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The Complete Field Guide to Butterflies of Australia
The Complete Field Guide to Butterflies of Australia
The Complete Field Guide to Butterflies of Australia
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The Complete Field Guide to Butterflies of Australia

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This is the first complete field guide to all butterfly species on Australia’s mainland and its remote islands.
Written by one of Australia's leading lepidopterists, it is stunningly illustrated with colour photographs of each of the 416 currently identified species. There is also a distribution map for each species on the Australian mainland.
It covers the five major family groups: Hesperiidae, Paplionidae, Pieridae, Nymphalidae and Lycaenidae, as well as the family Riodinidae, which has but a single species in Australia.
The introduction covers adult structure, classification, distribution and habitats, and life cycle and behaviour. This is followed by accounts of each of the 416 species, giving common name, scientific name, and other names (if any), as well as details of behaviour, habitat, status, and larval food plants. Accompanying each species is a distribution map, and photographs of the upperside and underside of both male and female specimens.
The book also contains a checklist of all species, a list of entomological contacts, a glossary, a bibliography, an index of common names and an index of scientific names.
LanguageEnglish
Release dateOct 25, 2004
ISBN9780643099685
The Complete Field Guide to Butterflies of Australia
Author

Michael F. Braby

Dr Michael Braby has been collecting and studying Australian butterflies for 35 years. He is a Visiting Fellow at the Australian National University and Chief Editor of Austral Entomology. He is particularly interested in the conservation, systematics, taxonomy, biogeography, biology and ecology of diurnal Lepidoptera and has published extensively in the field.

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    I want this book primarily to enable me to identify any butterfly that I may take photos of. It's really great, and I don't think I've found a butterfly that I've photographed yet that I haven't been able to find in here.

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The Complete Field Guide to Butterflies of Australia - Michael F. Braby

THE COMPLETE FIELD GUIDE TO

BUTTERFLIES

OF AUSTRALIA

Dedicated to Murray Upton, former Collection Manager

of the Australian National Insect Collection, for his wisdom,

practical knowledge and generous support over the years.

THE COMPLETE FIELD GUIDE TO

BUTTERFLIES

OF AUSTRALIA

MICHAEL F. BRABY

© CSIRO 2004

All photographs © Michael Braby, unless otherwise credited.

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

Braby, Michael F.

The complete field guide to butterflies of Australia.

Bibliography.

Includes index.

ISBN 0 643 09027 4.

1. Butterflies – Australia. 2. Butterflies – Australia -

Identification. I. Title.

595.7890994

Published by

CSIRO PUBLISHING

150 Oxford Street (PO Box 1139)

Collingwood VIC 3066

Australia

Distributed by

Steve Parish Publishing Pty Ltd

PO Box 1058,

Archerfield, Q 4108

Australia

Front cover illustration: Moonlight Jewel, Hypochrysops delicia.

Title page illustration: Northern Sword-grass Brown, Tisiphone helena.

Back cover illustration: Red-banded Jezebel, Delias mysis.

Cover and text design by James Kelly.

Typesetting and finished art by Roy Osborne.

Printed by Bookbuilders

Foreword

I can’t imagine what my life would have been if I had not been introduced to butterfly collecting at a summer camp when I was 13 years old. That started me into both a lifelong hobby and a career in science. Butterflies on the whole are just as beautiful as birds but in many ways easier to study. In most cases, adults can be collected without fear of threatening the viability of populations, and they can readily be marked with felt-tipped pens, released, and recaptured so that their movements may be tracked and their population sizes estimated. Many species can be cultured easily in the laboratory. They represent one of the most important and largest groups of organisms—plant-eating insects—and because of the activities of butterfly collectors, we often know what plants they eat. Above all, butterflies are readily identified to species because of the existence of field guides, of which this book is a magnificent example.

