03'S 152°24'E) Dalmorton, NSW (29°52'S 152°31'E) Dandabah, Bunya Mountains, Qld (26°51'S 151°34'E) Dandenong, Vic. (37°59'S 145°13'E) Dangar's Falls, Armidale, NSW (30°19'S 152°43'E) Darkes Forest, NSW (34°14'S 150°55'E) Darling Range, WA (31°31'S 115°59'E) Darwin, NT (12°27'S 130°50'E) Deepdene, WA (34°16'S 115°03'E) Denmark, WA (34°58'S 117°21'E) Depot Beach, NSW (35°38'S 150°19'E) Derby, Tas. (41°08'S 147°47'E) Derwent Bridge, Tas. (42°09'S 146°14'E) Devonport, Tas. (41°10'S 146°21'E)

Dimboola, Vic. (36°28'S 142°03'E) Doconing Road Beechina, WA (31°51'S 116°18'E) Donnybrook, WA (33°35'S 115°49'E) Don River, Tas. (41°10'S 146°20'E) Dorrigo, NSW (30°20'S 152°42'E) Dorrigo National Park, NSW (30°20'S 152°42'E) Drouin, Vic. (38°08'S 145°51'E) Drummond Cove, WA (28°40'S 114°36'E) Dryandra Forest, WA (32°46'S 116°58'E) Duaringa, Qld (23°44'S 149°40'E) Duggan Well, WA (33°09'S 118°09'E) Dulong, Qld (26°38'S 152°53'E) Dundas Rock, Norseman, WA (32°35'S 118°09'E) Dunwich, Qld (27°30'S 153°24'E) Dutson, Vic. (38°11'S 147°11'E) Dwellingup, WA (32°43'S 116°04'E) Ebor, NSW (30°24'S 152°21'E) Ellens Brook, WA (31°48'S 116°00'E) Eltham, Vic. (37°40'S 145°09'E) Emerald, Vic. (37°56'S 145°26'E) Eneabba, WA (29°50'S 115°11'E) Epping Forest, Tas. (41°46'S 147°22'E) Esperance, WA (33°51'S 121°53'E) Eucla, WA (31°43'S 128°52'E) Eungella, Qld (21°08'S 148°29'E) Evelyn Scrub, Qld (22°24'S 149°22'E) Fassifern, NSW (32°59'S 151°35'E) Fern Tree Gully, Vic. (37°53'S 145°18'E) Fingal Bay, NSW (32°45'S 152°11'E) Fitzroy Falls, NSW (34°39'S 150°29'E) Flinders Island, 5 km N Ranya, Tas. (40°10'S 148°08'E) Forrest, Vic. (38°31'S 143°43'E) Fowlers Bay, SA (31°59'S 132°34'E) Fremantle, WA (32°03'S 115°46'E) Frankston, Vic. (38°08'S 145°07'E) Fraser National Park, Qld (24°32'S 153°05'E) French's Forest, NSW (33°44'S 151°12'E) Freycinet National Park, Tas. (42°13'S 148°18'E) Fulham, Holmfirth, SA (37°10'S 140°57'E) Geraldton, WA (28°46'S 114°36'E) Gerroa, NSW (34°46'S 150°49'E) Gingin, WA (31°21'S 115°54'E) Girraween National Park, Qld (28°50'S 151°55'E) Gisborne, Vic. (37°29'S 144°35'E) Gladstone, Tas. (40°48'S 148°01'E) Glen Aplin, Qld (28°44'S 151°52'E) Glen Forest, WA (31°55'S 116°06'E) Glen Innes, NSW (29°44'S 151°44'E) Glenorie, NSW (33°36'S 151°01'E) Goombungee, Qld (27°19'S 151°42'E) Goondiwindi, Qld (28°33'S 150°19'E) Gosford, NSW (33°26'S 151°21'E) Goulburn, NSW (34°45'S 149°43'E) Grampians, Vic. (37°11'S 142°30'E) Groote Eyland, NT (14°00'S 136°40'E) Guildford, Perth, WA (31°54'S 115°58'E)

APPENDIX I

Halbury, SA (34°05'S 138°31'E) Halls Gap, Vic. (37°07'S 142°33'E) Hawkshead Lookout, Kalbarri National Park, 180 m, WA (GPS: 27°47'20'S 114°28'05'E) Hazelwood, Vic. (38°17'S 146°21'E) Healesville, Vic. (37°39'S 145°31'E) Heathcote, NSW (34°05'S 151°01'E) Heathmont, Vic. (37°50'S 145°15'E) Heifer Creek Road, Qld (27°40'S 152°10'E) Helensburgh, NSW (34°11'S 151°00'E) Herberton, Qld (17°24'S 145°23'E) Herberton, NSW (35°11'S 149°33'E) Hermies Garden Track, Big Desert, Vic. (35°48'S 141°47'E) Highbury, SA (34°53'S 144°55'E) Hobart, Tas. (42°55'S 147°20'E) Holyoake, WA (32°43'S 116°05'E) Hopetoun, WA (33°57'S 120°07'E) Hornsby, NSW (33°42'S 151°06'E) Hoskinstown, NSW (35°25'S 149°27'E) Howard, Qld (25°19'S 152°34'E) Howard Springs, NT (12°28'S 131°03'E) Hoyleton, SA (34°02'S 138°34'E) Huon-Picton River junction, Tas. (43°17'S 147°08'E) Huon Plains, Tas. (43°00'S 146°15'E) Huon River, Tas. (43°16'S 147°07'E) Inglewood, SA (34°50'S 138°46'E) Innaminna, NSW (31°33'S 147°09'E) Innisfail, Qld (17°32'S 146°02'E) Iron Range, Qld (12°42'S 143°18'E) Jamberoo Mountain, NSW (34°39'S 150°47'E) Jandakot, WA (32°07'S 115°50'E) Jarrnarm, Keep River National Park, NT (15°44' S 129°09'E) Jervis Bay, ACT (35°05'S 150°45'E) Jimberlana Hill, Norseman, WA (32°09'S 121°46'E) Jowalbinna Homestead, Qld (15°27'S 144°07'E) Jubilee Road, near Innisfail, Qld (17°32'S 146°02'E) Kalamunda, WA (31°58'S 116°03'E) Kalbarri, WA (27°42'S 114°12'E) Kalbarri Reserve, WA (27°40'S 114°15'E) Kallista, Vic. (37°53'S 145°22'E) Kanangra-Boyd National Park, NSW (34°02'S 150°03'E) Kanangra Walls, NSW (34°02'S 150°03'E) Kangaroo Island, SA (35°50'S 137°06'E) Karoola, Tas. (41°15'S 147°09'E) Karridale, WA (34°13'S 115°05'E) Katherine, NT (14°28'S 132°16'E) Katoomba, NSW (33°42'S 150°18'E) Katanning, WA (33°42'S 117°33'E) Kelmscott, WA (32°07'S 116°01'E) Kelso, Tas. (41°07'S 146°48'E) Kenmore, Qld (26°56'S 149°14'E) Kennedy River, Qld (15°21'S 144°02'E) Kiandra, NSW (35°53'S 148°30'E) Kiata, Vic. (36°22'S 141°48'E) Killarney, Qld (28°19'S 152°19'E)

