A Guide to the Katydids of Australia [Illustrated] 9780643095540, 0643095543

Ratyduls are among the most commonly seen Australian insects. They range in size from about 5 mm to well over 90 mm and

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Table of contents :
Title
Copyright
Contents
Preface
Acknowledgements
Introduction
Biology
Sound and hearing
Collecting and studying katydids
Habitats
Conservation
Guide to species
Lnistroscelidinae
Hexacentrinae
Tympanophorinae
Austrosaginae
Conocephalinae
Microtettigoniinae
Meconematinae
Tettigoniinae
Pseudophyllinae
Phyllophorinae
Mecopodinae
Phaneropterinae
Zaprochilinae
Phasmodinae
Key to subfamilies of Australian katydids
List of the Tettigoniidae (katydids) of Australia
Glossary
References
Appendix 1: Keeping katydids alive and preservation of specimens
Appendix 2: Special interest groups and entomological supplies
Index
Recommend Papers

A Guide to the Katydids of Australia [Illustrated]
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A GUIDE TO THE KATYDIDS OF AUSTRALIA

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A GUIDE TO THE

KATYDIDS OF AUSTRALIA

DAVID RENTZ

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© David Rentz 2010 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. A guide to the katydids of Australia / David Rentz. 9780643095540 (pbk.) Includes index. Bibliography. Katydids – Australia. Tettigoniidae – Australia. 595.7260994 Published by CSIRO PUBLISHING 150 Oxford Street (PO Box 1139) Collingwood VIC 3066 Australia Telephone: Local call: Fax: Email: Web site:

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

Front cover image: A rare pink morph of the Serrated Bush Katydid, Paracaedicia serrata. Photo by Jack Hasenpusch. Set in Minion 9.5/12 Cover and text design by James Kelly Typeset by Oryx Publishing Pty Ltd Printed in China by 1010 Printing International Ltd The paper this book is printed on is certified by the Forest Stewardship Council (FSC) © 1996 FSC A.C. The FSC promotes environmentally responsible, socially beneficial and economically viable management of the world’s forests. 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.

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Contents Preface Acknowledgements Introduction Biology Sound and hearing Collecting and studying katydids Habitats Conservation Guide to species Listroscelidinae Hexacentrinae Tympanophorinae Austrosaginae Conocephalinae Microtettigoniinae Meconematinae Tettigoniinae Pseudophyllinae Phyllophorinae Mecopodinae Phaneropterinae Zaprochilinae Phasmodinae Key to subfamilies of Australian katydids

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List of the Tettigoniidae (katydids) of Australia Glossary References Appendix 1: Keeping katydids alive and preservation of specimens Appendix 2: Special interest groups and entomological supplies Index

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This illustration of the ‘Great Green Gumleaf Grasshopper’ comes from McCoy’s Natural History of Victoria (1886). However, it is not a grasshopper at all but a katydid. Subsequent research has revealed the scientific name McCoy used was a synonym of Terpandrus horridus, a species found in the Sydney region. The species illustrated here is most likely Terpandrus endota. Source: Museum of Victoria

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Preface Welcome to the world of Australian katydids. These insects occur almost everywhere across the country from the highest mountains to the seashore, and on continental and oceanic islands. The loud calls of some katydids are often mistaken for the sounds of cicadas. With a little practice you will be able to distinguish one from the other without even seeing the singers! Katydids range in size from tiny species, 5 mm or so in length, to monsters that approach 130 mm. Many are secretive and have to be searched out in order to see them. At times, however, normally common species can be extremely abundant and cause public concern. We see this phenomenon in many Australian insects and in many grasshoppers. Some species can be agricultural pests and a number can cause problems to horticulturists because of their propensity to feed on developing flowers. This book is primarily an identification manual to be used in the manner that you might use a bird guide. As a result, information on the many other aspects of katydids is scant. The book by Daryl Gwynne, Katydids and Bush-crickets: Reproductive Behavior and Evolution of the Tettigoniidae, published in 2001, provides many details and references to the vast literature available on this subject. Naskrecki and Otte (1999) estimated that there are more than 6000 described species of katydids and that their diversity within the Orthoptera was second only to the grasshoppers which comprise some 12 000 species. There has been a trend in recent years to escalate subfamilies and tribes to family status. This has been accepted to some degree but a more conservative approach has been followed in the Orthoptera Species File (OSF). This catalogue is kept current and its scheme is what is followed in this book. Any serious student of Orthoptera taxonomy should become familiar with the OSF, a most useful tool. Currently 19 subfamilies are listed in the OSF. These form major, easily recognised groups of a few to many species. Of the 19 subfamilies five, the Saginae, Acridoxiinae, Bradyporinae, Hetrodinae and Lipotactinae do not occur in Australia. Five subfamilies are endemic to Australia. They are the Microtettigoniinae, Phasmodinae, Zaprochilinae, Austrosaginae and

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Tympanophorinae. In addition, all of the genera of the Tettigoniinae are endemic to Australia, as well most of the genera in the other subfamilies. Shared katydid genera (in the Mecopodinae, Conocephalinae, Phaneropterinae, Pseudophyllinae and Listroscelidinae) are usually found also in New Guinea. As far as we know, there are no introduced katydids in Australia. This book includes species that occur within mainland Australia and Tasmania, Norfolk and Lord Howe islands, and the islands of the Torres Strait. The fauna of the latter localities is poorly studied and we can expect to find many species there that remain unknown. Christmas Island species show no relationship with those of Australia and are not included in this book. The taxonomic status of the Australian species is only partially documented. Ten or fifteen years ago Australian katydids featured prominently in the fields of taxonomy, ecology, physiology of hearing and stridulation, behaviour and sexual selection. One species, undescribed at the time, was featured on the cover of two separate issues of the international journal, Nature. However, only three volumes of my monographic series on the Australian Tettigoniidae have been completed – at least another three would be needed to thoroughly cover the fauna. So much needs to be done. I hope that this book might stimulate more interest in the group than there is at present. David Rentz

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Serrated Bush Katydid, Paracaedicia serrata

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Acknowledgements The author is indebted to many people for providing photographs, information on types, and suggestions for the style of the book. First I would like to thank CSIRO Publishing and Nick Alexander for allowing the author a maximum of breadth regarding style and content and for helpful comments during the preparation of the text. Mr Peter Hudson of the South Australian Museum provided photographs of the types of several of Tepper’s species which have never been illustrated before. These helped in assigning names to some common species in southern Australia. CSIRO Entomology has allowed access to the Australian National Insect Collection which contains the bulk of the species described from Australia. Mr Eric Hines has been helpful in providing access to scanning electron micrographs made by the author when he was employed by CSIRO. Dr Darryl Gwynne has made helpful comments. The late Dr Fer Willemse provided a copy of Brunner’s rare Additimenta, a publication necessary for anyone seeking to unravel taxonomic problems in the Phaneropterinae. Mr Martyn Robinson, of the Australian Museum, Sydney, has been a source of information and his contacts with biologists have provided relevant information for several species. Mr Paul Brock has be a source of information and encouragement. Mr Ian Menkins has provided meticulous observations on the biology of the Mountain Katydid, Acripeza reticulata. Mr Patrick Honan of the Melbourne Zoo has provided some helpful comments regarding the biology of several species under his care in the live insect exhibit. Jack and Sue Hasenpusch have presented the author with a continuous stream of interesting katydids for more than two decades and have helped with suggestions and encouragement.

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Gary Wilson has helped with comments of a botanical nature on habitats and ecology. Finally I would like to thank my friend and colleague Mr You Ning Su, CSIRO Entomology, for many favours and friendship over many years.

The Striped Nicsara, Nicsara bifasciata, is a common species in northern Australia. Photo: Alan Henderson

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Biology Katydids occur everywhere on all continents except Antarctica and at all altitudes from sea level to high in the mountains. Many species occur in the driest parts of Australia. On one the most uncomfortable days spent in the Australian interior where temperatures were over the 50°C mark, there was little insect activity during the day. But after dark, the lights of a roadhouse attracted about a dozen species, some in large numbers, which had been sitting quietly during the day in trees and shrubs. The smallest katydids and some of the largest katydids occur in Australia. The smaller species tend to be found in habitats that have irregular rainfall such as deserts or heathlands. All of the species in the subfamily Microtettigoniinae are tiny – some of the males are only 5 mm in total body length. They have all the components of their bigger brethren and produce sound with tiny tegmina. Other small species are represented in the subfamily Tettigoniinae and these also occur in habitats that might be considered as marginal with regard to climate. Perhaps these species have sacrificed size for rapid development thereby requiring less food and resulting in smaller size. Their small size seems also to be correlated to their agility. All these species are very fast-moving and the collector will soon discover they are not easy to capture. On the other hand, they tend to be more common than many other larger species. It would be interesting to discover how many instars the smaller species go through as opposed to the larger ones.

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This katydid is only about 5 mm in length.

Reproduction Reproduction in katydids revolves around communication. The most common scenario is the male katydid produces a calling song heard by receptive females who come to the source of the song. Researchers have found that female katydids can be choosey about the males they mate with. They can detect the heaviest and, therefore, the fittest males and mate with them. Mating occurs with the male transferring a spermatophore which contains the sperm package. With the spermatophore, the male provides a nuptial meal (a gooey white portion called the spermatophylax) which is eaten by the female. This nourishes the mother and the eggs and, therefore, the male has contributed to the development of his offspring. The spermatophylax is often the most visible portion of the spermatophore with the sperm capsule hidden well within this mass. As the female eats the spermatophylax, the sperm move into her receptacle, the spermatheca. This is an internal structure near the tip of her abdomen. As the eggs are laid, they pass by the opening of the spermatheca and are fertilised. The subject of katydid mating biology has been extensively reviewed in Gwynne (2001) and should be

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consulted if you are interested in the details of this aspect of behaviour and evolution.

A female with a freshly transferred spermatophore.

A female Austrosalomona species is feeding on the spermatophylax portion of the spermatophore.

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A mating pair of Coptaspis species. The female is on the left.

A female Requena eating the spermatophylax after having removed it from the spermatophore.

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A female Polichne species with a small spermatophore. The spermatophylax has either been eaten or is a very small portion of the spermatophore in this species.

It is often said that all a taxonomist needs to make an identification is an accurate locality label and the male genitalia. This may seem absurd but there is some truth in the statement. The male and female katydid abdomen has unique characters. Those of the male are more obvious and used extensively in taxonomy. Reasons for this uniqueness are simple. Insects use the ‘lock-and-key’ mechanism to achieve effective mating. That is, the male’s genitalia are designed to fit exclusively into those of the female. If the structures are not precise, there will not be a successful mating. As a result, there are some very elaborate genitalic structures in the Tettigoniidae. With Indiamba malkini, the hook-like structures protruding from the tenth abdominal tergite (segment) and those of the cerci fit precisely into pockets on the abdomen of the female. The cerci seem to be very distinctive structures in most katydid species. Those of Terpandrus paruna grasp the female in appropriate pockets at the tip of her abdomen. The prongs of Yutjuwalia nyalma perform a similar function. For accurate identifications you need to be able to compare the cerci of males in most genera. In the Phaneropterinae, the tip of the cercus is also distinctive but not easily seen without a high-powered microscope.

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Male genital structures

Indiamba malkini

Yutjuwalia nyalma

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Terpandrus paruna

Caedicia mesochides

Eggs Katydids lay their eggs in a variety of substrates. The eggs are distinctive but not as obviously diverse as the eggs of stick insects, for example. There are a range of egg types – they often have subtle features and need to be studied under high magnification. The depressed area dorsally contains the micropyles. As the egg is laid the micropylar area passes by the opening of the spermatheca and the egg is fertilised. The micropyles of most species have a distinct appearance and their position on the egg is characteristic. Most katydids lay their eggs in the ground. The ovipositor is specially modified for this purpose. It is elongated and sharply pointed. If you watch a female in the egg-laying process you will notice that there are four parts to the ovipositor and they operate alternately; with each stroke the ovipositor goes slightly deeper into the ground. You can often predict the egg-laying site by the shape of the ovipositor. A sickle-shaped ovipositor indicates that the species oviposits in plant material, either dead wood, stems or living plant tissue. Elongate ovipositors of uniform dimensions often indicate egglaying in hollow grass stems. There are even katydids that lay their eggs in the tissues of plant galls. The most bizarre scenario known for an Australian katydid is that of Polichne argentata – a species of the dry interior of the continent. The female lays an egg on a twig at night and then flies to the ground where she picks up a small amount of soil in her mouthparts. She

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returns to the egg and coats it with the moist soil. It would seem that this protects the egg from desiccation. Eggs of some species undergo a diapause, that is, they suspend their development so that it can resume at times when it will be favourable for the hatchlings (called nymphs) when they hatch. The simple scenario for the life history of an Australian katydid is that the female lays the eggs during the summer. They develop during the winter and hatch the following spring. This single generation per year seems to be the norm in temperate regions. In southwestern Australia, 14 of the 16 species found in Eucalyptus woodland had a single generation per year (Gwynne et al. 1988).

Some katydid egg-types

The egg of Chlorobalius leucoviridis has a simple lattice-like pattern.

The eggs of the Tympanophorinae species have a cap at one end.

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The egg of Neophisis ecmurra lacks an obvious lattice pattern.

The cap on the top of the egg of Indiamba malkini is apparently for protection or to transport air to the egg which is laid in wood.

Most katydids lay their eggs in the ground. Photo: Alan Henderson

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This katydid has a sickle-shaped ovipositor. Photo: Alan Henderson

The female Polichne argentata coats her egg with moist soil.

