The Insect of Australia
The Insect of Australia
The Insect of Australia
Insects of
Australia
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The
Insects of
Australia
A textbook for students
and research workers
SECOND EDITION
K. VAN ACHTERBERG,
Rijksmuseum van Natuurlijke Historie, Leiden, The Netherlands (Chapter 42)
H. ASPocK,
Parasitologie, Hygiene-Institut der Universitat, Vienna, Austria (Chapter 33)
U. ASPocK,
Naturhistorisches Museum, Vienna, Austria (Chapter 33)
J. BALDERSON,
Division of Entomology, CSIRO, Canberra (Chapter 21)
E. B. BRITTON,
Division of Entomology, Canberra (Chapter 35)
G. W. BYERS,
Department of Entomology, University of Kansas, Lawrence, U. S. A. (Chapter 37)
J. H. CALABY,
Division of Wildlife and Ecology, CSIRO, Canberra (Chapter 29)
I. C. CAMPBELL,
Centre for Stream Ecology, Chisholm Institute of Technology, Melbourne (Chapter 16)
M. CARVER,
Division of Entomology, CSIRO, Canberra (Chapter 30)
G. CASSIS,
Division of Entomology, CSIRO, Canberra (Chapter 30)
R. F. CHAPMAN,
University of Arizona, Tucson, U.S.A. (Chapter 2)
D. H. COLLESS,
Division of Entomology, CSIRO, Canberra (Chapter 39)
I. F. B. COMMON,
32 Katoomba Crescent, Toowoomba, Queensland (Chapter 41)
B. CONDE,
Musee de Zoologie de l'Universite de la Ville de Nancy, France (Chapter 13)
P. S. CRANSTON,
Division of Entomology, CSIRO, Canberra (Chapter 4)
J. C. DEAN
Water Sciences Laboratories, Board of Works, Melbourne (Chapter 40)
viii Authors
V. A. DRAKE,
Division of Entomology, CSIRO, Canberra (Chapter 3)
G. M. DUNNET,
Department of Zoology, University of Aberdeen, United Kingdom (Chapter 38)
J. W. EVANS,
formerly Bellevue Hill, Sydney (Chapter 30)
R. A. FARROW,
Division of Entomology, CSIRO, Canberra (Chapter 3)
M. J. FLETCHER,
Biological and Chemical Research Institute, N.S.W. Department of Agriculture, Sydney (Chapter 30)
F. J. GAY,
Division of Entomology, CSIRO, Canberra (Chapter 20)
PENELOPE J. GREENSLADE,
Division of Entomology, CSIRO, Canberra (Chapter 11)
G. F. GROSS,
South Australian Museum, Adelaide (Chapter 30)
P. J. GULLAN,
Department of Zoology, Australian National University, Canberra (Chapter 4)
B. S. HEMING,
Department of Entomology, University of Alberta, Edmonton, Canada (Chapter 31)
L. HILL,
Tasmanian Department of Agriculture, Devonport (Chapter 30)
T. F. HOUSTON,
Western Australian Museum, Perth (Chapter 42)
G. IMADATl~,
Biological Laboratory, Konodai College, Tokyo Medical and Dental University, Japan (Chapter 12)
1. KATHIRITHAMBY,
Department of Zoology, University of Oxford, United Kingdom (Chapter 36)
D. K. McE. KEVAN,
MacDonald College, McGill University, Quebec, Canada (Chapter 23)
K. H. L. KEY,
Division of Entomology, CSIRO, Canberra (Chapter 25)
N. P. KRISTENSEN,
Zoological Museum, University of Copenhagen, Denmark (Chapter 5)
J. KUKALOVA-PECK,
Department of Geology, Carleton University, Ottawa, Canada (Chapter 6)
I. LANSBURY,
Hope Department of Entomology, University Museum, Oxford, United Kingdom (Chapter 30)
J. F. LAWRENCE,
Division of Entomology, CSIRO, Canberra (Chapters 1, 35)
D. K. McALPINE,
The Australian Museum, Sydney (Chapter 39)
I. M. MACKERRAS,
formerly Division of Entomology, CSIRO, Canberra (Chapter 1)
M. B. MALIPATIL,
Northern Territory Museum of Arts and Sciences, Darwin (Chapter 30)
D. K. MAROON,
Fagus, Manse Road, Killin, Perthshire, United Kingdom (Chapter 38)
E. N. MARKS,
Queensland Institute of Medical Research, Brisbane (Chapter 8)
C. D. MICHENER,
Department of Entomology, University of Kansas, Lawrence, U.S.A. (Chapter 42)
G. B. MONTEITH,
Queensland Museum, Brisbane (Chapter 30)
M. S. MOULDS,
Waitara, Sydney (Chapter 30)
L. MOUND,
Entomology Department, British Museum (Natural History), London, United Kingdom (Chapter 31)