Kings Park, Perth, WA (31°58'S 115°50'E) Kingston, Tas. (42°58'S 147°19'E) Kojarena, WA (28°44'S 114°51'E) Kuranda, Qld (16°49'S 145°39'E) Ku-ring-gai-Chase National Park, NSW (33°38'S 151°12'E) Lake Broadwater, Qld (27°21'S 151°07'E) Lake Loane, Tas. (41°48'S 146°15'E) Lake St Clair, Tas. (42°04'S 146°10'E) Lake Tyre, Tas. (41°49'S 146°20'E) Lamington National Park, Qld (28°14'S 153°08'E) Lancelin, WA (31°01'S 115°19'E) Lane Cove River, NSW (33°50'S 151°11'E) Latrobe River Survey Station 17 A, Tanjil Junction, Vic. (38°09'S 146°17'E) Launceston, Tas. (41°25'S 147°08'E) Leederville, WA (31°56'S 115°50'E) Leura, NSW (33°43'S 150°20'E) Lilyvale, NSW (34°11'S 151°00'E) Lion Mill, WA (21°56'S 120°38'E) Lismore Area, NSW (28°48'S 153°16'E) Little Desert, Vic. (36°30'S 141°30'E) Little Desert National Park, Vic. (36°30'S 141°30'E) Liverpool Range, NSW (31°45'S 150°45'E) Lizard Island, Qld (14°40'S 145°28'E) Lockerbie, Qld (10°47'S 142°28'E) Loddon Falls, NSW (34°17'S 150°54'E) Longford, Vic. (38°10'S 147°05'E) Longwood, Mt Lofty, SA (35°03'S 138°43'E) Lucas Heights, Sydney, NSW (34°03'S 150°50'E) Lucindale, SA (36°59'S 140°22'E) Macedon, Vic. (37°25'S 144°34'E) Macclesfield, SA (35°10' 138°50') Magnetic Island, Qld (19°08'S 146°50'E) Malua Bay, NSW (35°47'S 150°14'E) Mandurah, WA (32°33'S 115°42'E) Mangalore, Tas. (42°39'S 147°14'E) Manly, NSW (33°48'S 151°17'E) Marble Range, Eyre Peninsula, SA (34°27'S 135°30'E) Margaret River, WA (33°57'S 115°04'E) Marion Bay, Tas. (42°48'S 147°55'E) Maroochydore, Qld (26°39'S 153°06'E) Maroota, NSW (33°29'S 150°57'E) Martin, WA (32°05'S 116°03'E) Maryborough, Qld (25°32'S 152°52'E) Maryland, NSW (28°33'S 152°00'E) May Orchard, Brisbane, Qld (27°30'S 153°01'E) Meeniyan, Vic. (38°34'S 146°01'E) Megalong Valley, NSW (33°44'S 150°15'E) Melaleuca, Tas. (43°27'S 146°09'E) Melbourne, Vic. (37°50'S 145°00'E) Meningie, SA (35°42'S 139°20'E) Meringa, Qld (17°05'S 145°47'E) Miles, Qld (26°40'S 150°11'E) Milmed Track, Big Desert, Vic. (35°45'S 141°44'E) Millmerran, Qld (27°52'S 151°16'E) Millstream Falls, near Ravenshoe, Qld (17°41'S 145°17'E)

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Minnamurra Falls, NSW (34°38'S 150°43'E) Minyon Falls, near Rosebank, NSW (28°37'S 153°24'E) Mitcham, Vic. (37°49'S 145°12'E) Mittagong, NSW (34°27'S 150°27'E) Moe, Vic. (38°10'S 146°16'E) Moina, Tas. (41°27'S 146°07'E) Mole Creek, Tas. (41°33'S 146°24'E) Mooloolaba, Qld (26°41'S 153°07'E) Moraton Beach, Peel Island, Qld (27°30'S 153°22'E) Mordialloc, Vic. (38°00'S 145°05'E) Moresby Range, WA (28°40'S 114°40'E) Moonbar, Monaro, NSW (36°30'S 148°32'E) Moondarra Dam Area, Vic. (38°01'S 146°22'E) Mooney Mooney Creek, NSW (33°31'S 151°12'E) Moore River Bridge N, E Seabird, Lancelin Road, 30 m, WA (GPS: 31°14'51' 115°28'57') Moorina, Tas. (41°08'S 147°52'S) Moses Creek, near Mt Finnigan, Qld (15°47'S 145°17'E) Mosman, NSW (33°49'S 151°14'E) Mt Bellenden-Ker, Qld (17°16'S 146°54'E) Mt Buffalo, Vic. (36°44'S 146°47'E) Mt Canobolas, NSW (33°21'S 148°59'E) Mt Compass, SA (35°22'S 138°37'E) Mt Dale, WA (32°08'S 116°18'E) Mt Dromedary, NSW36°18'S 150°01'E) Mt Eliza, Tas. (42°57'S 146°24'E) Mt Evelyn, Vic. (37°47'S 145°23'E) Mt Franklin, ACT (35°28'S 148°49'E) Mt Gibraltar, Mittagong, NSW (34°27'S 150°27'E) Mt Hall, Eyre Peninsula, SA (33°04'S 134°29'E) Mt Irvine, Blue Mountains, NSW (33°29'S 150°28'E) Mt Kaputar, NSW (30°16'S 150°10'E) Mt Keira, NSW (34°24'S 150°51'E) Mt Kembla, NSW (34°27'S 150°48'E) Mt Kosciuszko, NSW (36°27'S 148°16'E) Mt Lewis, near Mt Molloy, Qld (16°35'S 145°17'E) Mt Lindsay Gate, Qld (28°21'S 152°53'E) Mt Lofty, SA (34°59'S 138°42'E) Mt Macedon, Vic. (37°23'S 144°35'E) Mt Mueller, Tas. (42°47'S 146°29'E) Mt Nelson, Tas. (42°55'S 147°20'E) Mt Pinnibar, Vic. (36°32'S 148°00'E) Mt Remarkable, SA (32°48'S 138°10'E) Mt Tamborine, Qld (27°44'S 153°10'E) Mt Tennyson Woods, near Mt Glorious, Qld (37°10'S 149°10'E) Mt Tibberoowuccum, near Beerburrum, Qld (26°56'S 152°56'E) Mt Tozer, Qld (12°45'S 143°13'E) Mt Vincent, NSW (32°55'S 150°20'E) Mt Wellington, Tas. (42°54'S 147°14'E) Mt Wilson, NSW (33°30'S 150°23'E) Muchea, WA (31°35'S 115°59'E) Mulgoa, NSW (33°50'S 150°40'E) Mulgrave River, near Gordonvale, Qld (17°05'S 145°47'E) Mundaring, WA (31°54'S 116°10'E) Munglinup, WA (33°50'S 120°51'E) Murramarang National Park, NSW (35°35'S 150°20'E) Murray Bridge, SA (35°07'S 139°16'E)