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The ovipositor of this katydid is elongate and is inserted into the hollow grass stem. Photo: Alan Henderson

Diapause can be very complex even within a species. In Europe a katydid species was found to have double diapause which was triggered by different day lengths at different latitudes. Some species can survive very long in the egg stage. Several European katydids require three winters in the egg before they hatch. On the other hand, there are some species that hatch without any dormant period, requiring only about 50 days to develop. This latter strategy results in overlapping generations where you find adults, nymphs and eggs of a given species in the same place at the same time. This is a common feature of tropical species.

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Katydid eggs hatch usually at dawn.

The nymphs of some species look very different from their parents.

Growth and development Katydid eggs hatch usually at dawn. The young often look like miniature versions of the adult but without wings. Many, though, are differently coloured and in some groups they are so different that you would not believe the two are the same species. They have a different appearance and different behaviour. Some species resemble floral parts and others, spiders, and still others can resemble Assassin Bugs (Reduviidae) in their appearance and behaviour. This mimicry lasts for only the first moult or two. Afterwards, when the nymphs are less susceptible to predation, they begin to resemble the adults.

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Our knowledge of the nymphs of most species is scant and incomplete. Reasons for this are varied but one might be that nymphs of most species emerge at times when collectors are not in the field. Or perhaps the nymphs of arboreal species are ‘out of reach’ of the general collector and they just never encounter one another. It seems that most species have a nymphal strategy for survival that is often quite different from the adults that they are to become. Katydid nymphs are under threat from predators from the moment they emerge from their eggs. The majority of species begin life attempting to free themselves from soil at a depth about that of the length of the ovipositor of their mother. Once atop the surface of the soil, they need to find a food source and, if a specific host plant, the quest is more difficult. If they are predaceous, then proper-sized prey must be located. At this point they are subject to predation from a host of enemies ranging from other insects to small vertebrates such as lizards, frogs, birds and small mammals. They are also at the mercy of the elements. Many die in wet years from fungal infections. In dry years they may die from heat or a dwindling food source. It is for all these reasons that, in many species, all the eggs do not hatch the following season. Some from the same batch may hatch over a period of years. Katydids whose eggs were laid in plant tissue, such as wood or dry grass, have a very different life scenario from those that emerge from the margins of leaves or from eggs laid in groups or singly on twigs. The katydids in this group are often protectively coloured with prominent reddish or orange colours so that vertebrate predators know that the katydids are toxic to some degree. Others from the former group are coloured to resemble something in their habitat such as green leaves or the colour of flowers. If the latter, they actually incorporate the pigments of the flowers into their own bodies through their metabolic processes. In this way they are somewhat concealed in or on the flowers. Remarkably, they can change their colour as they mature and the flowers fade. Even more remarkably, at the last moult the katydid changes completely and becomes the normal-looking adult. Many species are masters of camouflage. The combination of colour, structure and behaviour renders them virtually invisible in their habitats. Probably the best Australian examples are the species of Phricta (see pages 145–146). During the day the katydids rest with legs outstretched on tree trunks or branches perfectly concealed, moving only if greatly disturbed.

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How they avoid being taken by the ever-present marauding ants is unknown. The nymphs of the Queensland Palm Katydid, Segestidea queenslandica, are mottled greyish-white and resemble bird droppings on leaf surfaces (see pages 151–152). A number of katydid nymphs mimic ants, spiders or predaceous bugs as a strategy to avoid vertebrate predators. As the nymph matures, the mimicry changes to something a bit more benign, after all, the mimicry is only available to a creature that is the size of the animal that is mimicked. As the katydid outgrows its mimic it must rely on other strategies. This often is merely to resemble plants or plant parts.

This katydid nymph provides a wonderful example of protective colouration.

Nymphs of most Australian katydid species have not been associated with the adults they become. This is a fertile field for the observer. One can be certain that there is a unique life scenario with each katydid species. Raising katydid nymphs to maturity is not especially difficult. See page 206 for hints on how to do this.

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This tiny phaneropterine nymph lives on the ground among dry grasses, its white antennae resembling a grass stem and its body a seed or a bit of dead grass.

Katydids moult or shed their skins in order to grow. This is the common method of growth in insects with gradual development as opposed to those with complete development such as beetles or butterflies where there are three distinct states – egg, larva and adult. Katydid nymphs moult usually at night. There are good reasons for this. At night the humidity is generally high and this facilitates wriggling out of the old ‘skin’ and assuming the proper position for expansion and ‘growth’. Another reason for moulting at night is that the katydid is most vulnerable at this time and, under the cover of darkness, there is a reduced chance of predation. The moult has usually been completed by dawn.

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This phaneropterine nymph has a spot that obviously breaks up the insect pattern. Is this spotting just to break up the form of the insect so that it may be lost in the spotting of the surface of the leaf on which it sits? Or is there some other meaning to this pattern?

The moulting katydid usually hangs upside down in vegetation or at the end of a twig. You can see this with katydids, crickets, stick insects, mantids and some grasshoppers if you look around at night. Once it has determined the proper moulting site, the katydid increases its blood pressure and literally splits the old skin and crawls out headfirst. The nymphs moult four to nine times depending on the species and the climatic conditions.

This katydid has just moulted. It is hanging from the old skin which it will consume.

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An adult Austrosalomona katydid following the final moult. Blood pressure will inflate the wings.

This katydid is using blood pressure to inflate its wings to their proper shape.

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The adult katydid with wings fully developed. The skin will be consumed and the katydid will be ready for its life as an adult.

There seems to be no correlation between the number of moults and the taxonomic group to which the species belongs. Longer ‘growing seasons’ seem to harbour species that have more instars and a longer time between moults. In some species it has been found that males have fewer moults than females. Reasons for this are complex and probably vary from one species to the other. This is a wide-open field for study in Australia where very little is known about this aspect of katydid biology.

Food and feeding The food habits of katydids are very broad and include leaf and flower feeding and the highly specialised habit of nectar and pollen feeding. A number of species feed on seeds and fruit. Herbivorous species feed on a wide range of plants from grasses to the foliage of shrubs and trees. But many are opportunists and will feed on whatever food is available. Some even eat dead relatives! At night it is not uncommon to see wandering katydids feeding on road kills of their own kind or of other insects.

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Some katydid nymphs have food habits different from the adults they will become. Katydid foods range from species that are omnivorous to highly specialised feeders where their biology if finely tuned to the flowering of their host plants. The Pollen and Nectar-feeding Katydids and the Flower-feeding Katydids are example of this group. These species are ‘tuned’ to the flowering of native trees and shrubs although they sometimes short-circuit this and can be found in gardens feeding on flowering plants, both native and introduced.

A female Kawanaphila feeding on pollen and nectar at night.

Austrosalomona nymph feeding on a portion of a flower.

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A Phasmodes katydid consuming an entire flower at night.

A Phricta nymph after feeding on a bit of fresh leafy vegetation.

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These predaceous katydids are taking advantage of the nutrition provided by flowering eucalypts. Photo: Jack Hasenpusch

Several groups are primarily foliage feeders. These include the Mecopodinae, the Pseudophyllinae and the Phaneropterinae. Within these groups, some katydids prefer the more proteinaceous plant material such as flowers and floral parts but can develop quite successfully on green parts alone. A number of species eat seeds, especially grass seeds, and have mandibles that are modified for seed cracking. The heads of katydids often give a clue as to their feeding habits. Those with rounded heads are usually leaf eaters. Those with slanted heads either crack seeds or eat tough fruits or chew through wood. Others with projecting mouthparts are flower feeders, feeding on the flowers themselves, the pollen or the nectar within the floral parts.

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Katydids with slanted heads often have mandibles that permit them to dig in wood or consume tough seeds and fruit. Photo: Alan Henderson

Australia has two endemic subfamilies of the Tettigoniidae that specialise in flower-feeding. As a result, their mouthparts are also specially modified. Phasmodes ranatriformis, a stick-like species that feeds on all parts of the flowers in its habitat and has a robust mandible. The much more specialised species, Kawanaphila goolwa, feeds on pollen and nectar. The long groove alongside the incisor presumably transports the nectar to the crop. The series of spikes at the base of the mandible probably grinds and sorts the pollen grains since this area is often found clustered with pollen. Predaceous katydids have sharp incisors that are designed to grip and hold prey.

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This katydid with a rounded head is a leaf-eater.

This katydid is also a leaf-eater.

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The projecting mouthparts of this katydid allow it to insert its head deep into flowers to feed on pollen, nectar and other floral bits.

The forward-projecting mouthparts are called ‘prognathous’. This feature is characteristic of phamsmodine and zaprochiline katydids.

Katydid mouthparts can also provide clues to their feeding preferences. The mandible is usually modified for the food preferences of the species. This is often distinctive at the level of genus or higher. The standard mandible is short and robust and consists of stout teeth used for chewing and holding. Some species seem to be plant and animal feeders and have the mandibles thus adapted. The Austrosagine genus Psacadonotus may be an example of this category. Leaf eaters can easily be separated from carnivores and pollen eaters. To observe these differences, you must dissect

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the mandible from the specimen and study it under magnification. The mandible of carnivores tends to be elongate and to have the incisor elongated and hook-like whereas those of the seed crushers (the Copiphorini) have mandibles that have a groove on their inner edges, supposedly to hold the seed while the powerful jaw muscles crush. (I have learnt from personal experience that this group in particular can be dangerous to handle carelessly as their mandibles are very effective at cutting through the skin of a finger. If you are painfully bitten, it does not always indicate you have met up with a predaceous katydid; it may be a seed-crusher.) Predatory katydids are well represented in the Australian fauna. Where some species are omnivorous and will eat insects from time to time, there are others that appear to be wholly predatory. The most obvious are the Listroscelidinae which has a wide range of unrelated genera but all of which prefer insects as food. Hexacentrus mundurra has the mandible of a fullblown predator – it is elongate and bears a prominent incisor used for stabbing and holding.

Mandible structures

The standard mandible is short and robust.

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Kawanaphila goolwa has a more specialised mandible.

The mandible of Psacadonotus species is adapted to feed on plant and animals.

This is the mandible of a typical predator, Hexacentrus mundurra.

As you can see, a variety of feeding strategies is represented in the Australian katydids. Some katydids even produce special calls that lure prey

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to their end. Others capture insects in mid-air, on the fly, and others stalk prey stealthily as do praying mantids. There are ambush hunters and those that specialise on other tettigoniids for food. In Mexico there are katydids that steal larvae from wasp nests at night. There is a wide-open field of study in katydid ecology and behaviour. We have only begun to unravel the secrets of this group of insects. The majority of katydids are probably omnivores or opportunists. They are not specialised feeders. They may be seen feeding on flowers one day and eating caterpillars the next. In rare instances, you may observe a katydid feeding on a vertebrate such as a frog or lizard. A few species can be considered as biological control agents because they eat the eggs and caterpillars of moths. Like their relatives the grasshoppers, katydids probably have an important effect on local vegetation. It has been estimated that some species in North America can consume two per cent of the growth of a marsh grass and other species consume nearly 16 per cent of rush species (Juncus). Further, feeding of katydids reduces the production of seeds in certain plants. The feeding of katydids probably plays an important role in controlling vegetation in nature. In Australia, katydid numbers are usually low but there are several species that can be very abundant at times and have a distinct affect on vegetation, especially rangeland and pasturelands. Another negative effect of katydid feeding is the role these insects can have in horticulture. The controlled conditions of shade-house and glasshouse orchid culture can provide ideal habitats for a number of species that feed on developing flowers and new shoots. Management of live butterfly and insect zoos often complain about certain predatory species that ravage their stocks. These katydids can be difficult to control under such conditions where the use of pesticides is not possible. Economic damage results when katydids feed on developing citrus, thereby disfiguring the fruit and reducing its market value. With agriculture moving farther into the northern tropics and more exotic fruits and vegetables being propagated, we can expect local katydid species to take advantage on this new food source and become pests. The cut flower industry and specialty plants in horticulture such as orchids frequently suffer from the feeding of katydids.

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Katydid enemies Katydids are sought for food by many vertebrates such as frogs, lizards and birds. Bats are known to glean katydids from leaf surfaces at night homing in on the calling katydid. Many katydids have evolved techniques that tend to disadvantage these predators. The colour and pattern of camouflage in katydids is an example of this. Rainforest katydids in many parts of the world have evolved stridulation strategies that confuse bats. Many insects and other invertebrates find katydids an important food source. Internal parasites include Gordian Worms. These horse-hair-like worms are included in their own phylum, the Nematomorpha. They are large worms and to the astonishment of the collector, they occupy the greater part of the abdomen of the host. However, they seem to interfere very little with the physiology of the katydid, at least until late in the host’s life. The worms eventually emerge and have been proven to ‘direct’ the host to water where the worms congregate and tie other worms in a mating frenzy of ‘Gordian knots’! They lay their eggs in or near the water and are supposedly ingested by aquatic insects which are then eaten by the katydids and the cycle perpetuates. It is not unusual to find katydids, and other orthopteroids such as mantids, parasitised by Gordian Worms. More typical nematodes also attack katydids and some rival the Gordian worms in size. Insect predators include a variety of Digger Wasps (Sphecidae), and tachinid flies. Bugs (order Heteroptera) are often able to ambush and subdue katydids that are much larger than they are. Spiders are a constant threat. Many kinds of ants disable katydids and literally tear them to shreds. Some katydids are specialists at feeding on relatives. One species in particular specialises in eating other katydids.

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Parasitic worms can occupy the greater part of the abdomen of their host.