This means that while enjoying watching or collecting Australian butterflies, you also have the opportunity to add to scientific knowledge and contribute to the preservation of biological diversity. With millions of species and billions of natural populations of plants, animals, and microbes helping to run humanity’s life-support systems, there is no hope of understanding the details of the distribution and functioning of all of them. That’s why scientists have chosen butterflies of one of the major ‘model’ systems, an important sample from the panoply of life, to study in detail*. Butterflies are one large and diverse taxonomic group that science has a chance of understanding in depth—of virtually ‘completing’ in the near future. As Michael Braby notes, they already serve as a surrogate for many other groups of invertebrates and small vertebrates in conservation biology. For example, places with high butterfly diversity are also likely to have rich faunas of plant-eating insects of less well-studied groups. So taking careful notes of where and when you find each species, working out in different populations exactly what plants caterpillars eat and from which flowers adults sip nectar, and recording the behaviour of both caterpillars and butterflies, is not only fun, it’s important. Indeed, little or nothing is known about the life histories of some Australian species.

Some of my fondest memories, though, are just of the pleasure of knowing butterflies—of identifying beautiful and rare species in the field and, yes, enjoying adding specimens to my collection. And this guide is a perfect tool to help you to do this too!

Paul R. Ehrlich

Stanford University

* See, for example, Butterflies: Ecology and Evolution Taking Flight (2003) by C. L. Boggs, W. B. Watt and P. R. Ehrlich, and On the Wings of Checkerspots: A Model System for Population Biology (2004) by P. R. Ehrlich and I. Hanski.

Contents

Foreword

Preface

Acknowledgments

Introduction

Adult structure

Higher classification

Distribution and habitats

Life cycle and behaviour

How to use this book

How to identify Australian butterflies

Skippers Family Hesperiidae

Swallowtails Family Papilionidae

Whites and Yellows Family Pieridae

Nymphs Family Nymphalidae

Metalmarks Family Riodinidae

Blues Family Lycaenidae

Remote islands species

Checklist of Australian butterflies

Glossary

Bibliography

Index to common names

Index to scientific names

Further information

Preface

This book aims to provide a reliable way of identifying the adult stage of all butterflies in Australia, particularly when in the field. It includes all species recorded from the Australian continent, its continental islands and outlying political territories. Of the 416 butterfly species currently recognised, 398 are known from the mainland, while a further 18 species are found only on the more distant islands administered by various Australian governments.

The Introduction gives a brief overview of adult structure, higher classification, distribution and habitats, and provides notes on the life cycle and behaviour of the six families which occur in Australia. This is followed by a short chapter on how to use this book and identify specimens.

The species descriptions, which form the main body of the book (pages 32–309), provide a brief synopsis of each species. The butterflies are presented in a systematic sequence, which reflects current views on their higher classification, although within each subfamily the order of species may not follow their strict systematic order. To help with identification, there are notes for each species under the headings: similar species, variation, adult behaviour, habitat, status, larval food plants and larval attendant ants (where relevant). These notes are supplemented with distribution maps and charts of adult flight times. In some cases, line drawings of critical structures needed for identification are included with the species photographs.

Most of the information for the species accounts has been extracted and condensed from Butterflies of Australia: Their Identification, Biology and Distribution published in 2000. However, the text includes a considerable amount of new information, based on papers published in both the scientific and popular literature between 1999 and 2003, as well as new unpublished information, based either on my own observations or on personal communications from colleagues. Where particular facts published in Butterflies of Australia have since proven to be in error or inaccurate, these have been corrected.

At the end of the book there is a checklist of the Australian butterfly fauna, a list of entomological contacts, a short glossary, a bibliography, and indexes of the common and scientific names for all Australian butterflies.

I hope this field guide will help butterfly collectors, commercial breeders, biology students and professional entomologists, as well as being of value to conservation biologists, naturalists, tourists from overseas and those with only a casual interest in entomology. I trust it will also stimulate further interest in the natural history and conservation of these insects—especially in the conservation of their habitats—and find a place on the bookshelves of anyone who may simply wish to identify a butterfly in their garden.