Murwillumbah, NSW (28°19'S 153°24'E) Myponga, SA (35°24'S 138°27'E) Nambucca, NSW (30°38'S 152°59'E) Nannup, WA (33°59'S 115°45'E) Napier property, WA (34°50'S 118°00'E) Narangba, Qld (27°12'S 152°58'E) Narara, NSW (33°24'S 151°21'E) Narrabeen, NSW (33°39'S 151°17'E) Native Dog Flat, Buchan River, Vic. (37°31'S 148°16'E) Nedlands, WA (31°59'S 115°48'E) Nelson, Vic. (38°03'S 141°01'E) Nelson Bay, NSW (32°43'S 152°09'E) Nerriga, NSW (35°07'S 150°05'E) Newcastle, NSW (32°54'S 151°45'E) Newcastle Firth, NSW (32°54'S 151°46'E) New England National Park, NSW (30°00'S 151°50'E) New Norcia, WA (30°58'S 116°13'E) Nhill, Vic. (36°20'S 141°40'E) Niagara Park, NSW (33°23'S 151°21'E) Noongan, WA (31°20'S 118°58'E) North Durras, near Bateman's Bay, NSW (35°40'S 150°18'E) Numinbah, Qld (28°08'S 158°14'E) Oatley, NSW (33°59'S 151°04'E) Ocean Grove, Vic. (38°16'S 144°32'E) Oldfield, WA (33°53'S 120°51'E) One Tree Hill, Neerabup Lake National Park, 13 km S Yanchep, 50 m, WA (GPS: 31°39'28' S 115°44'15' E) Orford, Tas. (42°34'S 147°52'E) O'Sullivans Gap, near Buladelah, NSW (32°20'S 152°16'E) Otford, NSW (34°13'S 151°00'E) Oxford Falls, NSW (33°43'S 151°15'E) Palm Beach, NSW (33°36'S 151°19'E) Palm Island, Qld (18°42'S 146°36'E) Palmwoods, Qld (26°41'S 152°58'E) Paluma, Qld (19°05'S 146°20'E) Paluma Dam, Qld (18°23'S 146°12'E) Parkerville, WA (31°53'S 116°08'E) Peats Ridge, NSW (33°19'S 151°14'E) Pelican Point, Bateman's Bay, NSW (35°05'S 150°35'E) Pelion Hut, Tas. (41°51'S 145°59'E) Pelion Plains, Tas. ((41°51'S 145°59'E) Penrose State Forest, NSW (34°40'S 150°13'E) Perth, WA (31°56'S 115°50'E) Petersham, Sydney, NSW (33°53'S 151°33'E) Picton Lakes, NSW (34°14'S 150°33'E) Pilliga State Forest, NSW (30°21'S 148°54'E) Pine Creek State Forest, near Raleigh, NSW (30°26'S 153°01'E) Pisga Ridge, near Glenbrook, NSW (33°46'S 150°37'E) Port Arthur, Tas. (43°09'S 147°51'E) Port Darwin, NT (12°27'S 130°50'E) Port Douglas, Qld (16°29'S 145°28'E) Portland, Vic. (38°20'S 141°36'E) Port Lincoln, SA (34°44'S 135°52'E) Port Macquarie, NSW (31°26'S 152°55'E)

APPENDIX I

Port Macquarie, 3 mls S, NSW (31°28'S 152°55''E) Port Sorell, Tas. (41°10'S 146°33'E) Port Stephens, NSW (32°45'S 151°50'E) Port Victoria, SA (34°40'S 137°29'E) Prince Henry Heights, Toowoomba, Qld (27°34'S 151°57'E) Prospect, SA (34°53'S 138°35'E) Putty, NSW (32°57'S 150°40'E) Rainbow Beach, near Tin Can Bay, Qld (25°49'S 153°01'E) Razor Back, near Picton, NSW (34°11'S 150°36'E) Redcliffe, Qld (27°14'S 153°07'E) Red Hill, Vic. (38°07'S 145°49'E) Redlynch, Qld (16°53'S 145°42'E) Reynella, SA (35°06'S 138°32'E) Richmond River, NSW (28°53'S 153°35'E) Ridgeway, Tas. (42°56'S 147°17'E) Ringwood, Vic. (37°49'S 145°14'E) Robe, SA (37°11'S 139°45'E) Robertson, NSW (34°35'S 150°36'E) Rocky Range, Kangaroo Island, SA (35°50'S 137°06'E) Rooty Hill, NSW (33°46'S 150°50'E) Rosebank, NSW (28°40'S 153°24'E) Rose Bay, NSW (33°52'S 151°16'E) Rosebery, Tas. (41°47'S 145°31'E) Roseberry, Vic. (35°49'S 142°25'E) Rosedale, Qld (24°38'S 151°55'E) Roseville, NSW (33°47'S 151°11'E) Roseville Chase, NSW (33°47'S 151°11'E) Round Hill Fauna Reserve, NSW (32°58'S 146°09'E) Rous, NSW (28°53'S 153°23'E) Royal National Park, NSW (34°08'S 151°04'E) Russell, SA (35°26'S 139°43'E) Ryan's Creek, near Depot Beach, NSW (35°38'S 150°19'E) Saddle Back [Mountain], NSW (30°27'S 151°21'E) Saddletree Creek, near Maidenwell, Qld (26°48'S 151°42'E) St George's Basin, NSW (35°07'S 150°36'E) St Lucia, Brisbane, Qld (27°30'S 153°30'E) St Marys, Tas. (41°34'S 148°10'E) St Patricks Plains, Tas. (41°33'S 148°13'E) Sale, Vic. (38°06'S 147°04'E) Salmon Gums, WA (32°59'S 121°38'E) Samford, Qld (27°22'S 152°53'E) Sandgate, Qld (27°20'S 153°05'E) Sandringham, Vic. (37°57'S 145°00') Sawpit Creek, Snowy Mountains, NSW (36°30'S 148°20'E) Scottsdale, Tas. (41°09'S 147°31'E) Scottsville, Qld (20°34'S 147°49'E) Sea Lake, Vic. (35°29'S 142°50'E) Serpentine, WA (32°21'S 115°58'E) Seventeen Seventy, Qld (GPS: 24°09.342'S 151°53.096'E) Shady Creek, Vic. (38°07'S 146°03'E) Sheoak Road Scub, near Belair, SA (35°03'S 138°46'S) Shoal Cape, WA (33°53'S 121°07'E) Shoalhaven, NSW (34°53'S 150°53'E) Singleton, NSW (32°34'S 151°11'E) Slapes Gully, SA (34°57'S 138°41'E) Somersby, NSW (33°22'S 151°17'E)