This little pentatomid bug has subdued an adult katydid many times its own size by toxic injection. It will feed on the fluids of the katydid and then depart.

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Many katydids fall prey to spiders. Photo: Jack Hasenpusch

Green Tree Ants commonly kill insects attracted to lights.

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Terpandrus katydids are predators and other katydids are on the menu.

Flies of the family Tachinidae employ a variety of elaborate strategies to find their katydid hosts. Ormiine tachinids provide an extraordinary example. The flies are adapted to hear the calling songs of nocturnal male katydids. As one might expect female katydids are not parasitised by these flies. With the Western Australian agraeciine katydid Sciarasaga quadrata, the female fly sprays a barrage of living larvae up to 6 cm away from the potential host. Most of the larvae miss the target, but if they are successful, they burrow into the male katydid through the intersegmental membranes and begin feeding. After a few weeks, the male katydid succumbs and the adult flies emerge to infect another. The fly Homotrixa alleni is known to parasitise three katydid species in two subfamilies: Sciarasaga quadrata and Pachysaga croceopteryx of the Austrosaginae and Mygalopsis pauperculus of the Copiphorini. It was discovered that 90 per cent of the calling males of S. quadrata were parasitised by flies, some individuals by two species. The other is a sarcophagid fly of the genus Blaesoxipha. This fly does not use acoustical means to find its host. Parasitic wasps in a number of families attack the eggs of many katydid species. Gregarines and fungi are also encountered in Australian katydids. Katydids and other insects appear ‘frozen in time’ when attacked by fungi. They usually assume a prominent position on a twig or leaf, secure

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themselves and then die in place. It is assumed that the higher they are in the habitat, the easier it is for the fungal spores to spread and infect more insects. Quite often multiple infected insects can be found in a small area.

This adult katydid has been killed by a microorganism, probably by a fungus.

There are many parasites that attack katydids but do not either kill or seem to harm them. Tiny mites infest the auditory tracheae of a number of species. The blood red mites of a number of species are commonly seen on the intersegmental membranes of katydids or attached to the tegmina or wings where they feed on the blood of the host. The mites feed for a short period and then drop off the host. You will find these tiny mites on a number of the katydids illustrated in this book. Strepsiptera, the twisted-winged insects, are curious parasites. They are partially internal and external parasites. They slow the hosts down and infected katydids are easily recognised as such. Dark spots on the abdomen of female katydids often belies the presence of a myrmecolacid strepsipteran parasite. The dark spot is actually a portion of the parasitic female’s reproductive anatomy that permits the male strepsipteran to mate.

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Sound and hearing Katydid song All Australian katydids produce sounds. No Australian katydid ‘rubs its legs together’, to produce sound as popularly believed. In fact, there are no Orthopterans that produce sound that way. Certain grasshoppers produce sounds by rubbing the legs against the raised veins of adjacent tegmina. The most common method with katydids is stridulation by the ‘file and scraper’ method (see the illustration opposite). The stridulatory file of the male is located on the underside of the left tegmen (1). The male rubs this file against a raised vein on the right tegmen (2). A simple analogy is the rubbing of a thumbnail across the teeth of a comb. This method is used by males to attract females. There is a vestigial file on the right tegmen that serves no role in stridulation. (Rarely, there are individuals of some species that rub the right wing over the left. Right-overthe-left stridulation is common to crickets). The structure and shape of the tegmen is extremely precise since it produces the sound to which the females respond. The mirror (3) is important as its size and shape helps to determine the way the sound is heard by the recipient. There are a variety of stridulatory file types in the Australian Tettigoniidae. These range from the simple files seen in the Zaprochilinae to the more standard files of the Listroscelidinae to the highly specialised and complex files of the Tympanophorinae. To see the stridulatory file you must be able to see the underside of the tegmen. This involves either spreading the left tegmen or removing it from the insect. The file types illustrated on page 26 have been photographed using the scanning electron microscope.

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The ‘file and scraper’ mechanism of a katydid.

The shape, length and number of teeth in the file are useful taxonomic tools. The physiology of the male singer determines cadence, pulse rate, etc. Females of most, but not all, species do not have the ability to produce sound. They ‘answer’ the calling males by attempting to find them by following the sound signal. In a few examples sound is produced in other ways. In some Psacadonotus species there is a ‘femoral stridulatory comb’ which is used in defensive behaviour when the black fan-like hind wings are rubbed over the comb producing a raspy sound (see pages 79–82). Curiously, not all the species of Psacadonotus that have this comb use it in the same way. Other Australian katydids such as some Metaballus and Polichne species ‘shuffle the wings’ when disturbed. Some tropical species produce sound by rubbing their mandibles together. Vibration of the substrate and of vegetation by courting males has been observed in a few instances. The Phyllophorinae do not use the tegmina or wings to produce sound. They have hind coxae with ridges that produce a sound by stridulation. This seems to be a defensive behaviour. Females of most species are silent. But females of a number of phaneropterine species answer calling males. Recently this was discovered as well in the Chlorobalius leucoviridis (see page 56). The mechanisms

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females use to produce sounds are radically different from those of the males. They do not have a stridulatory file but rather have pegs or spines at various positions on the hind wings that are rubbed against certain veins on the tegmen. Female acoustic responses in Australian katydids are largely unstudied. Just as the songs of birds are species distinctive, so are the songs of katydids. Many Australian katydids have been documented on CDs which are an invaluable aid in identifying species. See Gwynne (2001); Rentz (1993b, 1996b).

Types of stridulatory files

Zaprochilus australis (Zaprochilinae)

Requena kerla (Listroscelidinae)

Chlorobalius leucoviridis (Listroscelidinae)

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Austrophlugis debaari (Listroscelidinae)

Tympanophora kalbarri (Tympanophorinae)

Hexacentrus mundurra has a slit opening at the base of its leg which serves as the entrance to the hearing organ.

Hearing It is often thought that katydids can only hear the calls of their own species since the structure of the file and scraper produces sounds of a rather narrow range of frequencies. But this is only true some of the time. Studies in Western Australia revealed that males of Mygalopsis marki hear one another but this can be interfered with by the singing of an unrelated katydid, Hemisaga denticulata. If there are other Mygalopsis marki males

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singing close by and a H. denticulata male calls, ‘masking’ these rival male calls, the M. marki males cannot hear each other. Once the H. denticulata stops singing, the M. marki males separate, moving as much as 10 metres to avoid the rivalry. Hearing is a most complex subject in katydids and has occupied the careers many researchers. The subject is too detailed to discuss here. See Gwynne (2001) for a comprehensive discussion of hearing in katydids. Suffice it to say that katydids hear sounds in two main ways: through the auditory tympana of the fore tibiae, and through the thoracic auditory opening. Thus they perceive vibratory sounds through receptors on both the legs and the abdomen. The fore leg of a predatory species, Hexacentrus mundurra has a slit opening at the base which is the entrance to the hearing organ. This is quite different from the tympanum-like structure typical of the phaneropterine katydids.

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The auditory tympanum of the fore tibia of a phaneropterine katydid.

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Habitats Katydids can be found in most habitats in Australia except the oceans and the tops of the highest peaks. Look for katydids on the ground, in shrubbery or high in trees. Many species are active during the day and many more are nocturnal. Most can be found during the day using specialised techniques which are discussed in relation to the tribes or subfamilies in this book. With some arboreal species, you can find nymphs on annual or perennial vegetation for most of their nymphal lives. With a little special care, these nymphs can be collected and raised to maturity without much difficulty. Choosing the time of year to collect is very important. If you look for Pollen Katydids, Pollen and Nectar-feeding Katydids or Stick Katydids in summer, you are not likely to find them. These insects are at their zenith in winter and spring and are largely gone by summer. Many Shield-backed Katydids are adult in mid to late summer but others are found early in the summer. Travel to remote locales will always yield species of interest. But you do not have to travel far to find katydids. At least two species can be seen and heard in the vegetation at Darling Harbour, Sydney. Even the most desolate of habitats harbour katydids. In the hot interior of the continent, the best time to look for katydids is after dark. However, it is not unusual to find katydids seeking shelter during the day on cliff faces or on large rocks. A locality near Bullfinch, Western Australia, yielded a number of katydids in the dead, dry leaf and twig litter under the eucalypts. Other species were found up in the trees in the foliage. Mixed eucalypt forests, such as in the vicinity of Balladonia, Western Australia, are productive in the spring following good rains, yielding nymphs of a number of species that begin life among the spring herbs and annuals. In summer, katydids occur in the dry leaf and twig litter as well as on shrubbery and in the trees. Areas of mulga vegetation offer many species but most are nocturna and not easily found during the day. Many can be attracted to lights after dark or found by searching vegetation.

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Collecting at a remote locale.

Darling Harbour, Sydney.

The coastal sand plain of Western Australia harbours some of the most interesting of Australian orthopteroids. Unfortunately, much of this habitat is under threat from agriculture and housing development. Many of the katydid species living there are highly localised and known from only a handful of specimens. These areas should be searched carefully and more should be done to conserve this unique and vanishing Australian heritage. The area around Margaret River and Cape Naturaliste in Western Australia has a stunning array of species, many of which are adult in late winter and spring. The Stirling Range, although appearing similar in many respects to Cape Naturaliste, has a different, but related katydid fauna. This and other similar areas are repeatedly burned from lightning strikes and

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‘control or prescribed’ burning activities. This affects the biota in one way or another. Heath habitats inland from the coast in south-western Western Australia are similarly diverse but with many different species. These areas should be searched in spring and summer. Many areas of South Australia are highly seasonal. Whether any katydids will be present at a given site is largely dependent on the rainfall patterns during the winter. If drought is in effect, there may be little if anything present. However, after good winter and spring rains, many species will be found. The eggs of many of the katydids seem to be able to remain dormant in the ground and hatch only when suitable climatic conditions prevail.

Balladonia, Western Australia.

Stirling Range, Margaret River, Western Australia.

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Heathland, Tutanning, Western Australia.

Kangaroo Island, South Australia.

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A billabong in central Australia.

Spinifex, Wilpena Pound, South Australia.

Lawn Hill, Queensland.

Kangaroo Island and the adjacent Eyre and Yorke peninsulas share species in genera that can be found in the southern coastal heaths of Western Australia. Coastal mixed sclerophyll forests of New South Wales harbour distinctive species that are present from spring until late summer. A small number of species occur during winter, but these are the exception. Prior to the devastating fires of 2003, the mountainous area in the Australian Capital Territory was a productive habitat for many distinctive species. Until the vegetation is restored, these species will remain rare.

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Every trip to the remote portions of the Kimberley region of tropical Western Australia always reveals species of interest. Some of the commonest katydid species of the Kimberleys are still undescribed. Billabongs can be interesting collecting sites, especially after dark. But the collector must be aware of the potential of crocodiles and snakes in the northern portions of the continent. Spinifex habitats can be productive for katydids at most times of the year. Many species spend the day well within the plants and can be very difficult to wrench from their habitats without destroying the plants. However, after dark most katydids become active and ascend the culms of the flowering plants to feed and stridulate. The glass collecting tube makes life more pleasant while collecting in this spiny habitat. Rather uniform habitats such as in the vicinity of Lawn Hill, Queensland, have interesting diurnal as well as nocturnal species. Much of the northern portion of the continent appears like this and interesting katydids can be expected from these places.

Cooloola wallum, Queensland.

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Mixed eucalypt forest, Mt Carbine, Queensland.

Tropical rainforest, Queensland.

Wallum is defined as a habitat of coastal vegetation in Queensland where Wallum Banksia (Banksia aemula) occurs. However, the term is used for coastal vegetation growing on sand dunes or flat country with acid soils and

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a high water table. Such is the situation in the Cooloola area. This is a prime habitat for katydids, especially in spring and summer. Much emphasis has been placed on rainforests in recent years. This habitat represents about one per cent of the total landmass of Australia but it contains great species diversity in both plants and animals. Each rainforest, whether in the tropics or the temperate portions of the continent, harbours unique species. Of course, tropical rainforests have greater species diversity. But these areas are difficult to sample because species are seasonal and many live high in trees and may be host specific. There are ways to get around this. Some of the tree-top species must venture to the ground to lay eggs. At that time they are vulnerable both to the insect-eating fauna and the collector! Once the eggs hatch, the nymphs can be collected and taken to the laboratory for rearing to maturity. The mixed eucalypt forests, adjacent to tropical rainforests, are often overlooked by zealous collectors anxious to sample rainforests. These habitats contain a wealth of species, some of which are widespread and others which are very localised.

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Conservation Because of the localised nature of the distribution of many species, katydids have been useful tools for habitat preservation. Many species are relicts of another biota of former times. Some seem to be clinging to their existence. As cities expand and rural lands become housing developments, we can expect to see the demise of much of the biota. Katydids provide excellent examples. The coastal heath of Western Australia has undergone extensive development as the population grows. Jandakot, a Perth suburb is an excellent example. This area of mixed coastal heath vegetation was extensively studied over the past several decades by ecologists, behaviourists and taxonomists alike. Many important studies have been published using examples from this area. Today little if any of the original habitat exists. In a few years time, there will be no trace of the biota of this area. Islands are especially vulnerable to introduced organisms and their fauna needs to be thoroughly documented. Some islands, such as Hinchinbrook in tropical Queensland, remain intact and without much disturbance by humankind. Islands such as this are important because they give us an idea of what the coastal vegetation may have been like before the depredations of agriculture, introduced organisms and development. Hinchinbrook seems to be untouched in this respect and deserves a faunistic survey. Other islands can be just the reverse. Phillip Island, adjacent to Norfolk Island, is the other extreme. Rabbits were introduced by sailors many decades ago and quickly overpopulated and reduced the vegetation to the extent that up to two metres of topsoil was lost to the ocean. Many species probably were decimated with the loss of the habitat. Ironically, introduced European olives provide a source of cover and food for a number of crickets that survive there today. Recent attempts to eliminate the rabbits from the island seem to be successful and, perhaps, some of the fauna will return.

