A Guide to the Cockroaches of Australia

A GUIDE TO THE

COCKROACHES

OF AUSTRALIA

This book is dedicated to

Kathy Hill

A GUIDE TO THE

COCKROACHES

OF AUSTRALIA

DAVID RENTZ

© David Rentz 2014

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

Rentz, David C., author.

A guide to the cockroaches of Australia / David Rentz.

9780643103207 (paperback)

9781486300372 (ePDF)

9781486300389 (epub)

Includes bibliographical references and index.

Cockroaches – Australia.

Cockroaches – Australia – Identification.

595.7220994

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CSIRO Publishing

150 Oxford Street

(PO Box 1139)

Collingwood VIC 3066

Australia

All photos are by the author unless noted otherwise.

Front cover (top left to right): Calolampra sp., Mediastinia australica, Leptozosteria prima (M Harvey); (main image) Ellipsidion simulans.

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Contents

Preface

Acknowledgements

1 Introduction

2 Cockroach ‘personalities’

3 Biology

4 Economic importance

5 Australian cockroaches in captivity (by Deanna Henderson)

6 Collection and preservation

7 Habitats and ecology

8 Morphology

9 Cockroach identification

Family Blaberidae

Family Nocticolidae (Delicate Cockroaches)

Family Corydiidae (Elusive Cockroaches)

Family Blattidae

Family Tryonicidae

Family Ectobiidae (formerly Blattellidae)

List of the cockroaches of Australia

Glossary

Australian entomological supplies

Websites and special interest groups

Orthopteran food mix

Bibliography

Index

The latest in photo imagery. This Giant Desert Cockroach, Megazosteria patula, has undergone special treatment. The image has been processed with Zerene Stacker software from source images taken with a Hasselblad H4D-50MS camera (120 mm Macro lens) in 4-shot, 50 megapixel mode on a Visionary Digital Bk Plus Laboratory System. The difference is in the detail. Digital photography by Geoff Thompson, Queensland Museum. Photos used with permission. © Queensland Museum.

Preface

Why publish a guide to Australian cockroaches? After all, the mere mention of the word ‘cockroach’ produces a negative reaction in most people. Most non-entomologists recoiled in horror when I told them I was working on a book on Australian cockroaches. Even some entomologists were not impressed. So how does one approach the subject and not lose the interest of prospective readers on the first page?

My friend Piotr Naskrecki in his recent book Relics calls them ‘blattodeans’ – which they are. He never uses the word ‘cockroach’ in his book. This is a novel concept but I have not adopted this idea for reasons that should become apparent.

Despite the revulsion that pervades the general community about cockroaches, they are often subjects of games and toys. Shops are replete with a variety of small toys, some more complimentary to the insects than others.

It is surprising that cockroaches can be topics of sculpture. The Australian native Giant Burrowing Cockroach (p. 87) seems to be the most popular Australian subject, perhaps due to the lack of familiarity of the artists with the range of species. Hopefully this book will remedy that void.

As you will discover in the pages of this book, the diversity and beauty of Australia cockroaches – or blattodeans – speaks for itself. I hope, therefore, that Guide to the Cockroaches of Australia goes some way towards elevating the reputation of these much-maligned insects, and promotes further study of them.

Macropanesthia frieze on a building at the University of Queensland. Photo: G Monteith

Cockroach games and toys are all designed to startle. At least the ‘Busy Bugs’ is more authentic in its appearance. It seems to be a representation of the Australasian Cockroach, Periplaneta australasiae.

This sculpture in South Australia is probably one of the many polyzosteriine cockroaches that roam the country (see p. 121). Photo: G Monteith

Acknowledgements

This book could not have been produced were it not for the generous cooperation of many individuals. I am in debt to those who have provided photographs, information on types and who have made suggestions about the coverage of the book. Dr W Humphreys, Dr M Harvey and Dr T Houston, Western Australian Museum, Perth, generously allowed use of many of their unique photographs – those of Tivia australica revealed the identity of this unusual species. Dr H Bohn provided notes on the type of Tivia australica. Dr D J Mann provided a photograph of A D Shelford from the collection of Oxford University. Dr J Hollier, Geneva Museum, provided a photograph of H de Saussure. Dr Lambert Smith permitted use of the photograph of the fossil cockroach wing.

Dr Alice Wells and Ms C Geromboux, of the Australian Biological Resources Study, Canberra, were most patient with my many requests. A fair number of changes were made to the Blattodea component of the faunal lists, some back and forth again, and the changes were made with prompt efficiency in a most pleasant manner.