Southport, Qld (27°58'S 153°24'E) Springbrook, Qld (28°13'S 153°17'E) Springvale, Vic. (37°57'S 145°09'E) Stannary Hills, Qld (17°19'S 145°13'E) Stansbury, SA (34°55'S 137°47'E) Stanthorpe, Qld (28°40'S 151°56'E) Stanwell Park, NSW (34°14'S 150°59'E) Stanwell Tops, NSW (34°14'S 150°59'E) Stoneyfell, Adelaide, SA (34°56'S 138°36'E) Stradbroke Island, Qld (27°35'S 153°28'E) Strahan, Tas. (42°09'S 145°20'E) Strickland State Forest, near Ourimbah, NSW (33°22'S 151°26'E) Styx River State Forest, near Armidale, NSW (30°31'S 151°39'E) South Black Range, NSW (35°26'S 149°31'E) Subiaco, Perth, WA (31°57'S 115°48'E) Swan River, WA (32°03'S 115°45'E) Sydney, NSW (33°53'S 151°12'E) Sydney, Bradfield Drive, NSW (33°53'S 151°12'E) Tabbimoble State Forest, near Woodburn, NSW (29°05'S 153°21'E) Tabulam, NSW (28°54'S 152°34'E) Tahmoor, NSW (34°13'S 150°35'E) Tallanalla, WA (33°06'S 116°07'E) Tallong, NSW (34°43'S 150°05'E) Tamala, WA (27°11'S 113°56'E) Tansey, Planted Creek, Qld (26°01'S 152°07'E) Taylor Range, Brisbane, Qld (26°35'S 152°07'E) Tea Tree Gully, SA (35°09'S 137°20'E) Tenterfield, NSW (29°03'S 152°01'E) Tewah Creek, via Tin Can Bay, Qld (26°05'S 153°02'E) Thruston Park, Mulga, Qld (26°07'S 145°58'E) Thumbs Forest Reservation, Tas. (42°36'S 147°52'E) Tidbinbilla, ACT (35°26'S 148°53'E) Tin Can Bay, Qld (25°55'S 153°00'E) Tinderry Mountains, NSW (35°42'S 149°16'E) Tintinara, SA (35°54'S 140°03'E) Tombstone Rocks, WA (30°42'S 115°15'E) Toronto, NSW (33°00'S 151°36'E) Toowoomba, Qld (27°34'S 151°57'E) Townsville, Qld (19°16'S 146°49'E) Traralgon, Vic. (38°12'S 146°32'E) Triabunna, Tas. (42°30'S 147°55'E) Tubrabucca Creek, Barrington Tops, NSW (31°53'S 151°25'E) Tuglow [Trail], NSW (31°53'S 151°27'E) Tully, Qld (17°56'S 145°56'E) Tully River Valley, Qld (17°57'S 145°45'E) Tuross [Heads], NSW (36°04'S 150°08'E) Tweed Heads, NSW (28°10'S 153°33'E) Tyringham, NSW (30°13'S 152°32''E) Ulladulla, NSW (35°21'S 150°29'E) Ungarra, SA (34°11'S 136°03'E) Upper Sturt, SA (35°01'S 138°42'E) Upwey, Vic. (37°54'S 145°20'E) Uralla, NSW (30°38'S 151°30'E)

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Valencia [Creek], Vic. (37°50'S 146°59'E) Virginia, Brisbane, Qld (27°23''S 153°04'E) Wacol, Qld (27°21'S 152°33'E) Wahroonga, Sydney, NSW (33°45'S 151°10'E) Walpole-Nornalup National Park, WA (35°00'S 116°49'E) Wandin, Vic. (37°47'S 145°26'E) Wannon Divide, Grampians, Vic. (37°34'S 142°23'E) Waratah, Tas. (41°42'S 145°33'E) Warburton, Vic. (37°45'S 145°42'E) Waroona, WA (32°51'S 115°55'E) Waroona Dam, WA (32°51'S 115°59'E) Warragul, Vic. (38°10'S 145°56'E) Warrah, NSW (31°39'S 150°40'E) Warren River, WA (34°27'S 116°09'E) Warwick, Qld (28°13'S 152°02'E) Warwick, WA (30°35'S 116°08'E) Watagan State Forest, NSW (32°57'S 151°14'E) Warrumbungle, NSW (31°15'S 148°48'E) Waterview Creek, near Ingham, Qld (18°55'S 146°06'E) Wedge, Eyre Peninsula, SA (33°30'S 135°08'E) Wedge Bay, Tas. (43°07'S 147°41'E) Wedge Island, WA (30°50'S 115°11'E) Wedge Island, 5 km E, N. Lancelin, WA (GPS: 30°49'04''S 115°14'16'') Wedge Island, 8 km E, vic. Duggan Well, WA (GPS: 30°47'44'' 115°51'51'')

Westwood, Qld (27°42'S 150°01'E) Whiskers, NSW (35°25'S 149°21'E) Willowie, SA (32°41'S 138°20'E) Willow Grove, Vic. (38°05'S 146°11'E) Windsor Tableland, Qld (20°21'S 146°10'E) Wirrimbirra, near Picton, NSW (35°21'S 150°23'E) Wilsons Promontory, Vic. (39°04'S 146°22'E) Wilton, NSW (34°14'S 150°42'E) Windsor, Vic. (37°52'S 144°59'E) Wollongong, NSW (34°26'S 150°53'E) Woodend, Vic. (37°22'S 144°32'E) Woodside, SA (34°57'S 138°53'E) Woolgoolga, NSW (30°07'S 153°12'E) Woy Woy, NSW (33°29'S 151°20'E) Wubin, WA (30°06'S 116°38'E) Wyong, NSW (33°17'S 151°26'E) Wyperfeld National Park, Vic. (35°34'S 142°06'E) Yanchep, WA (31°33'S 115°41'E) Yankalilla, SA (35°28'S 138°21'E) Yarragon, Vic. (38°12''S 146°21'E) Yinnar, Vic. (38°19'S 146°20'E) Young, NSW (34°19'S 148°18'E) Yule Brook Reserve, Kenwick, WA (32°02'S 115°58'E) Zeehan, Tas. (41°54'S 145°20'E)

Appendix II: CryoSEM figures of eggs and first instar larvae of Pollanisus subdolosa clara ssp. n.

Figures 436–448 were provided by Andreas Zwick (CSIRO, Canberra) who collected a female of Pollanisus subdolosa clara ssp. n. along the Western Distributor Road in the Budawang National Park (NSW) in early December 2003. He reared the species from eggs and managed to produce a detailed documentation of its early instars. As the typescript for the book was already finished at that date, these informative pictures are given as an appendix.

437

436

438

Figs 436–438. Eggs of Pollanisus subdolosa clara covered with the scales from the abdominal hairtuft of the female. NSW, Budawang National Park, Western Distributor Road. 436: Overview; 437: detail; 438: surface of egg, detail.