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Creeping development, such as this at Jandakot, a Perth suburb, threaten entire katydid populations.

Phillip Island, adjacent to Norfolk Island, has suffered significant biodiversity loss.

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Roadside verges provide a precarious habitat for katydids in agricultural areas.

Every field trip to an unusual location is bound to yield new species or rarities. It is important to document their presence, distribution and habitat preferences now. As agriculture and forestry extends more and more into the hinterlands, many habitats will either be totally destroyed or relegated to small patches. This is evident in the wheat belt of Western Australia. Some of the original vegetation is now to be found along roadside verges or at intersections of roads. This is a most precarious state. At the wrong time of the year, a fire can wipe out an entire population of insects. Misguided use of pesticides and weedicides can also result in detrimental effects. See Rentz (1993c) for a discussion of the situation.

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Guide to species Since we are dealing with a single family that is represented by potentially 1000 species or more, it is best to sort the species by subfamily. A subfamily is a unit comprising tribes and genera. Some are large and unwieldy, others small and homogeneous. But how does one get to the subfamily level? If you have a specimen in hand, go to the Key to subfamilies on pages 191–194 and see if you can reduce your choices to one of these units. If working with a photograph or from your memory, then go to the photos and plod through them, making comparisons and using the accompanying notes. Be sure to note the geographical distribution of your choice and the locality of your specimen. But also remember that distributions are sketchy for many species. And also remember, not all species may be included in this guide. There are many undescribed genera and species that are not to be found here. There is every possibility that you may have something new or unusual. The sequence of subfamilies in the body of the book follows a traditional phylogenetic scheme. That means, it is most likely that the more primitive groups are listed first and the more advanced ones follow. But this is subject to change and further research is most like to result in reordering the list. Some genera within a given subfamily have not been assigned to tribes and you will have to sort through the genera individually. A few tribes are not included here but can be found in the species lists on pages 195–200. Subfamilies and Tribes Listroscelidinae Conocephalomimini Phisidini Requenini

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Terpandrini Hexacentrinae Tympanophorinae Austrosaginae Conocephalinae Agraeciini Copiphorini Conocephalini Coniungopterini Microtettigoniinae Meconematinae Tettigoniinae Glyphonotini Nedubini Platycleidini Tettigonini Pseudophyllinae Phyllomimini Simoderini Phrictini Phyllophorinae Mecopodinae Mecopodini Sexavaini Phaneropterinae Ducetiini Holochlorini Phaneropterini Others Zaprochilinae

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Phasmodinae

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Hexacentrinae Fierce Predatory Katydids

This subfamily of highly predaceous katydids was considered a tribe of the Listroscelidinae by Rentz (2001) and is represented in Australia by two genera. Both are very distinctive and easily recognised. Hexacentrus is a pan-tropical genus, with 24 species listed in Eades and Otte (2009) but with only a single widespread species found in Australia. Alison is represented by a single species in Australia and three others described from New Guinea. Hexacentrus is strongly sexually dimorphic with the males having a balloon-winged shape and the females more typical in this respect. Alison is less sexually dimorphic with the females more closely resembling the males. In both genera males and females can fly short distances.

Genus Alison Rentz Alison’s Katydids This genus is related to Hexacentrus but is very different. It is probably most closely related to the Papuan genus Parateuthras, represented by two species in New Guinea. Alison has minute black markings on the base of the fore and middle tibiae and the tegmina of males and females are similar.

Alison roachae Rentz Roach’s Alison This species is known from a single locality, Shelburne Bay, Cape York Peninsula, Queensland. Females have a short, stout, falcate ovipositor, which is probably used to deposit eggs in wood or other plant tissue.

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Alison roachae (male)

Hexacentrus mundurra (female)

Genus Hexacentrus Serville Fierce Predatory Katydids Fierce Predatory Katydids are represented by species in Asia, the Pacific and central Africa. All have a remarkably similar appearance. The single Australian species is sexually dimorphic. The balloon-like male tegmina lends the males more to gliding than actually flying for any great distance.

Hexacentrus mundurra Rentz 70

Mundurra Fierce Predatory Katydid This species is common along the coast of Queensland from just north of Fraser Island to the tip of Cape York Peninsula. It also occurs along the Gulf of Carpentaria and into the Kimberley. The loud calling song of the male is one of the most distinctive nocturnal sounds of the tropics. This species lives in grassland, often along the margins of rainforests. It feeds on other insects with katydids a major portion of its diet. Females oviposit into the ground.

Hexacentrus mundurra (male)

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Conocephalinae Meadow, Spine-headed Forest, Snout-nosed and Gondwanan Katydids

Four distinctive tribes are represented in the Australian fauna. These are the Meadow Katydids (Conocephalini), the Snout-nosed Katydids (Copiphorini), the Spine-headed Forest Katydids (Agraeciini) and the Gondwanan Katydids (Coniungopterini). Some are very common. Several genera are common elsewhere in the Old World and others are virtually cosmopolitan. No comprehensive work has been done on this subfamily since that of Redtenbacher (1891). However, Ingrisch (1998) has made great advances on the reorganisation of the Agraeciini, and Pitkin (1980) included a number of common Australian species of Conocephalini in her revision of the species of the Pacific. Bailey (1979) reviewed the Australian Copiphorini. The Australian fauna is considerable and has many undescribed species. It should be noted that some species have examples with long and short wings. These can often be found at the same locality. The conocephalines are characterised by having the posterior margin of the lateral pronotal lobe with a swelling or ‘hood’ that overhangs the thoracic auditory spiracle. The boundary between the fastigium of the vertex and the frontal fastigium are closely pressed to one another. This group is very well preserved by the gutting technique described on pages 31–32. Key to the tribes of the Conocephalinae 1a Fastigium of head produced forward as a cone or spine; size variable but generally large katydids

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2 1b Fastigium of head produced not as a cone or spine 3 2a Fastigium produced forward as a distinct cone, very broad at its base, usually excavated and toothed below; slender to medium sized katydids Snout-nosed Katydids (Copiphorini) 2b Fastigium produced forward as a slender spine, usually very narrow at its base; size ranging from small to very large Spine-headed Forest Katydids (Agraeciini) 3a Pronotum surface normal, not thin or pliable; fastigium deltoid and convex when viewed from above, sides concave, surface not divided by median sulcus; male paraprocts not modified; slender to medium-sized katydids Meadow Katydids (Conocephalini) 3b Pronotum with surface very thin, pliable; fastigium produced as a peg-like projection or minute tubercle, sides not concave when viewed from above; surface divided or undivided by a sulcus; male paraprocts modified as flanges with lateral or ventral margins toothed; medium-sized, robust, very colourful species Gondwanan Katydids (Coniungopterini)

Tribe Agraeciini Spine-headed Forest Katydids and their relatives This tribe presents a disparate group of genera ranging from giants in the tropical genus Solomona to small delicate forms that occur in arid Australia. Attempts have been made in the past to separate out groups of genera into separate tribes or even assign them to new subfamilies. Ingrisch (1998) has recognised three subtribes in the Oriental Agraeciini. Some Australian genera can be assigned to his subtribes, others will doubtlessly require new subtribes, pending more comprehensive studies on the Australian components. For this guide, they will be placed in unassigned groups. Members of the Agraeciini have a spinelike projection between the base of the antennae and a narrow separation dividing the frontal fastigium and

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the fastigium of the vertex. They also have a ‘hood’ on the posterior margin of the lateral pronotal lobe that overhangs the thoracic auditory spiracle.

Subtribe Salomonina Genus Salomona Blanchard This genus is known from 73 species from the Indo-Malaysian region. These are large, formidable species that can cause damage to certain crops such as coconuts. Australia has a single species endemic to Iron Range, Queensland. Salomona is distinctive in Australia in its large size, with the pronotum with the disk and lateral lobes rounded, tegmina short to moderately long, the frons rough and a short, falcate ovipositor.

Salomona species

Salomona species Iron Range Salomona This species is uncommon in the Iron Range rainforests of northern Queensland where it lives in tree holes and emerges at night to prey on smaller insects. During the day, the katydids remain in the tree holes and defend themselves vigorously. From limited observations, individual katydids return to the same tree hole night after night. They have been seen tearing bits of wood to enlarge their habitat.

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Australian agraeciines not ascribed to subtribes Genus Austrosalomona Rentz Coastal Katydids Austrosalomona falcata (Redtenbacher) Olive-green Coastal Katydid This species is common along the east coast of Australia from Bateman’s Bay, New South Wales, to Iron Range rainforests of Queensland. It is easily identified by its olive-green colour and brown eyes. It is sympatric with several undescribed species with more restrictive geographic ranges. Like others in the genus, this species is at home in gardens and nurseries what it can cause damage to opening flowers and new shoots. The calling song of this species is commonly heard in Sydney’s parks and gardens.

Austrosalomona personafrons (male)

Austrosalomona personafrons Rentz Norfolk Island Katydid This species occurs in native forests on Norfolk Island. It would be a likely occupant of planted gardens. It is characterised by its robust body form, mottled greenish and brown pronotum and medium-brown body. The frons is olive-green.

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Austrosalomona falcata (male)

Austrosalomona species (female)

Austrosalomona species (male)

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Austrosalomona species (female)

Austrosalomona species Destructive Katydid This species is well known to horticulturists in the Kuranda–Cairns area, Queensland. It is a beautiful lime green, often with purple eyes. It is common in dense forests and gardens where it is probably an omnivore but irritates growers by feeding on the developing shoots and flowers of orchids and a wide range of other native and exotic plants. Like others in the genus, the species is found in adult or nymphal forms at all times of the year.

Austrosalomona zentae (male)

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Austrosalomona zentae (male)

Austrosalomona species (male)

Austrosalomona species Gwinganna Katydid This is a beautiful species related to A. falcata but very different in having the basal antennal segments black and the abdomen yellow. It is known along the coast from northern New South Wales to Iron Range, north Queensland.

Austrosalomona zentae Rentz Zenta’s Katydid This robust species is known only from Lord Howe Island. This is a dark greenishbrown species that, like other species in the genus, lives along

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rainforest margins. It will probably be found to occur in gardens on the Island.

Coptaspis brevipennis (female)

Genus Coptaspis Redtenbacher Woodland Katydids Woodland Katydids are found in coastal and heathland habitats from central New South Wales to Victoria. One species is known from Tasmania. They occur in mountainous as well as seaside habitats. They seem to be associated with Lomandra where they can often be found feeding on the flowers and seeds after dark. During the day they seek shelter under bark or in unfurling leaves or at the base of clumps of Lomandra, often with a number of katydids gathered together. All species are brachypterous and similar to one another in appearance. The male genitalia and calling songs are used to separate the species. At least four species remain undescribed.

Coptaspis brevipennis Redtenbacher Sydney Woodland Katydid This species occurs from Sydney south along the coast to Bateman’s Bay, New South Wales. Other undescribed species are known from the Australian Alps.

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Goodangarkia praesinus (female)

Goodangarkia praesinus (male)

Genus Goodangarkia Rentz Head squeakers Head sqeakers are robust, short-winged katydids, mostly green in colour and with powerful mandibles that permit them to pierce grass seeds and other tough plant material. They occur in meadows where they are subject to predation from a variety of organisms. Perhaps, as a result, these katydids have developed a vocalisation to startle predators. They produce an aubible squeak that emanates from the head from structures that have yet to be studied.

Goodangarkia praesinus (Karny) Cape York Head Squeaker This species is common in grassy woodlands in the vicinity of Cooktown and south to Cairns, northern Queensland. It is nocturnal and can be quite common. Both sexes emit a squeaking sound from the head when disturbed.

Undescribed genera and species 82

The following genera and species are as yet undescribed. They are relatively common where they occur and they are included here for that reason. These undescribed taxa reflect the slow progress in documenting and describing Australia’s abundant insect fauna. It is hoped that this book will be a stimulus for some to take an interest in this group of insects.

Undescribed genus Litter Katydids Litter katydids are very easy to over look. They are small and dark coloured and occur in the dark twigs and accumulated leaf litter under eucalypts in the arid regions of central and western Australia. Many undescribed species are known. There is a peculiar species found in the Pilbara, Western Australia, which spends the day on tree trunks living in depressions and knotholes where it ambushes passing prey.

Kalgoorlie Litter Katydid This species is not uncommon under eucalypts where it lives among the tangled dry twigs and dead leaves of the trees. It is very wary and not easy to see or capture. It seems to be diurnal and predaceous. Repetitive burning is likely to reduce the numbers of this and other species of the genus since they are only found in the dry litter under eucalypts. Burning of the habitat would surely eliminate the species.

Kalgoorlie Litter Katydid (male)

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Kalgoorlie Litter Katydid (female)

Purple-mouthed Emerald (female)

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Purple-mouthed Emerald (male)

Purple-mouthed Emerald (female)

Olive-green Katydid (female)

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Olive-green Katydid (female)

Undescribed genus Olive-green Katydids Olive-green Katydids are common in rainforests from the Daintree to the Atherton Tableland, Queensland, and like the Emerald Katydids, is known from a single species. This genus has relatively the same distribution as the Purple-mouthed Emerald.