Mr V Lee, California Academy of Sciences, San Francisco, and Dr Y N Su and Dr B Mantle, CSIRO Ecosystem Sciences, Canberra, provided information and other services concerning specimens and literature in the Australian National Insect Collection. Dr C Lambkin, Queensland Museum, verified identifications from photos with specimens in the collection. Dr P Naskrecki took photos of identified specimens in the Museum of Comparative Zoology, Harvard University, that were associated with the research of the late L Roth. Dr G Cowper and Ms C Flemming of the Academy of Natural Sciences of Drexel University helped find photos in the Academy Archives. Associate Professor J Neal, Purdue University, and Mr G White provided photos of the species of economic concern. Dr P Hudson, South Australian Museum, also helped to locate archival photographs. Dr U Kaly produced the graphic representations of the diversity of cockroaches in a Kuranda rainforest.

I have been a continual thorn in the side of Dr G Beccaloni, Natural History Museum, London, for information regarding listings in the Blattodea Species File (BSF). The replacement of the use of the name of the Blattellidae with Ectobiidae is a major change in the usage of the largest group of cockroaches in Australia.

Dr C Nalepa, North Carolina Department of Agriculture, was most helpful in answering questions about statements in her recent book.

Mr G Thompson, with the permission of the Queensland Museum, discovered photographs of the late M J Mackerras and provided the frontispiece (p. vi).

Drs James Walker and Harley Rose answered numerous queries on burrowing cockroaches of the families Blaberidae and Blattidae. They read sections of the manuscript and offered suggestions.

U Aspöck, Tom Bellas, O Béthoux, P Brock, C Dewhurst, E Edwards, G Cowper, R B Halliday, Jack and Sue Hasenpusch, Gary Wilson, D & A Henderson, S Ingrisch, B Mesibov, P Naskrecki, G Owen, M Upton, S Schmidt, F Spier, M Robinson, and P Shanahan made many helpful suggestions. Special thanks go to Mr Y N Su who provided the author with many PDFs of important publications and notes on specimens in the Australian National Insect Collection (CSIRO), the most comprehensive collection of Australian Blattodea. Buck Richardson accompanied the author on many field trips during the past three years that resulted in the discovery of many species of cockroaches that had never been photographed before.

A mating pair of Megazosteria patula at Bungle Bungle, WA. Photo: K Hill

The following persons provided photographic material that has greatly increased the scope of this book: P Agar, M Anthony, W Archer, T Bayliss, M Braby, P Brock, Neil Bryde, G Byrne, M Cermak, P Chew, D Clyne, G Cocks, S P Dhakal, K Edgerton, K Ellingsen, A Elliott, R Farrow, M Fletcher, P Gullan, P Glover, M Hanlon, M Harvey, M Hauser, J & P Hasenpusch, A Henderson, N Hewitt, D Hobern, P Honan, J & F Hort, P Hutchinson, S Humphries, W Humphreys, J de Jong, J Keeble, D & F Knowles, A Kristin, M Lenz, D Lightfoot, J McRae, M Macrae, T MacRae, D J Mann, D & K Marshall, J Matsui, S & D Mawson, C Miller, N Monaghan, J Neal, G & N Monteith, E Nielsen, C Nimienski, M Powell, B Richardson, R Richardson, M Robinson, C Rowan, B Seccombe, D Slaney, F Seow-Choen, M Smith, P Street, Y N Su, A Sundholm, G Tate, F Venter, J Waldock, D Wilson, G Wilson, G White, C Wu, P Zborowski. Those whose photos have been selected are identified where they are used. The photograph of Dr JWH Rehn was provided by the Academy of Natural Sciences of Drexel University, ANSP Coll. 457. The photograph of Mrs J Mackerras in the field was provided by the Queensland Institute of Medical Research through the goodwill of Ms Heather Matthews and G Thompson. Photographs not named with a photographer are those by the author.

Nick Alexander and Tracey Millen are thanked for their editorial efforts.

Ms B Foster Rentz and Ms T Turtle are thanked for their companionship during the three years gestation of this book.

1 Introduction

The Australian cockroach fauna is remarkable in its extent, colour and biology. Cockroaches occur in every habitat on the continent except in seas and lakes. The diversity of species is only just being appreciated. At my home in the Kuranda rainforest, I have recorded nearly 90 species ranging in size from 4 mm to 35 mm in virtually all habitats from the treetops to on, and in, the ground. They are major components of leaf-litter fauna. Their numbers suggest that they must be important in the return of the litter to soil, as well as food for other organisms. They occur along watercourses and there is at least one Australian aquatic cockroach whose water-loving habits has been observed but is not yet recorded in the literature.

