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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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Apr 1, 2016
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9781486301027
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Description

As fascinating as they are beautiful, butterflies are a pleasure to watch and an important group of invertebrates to study. This second edition of the award-winning book The Complete Field Guide to Butterflies of Australia is a fully updated guide to all butterfly species on Australia's mainland and remote islands.

Written by one of Australia's leading lepidopterists, the book is stunningly illustrated with colour photographs, many of which are new, of each of the 435 currently recognised species. There is also a distribution map and flight chart for each species on the Australian mainland, together with information on similar species, variation, behaviour, habitat, status and larval food plants.

The introduction to the book covers adult structure, higher classification, distribution and habitats, as well as life cycle and behaviour. A new chapter on collecting and preserving butterflies is included. There is also an updated checklist of all species, a glossary, a bibliography and indexes of common and scientific names.

Released:
Apr 1, 2016
ISBN:
9781486301027
Format:
Book

About the author

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

Dedicated to Ted Edwards, former Curator of Lepidoptera of the Australian National Insect Collection, for his encouragement, encyclopaedic knowledge and generous support over many years.

Text © Michael F. Braby 2016

All photographs © Michael F. Braby, unless otherwise credited

Butterfly plates and maps © CSIRO

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., author.

The complete field guide to butterflies of Australia /

Michael F. Braby.

Second edition.

9781486301003 (paperback)

9781486301010 (epdf)

9781486301027 (epub)

Includes bibliographical references and index.

Butterflies – Australia.

Butterflies – Australia – Identification.

595.7890994

Published by

CSIRO Publishing

Locked Bag 10

Clayton South VIC 3169

Australia

Telephone: +61 3 9545 8400

Email: publishing.sales@csiro.au

Website: www.publish.csiro.au

Front cover: (main image) Orange Alpine Xenica, Oreixenica correae.

Back cover: Bright Copper, Paralucia aurifera

Edited by Joy Window

Cover design by James Kelly

Typeset by Thomson Digital

Printed in China by 1010 Printing International Ltd

CSIRO Publishing publishes and distributes scientific, technical and health science books, magazines and journals from Australia to a worldwide audience and conducts these activities autonomously from the research activities of the Commonwealth Scientific and Industrial Research Organisation (CSIRO). The views expressed in this publication are those of the author(s) and do not necessarily represent those of, and should not be attributed to, the publisher or CSIRO. The copyright owner shall not be liable for technical or other errors or omissions contained herein. The reader/user accepts all risks and responsibility for losses, damages, costs and other consequences resulting directly or indirectly from using this information.

Original print edition:

The paper this book is printed on is in accordance with the rules of the Forest Stewardship Council®. The FSC® promotes environmentally responsible, socially beneficial and economically viable management of the world’s forests.

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 as one of the major ‘model’ systems, an important sample from the panoply of life, to study in detail.* Butterflies make up 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, Boggs CL, Watt WB, Ehrlich PR (2003) Butterflies: Ecology and Evolution Taking Flight, University of Chicago Press, Chicago, and Ehrlich PR, Hanski I (2004) On the Wings of Checkerspots: A Model System for Population Biology, Oxford University Press, Oxford.

Warm temperate rainforest, Minnamurra Falls National Park, New South Wales.

Contents

Foreword

Preface to the first edition

Preface to the second edition

Acknowledgments

Introduction

Adult structure

Higher classification

Distribution and habitats

Life cycle and behaviour

How to use this book

How to identify Australian butterflies

Collecting and preserving butterflies

Swallowtails Family Papilionidae

Skippers Family Hesperiidae

Whites and Yellows Family Pieridae

Nymphs Family Nymphalidae

Metalmarks Family Riodinidae

Blues Family Lycaenidae

Islands

Checklist of Australian butterflies

Glossary

Bibliography

Index to common names

Index to scientific names

Further information

Coastal heathland, Broadwater National Park, New South Wales.