Michael F. Braby

Canberra 2004

Acknowledgments

The introduction to this book relies heavily on the 1981 edition of Butterflies of Australia by Ian Common and the late Doug Waterhouse. I am very grateful for permission to incorporate many sections of text from that classic work into this book.

Most of specimens illustrated in this work are from the superb colour plates photographed by Dennis Crawford of Melbourne and originally published in Butterflies of Australia: Their Identification, Biology and Distribution in 2000. The plates have been supplemented with additional photographs prepared by the author, and I am most grateful to Ted Edwards and Vanna Rangsi (Australian National Insect Collection, Canberra) and Jan Forrest (South Australian Museum, Adelaide) for access to specimens in collections under their care; Cliff Meyer and Steve Brown also freely made available specimens in their extensive private collections. Digital images of specimens were kindly supplied by Kim Goodger and Phil Ackery (Department of Entomology, The Natural History Museum, London) and Steve Brown. Ted Edwards, Mike Couper and Peter Samson kindly provided additional images for inclusion in this book. All photographs in the introductory chapters are by the author unless otherwise indicated.

The distribution maps have been annotated from the base maps prepared by Murray Upton and published in Butterflies of Australia.

Andrew Atkins, David Britton, Steve Brown, Fabian Douglas, Rod Eastwood, David Lane, Russell Mayo, Cliff Meyer, Mike Moore, Chris Müller, Richard Weir, Matt Williams and Terry Woodger are thanked for clarifying or providing new information concerning the species text. Dr Don Harvey (National Museum of Natural History, Smithsonian Institution, Washington) helped with nomenclature of the higher classification of the Riodinidae, and Russell Cumming (Department of Environment) assisted with all things botanical.

I am very grateful to Nick Alexander and Murray Upton for their careful editing of the entire manuscript.

Finally, I thank my wife Lynette for her unfailing support and encouragement during the preparation of the manuscript.

Introduction

Butterflies are perhaps the best-known group of insects. Their brilliance and popularity have given them special significance and many people regard them with affection as the ‘birds’ of the insect world.

Australia does not have a particularly rich butterfly fauna by world standards, but nonetheless it has an interesting and unusual fauna, with nearly half of its species found nowhere else. The region also contains a number of distinctive endemic, or near endemic, groups that are crucial to our understanding of the origin and evolution of the world butterfly fauna.

It may come as a surprise to learn that Australia’s unique butterfly fauna has not yet been fully documented. New species continue to be discovered, and much remains to be learnt and recorded about the distribution, life histories, larval food plants and other aspects of the biology and behaviour of particular species.

Many earlier butterfly collectors reared adults from the immature stages but few recorded their observations, often in the mistaken belief that the biology and behaviour was well understood or that their observations were too trivial. Unfortunately, this is still true today. Although our knowledge has expanded rapidly in the last few decades, we still need to know much more about the taxonomy and ecological requirements of most species if we are to adequately conserve them.

Swamp Tiger, Danaus affinis.

Butterflies are playing an increasingly important role in conservation biology. They act as ‘flagships’ for the identification and preservation of critical habitats under threat, for the conservation of biodiversity, and as convenient ‘indicators’ for monitoring climate change or pollution. The more threatened species have become recognised as subjects worthy of protection and conservation in their own right. The focus on butterflies as flagships for conservation has given them special significance, as ‘ambassadors’ for the general conservation of invertebrate biodiversity, much of which is still poorly understood. With ongoing environmental degradation and deforestation the need to recognise, document and protect the remaining biological diversity on the planet has become more urgent than ever.

Amateur or private researchers have generated much of our scientific knowledge of Australian butterflies. Indeed, about 80% of all specimens in public museum collections have come from private collections donated by hobbyists. These specimens comprise an indispensable resource and form the very basis of our understanding of butterfly taxonomy, systematics, variation, seasonality and geographic distribution. Given the current lack of knowledge, collectors and naturalists can make a valuable contribution by establishing private reference collections, studying aspects of the biology and behaviour, and rearing the immature stages. Through careful observation and accurate recording, new discoveries can be published in news bulletins or local scientific journals. In this way private researchers can make a valuable contribution to the overall knowledge of Australian butterflies, at the same time gaining a deeper and richer satisfaction from their hobby.