M. D. MURRAY,
Pymble, Sydney (Chapter 29)
Authors ix
I. D. NAUMANN,
Division of Entomology, CSIRO, Canberra (Chapters 7, 42)
A. NEBOISS,
Museum of Victoria, Melbourne (Chapter 40)
T. R. NEW,
Department of Zoology, La Trobe University, Melbourne (Chapter 34)
E. S. NIELSEN,
Division of Entomology, CSIRO, Canberra (Chapters 1, 41)
K. R. NORRIS,
Division of Entomology, CSIRO, Canberra (Chapter 3)
A. F. O'FARRELL,
Department of Zoology, University of New England, Armidale (Chapter 17)
J. PAGES,
Faculte des Sciences de la Vie et de l'Environment, Laboratoire de Biologie animale et generale, Dijon, France
(Chapter 13)
W. L. PETERS,
Department of Entomology, Florida Agricultural and Mechanical University, Tallahassee, U.S.A. (Chapter 16)
J. T. POLHEMUS,
University of Colorado Museum, Englewood, U.S.A. (Chapter 30)
D. C. F. RENTZ,
Division of Entomology, CSIRO, Canberra (Chapters 22, 23, 24)
E. S. Ross,
California Academy of Sciences, San Francisco, U.S.A. (Chapter 26)
L. M. ROTH,
Museum of Comparative Zoology, Harvard University, Cambridge, U.S.A. (Chapter 19)
J. A. SLATER,
Systematic and Evolutionary Biology Section, University of Connecticut, Storrs, U.S.A. (Chapter 30)
G. B. SMITH,
Agricultural Division, Bayer Australia, Sydney (Chapters 14, 15)
C. N. SMITHERS,
The Australian Museum, Sydney (Chapters 27, 28)
P. STYS,
Department of Systematic Zoology, Charles University, Prague, Czechoslovakia (Chapter 30)
K. L. TAYLOR,
Division of Entomology, CSIRO, Canberra (Chapter 30)
R. W. TAYLOR,
Division of Entomology, CSIRO, Canberra (Chapters 4, 42)
G. THEISCHINGER,
20 Leawarra Street, Engadine, New South Wales (Chapters 18,32)
D. F. WATERHOUSE,
Division of Entomology, CSIRO, Canberra (Chapter 9)
J. A. L. WATSON,
Division of Entomology, CSIRO, Canberra (Chapters 14, 15, 17,20)
T.A. WEIR,
Division of Entomology, CSIRO, Canberra (Chapter 30)
M. J. WHITTEN,
Division of Entomology, CSIRO, Canberra (Chapter 10)
D. J. WILLIAMS,
C. A. B. International, London, United Kingdom (Chapter 30)
T. E. WOODWARD,
formerly Queensland Museum, Brisbane (Chapter 30)
Editorial Committee
P. B. Came (Co-ordinator)
J. F. Lawrence
E. S. Nielsen
J. P. Spradbery
R. W. Taylor
M. 1. Whitten
M. J. Littlejohn
Foreword
The first important general work on the insects of this continent, Australian Insects by W. W. Froggatt, was published in
1907. It was written primarily from the point of view of the field naturalist, and there is no doubt that it proved most
useful to the student of those days. Nineteen years later, the classical Insects of Australia and New Zealand by R. J.
Tillyard appeared, marking a major advance both in the content of accumulated knowledge and in the wealth of biology
and comparative morphology that it presented.
'Tillyard' had long been out of print when, in 1970, the first edition of the present book was published. It was a co-
operative venture, sponsored by the Division of Entomology, written largely by Divisional staff, and edited by Dr I. M.
Mackerras. That the book received international acclaim was due in no small part to Mackerras's scholarship and com-
mitment. The book has since found wide use as a student text and as an essential tool for professional entomologists.
By 1984 it was clear that the first edition (and material included in the 1974 Supplement) was in many respects out-
dated, and a revised edition was mooted. Where possible, the authors of chapters were invited to revise their 1970 texts
or to recommend specialists to assist or replace them in this task. Again CSIRO entomologists assumed a major role.
Specialists in other Australian and overseas institutions agreed to contribute and the book is all the more authoritative
for their efforts.