Preface to the first edition

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 continent, 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 that occur in Australia. A short chapter then follows on how to identify specimens.

The species descriptions, which form the main body of the book, provide a brief synopsis of each species. The butterflies are presented in a systematic sequence, which reflects current views on 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, the 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 Braby (2000) Butterflies of Australia: Their Identification, Biology and Distribution. 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 short glossary, a bibliography, indexes of the common and scientific names for all Australian butterflies and a list of entomological contacts.

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 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

Preface to the second edition

It is now 12 years since the first edition of this book was published, which sold more than 15 000 copies and is now out of print, and which won a Whitley Award (Certificate of Commendation) for the best book in the category of Field Guide in 2005. The extent and volume of new information on Australian butterflies that has been published during the last decade has been staggering – eight books and more than 700 research papers and popular articles on Australian butterflies were published between 2000 and 2014 (an average of approximately one article per week!). It was therefore decided that a completely new, revised edition was required to keep pace with this burgeoning literature, and to provide an up-to-date account of current knowledge. The number of butterflies formally recognised from Australia, for instance, has now risen to 435 species, of which 408 are known from the continent while a further 27 species are found only on the more distant islands administered by various Australian governments, as result of field exploration and inventory of remote areas of the country, together with ongoing taxonomic research. There have also been considerable advances in the theory and practice of biological systematics as a science, which has resulted in substantial changes to the higher classification of butterflies worldwide, mainly as a result of the advent of molecular biology and new methods for sequencing DNA, together with new developments in phylogenetic methods and an explosion of new tools (computer programs) to analyse large and complex datasets combining both morphological characters and molecular data. Consequently, the classification and order of species adopted in this work is quite different to that of the first edition.

The layout and scope of the book is similar to that of the first edition, but a new chapter on collecting and preserving butterflies has been included, and the sections on higher classification, distribution and habitats, and life cycle and behaviour in the Introduction have been substantially revised. The literature pertaining to Australian butterflies has, as far as possible, been reviewed up to the end of 2014 and this new information has been incorporated into the text for the species accounts. The distribution maps and flight charts have also been updated for almost every species, and 106 new colour images depicting set butterflies have been added to, or have replaced previous images in, the colour plates.

Dr Michael F. Braby

The Australian National University

Canberra 2016

Acknowledgments

The introduction to this book relies heavily on the 1981 edition of Butterflies of Australia by the late 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 the 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 (Australian National Insect Collection, Canberra), Gavin Dally (Museum and Art Gallery of the Northern Territory, Darwin), David Britton (Australian Museum, Sydney), Simon Hinkley and Catriona McPhee (Museum Victoria, Melbourne), Steve Brown (Bowral) and the late Lindsay Hunt (Adelaide) for access to specimens in collections under their care. Digital images of the following species were also generously supplied by Trevor Lambkin (Graphium codrus, Cephrenes moseleyi, Cyrestis achates, Taenaris myops, Hypochrysops chrysargyrus female, Arhopala philander), Grant Miller (Delias lara), Cliff Meyer (Euploea modesta), Geoff Thompson of the Queensland Museum (Croitana arenaria pilepudla) and Dick Vane-Wright of the Natural History Museum, London (Appias albina infuscata). Ted Edwards, Peter Samson, Mike Coupar and the late Ian Common kindly provided additional images for inclusion in the introductory chapters of this book. All photographs in the introductory chapters are by the author unless otherwise indicated.

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

Andrew Atkins, Deb Bisa, David Britton, Fabian Douglas, Rod Eastwood, Don Franklin, Trevor Lambkin, Cliff Meyer, Mike Moore, Chris Müller, Matt Williams and Andy Williams are thanked for clarifying or providing new information, or clarifying certain facts, concerning the species text. Malte Ebach helped with recent developments in the nomenclature and biogeographic regionalisation of Australia, and kindly provided access to unpublished material. Russell Cumming, Kym Brennan, Ian Cowie, Donna Lewis and John Westaway assisted with all things botanical; John Westaway provided expert advice on vegetation classifications and habitats and critically improved the text.