Adult structure

Adult butterflies, like all other insects, comprise three major divisions: head, thorax and abdomen (Fig. 1). These are usually readily distinguishable, even though most of the adults’ body and appendages are covered with special flattened hairs or scales. The exoskeleton is made up of a series of rather rigid plates or sclerites joined by flexible membranes.

Figure 1. The main parts of the body of an adult butterfly.

Head

The most obvious feature of the head is a pair of large compound eyes, the surface of each of which is made up of hundreds of hexagonal-shaped lenses or facets. The compound eyes enable the insect to recognise shapes, colours and movement.

Arising from between the eyes is a pair of long, three-segmented feelers or antennae. The thick basal segment of each antenna is called the scape, the smaller second segment the pedicel, and the remainder of the antenna the flagellum. In butterflies the flagellum is thickened towards its tip to form a club, the slender portion of the flagellum being called the shaft. In the Hesperiidae, the apical portion of the club tapers to form an apiculus (Fig. 2).

The mouthparts consist of a strongly coiled tongue or proboscis and a pair of upturned, three-segmented appendages or labial palps. Towards its tip the proboscis is provided with a series of small chemo-receptors. When the adult drinks the proboscis is uncoiled and extended to suck in water, nectar or other liquids.

Peter Samson

The endangered Bulloak Jewel, Hypochrysops piceata.

Thorax

The thorax is composed of three segments, each bearing a pair of legs. The first segment or prothorax is the smallest of the three. The much larger second and third segments or meso- and metathorax each bear a pair of membranous wings and have strong internal ridges or projections, known as apodemes, to which the stout muscles necessary to operate the wings are attached.

Figure 2. Antennal clubs of skippers, Hesperiidae: A, ochres, Trapezites maheta group; B, ochres, Trapezites sciron group; C, grass-skippers, Anisynta; D, grass-skippers, Signeta; E, sedge-skippers, Motasingha; F, grass-darts, Taractrocera; G, grass-darts, Ocybadistes; H, darters, Telicota. Source: after Common and Waterhouse (1981).

Legs

The legs (Fig. 3) consist of five segments; coxa, trochanter, femur, tibia and tarsus, but the basal two segments are small and the coxae of the mid- and hind legs are rigidly attached to the thorax. The tarsi are normally five-segmented (Figs 3A, 3F, 3H), with a pair of apical claws, and usually bear chemo-receptors. The fore legs are fully developed for walking in the Hesperiidae, Papilionidae and Pieridae, but are reduced to various degrees, at least in the males, in the Nymphalidae (Figs 3C, 3E) and Lycaenidae (Fig. 3G). This reduction usually involves the loss of the terminal claws and a reduction of the tarsal segments, which may be completely fused to form one elongate segment. The fore tibia in the Hesperiidae and Papilionidae bears a movable lobe known as the epiphysis (Fig. 3A). This usually has a marginal comb of hairs or bristles thought to be used to clean the antennae and proboscis. The tibia of the mid-and hind legs usually bears a pair of apical spurs, and in some Hesperiidae the hind tibia also has a pair of median spurs. The paired tarsal claws of the mid- and hind legs are usually simple hooks, but in the Pieridae each claw is forked or bifid (Fig. 3B).

In many nymphalid and some lycaenid species, the sexes cannot be distinguished unless the legs are examined. A simple, reliable method of separating males and females is to examine closely the tarsus of the fore leg using a ×15 hand lens or microscope. In these two families, the fore leg of the female is normal but in the male the fore leg is reduced with the terminal claw absent and the tarsal segments absent or fused.