A fossilised cockroach wing from Molteno Formation, South Africa, Triassic Age, 200 mya. Photo: L Smith

Cockroaches are blattodeans that are orthopteroids – most closely related to termites and mantids. They are well known in the fossil record and seem to have been especially abundant in the late Palaeozoic, some 300 million years ago. There is a very well-preserved fossilised blattodean wing from the Molteno Formation, which is situated in the heart of the semi-desert Karoo Basin of South Africa. This site has the richest known flora and insect fauna at about the time of origin of the mammals, dinosaurs and possibly the flowering plants in the Late Triassic. It is thought that blattodeans split from the common ancestor they shared with the mantids during the Lower Cretaceous about 140 million years ago. The early blattodean-like species have been dubbed ‘roachoids’. They had external ovipositors but the flattened body characteristic of cockroaches today.

This specimen of Polyzosteria fulgens from the Perth region is hardly a disgusting creature.It is attractive, sedate and moves about in its habitat by day. Photo: J & F Hort

A large native Australian cockroach, Anamesia serrata, a common inhabitant of the arid northern and central portion of Australia, p. 127. Photo: D Marshall

This colourful cockroach may represent an undescribed species. It is related to Balta hebardi (p. 264) but has very different pronotal markings. Usually these marks are diagnostic for species. This cockroach was photographed in a Perth suburb suggesting that you do not have to go far to find beautiful and interesting blattodeans. Photo: J & F Hort

Recent research, using DNA, biology and other characteristics indicates that termites are actually in a lineage of cockroaches and should not be allocated a separate order in the world of insects. They are merely modified Blattodea that are most closely related to the wood-feeding cockroaches of the genus Cryptocercus! It has been suggested that the initial oötheca-like packet of eggs in cockroaches were cemented together as is found in the Australian termite, Mastotermes darwiniensis. This termite extrudes eggs in packets of two and covers them with a tanned outer layer that is similar to the cockroach oötheca. It is believed that the mantids diverged first and the cockroaches and termites are nested within the Blattodea. The North American cockroach genus, Cryptocercus, represents the sister group to the termites.

The winged form of the Australian termite Mastotermes darwiniensis has had an important role in the discovery of the relationship between termites and cockroaches. Photo: M Lenz

Although termites are technically blattodeans, this book is only concerned with the families of ‘true’ cockroaches: Nocticolidae, Corydiidae, Blattidae, Ectobiidae, Tryonicidae and Blaberidae that occur in Australia.

Cockroach diversity in Australia

The cockroach component of the order Blattodea is of moderate size with approximately 4500 species in 460 genera worldwide. It is expected that the number of blattodean species will increase dramatically because some of the diverse families, such as the Ectobiidae (previously known in most studies of the Australian fauna as the Blattellidae) are, as yet, very poorly studied. More than 530 species are now recognised from Australia and in excess of 90% of these are considered endemic. At least twice that number of Australian species are yet to be described.

Cockroach taxonomy and biology can be a rewarding pursuit in Australia because many groups have been little studied.

Some of the world’s smallest and largest cockroaches are found in Australia. Adults of Nocticola can measure as little as 3 mm in length, while those of Macropanesthia rhinoceros are up to 80 mm in length and rank among the heaviest of insects, sometimes weighing 33 grams.

Intensive collecting and trapping at my home at Kuranda in the tropical rainforest of north Queensland has revealed a rich diversity of species not seen elsewhere. An array of families is present and 88 species have been identified to date. Most have been determined on the basis of several characters with the most distinguishing being the minute details of the male genitalia.

Of the 88 species, 70 (79%) are in the Ectobiidae, the most diverse family in Australia. The Blaberidae follow well behind with only 8 species (9%). Others are illustrated in the graphs. This further suggests the importance of cockroaches in the environment since almost all of the ectobiids occur in leaf litter. They must be very important in the decomposition process. Virtually every handful of leaf litter contains cockroaches, often several species in various stages of development. It would be educational to perform detailed census of cockroaches in other habitats around Australia. A similar result may be obtained, although the total number of species may not be so great. Collections in coastal mixed closed forests in coastal New South Wales (including Bawley Point) during the 1990s revealed about two dozen species. Cockroaches may be the most abundant of orthopteroids, both in numbers and diversity in all habitats in Australia.

Origins and affinities

The Australian cockroach fauna contains endemic elements as well as those derived from other regions of the world. There are many endemic genera, mostly in the Blattidae, in the subfamily Polyzosteriinae (p. 121) and many Blaberidae (p. 72), especially the Epilamprinae, Panesthiinae. Genera in several subfamilies of the Ectobiidae (p. 196) are endemic to Australia.