228


439

440

441

442

443

444

Fig. 439. Abdominal scale of female’s hairtuft on eggs of Pollanisus subdolosa clara, detail, ultrastructure. NSW, Budawang National Park, Western Distributor Road. Andreas Zwick (CSIRO, Canberra). The ultrastructure of these scales shows primitive characters in the form of ‘transverse striae’ as described by Davis (1978) and discussed on p. 27. Fig. 440. Abdominal scale of female’s hairtuft on eggs of Pollanisus subdolosa clara, detail. NSW, Budawang National Park, Western Distributor Road. Andreas Zwick (CSIRO, Canberra). Figs 441–444. First instar larva of Pollanisus subdolosa clara, NSW, Budawang National Park, Western Distributor Road. Andreas Zwick (CSIRO, Canberra). 441: dorsal view; 442: dorsal view, larger magnification; 443: lateral view; 444: dorsal view, last segments.

APPENDIX II

445

446

447

448

Figs 445–448. First instar larva of Pollanisus subdolosa clara, NSW, Budawang National Park, Western Distributor Road. Andreas Zwick (CSIRO, Canberra). 445: detail, setae; 446: detail, seta, high magnification; 447: detail, unispined microtubercles on cuticula; 448: detail, thoracic leg.

229


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Index to Scientific Names Valid names given full treatment in the present volume are in boldface. Italicised page numbers indicate a figure. Pl. refers to plates. Plant names are listed alphabetically under genus.

Acacia 45 acharon, Adscita 95 acharon, Pollanisus 59, 67, 86, 88, 89, 92, 94, 95–96, 98, 99, 101, 129 acharon, Zygaena 59, 95, Pl. 58 acheron, Pollanisus 59, 95 Acoloithus 35 Acoloithus falsarius 42 Acoloithus novaricus 42 Acraea 43 adelaidae, Pollanisus 67 Adscita 27, 33, 41, 45, 46, 49, 50, 51, 53–55, 71 Agalope 38, 41 Agalopini 34 Aglaope 3, 9, 37, 38, 41 Aglaopini 34, 46 americana, Harrisina 42 amethystina, Pollanisus 59, 79 amethystina, Procris 59, 79, 80, 81 amethystinus, Pollanisus 79, 95 ampellophaga, Theresimima 33, 39, 42, 45, 46 Amuria 4, 34, 49, 53–55, 186, 187, 191, 201, 202, 207–208, 209 Anastatoidea brachartonae 210 anelia, Onceropyga 14, 59, 133, 134–35, 136, 137, 218, Pl. 33 Angophora 140 angustifrons sp. n., Pollanisus 59, 66, 86, 88, 89, 92, 94, 96, 98, 99, 100–101, 123, Pl. 23 Anomoeotidae 43 Apanteles 42 Apiaceae 41 apicalis, Adscita 81 apicalis, Pollanisus 38, 39, 42, 42, 59, 65, 67, 70, 76, 81–84, 111, 112, 120, 128, Pl. 9, Pl. 10, Pl. 58, Pl. 59, Pl. 60, Pl. 64 apicalis, Procris 59, 81 Apocyanaceae 85 Araeocera 27 Arbudas 37, 38 Arctiidae 27 Arctotheca 45 Arctotheca calendula 64, 74, 76, Pl. 60 Areca 191

Arecaceae 41, 42, 44, 191, 207, 208, 210 Artemisia 44 Artona 4, 33, 34, 47, 49, 51, 53–55, 57, 59, 61, 186, 187, 191, 195, 201, 202, 207, 208 Artona-group 4 Artona-Levuana-type 191 Artonini 1, 3, 4, 9, 10, 27, 31, 33, 34, 35, 36, 38, 39, 40, 41, 42, 45, 47, 49, 50, 51, 52, 53, 57, 58, 59, 61, 139, 187, 216 artonoides sp. n., Homophylotis 52, 60, 192, 193, 194, 195–96, 197, 198, 199, 202, 203, 219, Pl. 54 Asteraceae 41, 44, 45, 64, 74, 76, 187, Pl. 60 Atraphaxis 44 Atriplex paludosum 175 Atucia 215 Atucia bidens 215 Australartona gen. n. 1, 4, 37, 41, 50, 51, 52, 53–55, 60, 62, 185–89 Australartona mirabilis sp. n. see mirabilis sp. n., Australartona Balataea 208 Balataea-group 4 Banksia ilicifolia 76 Beauveria bassiana 211 Boradio 9 Brachartona 207, 208 Brachymeria apicornis 210 Brachymeria lugubris 210 Brachymeria punctiventris 210 Braconidae 42 brillians, Harrisina 42, 45, 47 brueckneri, Pyromorpha 46 Buckinghamia celsissima 140 budensis, Jordanita (Roccia) 38 Cadurcia leefmansi 210 calliceros azurea ssp. n., Pollanisus 59, 65, 107, 116, 125, Pl. 32 calliceros calliceros, Pollanisus 59, 65, 106–107, 116, 125, Pl. 31 calliceros, Pollanisus 13, 14, 41, 47, 59, 61, 63, 67, 105, 106, 107 Callimera arcufer 211 Callizygaena 139 Callizygaeninae 1, 3, 33, 34, 35, 36, 47, 216 Callizygaenini 4, 139, 216

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Campylotes 41 Carduus 44 catoxantha, Artona 208 catoxantha, Brachartona 60, 207, 208 catoxantha comb. n., Palmartona 1, 4, 31, 42, 47, 58, 60, 203, 208–11, 212, 213, Pl. 56 catoxantha, Zeuxippa 209 Cayratia trifolia 216 Celastraceae 41 Centaurea 45 Cephalaria 44 Ceraphron 211 Chalcididae 42 chalcochlora, Zygaenoprocris 31 Chalcosiinae 1, 3, 4, 9, 10, 27, 31, 33, 34, 35, 36, 37, 38, 39, 41, 43, 44, 46, 47 Chalcosiini 31, 33 Charops 42 chorista, Artona 42, 47 Chrysomelidae 43 Cicindelidae 43 Cirsium 45 Cissus antarctica 134, 135 Clelea 61 Clossiana 43 Coccinellidae 176 Cocos nucifera 210 Coleoptera 43, 176 commoni sp. n., Pollanisus 59, 66, 67, 86, 88, 89, 92, 94, 96, 98–99, 101, 122, 123, 130, Pl. 21 continentalis sp. n., Hestiochora 59, 140, 141, 142, 143, 145–48, 149, 151, 152, 154, 155, 159, 161, Pl. 37 contrastus sp. n., Pollanisus 59, 66, 67, 86, 88, 91–94, 96, 98, 99, 101, 102, 103, 114, 121, 129, Pl. 17 cf. contrastus 1, Pollanisus 59, 92, 93, Pl. 18 cf. contrastus 2, Pollanisus 59, 92, 93, Pl. 18 controversus, Zygaena 47 Coprosoma 187 coronias, Adscita 170 coronias comb. n., Myrtartona 60, 165, 169, 170–72, 173, 178, 180, 183, 186, 218, Pl. 45 coronias, Pollanisus 170 coronias, Procris 60, 170 coronias, Turneriptocris 170 Cotesia 42 Cotesia artonae 210, 211 Ctenuchinae 27 cuprea, Pollanisus 72 cupreus, Pollanisus 27, 59, 65, 67, 72–74, 75, 76, 80, 109, 117, 126, Pl. 4, Pl. 60, Pl. 63, Pl. 64 cyanota, Pollanisus 59, 63, 66, 67, 104–105, 115, 124, 131, Pl. 30 cyanota, Procris 59, 104