Olive-green Katydid This katydid is distinctive in its colour and body form. It is predaceous and frequently visits lights where it can be observed subduing small insects. The short, falcate ovipositor is probably used to insert eggs into dead wood or cracks in tree bark.

Undescribed genus Emeralds Purple-mouthed Emerald This species is a fairly common resident of rainforests from the Daintree to the Atherton Tableland. It is small and compact, and emerges after dark to prey on small insects. Only a single species is known.

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Small Forest Gem (female)

Undescribed genus Small Forest Gems Small Forests Gems are known from the Daintree south to the Atherton Tableland.

Small Forest Gem The short-winged species is nocturnal as are all other rainforest agraeciines. It sits quietly on leaf surfaces during the day with legs outstretched, much in the same manner as the unrelated phisidine species of the Listroscelidinae. After dark it forages for small insects in the understorey vegetation.

Undescribed genus Heath Katydids This genus is known from a number of undescribed micropterous species. All but one are straw-brown in colour and many have a dark stripe running down the middle of the body. They are usually common but a great deal of searching is necessary to locate specimens. All species live or near the ground and are often associated with spinifex (Triodia).

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South Australian Heath Katydid (male)

South Australian Heath Katydid This species occurs in heath communities where it is easily recognised by its seemingly wingless body form and straw-brown colour. It could be overlooked as a nymph of another species. Checking the male for the presence of tegmina under the pronotum is the sure way to check on maturity. Females are more difficult because they are wingless. The toughness of the body and the condition of the ovipositor is the best way to determine if they are mature.

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Armadillagraecia mataranka (male)

Genus Armadillagraecia Rentz, Su, Ueshima & Robinson Armadillo Katydids This genus is known from three species over a broad range of Western Australia, the Northern Territory and Queensland. The shape of the head and pronotum is distinctive. The predominant colour form is green in males, brown in females. However, a single species from the Darwin area is dark grey or black in both sexes.

Armadillagraecia mataranka Rentz, Su, Ueshima & Robinson Mataranka Armadillo Katydid This species is known in Aboriginal folklore in the Northern Territory where it can be fairly common. It occurs in mixed forests where it lives in ground cover, on or near the ground. It is difficult to locate during the day but after dark males ascend shrubbery and sing incessantly and in synchrony with one another. They are very easy to approach and resume singing shortly after being disturbed.

Armadillagraecia mataranka (female)

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Kapalgagraecia nauma (female)

Genus Kapalgagraecia Rentz, Su, Ueshima & Robinson This peculiar genus is obviously related to Armadillagraecia but differs in a number of important characters such as the absence of fixed recurved spines on the fore tibiae and the uniformly brown colour of both males and females. Both sexes have a dark triangular patch on the frons. Two species are known; a small species, K. brayi has been found only on Groote Eylandt, Northern Territory, and the other species is widespread from the Kimberley region to Kakadu National Park.

Kapalgagraecia nauma Rentz, Su, Ueshima & Robinson Kakadu Katydid This species seems widespread but uncommon in grasslands and mixed woodlands over a broad expanse of the Northern Territory and adjacent northwestern Western Australia. The katydids have been found wandering on the ground. It is assumed that they feed on seeds and flowers like their relatives, Armadillagraecia.

Genus Nicsara Walker Nicsaras Nicsara presently includes 36 species described from Australia and the Indo-Malaysian regions. At least ten species have been recognised in the Australian fauna. Most, but not all, are long-winged. Once revisions are made, it seems likely that some described Nicsara species will be transferred to other genera. The Australian species are distinctive in colour and form and seem to be gregarious. They are nocturnal and are

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omnivorous, feeding on a wide range of flowers and fruits. They do not seem to be predaceous.

Nicsara bifasciata (Redtenbacher) Striped Nicsara This species is widespread in the northern portion of Australia where it frequents the Eucalyptus woodland community. The black and light grey colour and pattern are characteristic. Males have elongate, fingerlike styles on the subgenital plate and females an elongate ovipositor. These nocturnal katydids often aggregate in suitable habitats under bark or deep in the axils of Pandanus to which they return night after night. During the day they remain quiet with the legs outstretched.

Nicsara cornuta (Redtenbacher) Spine-headed Nicsara This species lives in inland forests in southern Queensland. It is distinctive in its grey and black colouration and the prominent fastigial spine that protrudes from the head. This species spends the day under bark and emerges at night to feed and mate. It often returns to the same site after nocturnal foraging.

Nicsara spuria (Redtenbacher) False Nicsara This species is well camouflaged where it lives on bark. It often returns to the same place night after night. It is known from coastal forests from northern New South Wales to Bowen, Queensland.

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Nicsara bifasciata (male)

Nicsara cornuta (female)

Nicsara trigonalis Walker Tropical Nicsara This species was described from the vicinity of Darwin, Northern Territory. Like other species in the genus, the colour and pattern are species distinctive. It is widespread and appears to have populations that are distinctive in size and genitalic configuration. Further work needs to be done on this group.

Nicsara spuria (male)

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Nicsara trigonalis (male)

Genus Secsiva Walker Secsivas This genus was originally described to contain a single species from the Australian tropics. Another species originally described in Agraecia was easily referable to this genus. Several undescribed species are now known. There are two body forms within the genus. The slender form contains species that live in grasses. They straddle the stems and actively move about after dark. The other is a more robust form that seems to prefer life in low shrubbery and woody herbs adjacent to rainforests. All known species produce low, soft sounds that are almost inaudible to the human ear.

Secsiva differens (male)

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Secsiva univitta (male)

Secsiva differens (Redtenbacher) Cape York Secsiva This is a light brown species with bluish-green marks on the frons, thorax and the tegmina. It lives in shrubbery adjacent to grasslands in rainforests habitats on Cape York Peninsula, Queensland.

Secsiva univitta Walker Victoria River Secsiva This is a slender, uniformly light brown species with a few bluish-green stripes on the head. It seems to prefer dry grasses where it feeds after dark on the grass flowers and developing seeds. It has been found in small groups of individuals aggregating under bark and in Pandanus leaf axils.

Secsiva univitta (male)

Tribe Copiphorini Snout-nosed Katydids 94

The Snout-nosed Katydids are so-called because they have a broad peg between the antennae instead of a spine as in the Agraeciini. In some examples, the difference between a ‘spine’ and a ‘peg’ is not clear. This will cause some difficulties in ascribing some species to the appropriate tribe. This is an example of a group that needs a great deal of work. The lateral lobes of the pronotum may, but not always, lack the swelling or hood that overhangs the thoracic auditory spiracle. Some of the loudest katydids heard in Australia belong to this group. Some can easily be heard from a moving vehicle.

Genus Mygalopsis Redtenbacher Sluggish Snout-noses This genus is endemic to Western Australia where four species occur in the coastal heath community. They are large, but not often seen as they are very cryptic and secretive in their habits. All species have short hind legs and peculiar development of the hind wings where short spines occur on the undersides. These produce a rasping sound when the katydids are disturbed.

Mygalopsis marki Bailey Mark’s Sluggish Snout-nose This species occurs in two size morphs depending on the season. The snout is grey or black ventrally. This species is known along the coastal sand plain north of Cape Naturaliste.

Mygalopsis pauperculus (Walker) South Coast Sluggish Snout-nose This is the smallest known species in the genus and occurs from Augusta south to the vicinity of Albany.

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Mygalopsis marki Photo: William Archer

Mygalopsis pauperculus (female)

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Mygalopsis sandowi (male)

Mygalopsis sandowi Bailey Sandow’s Sluggish Snout-nose This is a larger, more robust species that occurs inland in woodland habitats where males sing from the ground or low in shrubbery.

Genus Euconocephalus Karny Euconocephalus broughtoni Bailey Broughton’s Snout-nose This is a sluggish species with rather peculiar habits. Males sing at night from low shrubbery adjacent to rainforests. This robust species is highly vagile and individuals fly freely and come to lights in the middle of dense forests. They are seemingly distracted by the lights from their travels either through or over the forests. Like other species in the tribe, Broughton’s Snout-nose feeds on grass seeds. It is distinguished in having a long wings and a short fastigium with a notch between the fastigium and the vertex clearly open.

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Euconocephalus broughtoni (male)

Genus Pseudorhynchus Serville Mimicking Snout-noses Pseudorhynchus is the Old World equivalent of the New World Neoconocephalus. In fact, it is very difficult to distinguish the two genera, if they can be told apart at all. Australian species have the same habits of many of their northern counterparts. They live in grasses where they use the powerful mandibles to pierce grass seeds which seem to be their sole diet. Both sexes have a sharply pointed fastigium of the vertex. Males have distinctive calls and genitalia, features useful in identification. Green and brown morphs can be encountered at any locality. At some localities, more than one species can be found.

Pseudorhynchus mimeticus Redtenbacher Mimicking Snout-nose This species is found in grassy habitats along the coast from southern Queensland to Bateman’s Bay, New South Wales. Its loud, incessant buzz can be heard along the shoreline at Bondi Beach, Sydney. There are long and short-winged forms with the latter being more common in the southern portion of the range of the species.

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Pseudorhynchus mimeticus (male)

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Pseudorhynchus mimeticus (female)

Pseudorhynchus lessonii (male)

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Pseudorhynchus lessonii (male, brown morph)

Pseudorhynchus lessonii Serville Lesson’s Mimicking Snout-nose This is a common, loudly buzzing katydid of the tropics. It occurs throughout the Pacific in grassy habitats. In the west it is known from as far south as Kununurra and in the east to the vicinity of Sydney as well as across the northern portion of the continent. It is also found on both Norfolk and Lord Howe islands. Over much of its range, this species is sympatric with P. selonis. This species is recognised by the bulbous and stout stridulatory vein when viewed in dorsal aspect. Females of the two are virtually indistinguishable.

Ruspolia marshallae (male)

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Genus Ruspolia Schulthess Schindler Robust Snout-noses The species in this genus have a broad fastigium of the vertex (snout) with the notch between the fastigium of the vertex and the vertex closed and without a tubercle at the base. This genus often occurs alongside Pseudorhynchus species but is not as common. Many species occur in temperate and tropical regions of both the Old and New Worlds.

Ruspolia marshallae Bailey Marshall’s Snout-nose This species occurs in Queensland where it is widely distributed. The calling song is relatively soft when compared with others in the tribe.

Tribe Conocephalini Meadow Katydids The Meadow Katydids are an important part of the Australian fauna. They occur in grassy habitats everywhere from the coast to the high mountains and from the deserts to the tropics. The dominant genus is Conocephalus which is cosmopolitan in its distribution. The author has identified about 40 undescribed species in Australia. This group was divided into subgenera beginning in 1915 by the North American orthopterists, James Rehn and Morgan Hebard. This was probably as a result of trying to deal with an increasing number of species easily referred to Conocephalus from all continents. A few other subgenera have been added from time to time. Eades and Otte (2009) list nine subgenera. The Australian fauna contains described species in three of these subgenera. However, many species remain undescribed and may be ascribed to other subgenera.

Genus Conocephalus Thunberg Conocephalus species This species is widespread in the Kimberley region where it lives in tall Sorghum grasses. It is as yet undescribed.

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Subgenus Conocephalus Thunberg Conocephalus (Conocephalus) willemsei Pitkin Willemse’s Meadow Katydid This species is known from the Northern Territory as well as Papua New Guinea. It has the tegmina concolorous and the internal spine of the male cerci has a pointed tip. The ovipositor is slightly curved and a little thicker beyond the middle. This species is known only from long-winged forms.

Subgenus Chloroxiphidion Hebard Conocephalus (Chloroxiphidion) upoluensis (Karny) Upolu Meadow Katydid This is Australia’s most widespread Conocephalus species. It is distinctive in the shape of the male cercus and the tenth tergite not having minute projections. There are some spots on the tegmen. This species is known only from long-winged forms. It occurs in every state and territory wherever there are grasses. It is nocturnal and has a distinctive call. It is commonly attracted to lights and can be present in large almost plague numbers at times. Key to the known subgenera of Australian Conocephalini 1a Prosternum armed with two spines 2 1b Prosternum unarmed. Hind tibia with three pairs of apical spurs; fore and middle tibiae armed ventrally with 5–7 spines; male subgenital plate with apex incised Subgenus Conocephalus 2a Hind tibia with two pairs of apical spurs; fore and middle tibiae armed with 5–6 spines on ventral surface; male subgenital plate with apex truncate, not emarginated Subgenus Chloroxiphidion

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2b Hind tibia with three pairs of apical spurs; fore and middle tibiae armed with 5–10 spines on ventral surface; male subgenital plate feebly emarginated Subgenus Anisoptera

An undescribed species of Conocephalus that is common in grasslands of the Kimberley region

Conocephalus (Conocephalus) willemsei

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Conocephalus (Chloroxiphidion) upoluensis

Conocephalus (Chloroxiphidion) albescens

Conocephalus (Chloroxiphidion) albescens (Walker) Whitish Meadow Katydid This species is easily identified by the distinctive tenth tergite of the male’s abdomen. There are short projections that point downwards. This species is common in south-eastern Australia. Like other members of the genus, the calling song is distinctive. Both long- and short-winged forms are known. However, the long-winged examples are most common.

Conocephalus (Chloroxiphidion) laetus (Redtenbacher) 105

Sorghum Meadow Katydid This species is easily identified. It is slender with a yellow tip to the male’s abdomen. The tegmina are concolorous and the ovipositor is at least 13.7 mm in length. It is associated with a number of grasses, native and introduced, tall and short. It is diurnal with the males producing an audible sound. This species has a spotty distribution in northern Australia and extends to New Guinea and the Pacific as well as Africa.