The result of the revision is almost a new book. Some text and many illustrations have been retained unaltered from
the first edition. However, most chapters have been rewritten entirely to incorporate the wealth of new information now
available. Many new illustrations have been added, some illustrations from the first edition have been amended, and
most of the plates have been rearranged and relabelled. Material of a physiological and cytological nature, or dealing
with aspects of reproduction and metamorphosis, has either been incorporated into other chapters (especially an expand-
ed Chapter 2) or omitted. In its place new chapters deal specifically with the impingement of insects on human history
and economy, with the use of insects in scientific research, and with the history of Australian entomology. Another new
feature is a key to the insect orders in Chapter 1.
The task of the authors/revisers of chapters was to give an account of the insects of Australia primarily from the sys-
tematic point of view, but with some account of their morphology, where they live, and what they do. To achieve even
this limited objective in two volumes of reasonable size has involved rigorous selection of subject matter and imposed
severe limitations on its presentation. We expect that, in consequence, few authors of the systematic chapters are really
satisfied with the adequacy with which the allocated space has permitted them to cover their orders. Apart from consid-
erations of space it has been necessary for all authors to conform rather closely to an established pattern; this they have
accepted for the most part cheerfully, although they would sometimes have preferred to deviate to suit their own particu-
lar needs.
The expansion in our knowledge of Australian insects since 1926 is reflected in the increase in the number of known
species and families. Tillyard estimated 37 300 species and listed 401 families. The first edition of The Insects of
xi
xii Foreword
Australia recorded 54071 species (an increase of 45% over Tillyard) and 574 families. In the present two volumes we
see the figures escalate to 85920 (an increase of 59% over the first edition) in 661 families. These numbers reflect
increased specialist exploration of the continent but alone do not depict the magnitude of the work still to be done. At
least some entomologists estimate that more than half of Australia's insect species have yet to be named. The proportion
of species recognisable from the literature, and for which we have even rudimentary biological and distributional data, is
certainly smaller. The increases in the number of families of insects known from Australia emphasise that this continent
remains an entomological frontier. Some of the new families have come from the subdivision of existing families and
others result from the discovery in Australia of existing families previously not known to occur here. There have also
been a few notable discoveries of entirely new, exclusively Australian families.
In the teaching of Zoology the trend for some years has been away from a taxonomic approach as a major founda-
tion on which to build other studies. Although this may mean that books like our volumes will progressively occupy a
less and less central position in the formal training of students, they will continue to be invaluable sources of informa-
tion and to provide an entry into the relevant but widely scattered literature. Indeed it is not too much to hope that a
work of this sort will stimulate interest in a more broadly based taxonomy than has been general in Australia in the past.
The keys-many of which are new-the abundance of illustrations, and the wealth of previously unpublished biological
information certainly provide the student with a broad, firm foundation to the subject.
D. F. Waterhouse,
P. B. Came,
Canberra I. D. Naumann,
January 1990 Division of Entomology CSIRO
Acknowledgments
The indebtedness of individual authors to those who helped them in their work is acknowledged separately in the appro-
priate chapters. Here we wish to record our appreciation and thanks to those who contributed in more general ways.
The illustrations are as important as the text in a work of this kind. Most of those in these volumes are original and
were made, under the guidance of the authors or editors, by the artists whose names appear in square brackets in the leg-
ends. We owe a special debt to Frank Nanninga, who painted the coloured plates and drew a great many of the line illus-
trations. Anne Hastings, Se Kim, Terry Nolan and Sandy Smith contributed most of the new line illustrations included in
this edition. Ms Hastings, Mr Kim, Mrs Smith, Chris Hunt and Andrew Carter were responsible for the layout of illus-
trations and for preparation of these for the Press. John Green and Alan Edward provided the photographic prints and
Colin Beaton and Helen Geier the plates of electron micrographs. Where illustrations have been redrawn from published
work, this is acknowledged in the appropriate legends. We thank the Elsevier Publishing Company, Amsterdam, for per-
mission to make direct use of Fig. 3.15 (from Berg 1975), Kluwer Academic Publishers, Dordrecht, for Fig. 35.40 (from
Lawrence 1985a), and the Editors of the Memoirs of the Queensland Museum for Fig. 24.7 (from Rentz 1980). Several
other illustrations are reproduced with the permission of the Editors of The Australian Journal of Zoology and
Invertebrate Taxonomy.
Preparation of such a large manuscript in good order is an arduous business, and we are indebted to Irma Pumpurs, on
whom the greatest burden fell, for the care and accuracy with which she completed her work. Eric Zurcher processed the
many text files into final format for the Press. Judith Olditch, Judith Hull, Heather Baker, Patsy Gillison, Eva Bugledich
and Jonathan McCabe also assisted in the keying and correcting of text. Anne Frodsham and Patricia Wellisch (Press
liaison) and Ray McInnes provided invaluable support to the Editorial Committee.