The red columns illustrate the families found in the kuranda rainforest. The blue columns represent the subfamily against the number of species on the vertical axis.

Habitat preferences of species in kuranda rainforest. Some species overlap, at least during part of their lifetime. Most of the standing trees in the rainforest have bark that does not flake or dehisce. As a result it is generally a habitat not used by cockroaches as it is in eucalypt woodlands. This suppresses the number of species that may be found under bark in the rainforest. The great majority of species live in leaf litter and are probably very important in its decomposition.

There are many genera in all of the families that are shared with other regions of the world, especially the tropical regions to the north. Genera such as Shelfordina and Balta in the Ectobiidae occur in New Guinea, Indonesia, Malaysia and other parts of south Asia. Methana, here placed in the Blattidae (p. 112), occurs in the Oriental Region as well as Australia and Norfolk Island. Blattids like Polyzosteria, occur in New Zealand as well as the Oriental Region and New Caledonia. Celatoblatta (and its synonym Austrostylopyga) occurs in both Australia and New Zealand. The genus Tryonicus occurs at high elevations in northern Australia and New Caledonia. It is a relict that requires cool temperatures and could be threatened by the warming of climate change as are other organisms living on mountaintops in the northern tropics.

The introduced cosmopolitan species are generally not found in nature. Some exceptions are the Australasian Cockroach, Periplaneta australasiae, and the Dusky Cockroach, P. fuliginosa, which can be found short distances away from human habitation. The Surinam Cockroach, Pycnoscelus surinamensis (p. 104), seldom enters homes but can be found outside in tropical and subtropical regions in compost or in roof gutters. Oddly enough it is not found far from human habitation. An Australian species, The Coulon, Paratemnopteryx (formerly Shawella) couloniana, has been introduced from Australia to New Zealand where it can be a pest. It can also enter dwellings in Australia.

Cave-dwelling cockroaches

Australia has an abundance of cave systems. Some have been studied more than others, but in general the biota of these systems is considerable but largely undescribed. The cockroaches are a notable exception, in that several species have been described. Cockroaches can be found in all of the cave systems.

Australian caves can be found in the tropics, subtropics, subtropical dry zones and transitional zones with winter rain. Both karst limestone caves and tunnels and lava (lava tubes) occur in Australia. Examples include the Cape Range of north-western Western Australia which is an extensive karst system and the Undara and Chillagoe Caves of tropical north Queensland which are lava tubes.

Cave systems in Australia harbour some of the world’s most interesting dark-adapted cockroaches. The arid Cape Range supports one of the world’s richest cave biota. The Cape Range attains an altitude of only 330 m but there are more than 600 limestone caves to 100 m depth. The fauna there is entirely endemic and is derived from rainforest species that used to occupy the area. One of the most spectacular is Nocticola flabella (p. 107), the most cave-adapted cockroach in the world. It is extraordinarily long-legged and lacks pigmentation and eyes. Its exoskeleton is so thin that only the mandibles and genitalia are normally sclerotised.

Blattelline ectobiids (p. 203) contribute the majority of species found in Australian caves. They can be large and well sclerotised, like Trogloblattella nullarborensis or small and delicate like Paratemnopteryx stonei. This species exhibits considerable morphological variation and occurs in caves some 150 km apart. Approximately half of the known species of Neotemnopteryx are cave dwellers, while the terrestrial species are presently found along the east coast of the continent.

The spectacular Nocticola flabella is the most cave-adapted species in the world. Male (left) and female (right). Photos: J McRae

The family Nocticolidae (p. 105) has an Old World distribution but Nocticola species can be found in the Philippines, Vietnam, Ethiopia, South Africa and Madagascar as well as Australia. Christmas Island harbours the only known species of Metanocticola, the only species from that locality covered in this book.

Cave cockroaches feed on a variety of material depending on how far they are found from the entrance of their caves. The Nullarbor Cave Cockroach, Trogloblattella nullarborensis, is known to feed on tree roots. Other species found in the presence of bats feed on dead bats or guano from bats or swiftlet birds.

What is a cockroach?

Cockroaches have a dorso-ventrally flattened body with the head directed downwards, and bearing chewing mouthparts. Cockroaches undergo gradual metamorphosis and usually have short lifecycles. Males and females of many species are dissimilar. Many have highly specific ecological preferences and elaborate biological lifestyles. They involve activities as diverse as parental care, Müllerian mimicry, host preferences and chemical defence.