Conocephalus (Chloroxiphidion) dubius Willemse Doubtful Meadow Katydid This species is known from only the type from Burnside, Northern Territory. It is of doubtful status. Conocephalus species should be collected from the type locality to see if this is a distinct species.

Conocephalus (Anisoptera) semivittatus

Subgenus Anisoptera Latreille Conocephalus (Anisoptera) semivittatus (Walker) Blackish Meadow Katydid This species has a broad distribution across the Pacific and is common in rank grasses from Cape York south to Bateman’s Bay, New South Wales. It also occurs on Lord Howe and Norfolk islands as well as New Zealand.

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Both long- and short-winged forms are known. Short-winged individuals seem to be more common in the northern portion of the range of the species. There appears to be a cline for colour in this species with very light coloured individuals occurring in the north and the species becoming darker as one proceeds south from Cape York. The darkest individuals come from the south coast of New South Wales. Two subspecies have been described from Australia but these need to be re-examined in light of the large amount of material that has been collected recently.

Conocephalus (Anisoptera) bilineatus Photo: Peter Street

Conocephalus (Anisoptera) bilineatus (Erichson) Small Meadow Katydid This species shares characters with both C. albescens and C. upoluensis but is in a different subgenus. It has a narrower fastigium of the vertex and the tegmen has some spots. Both long- and short-winged forms are known with the latter being the most commonly encountered. This species occurs in the south-eastern portion of the country including Tasmania and is also known from New Zealand.

Tribe Coniungopterini Gondwanan Katydids Three genera of this tribe are known. One occurs in Chile, the others are from Australia and New Guinea. The Australian genera are more

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widespread than is usual for many tettigoniids. Each is known from a small number of described species. The Coniungopterini are remarkably coloured and have wasp-like jerky movements, flicking the antennae and displaying bright colours. They stridulate both day and night and seem to be predaceous. The falcate ovipositor is for placing the flat eggs in cracks in tree bark.

Genus Metholce Walker Black-footed Gondwanan Katydid This genus comprises one described Australian species and a large, differently coloured, undescribed species from New Guinea.

Metholce nigritarsis Walker Black-footed Katydid This is a stunning species. Young nymphs are often blood red and may resemble spiders or ants. The adults combine an array of colours which provide protective colouration in the trees in which they live. The hind wing is bright pink. The pronotum is unusually thin and quite pliable. The Black-footed Katydid is found in Australian trees and shrubs as well as introduced street trees. It ranges in eastern Australia from the Australian Capital Territory to central Queensland. Populations have also been found at Cape Naturaliste, Western Australia. Its fast-paced call can be heard from a considerable distance and is uttered during the day and on warm nights. It does not come to lights and is difficult to collect. Climbing is often the only way to obtain specimens.

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Metholce nigritarsis (male)

Veria colorata (male)

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Veria colorata (male, black form)

Veria colorata (female)

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Veria colorata (female)

Genus Veria Walker Colourful Gondwanan Katydid Veria colorata Walker Colourful Gondwanan Katydid This is another stunning katydid. There are two colour morphs. The most commonly collected form is a polka-dot black and white pattern, the other is entirely black or with very faint traces of a pattern. This is a tropical and subtropical species with populations on both sides of the continent. They are not usually collected because the katydids occur in trees, usually the tallest trees in the area. They are active during the day and the stridulations can continue on warm nights.

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Microtettigoniinae Micro Katydids

This Australian endemic subfamily comprises a single genus, Microtettigonia, with seven described species. The subfamily is characterised by the small size of nearly all species. Males of some species are only 5 mm in total body length. All known examples are micropterous and live on or near the ground in heath habitats in the southern part of Australia. Species are known from both sides of the Nullarbor Plain. They are herbivorous and seem to be associated with grasses and other monocotyledonous plants such as Lomandra, Gahnia and the like. One species, M. kutyeri, perches by day in the seed heads of a rush, head downwards, in meadows at Windy Harbour, Western Australia. Another species, M. tunte, occurs with it but is not found on the rush seed heads. These katydids are extremely fleeting and very difficult to capture. A glass tube seems to be the best technique used to capture specimens.

Genus Microtettigonia Rentz Micro Katydids Microtettigonia seems to be related to Conocephalus and its allies. All known species are diurnal. Male and female genitalia are very species distinctive. There are no sclerotised portions of the male genitalia but the cerci and tenth abdominal tergite are very important diagnostic characters. These katydids are best preserved first in fixative and then transferred to 75 per cent ethanol since genitalic examination is so important in making a positive identification. This provides the best preservation of these characters. Microtettigonia species seem to be relatively restricted in their distributions and as a result many other species are to be expected as remote

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areas are surveyed.

Microtettigonia alleni Rentz Allen’s Micro Katydid This species occurs in the Eneabba district of Western Australia where it is a member of the coastal sand plain heath community. The male cercus has a proximal internal tooth and the projections of the tenth tergite are convergent and directed downwards. The ovipositor is short and curved upwards and not serrate.

Microtettigonia alleni (male)

Microtettigonia tachys (female)

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Microtettigonia tachys (female)

Microtettigonia tachys Rentz Speedy Micro Katydid This species is known from many localities along the south coast of Western Australia from Mt Ragged west to Albany. It is a member of the coastal sand plain heath community and its range extends inland in similar habitats in appropriate areas. The distribution of this species has probably been reduced by the extensive destruction of the coastal sand plain community in the interests of agriculture.

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Meconematinae Leaf Marauding Katydids

This large, worldwide subfamily is primarily tropical and subtropical in distribution. It seems to be a disparate assemblage of genera which will undoubtedly be broken up and assigned to other higher taxa. Of the several tribes that have been identified, only one, the Phulgidini, occurs in Australia. The majority of genera have not been ascribed to any tribe, but fortunately all the known Australian genera fit neatly into the Phlugidini.

Tribe Phlugidini Phlugidine katydids are small, long-legged species that are built for speed. The head has the fastigium of the vertex flat and not projecting. The thoracic auditory opening is small and not much larger than the thoracic spiracle. The pronotum is often truncate and not much produced posteriorly. The ovipositor has the basal portion swollen. Many species seem to be both nocturnal and diurnal in their activity.

Genus Austrophlugis Rentz Australian Swayers A suite of Austrophlugis species has been recorded from Darwin and vicinity, and the Victoria River. Other species are known from the east coast of Queensland from Brisbane to Cape York. Undescribed species are to be expected in this genus because many are localised and are not likely to be encountered by general collecting. The mini bat detector is of considerable use in locating singing males. Many are active during the day and are very wary and take to flight or actively escape by jumping when approached. They are not usually attracted to lights but may be attracted to the small insects that remain on

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vegetation after light-trapping. In this genus both sexes are fully winged. The middle tibia bears two spines on the ventral surface.

Austrophlugis debaari Rentz Debaar’s Swayer This species is fairly widespread and is known from Magnetic Island in the north and south to Gympie and suburbs in Brisbane where it is found sympatrically with A. orumbera. It is distinguished by the pronotum that is not or slightly produced posteriorly. Males have the tenth tergite apically produced as a pair of divergent, lobiform appendages with the cercus elongate with the ventral margin acute and directed inwards. Females have the tenth tergite not bearing angulate appendages and not spiniform in appearance. This species has been observed to catch small insects on the hop in mid-air.

Austrophlugis debaari (female)

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Austrophlugis malidupa (male)

Austrophlugis kumbumbana Rentz Perth Swayer This is the only known Western Australia example of the genus. It occurs in the vegetation around King’s College, Perth. This species appears to be parthenogenetic but this feature is worthy of further study. Why is it known only from this southern locality when all the other species are separated from it by some thousands of kilometres?

Austrophlugis malidupa Rentz Malidupa Swayer This is a diurnal species common on the Atherton Tableland and from the Mt Garnet vicinity as well as Forty Mile Scrub National Park, Queensland. An odd variant has been found at Kuranda, Queensland. It is similar to typical Malidupa Swayers but differs in the female abdomen not being apically divided and males having the tenth tergite with an elongate, decurved median process. The cercus has the dorsal margin produced.

Genus Lucienola Gurney Short-winged Swayers All known species in this genus are short-winged with the tegmina either partially or completely hidden by the pronotum. The three Australian species were originally assigned to Tenuiphlugis, a genus later considered a

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synonym of Lucienola. Others are known from New Guinea and the Solomon Islands. The middle tibia is unarmed on the ventral surface. Males are highly distinctive in having styliform cerci that are not modified for grasping.

Lucienola pitti (Rentz) Striped Swayer This species inhabits Cape York Peninsula from Lockerbie south to the Iron Range and Heathlands, Queensland. It is easily recognised by the brown median stripe on the surface of the body. The projections of the male tenth tergite are widely separated and are divergent. They are positioned over or beyond the internal margin of the insertion of the cercus beneath. The male cercus is straight on the outer margin. Females have the interocular distance less than the width of one eye. These katydids seem to be very common as indicated by the amount of stridulation heard through the mini bat detector. They are active by day but courting has been observed in the evening.

Lucienola pitti (male)

Lucienola tiwiwarrina (Rentz) Speedy Swayer This species is known from many localities in the western Kimberley Region, Western Australia. Almost all of the comments on the biology of L. pitti pertain to this species as well. It seems to prefer low vegetation and broad-leafed shrubs and trees. This species is readily distinguished from L. pitti by the projections of the male’s tenth tergite being close together,

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parallel or nearly parallel and positioned in the centre of the tergite, well within the internal margin of the insertion of the cercus beneath. The male cercus is concave on the outer margin when viewed dorsally. The female interocular distance is greater than the width of one eye.

Lucienola tiwiwarrina (male)

Genus Indiamba Rentz Glossy Swayers This distinctive genus is known from three species from the Northern Territory. Both sexes are short-winged. This genus is noteworthy because of the extreme development of the male genitalia. The lateral extensions of the reduced tenth tergite fit into grooves at the base of the cercus on the internal margin. Males also possess a hook at the base of the cercus on the internal margin. Females are less specialised and are slightly more robust than those of the other genera.

Indiamba malkini (Jin) Malkin’s Swayer This species has extraordinary incurved, hook-like projections of the tenth tergite that fits into the internal margin of the enlarged cercus (see page 9). The female subgenital plate is elongate; much longer than broad.

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Indiamba quamara Rentz Quamara Swayer This species occurs in the Darwin vicinity south to Mataranka, Northern Territory. Males have small intercercal hooks with the median process very small and not hood-shaped or internally setose. Females have a short, broad subgenital plate. This species lives on the upper surface of the leaves of Terminalia carpentariae from one to three metres in height during the day. When alarmed the katydids run to the undersides of the leaves and assume a prone position. Females have been observed ovipositing at the base of another small tree, Erythrophloeum chlorostachys, about 1.5 metres from the ground.

Indiamba malkini (male)

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Indiamba quamara (male)

Indiamba quamara (female)

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Indiamba wirrawilla (female)

Indiamba wirrawilla Rentz Arboreal Swayer This species is known from Arnhem Land and Kapalga, Northern Territory. The male cercus enclosed in sheath formed by the elongation of the tenth tergite. The internal hooks are very large and the median process of the genitalia is hood-shaped and setose internally.

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Phyllophorinae Hooded Katydids

The Phyllophorinae is a peculiar group of mostly Indo-Malaysian or East Asian katydids. They are highly distinctive in the following characters: males without a typical tegminal stridulatory apparatus; pronotum extended hood-like over the bases of the tegmina with the margins dentate or spinose; tarsi depressed; fore tibiae with an open, slit-like foramen. Hooded Katydids, where known, produce sound as a defensive mechanism by movement of the hind coxae against the outer margin of the mesosternal lobes. The coxae bear a series of parallel transverse plates that produce sound. Two genera, each with a single species are known from Australia. They are confined to the wet tropics of Cape York Peninsula, Queensland. Little is known of their habits other than that they are herbivorous.

Genus Siliquofera Bolîvar Giant Katydids Siliquofera contains a single species that has been recorded from northern Cape York and New Guinea. It is distinguished by its large size and the pronotum that has the margins uniformly denticulate.

Siliquofera grandis (Blanchard) Giant Katydid This species is widespread in Indonesia and New Guinea. It has been seen rarely in the vicinity of Bamaga and Lockerbie, Cape York, Queensland. It is easily recognised by its large size. No other known katydid species in that area approaches it in size. It probably lives high in trees and this is the reason it is seldom seen. Paradoxically, it is apparently more commonly

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encountered in New Guinea and Indonesia since there are ample specimens of it in collections.

Siliquofera grandis Photo: Rolf Oberprieler

Genus Phyllophorella Karny Small Hooded Katydids Phyllophorella comprises 12 species with only a single species represented in Australia. It is easily distinguished from Siliquofera by the shape of the pronotum. There are prominent spine-like teeth that protrude from each side of the margin of the pronotum in the middle in Phyllophorella but the pronotum in Siliquofera is uniformly dentate along the margin.

Phyllophorella queenslandica Rentz, Su & Ueshima Queensland Small Hooded Katydid This species has a much broader range than its relative S. grandis. It occurs from the vicinity of Kuranda north to the Lockerbie area. It is not known from the Atherton Tableland. The specimens from the north are smaller than those from the south. Nymphs have a distinctive stance when they are at rest during the day. The colour of the tegmina is variable. About half of the individuals are uniformly green, the rest have brown spots that presumably simulate dead patches on leaves. When these spots are present, they are bilaterally

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symmetrical, that is they are in the identical position on either the right or left tegmen. Females oviposit in the ground late in the wet season.