The bibliography was checked by Josephine Cardale. Almost all references were confirmed against original publica-
tions and we are grateful for the help in this sometimes difficult task which we received from the staff of the CSIRO
Library, Black Mountain, Canberra. Ms Cardale also compiled the index.
Many referees and some of the staff and students of the Department of Entomology, University of Queensland, com-
mented on chapters and tested some keys. We thank them for their unselfish, anonymous contributions.
The Utah Foundation, the Ian Potter Foundation, Wellcome Australia and the University of Melbourne generously
provided funds to enable the production of many new illustrations. We thank them for their continuing interest in the
production of these volumes.
Finally the Editorial Committee would like to record with particular appreciation the cordial and understanding rela-
tionship that has existed between them and the Directors and the staff of the Press over what has proved to be quite a
long period. Sally Paxton and Kevin Jeans of CSIRO Editorial Services and Susan Keogh, among others of the Press,
were responsible for typesetting and page layout.
This work may be cited as: CSIRO (ed.). The Insects of Australia. A textbook for students and research workers. 560
+ 600 pp., 2 volumes (Carlton: Melbourne University Press).
xiii
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Contents
Foreword xi
Acknowledgments xiii
Introduction 1
VOLUME I
Skeletal anatomy and key to orders, by J. F. LAWRENCE, E. S. NIELSEN and I. M. MACKERRAS 3
2 General anatomy and function, by R. F. CHAPMAN 33
3 General biology, by K. R. NORRIS with contributions by R. A. FARROW and V. A. DRAKE 68
4 Principles and practice of systematics, by P. S. CRANSTON, P. J. GULLAN and R. W. TAYLOR 109
5 Phylogeny of extant hexapods, by N. P. KRISTENSEN 125
6 Fossil history and the evolution of hexapod structures, by J. KUKALOVA -PECK 141
7 Biogeography, by P. S. CRANSTON and I. D. NAUMANN 180
8 Biographical history, by E. N. MARKS 198
9 Insects and humans in Australia, by D. F. WATERHOUSE 221
10 Australian insects in scientific research, by M. J. WHITTEN 236
11 Collembola, by PENELOPE J. GREENSLADE 252
12 Protura, by G. IMADATE 265
l3 Diplura, by B. CONDE and J. PAGES 269
14 Archaeognatha (Microcoryphia), by J. A. L. WATSON and G. B. SMITH 272
15 Thysanura (Zygentoma), by G. B. SMITH and J. A. L. WATSON 275
16 Ephemeroptera, by W. L. PETERS and I. C. CAMPBELL 279
17 Odonata, by J. A. L. WATSON and A. F. O'FARRELL 294
18 Plecoptera, by G. THEISCHINGER 311
19 Blattodea, by L. M. ROTH 320
20 Isoptera, by 1. A. L. WATSON and F. J. GAY 330
21 Mantodea, by J. BALDERSON 348
22 Grylloblattodea, by D. C. F. RENTZ 357
23 Dermaptera, by D. C. F. RENTZ and D. K. McE. KEVAN 360
24 Orthoptera, by D. C. F. RENTZ 369
xvi Contents
2 Blattoid-Orthopteroid orders
3 Hemiptera
4 Coleoptera
6 Batesian mimicry
7 Lepidoptera
8 Lepidoptera
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Introduction
Insects are among the most abundant and successful of impose some hardship on the amateur entomologists who
terrestrial animals. They include about three-fourths of all may wish to use the book, but it is hoped that they will
the described species of animals, and they have become find sufficient explanation in the text and illustrations to
adapted to a great range of environments, from high lati- meet their needs. There is no glossary. Terms with which
tudes to the equator, from rainforest to desert, from the reader might not be familiar are defined in the text
mountains to the shore, and to varied ways of living- and entered in the index.
phytophagous, carnivorous, saprophagous, parasitic. Many 2. Chapters 1 to 10 have been reduced to the minimum
have become aquatic during part or the whole of their life that is considered essential for the general student of ento-
history. The breadth of their adaptive radiations has mology as a background for the chapters that follow.