I am very grateful to Joy Window for her 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 size and brilliance have given them special significance and unrivalled popularity, and many people regard them with affection as the ‘birds’ of the insect world.

Australia does not have a rich butterfly fauna by world standards, but nonetheless it has an interesting and unusual fauna, with half of its species found nowhere else. The region also contains several distinctive endemic, or nearly endemic, groups that are crucial to our understanding 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 catalogued. 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. Moreover, in contrast to the Northern Hemisphere, national longterm recording schemes to monitor broad changes in distribution, relative abundance and/or assemblages at particular reference sites over time are currently not in existence in Australia.

Tawny Coster, Acraea terpsicore, a new butterfly for Australia which arrived from South-East Asia in 2012.

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 were well understood or that their observations were too trivial. Unfortunately, this is still true today. Although our knowledge has expanded rapidly in the last four decades, we still need to know much more about the taxonomy, distribution and ecological requirements of most species if we are to adequately conserve them.

Butterflies are now playing an increasingly important role in conservation biology worldwide (New 1997, 2014). They act as ‘flagships’ for promoting awareness of conservation need for insects generally or focusing attention on particular threatened species and their critical habitats in need of protection and conservation management, and as convenient bioindicators for monitoring the quality and health of our natural environment, such as the effects of habitat loss through deforestation and urbanisation, climate change or pollution. The more threatened species have become recognised as subjects worthy of protection and conservation management 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 has not been formally described. 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, therefore, 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 adult’s 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 features of the head are 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 chemoreceptors. When the adult drinks the proboscis is uncoiled and extended to suck in water, nectar or other liquids.

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).

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, respectively, 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.

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 chemoreceptors. 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). The epiphysis 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 to form an elongated segment.

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

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 third 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 dorsum or inner margin. 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.

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 (C). 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 (A) veins. 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 R1 is entirely fused with Sc forming a composite vein Sc+R1. 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 discocellulars. 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 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 Fig. 5.

Figure 4. Structure of wings of adults: (A), areas and (B), venation.

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

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. The patterns primarily serve to attract mates, advertise warning colouration to potential predators or avoid detection through camouflage or crypsis. Bands of colours may be complete, consist of a series of spots separated only by the veins, or of 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.

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.

Wing scales of the Scarlet Jezebel, Delias argenthona.

Mike Coupar

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 reproductive organs or 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 and superfamily. Each taxonomic category is given a standard suffix: ina for subtribe, ini for tribe, inae for subfamily, idae for family and oidea for superfamily.

Butterflies and moths belong to the order Lepidoptera (which means scaled wings), a large group of insects containing more than 165 000 described species (Kristensen et al. 2007) that arose in the late Triassic and then diversified in the Cretaceous (Misof et al. 2014). They are distinguished from all other orders of 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.

Until fairly recently butterflies were classified into three of these superfamilies; the Hedyloidea, Hesperioidea and Papilionoidea (Scoble 1986; Ackery et al. 1999; Vane-Wright 2003; Wahlberg et al. 2005). However, four independent, molecular phylogenetic or phylogenomic studies based on large numbers of genetic markers and comprehensive taxon sampling across the Lepidoptera (Regier et al. 2009; Mutanen et al. 2010; Heikkilä et al. 2012; Kawahara and Breinholt 2014) have revealed that this classification does not accurately reflect the evolutionary history of the group and therefore is unsatisfactory. Consequently, the systematics and higher classification of the butterflies has been completely revised to now comprise just a single superfamily, the Papilionoidea, with seven families: Papilionidae, Hedylidae, Hesperiidae, Pieridae, Nymphalidae, Riodinidae and Lycaenidae (van Nieukerken et al. 2011; Simonsen et al. 2012).