Wings

Adult butterflies have two pairs of fully developed wings. Each wing is essentially a flattened membranous sac, with the upper and lower membranes pressed closely together and strengthened by a series of tubes or veins. For descriptive purposes each fore or hind wing may be regarded as being triangular in shape (Fig. 4A). One corner of the triangle, attached to the thorax, forms the base of the wing, one the apex, and the other the tornus. The leading edge of the wing running from the base to the apex is the costa; the outer edge joining the apex to the tornus is the termen; and the trailing edge running from the base to the tornus is the inner margin or dorsum. Names are also given to the main areas of the wings: basal, subbasal, submedian, median, postmedian, subterminal, terminal, subcostal, costal, subapical, apical, subtornal and tornal.

Figure 3. Structure of fore legs of adults: A, male Hesperiidae (grass-skippers, Toxidia); B, tarsal claws of Pieridae (jezebels, Delias); C, male and D, female Nymphalidae (tigers, Danaus); E, male and F, female Nymphalidae (nymphs, Vanessa); G, male and H, female Lycaenidae (hairstreaks, Jalmenus).

Source: after Common and Waterhouse (1981) and B. Murray.

The currently accepted nomenclature of the wing veins of Lepidoptera is shown in Fig. 4B and detailed below. The costa of the fore wing is strengthened by the first main longitudinal vein or costal vein. The other veins of both fore and hind wings follow a characteristic basic pattern which includes a further five sets of longitudinal veins. These are the subcostal (Sc), radial (R), median (M), cubital (Cu) and anal veins (A). The subcostal vein is never branched in butterflies and always terminates on the costa. The radial vein is branched but the pattern differs in the fore and hind wings. In the fore wing, the radial vein has as many as five branches (R1 to R5 ) but some of these may be fused so that there are only four or even three branches. The branches of the radial vein may be separate, as in the Hesperiidae, or they may be ‘stalked’ and divide from one another. Sometimes R1 is partly confluent with the subcostal vein. In the hind wing, the radial vein has only two branches, R1 and the radial sector (Rs), but 1 R1 is entirely fused with Sc forming a 1 composite vein Sc+R . The basal sections of the radial and cubital veins together with weak transverse veins or discocellulars enclose the area known as the discal cell, or simply the ‘cell’. In this form the cell is said to be closed, however, if one or more of the discocellulars are absent, it is said to be open. The basal sections of the median vein have been lost in adult butterflies, and the three branches (M1, M2 and M3) usually begin at the disco cellulars. There are two main branches of the cubital vein, the anterior branch (CuA) and the posterior branch (CuP), but the latter has been almost entirely lost in butterflies, being found only in the fore wing of the Papilionidae. The anterior branch of the cubital vein has two further branches (CuA1 and CuA2 ). Of the three anal veins (1A, 2A and 3A), there are never more than two in butterflies, but often one of these is missing, and in the fore wing they may be either completely fused (i.e. 1A+2A) or separate for only a short distance near the base. An additional small vein, the humeral vein, is sometimes present near the base of the hind wing, running from the subcostal vein towards the basal end of the costa. Differences in wing venation between the butterfly families are shown in Figure 5.

Figure 4. Structure of wings of adults: A areas and B, venation. Source: after Common and Waterhouse (1981).

Figure 5. Wing venation of adults: A, Hesperiidae (Regent Skipper, Euschemon rafflesia); B, Hesperiidae (ochres, Trapezites); C, Papilionidae (kite swallowtails, Graphium); D, Pieridae (whites, Pieris); E, Nymphalidae (nymphs, Vanessa); F, Lycaenidae (hairstreaks, Jalmenus).

Source: modified after Nielsen and Common (1991).