brought some of them, though only a very small propor- Chapter 1 has presented particular difficulty in this
tion of the total, into close contact with human popula- respect, because the anatomical terminology used in it has
tions. On the one hand, they damage or destroy crops and been based, as far as possible, on usage among morpholo-
domestic animals, transmit diseases of plants and animals, gists, whereas the writers on many of the orders carry the
including people, and damage or destroy human habita- historical burden of a terminology that has grown up with
tions. On the other hand, humans have added bees and little or no reference to what has been done outside those
silkworms to their flocks, and have learned to use other orders. There seems to be no way to avoid this unfortu-
species to control noxious insects and weeds. Moreover, nate situation. There is the further minor difficulty that
insects have provided some of the most valuable tools of the earlier chapters must anticipate Chapters 5 and 6, but
biological research in many fields from the broadest a general picture of the classification used may be
aspects of evolution to the de'tailed mechanisms of inheri- obtained by referring to Figs 5.1 and 5.5. Chapter 6 dif-
tance. fers from other chapters in a few interpretations of mor-
It is natural, then, that entomology should have shared phology.
in the steadily increasing volume of research that has 3. The presentation of the remaining chapters is essen-
made it so difficult to be both brief and comprehensive tially systematic, an arrangement that serves to emphasise
when reviewing any scientific discipline. Selection is the evolutionary perspective into which any study of
unavoidable, and the purpose of these notes is to set out insects should be fitted. The systematic statements have
the scope of this book and the ways in which problems of been used as pegs on which to hang short accounts of
presentation and compression have been met. where the insects live and what they do. Wider problems
1. It has been necessary to assume that the reader are touched on when describing insects that illustrate
would have a background of general knowledge equiva- them particularly well.
lent to what might reasonably be expected at the end of 4. The book is designed to cover Australian insects
the first year of a university course in Science. This may only. The insects of New Zealand, New Guinea and the
2 The Insects of Australia
Pacific are referred to here only when they are relevant to DIPL Diplura;
particular problems that are being discussed. However, an ARCH Archaeognatha;
attempt has been made to put the Australian fauna into THNU Thysanura;
perspective with that of the rest of the world by including EPHM Ephemeroptera;
at least a mention of important groups that do not occur ODON Odonata;
here. There are abbreviated chapters dealing with the PLEC Plecoptera;
small orders not represented in Australia. BLAT Blattodea;
5. In the systematic chapters the censuses include unde- ISOP Isoptera;
scribed species and unrecorded synonymy known to the MANT Mantodea;
authors, but the completeness of this information differs GRYL Grylloblattodea;
from order to order, so the figures should be taken as DERM Dermaptera;
approximations. Their reliability as indices of the total ORTH Orthoptera;
number of species in the country varies greatly. Thus, it is PHAS Phasmatodea;
reasonable to suppose that most of the species of EMBI Embioptera;
Australian butterflies and mosquitoes are already known; ZORA Zoraptera;
the same cannot be written of the less-studied families of PSOC Psocoptera;
Lepidoptera and Diptera. PHTH Phthiraptera;
6. The keys to families and higher taxa have been HEMI Hemiptera;
based primarily, though not exclusively, on Australian THPT Thysanoptera;
material. Some of them may have a wider application, but MEGA Megaloptera;
they should be used with reserve for placing specimens RAPH Raphidioptera;
from other regions. There has not been room for more NEUR Neuroptera;
than a few keys below the family level. COLE Coleoptera;
7. Common names have been used sparingly, and only STRP Strepsiptera;
for widely known species. A full list, with standard abbre- MECO Mecoptera;
viations of the names of authors of species, is given by SIPH Siphonaptera;
Carne (1987). The techniques of collecting, preserving, DIPT Diptera;
and studying insects have been described by Upton TRIC Trichoptera;
(1991). LEPI Lepidoptera;
8. The following abbreviations have been used, where HYMN Hymenoptera.
appropriate, for political divisions of Australia: 9. References have presented a particular problem. It is
Qld Queensland; impossible to give extensive bibliographies in the space
N.S.W. New South Wales; available, and reliance has been placed as much as possi-
Vic. Victoria; ble on references to reviews and monographic works
S.A. South Australia; which provide a recent, readily available entry into the
W.A. Western Australia; major literature. It must be stressed that this in no way
N.T. Northern Territory; implies any lack of appreciation of earlier classical
A.c.T. Australian Capital Territory. papers. Direct study of these works is indispensable to
Tas. Tasmania any serious student, and some of them have been referred
T1 denotes the first abdominal tergum (or tergite), T2 to in general terms when the full references are included
the second, etc.; SI, S2, etc. denote the corresponding in a publication that is cited. References in round brackets
abdominal sterna (or sternites). relate to the statements in which they occur, those in
Where appropriate, ordinal names are abbreviated as square brackets usually to the group as a whole. Papers on
follows: Australian insects up to 1930 have been listed in the
COLL Collembola; bibliography by Musgrave (1932).
PROT Protura;