The relationship of the Papilionoidea to the rest of the Lepidoptera remains unclear, but the Calliduloidea (Old World butterfly moths) and Thyridoidea (picture-winged leaf moths) appear to be the closest relatives to the butterflies. 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, the 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, callidulids, castniids and zygaenids are brightly coloured and day-flying and some have clubbed antennae. Since butterflies represent only a small fraction of the Lepidoptera and belong to the suborder 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 evolutionary tree showing relationships of butterfly families. Numbers refer to the number of species in each family in Australia, including its remote islands.

Relationships within the Papilionoidea are now fairly well established (Regier et al. 2009; Mutanen et al. 2010; Heikkilä et al. 2012) (Fig. 6). The Papilionidae (swallowtails) comprise a lineage that diverged very early from the remaining families in the evolutionary history of the butterflies. The six other families descended from two other lineages or ‘branches’ in the evolutionary tree. One branch gave rise to the Hesperiidae (skippers) and the Hedylidae, an unusual group of nocturnally active butterflies from Central and South America – these two families are closely related and evolved from a common ancestor. The other branch gave rise to the remaining families, which include the Pieridae (whites and yellows), Nymphalidae (nymphs), Riodinidae (metalmarks) and Lycaenidae (blues).

In Australia, all but the Hedylidae occur on the continent and its remote islands, although the numbers of species in each of these families varies greatly. In Australia, the blues and skippers are the most numerous groups, representing approximately two-thirds (64% or 279 species) of the total number of butterfly species, whereas the swallowtails, whites and yellows, and metalmarks are less well represented.

The checklist (near the end of this book) of the 435 species currently recognised from Australia includes 27 species from remote islands (see pages 328–341) and is arranged according to their higher classification. In this work, the higher classification of each family (i.e. subfamily, tribe, subtribe) primarily follows Nazari et al. (2007) and Simonsen et al. (2010) for the Papilionidae; Warren et al. (2008, 2009) for the Hesperiidae; Wahlberg et al. (2014) for the Pieridae; Wahlberg and Wheat (2008) and Wahlberg et al. (2009) for the Nymphalidae; Harvey (1987) and Vane-Wright (2003) for the Riodinidae; and Pierce et al. (2002) and Vane-Wright (2003) for the Lycaenidae. Braby (2010) provided a global overview of most of these and other recent studies in relation to the Australian fauna, including a detailed review of systematic revisions of subfamilies, tribes and subtribes relevant to Australia.

Distribution and habitats

Of the 435 species of butterflies recorded from Australia and its remote islands, 408 are known from the continent. Of the species recorded from the mainland and adjacent islands, nine (Borbo cinnara, Delias lara, Danaus chrysippus, Euploea leucostictos, E. climena, Vanessa cardui, Junonia erigone, Hypolimnas anomala, Charaxes andrewsi) are not permanently established, comprising either rare vagrants or breeding populations that appear to be temporary, dependant on immigrants from elsewhere. The residential/breeding status of a further three species (Appias celestina, Lexias aeropa, Apaturina erminia) is currently uncertain. Thus, the resident fauna that is permanently established in Australia comprises at least 396 species.

Butterflies are not evenly distributed across the Australian continent. They are primarily a tropical group of insects and often depend on mesic habitats and the plants which they support 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 along the northern and eastern coastal areas of Australia, especially in north-eastern Queensland where conditions of temperature, humidity and rainfall are optimal and patches of rainforest still remain.

A recent review of biogeographical regions of Australia (Ebach 2012), together with new analyses of plant distributions (González-Orozco et al. 2014; Ebach et al. 2015), has identified five broad phytogeographical regions: Northern, Euronotian, Northern Desert, Eremaean and South-western (Fig. 7). These biogeographical regions are further subdivided into subregions and are also correlated with climatic and environmental factors, particularly the seasonal distribution and variation in mean annual temperature and rainfall.