The wing surfaces of most Lepidoptera are densely clothed with overlapping scales arising from minute sockets, which in butterflies are arranged more or less in transverse rows. The wing margins, especially between the apex and tornus, may be furnished with projecting marginal scales forming a scale-fringe. The colours of butterflies are due either to light being refracted by the physical structure of the scales or to pigments deposited in the scale wall. The colours of the scales are distributed to produce complex and beautiful patterns of bands, spots and rings. Bands may be complete, consist of a series of spots separated only by the veins, or as a broken series of spots with the constituent spots being more widely separated from one another. In addition to the normal clothing of wing scales, males of many species possess specialised sex-scales, or scent-hairs, known as androconia, which are associated with scent- or pheromone-producing glands and used during courtship. These sex-scales may terminate in a series of fine hair-like projections, and are either scattered among the normal wing scales or arranged in characteristic patches, sometimes known as sex-marks, sex-brands or scent-pouches, where they are concentrated along the costa of the fore wing or, where they are concentrated as dense tufts of long hairs, as hair-pencils. The presence of these patches of sex-scales is a useful feature for distinguishing males from females, and their shape, position and colour are a useful means of separating otherwise similar species.

Mike Coupar

Wing scales of the Scarlet Jezebel, Delias argenthona.

Wing venation provides important clues to the identity and relationships of most butterfly species and the main features can usually be examined without removing the wings from the specimen. If only a cursory examination is to be made the wings should be examined with a lens or dissecting microscope from the underside, while the wing is held at an oblique angle to a strong light source. The venation will stand out more strongly if the area to be examined is wetted with a drop of 70% ethyl alcohol or the scales brushed up into a vertical position.

Abdomen

The abdomen has 10 segments each of which consists of a dorsal plate or tergum and a ventral plate or sternum joined laterally by a membranous pleural area; the spiracles are found on the pleural area of the first seven segments. A thickened sternum is absent on the first segment. The last two or three segments are greatly modified to form the genitalia. In males, the ninth and tenth segments of the abdomen form the genitalia. Females have two genital openings: one opening, situated at the posterior end of the abdomen, is used for egg-laying, and the other, situated on the mid-ventral surface between the seventh and eighth sterna, is used for mating.

Higher classification

Biological classification involves the arrangement of groups of organisms into a hierarchical framework that reflects the evolutionary history of those organisms. Closely related species are classified into the same genus; related groups of genera are then classified into tribes, then families, then orders and so on. Sometimes subdivisions are made within the higher taxonomic categories, for example subtribe, subfamily, superfamily, etc. Each taxonomic category is given a standard suffix: iti for subtribe, ini for tribe, inae for subfamily, idae for family and oidea for superfamily.

Butterflies and moths belong to the large insect order Lepidoptera (which means scaled wings). They are distinguished from all other insects by the possession of two pairs of membranous wings clothed with overlapping scales. In Australia, there are around 10 500 scientifically named species of Lepidoptera arranged in about 30 superfamilies.

Butterflies are usually classified into two superfamilies, the Hesperioidea (generally known as skippers) and the Papilionoidea (all other butterflies). Malcolm Scoble, from the Natural History Museum in London, has provided convincing evidence that the Hedyloidea comprises a third superfamily of butterflies. The Hedyloidea are an unusual group of nocturnal butterflies from Central and South America.

The Hesperioidea and Papilionoidea are distinguished from one another by the shape of the head and antennae, and the venation of the fore wing. In the Hesperioidea, the antennae, which arise farther apart on the head, are thickened subapically with the basal portion of the club gradually thickened while the apical portion, known as the apiculus, is curved or sharply bent. In the Papilionoidea, the fore wing has the peripheral veins stalked, but not in the Hesperioidea.

The relationship of the three butterfly superfamilies to the rest of the Lepidoptera remains unclear, although the Uranioidea (swallowtail moths) and Calliduloidea (old world butterfly moths) appear to be the closest relatives. Moreover, there is no single character that will always distinguish butterflies from moths. Traditionally, butterflies have been distinguished from other Lepidoptera by their day-flying habits, bright colours, ability to close their wings dorsally over the thorax, the presence of a large humeral lobe on the hind wing, absence of a frenular wing-coupling device, and possession of clubbed antennae. However, some butterflies do not meet all of these requirements, for example, the male of the Australian Regent Skipper, Euschemon rafflesia, has its wings coupled together by means of a frenulum and retinaculum (Fig. 5A), a wing-coupling device found in many groups of moths, while some moths share ‘butterfly’ characteristics, for example, many castniids have clubbed antennae and are day-flying. Since butterflies represent only a small fraction of the Lepidoptera and belong to the Suborder Glossata, series Ditrysia, which includes about 98% of all species of moths, it is probably more accurate to consider butterflies as a group of day-flying ditrysian moths!