cyanotas, Pollanisus 59, 104 cyanotus, Pollanisus 104 cyclops, Amuria 207 Cyclosia 38 Cyclosiini 33 Dillenia 41 Dillenia alata 64, 98 Dillenia pentagyna 41 Dilleniaceae 41, 46, 57, 64, 85, 90, 98, 100, 103, Pl. 59, Pl. 62 Diplopoda 43 Dipsacaceae 33, 44, 45 Diptera 42, 50 Dirhinus banksi 210 discivitta, Artona 27 doldoides, Homophylotis 192, 194, 198, 199 dolens, Neoprocris 164 dolens, Pollanisus 164 dolens, Procris 60, 163, 164 dolens, Turneriprocris 16, 29, 32, 60, 164–65, 166, 167, 169, 170, 171, 218, Pl. 44 Doryphora sassafras 187 duskei, Zygaenoprocris 44 edwardsi sp. n., Pollanisus 59, 66, 67, 86–87, 88, 89, 92, 93–94, 96, 98, 99, 101, 113, 121, 128, Pl. 12 empyrea, Pollanisus 59, 65, 67, 73, 74, 75, 76, 79–81, 111, 119, 127, Pl. 8, Pl. 64 empyrea, Procris 59, 79, 80, 81 empyreus, Pollanisus 79 Ephemeroidea 27 Erasmia 38 Eriocranidae 27 erythrota, Hestiochora 15, 59, 139, 141, 142, 143–44, 154, 158, 161, Pl. 35 Eterusia 41 Eucalyptus 41, 57, 153, 170, 174, 175, 187 Eucalyptus calophylla 152 Eucalyptus fasciculosa 140, 147 Eucalyptus gittinsii 140, 146, 147 Eucalyptus marginata 152 Eucalyptus odorata 140 Eucalyptus rudis 152 eumetopus, Pollanisus 59, 66, 67, 86, 88, 89, 92, 94, 95, 96, 97, 98, 99, 101, 114, 122, 130, Pl. 20 eungellae sp. n., Pollanisus 59, 66, 67, 86, 88, 89, 92, 94–95, 96, 98, 99, 101, 102, 114, 122, 129, Pl. 19 Eupelmus catoxanthae 210 Euphorbiaceae 74, 76 Euplectromorpha artonae 210 Euplectromorpha viridiceps 210 Eurytoma albotibialis 210 exigua, Spodoptera 43 exulans, Zygaena 41

INDEX TO SCIENTIFIC NAMES

Fabaceae 41, 42, 102 falsarius, Acoloithus 42 filipendulae, Zygaena 31 funeralis, Artona (Balataea) 47 furcata sp. n., Hestiochora 59, 140, 141, 142, 143, 146, 148–49, 152, 156, 159, 161, Pl. 38, Pl. 39 Goryphus bituberculatus 210 Goryphus fasciatipennis 210 Goryphus infernus 210 Goryphus javanicus 210 Goryphus rufibasalis 210 gracilis, Leptozygaena 52 Gynautocera 38 Hadeninae 43 Hakea salicifolia 187 Harrisina 33, 41, 63 Helichrysum 45, 64, 71, 83, 187 Helichrysum viscosum Pl. 61 Heliconiinae 43 Heliconius 43 Heliothinae 43 Hemiteles 210 Hestiochora 1, 4, 5, 27, 33, 36, 37, 39, 41, 50, 51, 52, 53–55, 57, 59, 62, 133, 139–62, 185, 186, 187 Hestiochora continentalis sp. n. see continentalis sp. n., Hestiochora Hestiochora erythrota see erythrota, Hestiochora Hestiochora erythrota-tricolor-group 140, 142, 143, 148 Hestiochora furcata sp. n. see furcata sp. n., Hestiochora Hestiochora intermixta sp. n. see intermixta sp. n., Hestiochora Hestiochora occidentalis sp. n. see occidentalis sp. n., Hestiochora Hestiochora queenslandensis sp. n. see queenslandensis sp. n., Hestiochora Hestiochora queenslandensis-group 149 Hestiochora tricolor see tricolor, Hestiochora ‘Hestiochora tricolor auctt.’ 141–43 ‘Hestiochora tricolor habitus’ 143 Hestiochora xanthocoma see xanthocoma, Hestiochora Hestiochora xanthocoma-occidentalis-group 142, 150 Heterogynidae 43 Heteropanini 34, 35 Heteroptera 42, 43 Hibbertia 41, 46, 64, 71, 72, 74, 76, 80, 83, 85 Hibbertia dentata 84, 85 Hibbertia hypericoides 74, Pl. 62

Hibbertia obtusifolia 37, 38, 68, 72, 82, 84, 187, Pl. 59, Pl. 62 Hibbertia scandens 84, 85, 90, 94, Pl. 62, Pl. 63 Hibbertia sericea 72 Hibbertia spicata 74, 76 Hibbertia stricta 69, 72, Pl. 62 Hibbertia subvaginata 76 Hibbertia synandra 103 Hibbertia virgata 84 Histia 38 Histiochora 59, 139 Homophylotis 4, 31, 50, 51, 52, 53–55, 58, 60, 62, 186, 191–99, 201–202, 209 Homophylotis artonoides sp. n. see artonoides sp. n., Homophylotis Homophylotis pseudothyridota sp. n. see pseudothyridota sp. n., Homophylotis Homophylotis thyridota see thyridota, Homophylotis Hymenoptera 27, 42, 50 Ichneumonidae 42 ignorata, Neoilliberis 46 Illiberis 33, 35, 36, 45 incertus sp. n., Pollanisus 59, 66, 67, 86, 88, 89, 92, 94, 96, 98, 99–100, 101, 114, 123, 130, Pl. 22 inconcinna, Thyrassia 21, 22, 27, 60, 216–17, 217, 219, Pl. 57 Incurvariidae 27 infausta, Aglaope 36, 46 Insecta 43 inserta, Parthenocisus 46 intermixta sp. n., Hestiochora 59, 141, 142, 143, 146, 148, 149, 150, 152–53, 158, 161, 162, Pl. 43 iridescens, Levuana 36, 42, 47, 72, 188, 208 isolatus sp. n., Pollanisus 59, 66, 105–106, 115, 125, Pl. 31 Issoria 43 Jordanita 27, 33, 35, 41, 44, 45, 46, 51, 63, 71 Kennedia 41 Kennedia rubicunda 64, 102 Knautia 44 Kunzea 41, 57, 170 Kunzea ambigua 171, 172 labasi, Aglaope 46 Lamiaceae 41 Lepidoptera 9, 34, 36, 43 Leptospermum 41, 57, 164, 170, 174 Leptospermum juniperinum 164, 165, 170, 171, 172 Leptospermum myrsinoides 164, 165, 187, Pl. 62 Leptozygaena 52 Leptozygaena gracilis 52