Because most butterflies depend on plants for their survival (in the larval stage), it is instructive to compare the diversity within each of these five phytogeographical regions – 292 butterfly species have been recorded from the Northern region, which encompasses much of the monsoon tropics, while 284 species have been recorded from the Euronotian region, which includes the mesic subtropical and temperate areas along the eastern and southern seaboard. In contrast, two of the three other areas are characterised by semi-arid and arid climates and support substantially fewer butterflies: 67 have been recorded from the large Eremaean region, which receives an annual rainfall of less than about 500 mm; and 59 species from the Northern Desert region, which has a low summer rainfall. Fifty-eight species have been recorded from the South-western region, which has a mediterranean climate. Several species range into the arid interior only marginally or temporarily. Species richness is thus highest along the moist, eastern and northern coast and declines rapidly west of the Great Dividing Range (Kitching and Dunn 1999).

Within the Northern and Euronotion regions, three areas support exceptionally high numbers of species of butterflies: Cape York Peninsula (particularly Iron Range-McIlwraith Range); the Wet Tropics (between Cooktown and Townsville, including the Atherton Tableland, QLD); and a subtropical area in south-eastern Queensland and north-eastern New South Wales where the tropical and temperate faunas overlap. By far the richest part of Australia is northern Queensland where 277 species (68% of the mainland fauna) are found north of 20°S latitude (Bowen), of which 228 occur on Cape York Peninsula (i.e. the area north of Princess Charlotte Bay at ~14.5°S latitude).

Figure 7. Major biogeographical regions within Australia, showing the number of butterfly species in each.

In terms of endemism, 200 (49%) of the 408 species recorded from the Australian continent are endemic to the mainland and Tasmania – that is, half of the total number of species are unique to Australia and occur nowhere else in the world. Within the five biogeographical regions, 100 species are restricted to the Northern region (of which 25 species or 8.5% of the total are endemic), 75 species are restricted to the Euronotion region (of which 73 species or 25.7% of the total are endemic), while 14 species are restricted to the South-western region (of which 13 species or 22.4% of the total are endemic). Only one species is endemic to each of the Eremaean (Desert Sand-skipper, Croitana aestiva) and Northern Desert (Dark Opal, Nesolycaena medicea) regions. The Euronotion and South-western regions are thus distinctive in that they contain highest levels of endemism (25.7% and 22.4%, respectively), indicating that they support relatively large proportions of butterflies that are found nowhere else.

Although political boundaries within Australia bear little relationship to the natural distribution of the fauna, it is sometimes useful to know the number of species found in each state and territory. There are 343 species recorded from Queensland, 217 species from New South Wales, 135 from the Northern Territory, 128 from Victoria, 127 from Western Australia, 86 from the Australian Capital Territory, 73 from South Australia, and 42 from Tasmania. Only three species, all in the subfamily Satyrinae, are endemic to Tasmania.

The photographs on pages 14–18 illustrate the variety of habitats in Australia where butterflies breed and their larval food plants grow. These habitats are based largely on natural vegetation types and structural classification, according to the national vegetation information system (Executive Steering Committee for Australian Vegetation Information 2003), together with topographic features and climatic factors. Most species make use of several or even many different habitats, but some species live in just one type of habitat, often within a particular microhabitat, especially those with very specialised ecological requirements in which the larvae feed on just one plant species.

An example of a habitat specialist is the Black Grass-dart, Ocybadistes knightorum, which breeds in sheoak (Casuarina) open-forest with an open grassy understorey dominated by its sole larval food plant, the grass Alexfloydia repens, in semi-saline areas of the king tide zone above mangroves in subtropical coastal lowlands. In contrast, the closely related Green Grass-dart, Ocybadistes walkeri, is a habitat generalist because it breeds on at least 13 species of Poaceae, including several introduced weeds, that grow in a variety of habitats, from tropical savannah woodland to temperate

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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.