Phyllophorella queenslandica (male)

Phyllophorella queenslandica (male) Photo: Gary Wilson

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Phyllophorella queenslandica (nymph) Photo: Jack Hasenpusch

Phyllophorella queenslandica (nymph) Photo: Jack Hasenpusch

Like the nymphs of the unrelated Chloracantha genus, the nymph of Phyllophorella queenslandica has a small white spot at the end of the body, but not at the first instar.

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Mecopodinae Long-legged Katydids

The Mecopodinae is a rather large assemblage of disparate-looking species. It has primarily an Old World tropical distribution. Within Australia two tribes are represented: the Mecopodini and the Sexavaini. Both are confined to the tropics. This subfamily is recognised by the reduced rims of the antennal sockets and the tibial auditory structure which is open on both sides. The members of this tribe seem to be herbivores.

Tribe Mecopodini These katydids are easily recognised by the shape of the fastigium of the vertex which is rounded or truncate. A single genus is endemic to Australia. The four species live in similar habitats on both sides of the continent. They are grassland inhabitants and nocturnal in their activities. They are adult during the dry season. The species are difficult to distinguish with the male genitalia providing the best differential characters.

Genus Austromecopoda Rentz, Su & Ueshima Grass Mecopodes This genus should not be confused with any other with the possible exception of the Grass Straddlers, Antipodectes (page 122–123). It is easily distinguished from that genus on the structure of the tibial auditory structure, the much broader tegmina and the reduced plantula of the hind tarsus in Austromecopoda. Four species have been described. One is widespread and occurs at disjunct localities from the Kimberleys, Western Australia, Darwin, and inland, including Arnhem Land, Northern Territory, to the vicinity of Cooktown and Cape York, Queensland. Three others are

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from Cape York Peninsula. Specialised searching is necessary to locate these katydids since they are cryptic and not easily prompted to jump.

Austromecopoda kayaman (female)

Austromecopoda kayaman Rentz, Su & Ueshima Frosty Grass Mecopode This species is known from Iron Range, Queensland, where it was found in disturbed habitats adjacent to rainforests. It often occurs among dry leaves under trees. The tegmina have frosty white blotches which distinguishes it from other Austromecopoda.

Tribe Sexavaini Palm and Banana Katydids This tribe comprises many genera mostly from the Old World tropics. Three closely related genera are often known locally in the Pacific Islands as Tree Hoppers or Coconut Treehoppers. Some can cause considerable damage to coconuts, bananas and oil palms. It is to one of these genera to which the single Australia representative belongs.

Genus Segestidea Bolîvar Palm Katydids 130

Segestidea queenslandica Rentz, Su & Ueshima Queensland Palm Katydid This species occurs along the east coast from the vicinity of Innisfail north to the Daintree region and east to the Atherton Tableland, Queensland. It is among the largest of Australian katydids, with some females measuring more than 90 mm in body length. It is not uncommon where it occurs and can often be seen after dark feeding on a wide range of palms, both native and introduced. This species is facultatively parthenogenetic. Males are extremely rare. Unmated females lay large numbers of eggs in the ground late in the rainy season that eventually hatch into females. During their mission to oviposit, they are encountered on low vegetation at night. Nymphs occur on small palms where their colour and posture convey the impression of a bird dropping or organic matter on the leaf as they sit motionless in full view during the day.

Segestidea queenslandica (female)

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Segestidea queenslandica (male)

Palm Katydid nymphs are very soft-bodied and vulnerable to vertebrate predators. The strategy during the day is to remain motionless on leaf surfaces. The colour pattern resembles a bird dropping or blotch on the leaf surface.

Segestidea queenslandica (nymph)

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Phaneropterinae Bush Katydids

This is the most diverse and most difficult of the katydid subfamilies to deal with because there are so many species and so little taxonomic work has been done with the Australian species. The generic groups have been ranked as tribes by some specialists and as groups by others. Many genera found in the Orthoptera Species File (OSF) will be shown to not be assigned to any tribe or group. That is, in part, as a result of taxonomists describing genera and species without much reference to their relationships. The names of many species date back to specimens collected on the early European scientific expeditions to Australia. As a result, most of the type material resides in European collections. In addition, the early describers did not realise the value of precise locality data and their descriptions were often brief and not very informative. Couple that with the general lack of illustrations in many early works and the lack of appreciation of the most informative taxonomic characteristics of a species, and you can see the problem. The efforts of those who maintain the OSF are helping to reveal the identity of the old specimens by adding photographs and important taxonomic characters on the website. However, much more needs to be done before we know what each taxon represents. Bush Katydids occur in most habitats. The majority of species live in shrubbery or trees, but there are suites of species that live on or near the ground and in grasses and annual plants. All known species are herbivorous. They can be found from sea level to just below the tree-line in the mountains. They are common in rainforests but there may be more species per unit area in eucalypt woodland habitats. Many species seem to prefer certain hosts or host associations. This subfamily has many interesting biological properties. One is that the nymphs are often mimics of other insects, for example, spiders, ants, or

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bugs. They can advertise aposematic colouration. In some species, the model of the mimic changes with the instar. The adult often looks nothing like the nymph. In at least one species, the timing of the hatching of the eggs is correlated with the flowering of the host Acacia and the first and second instars resemble Acacia flowers as a mode of protective colouration. As the flowers fade and dry up, so the colour of the nymph changes from bright yellow to yellow-brown and at the final moult, a greenish katydid emerges resembling an Acacia leaf. There is ample opportunity for the interested person to make original observations of the habits of even the commonest species. A variety of ovipositional strategies are used by this subfamily. Some species use the modified ovipositor to lay flat, disk-like eggs between top and bottom layers of a leaf. Others deposit rows of eggs on twigs and stems, and still others oviposit in the ground. There is one example of a phaneropterine that lays its egg on a twig and then flies to the ground and picks up a bit of soil with which it coats the egg. The calling songs of most species are very complex and in some, the females answer the males. For the most part, the great majority of species have not been sound-recorded, so there is a great opportunity to document this area of their natural history. As with other tettigoniid groups, there are many well-known, but as yet, undescribed species. Some of the commonest species still do not have names. And the ‘taxonomic impediment’ noted above casts doubt on the true identity of some well-known forms. So some of the identifications here may be tentative. The Phaneropterinae is characterised by the following combination of characters: head usually round, globular, not frontally flattened; tibial auditory structure open and tambourine-like in all but a few genera; legs rectangular in cross-section and armed on all four corners; ovipositor of many species short and broad, and laterally compressed and partially hidden by the abdomen. The list below follows the scheme in the OSF. Where tribes have been assigned, they are used here. Groups will be listed as such. But where no tribe or group has been noted, the genera are placed here alphabetically.

Tribe Ducetiini 134

Ducetias This tribe contains species that occur in the Ethiopian and Oriental regions as well as New Guinea, the Solomon Islands and northern Australia. The single species is distinctive and not easily confused with any others with the possible exception of some Polichne species.

Ducetia japonica (Thunberg) Pacific Ducetia This species is common and widespread over much of South-East Asia, Indonesia and the Pacific. It lives in disturbed habitats such as roadside verges and rainforest margins throughout the Australian tropics. There are two colour morphs: brown and green, with the latter much more common than the former.

Ducetia japonica (male, green morph)

Tribe Holochlorini White-footed Katydids This South-East Asian and Oriental tribe contributes a single endemic genus and species to the Australian fauna.

Leucopodoptera eumundii Rentz & Webber Eumundi

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This nocturnal species is known from rainforest sites from Cooktown to Bartle Frere as well as the Atherton Tableland. It occurs in lowland vine forests through to mesophyll and notophyll vine forests in the cooler regions. It feeds on a variety of plants with low foliage in canopy gaps. Adults are known from the end of the wet season (February–May).

Ducetia japonica (male, brown morph)

Nymphs have been seen from November to January. The Eumundi is very easily identified and cannot be confused with any other rainforest katydid.

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Leucopodoptera eumundii (male) Photo: Alan Henderson

Tribe Phaneropterini This tribe comprises genera from the Old and New Worlds. It contributes a single species to Australia in the very widespread genus that is the namesake of the tribe. Phaneroptera comprises species that occur in Asia, the Pacific and Africa. None is known from South America.

Phaneroptera gracilis (female)

Phaneroptera gracilis Burmeister Slender Bush Katydid

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This species is widespread across the Old World and the Pacific region. It is delicate and seemingly fragile. It occurs in grassland and marginal or ecotone communities.

Genus Group Cosmophylla This genus group is known from two genera, one from Chile, with two species, and the other from Australia, also known from two species. The group is characterised by the robust appearance of all its members, similarity of tegminal venation and the similarity of the male genitalia and the female ovipositor.

Genus Ozphyllum Rentz, Su & Ueshima Australian Bush Katydids Ozphyllum is known from two species that live in the understorey of rainforests on the east coast of the continent. They have a characteristic appearance and are very soft-bodied. The tegmina are more pliable than in many other phaneropterines. These katydids are nocturnal and spend the day at rest, in cryptic posture on leaf surfaces.

Phaneroptera gracilis (male)

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Ozphyllum naskreckii (male)

Ozphyllum naskreckii Rentz, Su & Ueshima Naskrecki’s Bush Katydid This species is known from wet sclerophyll forests from Woodenbong, New South Wales, north to Mt Glorious, Queensland. At times this katydid can be very common and is especially visible on dark when males are calling. It has been observed feeding on introduced ivy.

Ozphyllum kuranda Rentz, Su & Ueshima Kuranda Bush Katydid This species ranges from Baldwin Swamp in the Bundaberg region north to Kuranda, Queensland. It has not been found in the Daintree region or on the Atherton Tableland. It is sporadic in its occurrence, being common one year and absent the next. It is distinguished from O. naskreckii by the shape of the male cercus. In this species the apex is spoon-shaped; in O. naskreckii it is hooked.

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Ozphyllum kuranda (male)

Genus Group Ephippithytae This group comprises at least ten genera, all of which are endemic to Australia and New Guinea. The majority of Australian Bush Katydids belong to this group with Caedicia being among the largest known Australian tettigoniid genus. Rentz et al. (2008) presented a key to the genera belonging to this group.

Genus Ephippitytha Serville Spotted Katydids This genus consists of five rather similar species, from inland arid habitats and northern Queensland rainforests. Several species listed in the OSF are undoubtedly synonyms. Ephippitytha species are easily recognised by their large size, spectacular appearance and spotted tegmina. Ephippitytha nymphs have an ant-like appearance at first. Later, this gradually changes and both E. trigintiduoguttata and E. kuranda assume the appearance and behaviour of the adults.

Ephippitytha trigintiduoguttata (Serville) 32-spotted Katydid This large, slender species has more-orless 32 spots. The fore and middle femora are not flanged and the hind tibia bears three narrow bands, including the tarsi. This species has a widespread distribution and is often

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found with Alectoria superba. It is frequently attracted to lights at petrol stations. The short ovipositor is used to glue eggs to dead twigs.

Ephippitytha trigintiduoguttata first instar nymph

Ephippitytha trigintiduoguttata late instar nymph

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Ephippitytha trigintiduoguttata (female)

Ephippitytha kuranda (female)

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Alectoria superba (female, yellow morph)

Alectoria superba (male, green morph)

Ephippitytha kuranda Rentz, Su & Ueshima Kuranda Spotted Katydid This species is more robust than its relative. The fore and middle femora are flanged at the apex. The hind tibiae have four broad bands, including the tarsi. This species is attracted to lights in the rainforest. The ovipositor is short and used to glue eggs to dead twigs.

Genus Alectoria Brunner von Australian Crested Katydids This genus with its single species is among the most easily recognisable of Australian Orthoptera. Only a single species is known.

Alectoria superba Brunner von Wattenwyl Australian Crested Katydid No other katydid anywhere else in the world has the appearance of this spectacular creature. The crest is actually A-shaped and is hollow within. Its function is unknown and may have to do with amplification or directing stridulation. Females have the crest developed equal to that of the males. This species is widespread but never common in the arid and semi-arid parts of the continent. Yellow and green colour morphs occur in nearly equal numbers.

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Alectoria superba (final instar nymph)

The last instar of Alectoria superba is not greatly unlike that of the adult. The wing pads are small and separated. This inicates that the katydid is ready to undergo its final moult. It will do this after dark when winds subside, the humidity is relatively high and the danger from predators is reduced. It will harden by morning and be ready to commence adult life.

Genus Protina Brunner von Wattenwyl Semi-crested Katydids Like Alectoria, this genus is represented by only a single species. The distribution of the two genera probably overlaps in some areas but Protina is much less common than Alectoria.

Protina guttulata Brunner von Wattenwyl Semi-crested Katydid This species bears some resemblance to Alectoria superba but is more robust and the crest is much more reduced. It is unusual in several respects. No other katydid has the posterior margin of the pronotum developed as it is here and few katydids are pinkish in colour as adults. This species is very uncommon and is known from only a few localities. Like A. superba it is nocturnal. The specimen in the photo below has the egg of a fly parasite on its tegmen.

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Protina guttulata (male, showing the well-developed posterior margin of its pronotum) Photo: Jack Hasenpusch

Protina guttulata (male) Photo: Jack Hasenpusch

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Currimundria delicata (male)

Genus Currimundria Rentz, Su & Ueshima Flying Fox Katydids The Flying Fox Katydid is so-called because of the appearance of the tegmina of both sexes; it is bowed or arched in the middle resembling somewhat the appearance of a flying fox at rest. Only a single species is known in the genus.