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246


leucopleura, Adscita 172 leucopleura comb. n., Myrtartona 17, 30, 60, 169, 170, 172–74, 178, 180, 181, 183, Pl. 46, Pl. 61, Pl. 63 leucopleura, Pollanisus 172 leucopleura, Procris 60, 172 leucopleurus, Pollanisus 173 Levuana 53 Licuala 191 Ligustraum 45 lithopastus, Pollanisus 59, 65, 67, 70, 76, 77–79, 105, 110, 118, 127, 218, Pl. 6, Pl. 63 Lithosiinae 27 Livistona 210 Lomographa temerata 43 Lophosoma 27 Macrozamia communis 91 mannii, Adscita (Tarmannita) 38 mariannae sp. n., Myrtartona 60, 135, 170, 175, 176–77, 179, 182, Pl. 48 martini, Artona (Balataea) 47 Megalopygidae 43 Melaleuca 41, 57, 170, 171, 172, 174, 175, 187, Pl. 63 Melaleuca lanceolata 175 melaleuca, Homophylotis 4 melaleuca comb. n., Pseudoamuria 201, 205, 206 Mellitobia hawaiiensis 211 metallica, Harrisina 42, 45, 47 Metroxylon 210 Microgaster indicus 210 Micropterigidae 27 Micropterigoidea 50 mimetica, Monoschalis 60, 216 Mimosaceae 45 miocaenica, ‘Zygaena’ 47 mirabilis sp. n., Australartona 4, 17, 18, 32, 37, 38, 58, 60, 171, 172, 185, 186–88, 188, 189, 218, Pl. 48, Pl. 49, Pl. 50, Pl. 64 modestus sp. n., Pollanisus 59, 65, 76–77, 109, 117, Pl. 3 Monimiaceae 187 Monoschalis mimetica 60, 216 Musa 191, 201, 210 Musaceae 41, 44, 191, 201, 210 Myriapoda 43 Myrtaceae 41, 57, 140, 147, 163, 164, 165, 169, 170, 172, 174, 177, Pl. 62 Myrtartona gen. n. 37, 41, 50, 51, 52, 53–55, 57, 60, 62, 163, 169–83 Myrtartona coronias comb. n. see coronias comb. n., Myrtartona Myrtartona leucopleura comb. n. see leucopleura comb. n., Myrtartona Myrtartona mariannae sp. n. see mariannae sp. n., Myrtartona

Myrtartona rufiventris comb. n. see rufiventris comb. n., Myrtartona Naufockia 35 neglecta sp. n., Pseudoamuria 60, 202, 203–204, 206, Pl. 50 Neoalbertia 35 Neoilliberis 35 Neoplectrus bicarinatus 210 Neoplectrus maculatus 210 Neoprocris 27, 34, 60, 163 Nicolaia-type 191 nielseni sp. n., Pollanisus 59, 64, 65, 67, 73, 74, 75–76, 80, 108, 109, 126, Pl. 5, Pl. 60, Pl. 63 novaehollandiae, Procris 59, 81 novaricus, Acoloithus 42 Nymphalidae 43 occidentalis sp. n., Hestiochora 47, 59, 141, 142, 146, 150, 151–52, 153, 154, 157, 160, 162, Pl. 42 Ocneropyga 59, 133 Oleaceae 45 oligocenica, Neurosymploca? 47 Onceropyga 4, 33, 34, 36, 37, 41, 50, 51, 52, 53–55, 57, 59, 62, 133–37, 139, 147, 170 Onceropyga anelia see anelia, Onceropyga Onceropyga pulchra sp. n. see pulchra sp. n., Onceropyga Orna 10, 27 Oxyopes 176 Oxyopidae 176 Palmartona gen. n. 4, 31, 41, 50, 51, 52, 53–55, 60, 61, 62, 186, 191, 201, 207–13 Palmartona catoxantha comb. n. see catoxantha comb. n., Palmartona Parthenocisus inserta x P. quinquefolia 46 paupera, Jordanita (Roccia) 44 persepolis, Zygaenoprocris 44 Phacusa 27, 41 Photoptera erythronota 210 Phyllanthus calycinus 74, 76 Pinaceae 41 Pinus 41 Pipturus 41 Pipturus argenteus 64, 97 Pleurotropis detrimentosus 210 Pleurotropis ptychomyiae 210 plumbeola, Clelea 47 Poaceae 41, 58, 187, 208, 210, Pl. 62, Pl. 64 Pollanisus 1, 4, 5, 27, 33, 34, 35, 36, 37, 41, 46, 50, 51, 52, 53–55, 57, 59, 61, 63–131, 133, 135, 139, 147, 149, 170, 185, 187


Pollanisus acharon see acharon, Pollanisus Pollanisus acharon-group 85 Pollanisus angustifrons sp. n. see angustifrons sp. n., Pollanisus Pollanisus apicalis see apicalis, Pollanisus Pollanisus calliceros see calliceros, Pollanisus Pollanisus calliceros azurea ssp. n. see calliceros azurea ssp. n., Pollanisus Pollanisus calliceros calliceros see calliceros calliceros, Pollanisus Pollanisus commoni sp. n. see commoni sp. n., Pollanisus Pollanisus contrastus sp. n. see contrastus sp. n., Pollanisus Pollanisus cf. contrastus 1 see contrastus 1, Pollanisus Pollanisus cf. contrastus 2 see contrastus. 2, Pollanisus Pollanisus cupreus see cupreus, Pollanisus Pollanisus cyanota see cyanota, Pollanisus Pollanisus edwardsi sp. n. see edwardsi sp. n., Pollanisus Pollanisus empyrea see empyrea, Pollanisus Pollanisus eumetopus see eumetopus, Pollanisus Pollanisus eumetopus-group 64, 85, 99 Pollanisus eungellae sp. n. see eungellae sp. n., Pollanisus Pollanisus incertus sp. n. see incertus sp. n., Pollanisus Pollanisus isolatus sp. n. see isolatus sp. n., Pollanisus Pollanisus lithopastus see lithopastus, Pollanisus Pollanisus modestus sp. n. see modestus sp. n., Pollanisus Pollanisus nielseni sp. n. see nielseni sp. n., Pollanisus Pollanisus sp. 1 59, 64, 79, 111, 118, 127, Pl. 7 Pollanisus sp. 2 59, 64, 102, Pl. 24 Pollanisus sp. 3 59, 64, 102, 103, Pl. 10, Pl. 25 Pollanisus sp. 4 59, 64, 102, Pl. 26 Pollanisus sp. 5 59, 64, 102–103, Pl. 27 Pollanisus sp. 6 59, 64, 102, 103, 115, 124, 131, Pl. 28 Pollanisus sp. 7 59, 64, 103–104, Pl. 29 Pollanisus sp. 8 59, 64, 104, Pl. 29 Pollanisus subdolosa see subdolosa, Pollanisus