Figure 6. A simple family tree of butterfly superfamilies and familes in Australia. Numbers refer to the number of species in each family on mainland Australia.

The Hesperioidea contains a single family, the Hesperiidae, whereas the Papilionoidea is generally divided into four or five families: Papilionidae (swallowtails), Pieridae (whites and yellows), Nymphalidae (nymphalids), Riodinidae (metalmarks) and Lycaenidae (blues)(Fig. 6). All of the six families of butterflies occur in Australia and its remote islands although the proportion of species in each of these families varies greatly. In Australia, the lycaenids and hesperiids are the most numerous groups, representing two-thirds of the total number of butterfly species, whereas the papilionids, pierids and riodinids, are very poorly represented.

The checklist of the 416 species currently recognised from Australia includes 18 species from remote islands (see pages 302–309) and is arranged according to their higher classification. The higher classification of each family (i.e. subfamily, tribe, subtribe) primarily follows Evans (1949), Ackery (1984) and Vane-Wright (2003) for the Hesperiidae; Miller (1987), de Jong et al. (1996) and Ackery et al. (1999) for the Papilionidae; Braby and Pierce (in press) for the Pieridae; Vane-Wright (2003) for the Nymphalidae, including Miller (1968) for the Satyrinae and Penz and Peggie (2003) for the Heliconiinae; Vane-Wright (2003) for the Riodinidae; and Pierce et al. (2002) and Vane-Wright (2003) for the Lycaenidae.

Distribution and habitats

Butterflies are primarily a tropical group of insects and often depend on rainforest plants for larval food. They are not well adapted to arid environments of low moisture and extremes of high temperature. Hence the best place to find butterflies in Australia is in the northern, coastal areas of north-eastern Queensland where conditions of temperature, humidity and rainfall are optimal and large tracts of rainforest still remain.

Chocolate Argus, Junonia hedonia.

Australia is divided into five major ‘faunal provinces’—biogeographical regions based on climatic and environmental factors (Fig. 7). Within these five provinces, 314 butterfly species have been recorded from the tropical-subtropical Torresian province. In more temperate areas, 193 species have been recorded from the Bassian province and 56 species from the South-western province. In the Timorian province there are 126 species, but only 92 have been recorded from the large arid Eyrean province, which receives an annual rainfall of less than about 500 mm. Several species range into this arid area only very slightly or temporarily. Species richness is highest along the moist, eastern coast and declines rapidly west of the Great Dividing Range.

Three areas support exceptionally high numbers of species of butterflies: Cape York Peninsula; the Wet Tropics (between Cooktown and Townsville, including the Atherton Tableland, QLD); and a subtropical area in south-eastern Queensland and northeastern New South Wales where the Bassian and Torresian faunas overlap. By far the richest part of Australia is northern Queensland where 271 species (68% of the total fauna) are found north of Townsville, of which 226 occur on Cape York Peninsula north of Coen.

Figure 7. Major faunal provinces showing the number of butterfly species in each. Source: C. Hunt.

The photographs on pages 11–13 illustrate the range of habitats or natural vegetation types in Australia where butterflies typically occur. Most species make use of several or even many different habitats, but some species live in just one type of habitat, particularly those with very specialised ecological needs.

Nearly half of the butterflies (173 species) in Australia are associated with rainforest (i.e. forests with a closed canopy) and of these 114 species are found only in these habitats. Rainforest habitat types include monsoon forest (tropical semi-evergreen/semi-deciduous dry rainforest), littoral rainforest (rainforest close to the shore line), gallery forest (narrow corridors of tropical rainforest along the banks of

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