Interest in blattodean classification has been very popular over the years partly because there is a rich fossil history of the group, a considerable diversity of extant species and the economic importance of some of them. Until recently, three classification schemes were in practice at the same time. This was very confusing even to an experienced entomologist. Which classification was the one to use? There was that of Princis (1960) who used classical morphological characters such as wing venation and wing-folding, the patterns of the spines on the legs and the shape of both male and female genital appendages. He integrated Handlirsch’s (1926–1930) system and recognised four suborders, 28 families and 21 subfamilies. McKittrick (1964) used ovipositional behaviour in combination with male and female genitalic structure and musculature. She used the morphology of the proventriculus – the toothed structure at the end of the foregut that sorts and aligns food as it enters the midgut for digestion. Although this structure is very informative, it is not used in this guidebook. The proventriculus, however, is easily studied after removal from the gut. McKittrick’s classification system resulted in two superfamilies, five families and 20 subfamilies. (The Tryonicinae was added later.) Her classification was strongly supported for years. Subsequently, Roth, Grandcolas and others made important changes. McKittrick’s classification system was followed in Insects of Australia and modified slightly in the second edition of that work. For years, this has served as the reference for blattodean classification for Australia.

Isopods are often mistaken for cockroaches, especially at night when they wander in search of food. Examination of the legs quickly reveals that they are not insects.

Some beetles can be confused with cock– roaches. This carabid beetle of the subfamily Pseudomorphinae often occurs under bark with similar-appearing cockroaches. The resemblance is startling.

However, considerable taxonomic changes have been made recently that must be followed according to the system of zoological nomenclature. Beccaloni (2007) pointed out that the family name ‘Blattellidae’ had been incorrectly used since 1987 when it was officially ‘abolished’ in favour of the older name Ectobiidae. The only problem that this will cause Australian entomologists is to remember the new name. The subfamily ‘Blattellinae’, which contains the majority of Australian cockroach species, remains unchanged. Another change involves the Polyphagidae. This family has been renamed the Corydiidae. And finally the subfamily Tryonicinae has been elevated to family rank. The Tryonicidae are an Australian, New Zealand and New Caledonia group. Several genera have been transferred to this family. They include Methana, Scabina and Temnelytra. However, because of the diversity of opinion on this transfer, the ‘Methanini’ are retained in the Blattidae in this book. (Names of tribes end in ‘ini’.) The Methanini and the Parcoblattini have been used extensively in the past but are not now recognised in the various catalogues and will not be dealt with as such beyond this point in this book.

A map illustrating the communities that define the distribution of species.

The classification of the Blattodea.

The big change involves the inclusion of the termites (formerly the Isoptera) in the order Blattodea. Termites have been discovered to be merely highly modified cockroaches! The present classification of the Blattodea is represented in the diagram on page 8. Time will tell if this classification is accepted by the entomological community but there is good evidence that it is the best scheme to use now.

A problem with the identification of blattodean species is often associated with their rather conservative external features. But the more one looks, the more distinctive characters one can find on a cockroach. Colour features are frequently overlooked because they may be changed in old museum material or discoloured from poor initial preservation. These ‘colour characters’ include leg banding and the patterns of the coxae and ventral surface of the abdomen, the colour of the antenna and whether it is hairy or not. The frons (face) and top of the head often bear colours and patterns that are species distinctive. But many investigators have not used these characters in their descriptions. As a result, some very distinctive features that would aid in identification have been missed. One possible reason for this might be early descriptions have been made from pinned specimens that had been originally preserved in alcohol. Almost all colouring and some patterning disappear when this happens. Another character of importance is the armature hairs and spines on the anterior margin of the ventral surface of the fore femur – often called the femoral comb. This is used extensively in the higher classification of some groups and at the specific level in others. (Cockroaches should be mounted so that this character can be seen.)

Because cockroaches are orthopteroid insects, they have traditionally been included in faunistic studies along with mantids and stick insects. They are frequently relegated to ‘appendices’ in these treatments. Many of the authors of such works have been specialists on one or more orthopteroid groups but generally not Blattodea. As a result, they felt compelled to provide names for the supposed new species included in their studies. Thus, generic placements were often incorrect and species were often described but without noting the most characteristic features. Thus, convergence and parallel evolution can easily go unnoticed. Many cockroach species, therefore, have been assigned to incorrect higher taxa or even the wrong family. Males have been placed in one genus and the females in another.

Most cockroaches have highly complex, but species-specific, concealed genitalia. Until about 30 years ago, concealed genitalia were not routinely examined, and recent assessment of