Pollanisus subdolosa clara ssp. n. see subdolosa clara ssp. n., Pollanisus Pollanisus subdolosa conspecifics 91 Pollanisus subdolosa subdolosa see subdolosa subdolosa, Pollanisus Pollanisus subdolosa-eumetopus-group 102, 103, 104 Pollanisus trimacula see trimacula, Pollanisus Pollanisus viridipulverulenta see viridipulverulenta, Pollanisus Pollanisus viridipulverulenta-group 64 Pollanisus-Hestiochora-group 175 Polygonum 44 Pomaderis apetala 187 pomonella, Cydia 46 postvitta, Artona 27 Praezygaena 27 Praviela 35 Primilliberis 35 Procridinae 1, 3, 4, 9, 10, 27, 31, 33, 34, 35, 36, 37, 38, 39, 40, 41, 42, 43, 44, 45, 46, 47, 49, 50, 51, 52, 57, 59, 61, 64, 72, 139, 140, 187, 216, Procridini 1, 3, 4, 9, 10, 23, 24, 25, 26, 27, 31, 34–35, 38, 40, 42, 45, 46, 49, 50, 57, 61, 71, 139 Procris 49, 51, 61 procumbens, Thyrassia 216 Proteaceae 140, 187 pruni, Illiberis (Primilliberis) 9, 10, 23, 24, 31, 47, 50 pruni, Rhagades (Rhagades) 38, 41 Pryeria 9, 10, 27 Pryeria sinica 46 Pseudoamuria gen. n. 4, 31, 34, 50, 51, 52, 53–55, 58, 60, 61, 62, 186, 191, 192, 201–206 Pseudoamuria neglecta sp. n. see neglecta sp. n., Pseudoamuria Pseudoamuria uptoni sp. n. see uptoni sp. n., Pseudoamuria pseudothyridota sp. n., Homophylotis 60, 192, 193, 194–95, 197, 198, 199, 202, 203, Pl. 52, Pl. 53 Ptychomyia 42 Ptychomyia remota 210, 211 pulchra sp. n., Onceropyga 59, 134, 135–36, 137, 175, 176, Pl. 34 pyramidalis, Anacamptis 45 Pyromorpha 27, 34 quadrimaculata, Artona 207 queenslandensis sp. n., Hestiochora 59, 141, 142, 146, 148, 149–50, 152, 153, 156, 159, 160, 162, Pl. 40 quinquefolia, Parthenocisus 46 Reduviidae 42 Rhagades 9, 33, 35, 36, 41, 45, 46

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Rhamnaceae 187 Rosaceae 41 rotundata, Illiberis (Primilliberis) 38 Rubiaceae 187 rufiventris comb. n., Myrtartona 27, 60, 135, 169, 170, 174–76, 179, 181, 182, 183, Pl. 47 rufiventris, Hestiochora 174 rufiventris, Neoprocris 174 rufiventris, Procris 60, 139, 163, 169, 174 Saccharum 210 Scabiosa 44 Scapha auriflua 143 Senecio 187 sequens, Pollanisus 59, 81 sinica, Pryeria 46 Soritia 38, 41 statices, Adscita (Adscita) 9, 10, 24, 25, 26, 31, 38 straminea, Parsonsia 84, 85 subcordata, Syntomis 60, 215 subdolosa clara ssp. n., Pollanisus 38, 59, 66, 86, 88, 89–91, 113, 121, 128, 227, 227, 228, 229, Pl. 14, Pl. 15, Pl. 59 subdolosa conspecifics, Pollanisus 91 subdolosa, Pollanisus 59, 67, 87, 88, 91, 92, 93–94, 95, 96, 98, 99, 100, 101, Pl. 64 subdolosa subdolosa, Pollanisus 59, 66, 86, 87–89, 128, Pl. 13 subdolosa, Procris 59, 87 subdolosus, Pollanisus 87 subolosa cf. subdolosa, Pollanisus 113, 129, Pl. 15, Pl. 16 subsolana, Jordanita (Solaniterna) 38, 41 Syncarpia 140, 146, 147 Syntomis subcordata 60, 215 Syntomosphyrum 211 Syntomosphyrum nigricoxis 210 Syntomosphyrum obscuriceps 210 Syntomosphyrum zygaenarum 210 Tachinidae 42, 84 temerata, Lomographa 43 tenius, Illiberis 47 Tetracera 41 Tetracera nordtiana 64, 99, 100 Tetrarrhena juncea 187, Pl. 62, Pl. 64 Theaceae 41 Theresimima 9, 45 Thibetana 34, 51, 57 Thyrassia 27, 31, 41, 50, 51, 53–55, 57, 58, 60, 61, 215–17, 219 Thyrassia inconcinna see inconcinna, Thyrassia

thyridota, Homophylotis 18, 19, 60, 191, 192–94, 195, 197, 198, 202, 203, 219, Pl. 51 Tortricidae 27, 46 Trichospilus pupivora 210 tricolor, Hestiochora 59, 141, 142, 143, 144–45, 146, 147, 149, 154, 158, 161, Pl. 36, Pl. 58 tricolor, Procris 59, 139, 144 tricuspidata, Parthenocisus 46 trifolii, Zygaena 36 trimacula, Pollanisus 59, 66, 67, 84–86, 112, 120, 128, Pl. 11, Pl. 64 trimacula, Procris 59, 84 trimaculus, Pollanisus 84 Triprocris 25, 35 Turneriprocris 41, 50, 51, 52, 53–55, 57, 60, 62, 163–67, 170, 186 Turneriprocris dolens see dolens, Turneriprocris Turneriptocris 60, 163 turolensis, Zygyaena? 47 ulmivora, Illiberis 27 uptoni sp. n., Pseudoamuria 20, 60, 193, 195, 201, 202–203, 205, 206, 219, Pl. 55 Urticaceae 41, 97 variabilis, Neoalbertia 46 viridipulverulenta, Pollanisus 11, 12, 28, 31, 32, 37, 38, 59, 65, 67–72, 76, 82, 83, 108, 116, 117, 126, Pl. 1, Pl. 2, Pl. 3, Pl. 59, Pl. 60, Pl. 61, Pl. 64 viridipulverulenta cupreus, Pollanisus 72 viridipulverulenta, Procris 59, 63, 67 viridipulverulentus, Pollanisus 67 Vitaceae 41, 44, 46, 135, 216, 217 Vitis 41, 217 Wiegelia 35 xanthocoma, Hestiochora 59, 139, 140, 141, 142, 150–151, 152, 153, 157, 160, 162, Pl. 41, Pl. 58 xanthosoma, Hestiochora 59, 150 Yponomeutidae 43 Zamiaceae 91 zea, Helicoverpa 43 Zingiberaceae 41, 44, 191, 208 Zygaena 33, 40, 42, 44, 46 Zygaenidae 1, 3, 27, 31, 34, 36, 38, 39, 41, 43, 45, 49, 50, 57, 58, 59, 61 Zygaeninae 1, 3, 4, 9, 10, 27, 31, 33, 34, 35, 36, 37, 38, 39, 41, 43, 44, 45, 46, 47, 49, 61, 64, 139 Zygaenoidea 3, 37, 39, 49 Zygaenoprocris 27, 44, 45, 50


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