Currimundria delicata Rentz, Su & Ueshima Flying Fox Katydid This species occurs in rainforests from the vicinity of Cooktown south to Kuranda, Queensland. It has not been found in the Atherton Tableland. It has a very delicate appearance and white ‘feet’ which are actually the tarsi. It shares this latter feature with many rainforest katydids. The ovipositor is short and very sharp suggesting the eggs are laid within plant tissue.

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Currimundria delicata (male)

Genus Diastella Brunner von Wattenwyl Speckled Katydids Diastella is known from three species which occur in Australia and another from New Guinea. The tegmen is narrower in the basal third than it is in the distal third and it is slightly bowed in the middle on the posterior margin, but never to the extent as seen in Currimundria. Some species have speckled tibiae.

Diastella kuranda Rentz, Su & Ueshima Kuranda Speckled Katydid This is a rather plain, green katydid with speckled fore and middle tibiae and often with a small spot on the thorax. The nymphs also have this spot enabling us to distinguish them from many other phaneropterines. The ovipositor is short and barely protrudes from the abdomen. At rest on leaf surfaces, the legs are tucked close to the body and the tegmina are slightly tilted upwards.

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Diastella kuranda (nymph)

Diastella kuranda (male)

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Diastella kuranda (female)

Genus Caedicia Stål Caedicias With nearly thirty species attributed to Caedicia, this is one of the largest and most complex of Australian katydid genera. A perusal of the type specimens on view in the OSF suggest that there are species included here that actually belong to other genera. Caedicias can be found in many habitats in Australia from deserts to rainforests. The Garden Caedicia lives in gardens in Melbourne, Sydney and Canberra. Other species have been found on both Lord Howe and Norfolk islands and New Guinea as well. Colourful species as well as rather plain green ones attest to the diversity of Caedicia. Generic revisions will probably remove some species currently residing in Caedicia but we know of at least 30 undescribed species that suggest the genus will remain a large one. The microstructure of the male cercus as well as the number and spacing of teeth on the stridulatory file are all characters that have been little used in the past and need to be incorporated into modern classifications. Those coupled with colour and leg armature will serve to distinguish the species. Caedicia nymphs are usually very different from the adults that they become. They often resemble plant material, such as flowers or floral parts, leaves and even other insects that vertebrate predators avoid. Rearing nymphs and maintaining photographic records are all part of understanding a species and its role in its environment. Most Caedicias are very easily reared on the Orthoptera food mix (see page 205). They do not seem to be cannibalistic.

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A Caedicia nymph.

Caedicia pictipes Stål Bandy-legged Caedicia This is the most important species to the taxonomist because it is the type species of the genus. That is, it is this species to which all other members have to be compared to belong to Caedicia. The type locality is a rather vague ‘Cape York’, although there is no precise locality of that name. Cape York is the regional name of the northern peninsula in Queensland. The type is a female and that adds a bit of uncertainty because females usually have fewer more subtle characters but it clearly has banded legs. With this species, this character is variable. Some individuals from a given locality can have the hind legs entirely dark brown while others can have the distinctive bands of the type, while still others can have the hind tibiae green and lacking any brown colour at all. One specimen that I have seen had the hind tibiae banded on one side and green on the other. The most important taxonomic character to note is the male cercus. In this species it is more robust than most and the tip is blunt and bears a small row of more than two tiny teeth. This species occurs in non-rainforest habits such as eucalypt woodlands and mixed forests bordering on rainforests. It feeds on a variety of shrubs and grasses, and seems especially fond of flowers.

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Caedicia pictipes nymph

Caedicia pictipes (male)

Nymphs of this species (and other species too) often have striped hind legs and some patches on the dorsal surface of the abdomen. This may just break up the form of the insect or may aid in the deception of a bird dropping.

Caedicia marginata Brunner von Wattenwyl Colourful Caedicia This species was described from Kangaroo Island, South Australia. It has been called the ‘Queensland Caedicia’ from time to time but this species may not occur in Queensland. It is related to the type species as well as a complex of other species with the combination of narrow tegmina, colourful

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abdominal tergites and some colour on the pronotum and dorsal margin of the tegmina.

Caedicia araucariae Rentz Araucaria Caedicia This species is known only from Norfolk Island where it lives on windswept bluffs among the leaves and branches of Norfolk Island Pines. Its short and robust form probably aids in stabilising the katydid in its precarious environment. The large white spots, present on several unrelated tettigoniids, probably serve as disruptive colouration and may appear to potential predators are sun spots on leaves.

Caedicia marginata (male)

Caedicia araucariae (female)

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Caedicia gracilis (male)

Caedicia gracilis (male)

Caedicia noctivaga (male)

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Caedicia simplex (male)

Caedicia gracilis Rentz Norfolk Island Caedicia This species is similar to the Araucaria Caedicia but lacks large spots on the tegmina. There may be scattered, small light-coloured spots. The male has slightly different genitalia. It occurs on trees and shrubs (both native and introduced) on Norfolk Island. Like its relative, it is small and robust.

Caedicia noctivaga Rentz Wary Caedicia This species also occurs on Norfolk Island. It is very different from all other known species in the genus because males have the tenth tergite divided. Males sing from perches about two metres from the ground, moving frequently after a bout of singing. In other respects, this is a very plainlooking katydid.

Caedicia simplex (Walker) Garden Katydid This is an interesting, although plain-looking species. It occurs broadly throughout the temperate areas of the east coast. Some specimens identified as C. simplex have been recorded from Perth, Western Australia. It also occurs in New Zealand. The Garden Katydid is a remarkable species. It has been observed in Canberra feeding and singing on Viburnum shrubs in midwinter in the late afternoon at temperatures of 5°C. The call is a simple lisp and is heard emanating from gardens in Sydney and Melbourne day and

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evening. Concealed in foliage, this katydid is difficult to see. It lays its eggs on branches in rows and this may account for part of its distribution. It could easily be transported in nursery stock in the egg stage.

Caedicia simplex (early stage nymph)

Caedicia simplex (final stage nymph)

Caedicia simplex nymphs often attract the attention of gardeners when they find them in their rose flowers. Early instars are often bright red, purple or pink. This gradually fades as they mature and at the final moult they turn the normal plain green colour characteristic of the species.

Caedicia flexuosa Bolîvar Curvy-tailed Caedicia

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This is a slender, grey-green species that occurs in considerable numbers in rainforests habitats from Townsville to New Guinea. It is also common on the Atherton Tableland. Males have relatively long, curvy cerci, hence its name.

Caedicia kuranda Rentz, Su & Ueshima Kuranda Caedicia This species has been recorded in the rainforests around Kuranda where it is frequently attracted to lights. The lustre of both the pronotum and the tegmina contrast. The male cerci are simple and gently upcurved and are apically blunt with a pair of minute teeth. The male subgenital plate has a pair of short, peg-like protuberances. Females have the supra-anal plate divided with a pair of style-like pegs at each corner. The ovipositor is short and upcurved.

Caedicia flexuosa (female)

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Caedicia kuranda (male)

Caedicia webberi Rentz, Su & Ueshima Webber’s Caedicia This is one of the most distinctive species in the genus. The tegmina are covered in small white or creamish-white spots. The abdomen has a series of small white spots on the side. This species is common in rainforests along the coast from Palmerston Gorge north to Lockerbie, Queensland. The male cercus is hammer-shaped and with teeth and brushes apically. Females have a short unarmed ovipositor and a colour pattern identical to that of the male.

Genus Symmachis Brunner von Wattenwyl White-legged Katydids This genus currently comprises three species. The type species, S. lateipennis, is distinctive. The other two may actually represent other genera. Symmachis is distinctive in having the fastigium of the vertex obtuse and touching the frontal fastigium. The tegmen is of uniform width throughout.

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Caedicia webberi (male)

Symmachis lateipennis Brunner von Watttenwyl White-legged Katydid This is a slender, small katydid with ivory legs and a broad stance. It is very easily distinguished. This species has a tropical distribution and is known from mixed eucalypt forests often bordering rainforest, but this katydid does not occur in rainforests.

Symmachis lateipennis (male) Photo: Alan Henderson

Genus Polichne Stål Polichnes

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Eleven species of Polichne are listed in the OSF, with an African species as a doubtful inclusion. At least 20 undescribed species are known throughout Australia and New Guinea. This genus is characterised by its slender appearance, lateral lobes of the pronotum usually longer than deep, the narrow tegmina and the short, bud-like ovipositor. Polichnes are common in grassy habitats. Their long legs serve them well in straddling grasses. Some species appear to be more widespread than many other phaneropterines but this might reflect the poor knowledge of the group and, perhaps, more than one species is represented in these distributions. One of the most outstanding examples of nymphal adaptation is seen in one species of Polichne. The early nymphal stages strongly resemble an ant, but as it matures and grows it becomes larger and the ‘ant cover’ just would not work. As an adult, one would never suspect the association at all.

A first instar Polichne sp. nymph, similar in shape, colour and behaviour to a Green Tree Ant, Oecophylla smaragdina.

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The same specimen a little later in life after a moult or two. Notice the shape and colour changes.

The adult of the same Polichne specimen as above. Without rearing this specimen to maturity, you would never know it was the same species as represented by the first instar.

Polichne parvicauda (male)

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Polichne spinulosa (female)

Polichne angustiloba (male)

Polichne argentata (female)

Polichne parvicauda (Stål) Short-tailed Polichne As presently understood, this species probably has the broadest distribution of any Australian katydid. It seems to occur wherever there are grassy habitats. The body form is grass-like and this probably affords protection in deep grasses. Two colour morphs, brown and green, are known at present in nearly equal numbers. Females glue tiny eggs singly to grass stems.

Polichne angustiloba Brunner von Wattenwyl White-margined Polichne 161

This is a large, easily-recognised species that is uniformly green but with the costal region of the tegmen frosty white. It is widespread in the tropics.

Polichne spinulosa Brunner von Wattenwyl Black-lined Polichne This is a more robust species than most other members of the genus. The dorsal surface of the pronotum bears a pair of parallel black stripes; otherwise the katydid is rich green. This is a very widespread species that is encountered mostly in the tropics on the east coast of the continent.

Polichne argentata Brunner von Wattenwyl Striped Polichne This is an inland species most commonly found in arid regions dominated by Mulga, Acacia aneura. The lower lobe of the pronotum is white and the costal region of the tegmen is white with a mottled black overlay bordering the subcostal vein. The abdomen is striped. Females of this species lay eggs singly on twigs and fly to the ground and retrieve bits of soil with which they coat each egg.

Polichne argentata (female with eggs)

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Spinifex Polichne

Polichne sp. Opalescent Polichne This species is known from inland, eucalypt woodland habitats in the tropics where it lives on shrubs close to the ground. It has been observed feeding on Acacia flowers. The lower margins of the pronotum reflect light in a manner similar to that of an opal. This is very difficult to reproduce in a photograph.

Polichne sp. Spinifex Polichne This species finds protection in the spiny leaves of spinifex, Triodia. This affords it protection from enemies and helps it to cope with the desiccation of the dry habitats in which the host plants occur. After dark, the katydids emerge to feed on the seed heads of the plants, when present.

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Opalescent Polichne

Kurandoptera purpura (male)

Genus Kurandoptera Rentz, Su & Ueshima Purple-winged Katydids Kurandoptera is known from a single, distinctive species that lives in the wet tropical rainforests of north Queensland. It resembles Polichne in its slender shape but differs in many critical aspects. The entire body is highly polished, the MA vein of the tegmen is poorly defined or absent and the ovipositor is unusually large for the size of the katydid.

Kurandoptera purpura Rentz, Su, Ueshima 164

Purple-winged Katydid This is a small species living in the rainforest understorey from Kuranda and the Atherton Tableland, Queensland. Females have a large, sickleshaped ovipositor that is serrated at the tip on both margins. This species is frequently attracted to lights.

Kurandoptera purpura (female)

Genera not assigned to tribe or group Genus Acripeza Guérin-Méneville Mountain Katydids This distinctive genus is one of Australia’s insect icons. It combines a number of characteristics not seen in any other katydid. Both sexes are predominantly black and are strongly aposematic, displaying red, white, blue and orange when threatened. The texture of the male tegmina is flimsy and pliable, whereas, those of the female are hardened and shell-like. This genus shows no close relations with any other.

Acripeza reticulata Guérin-Méneville Mountain Katydid The common name of this katydid is a misnomer. It was first discovered in the southern mountains in relatively large numbers but it is not restricted to montane habitats. It has been found in paddocks near Nyngan, New South

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Wales, and in similar conditions at Moonie, Queensland. It is also known from Herberton in northern Queensland. The Moutain Katydid is often found in large numbers and is frequently seen crossing roads and walking tracks. Early instar nymphs behave much like spiders or ants and display their colours, including the bright orange ‘neck’. Adults are sexually dimorphic. The males are long-winged and capable of flight while the females are incapable of any flight. The colour pattern of the abdomen is similar in both sexes but much more obvious in females. When threatened, droplets of liquid exude from the intersegmental membranes of the abdomen. This species feeds on a variety of plants, including forbs and shrubs. Eggs are laid either glued to twigs on shrubbery or in little pits in the ground where they are cemented into place by saliva from the mother.

Acripeza reticulata (female)

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Acripeza reticulata (male, displaying)

Acripeza reticulata (female, displaying) Photo: Ian Menkins

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Acripeza reticulata (first instar nymph)

Acripeza reticulata (nymph, prior to moulting)

Acripeza reticulata (nymph, after moulting)

Mountain Katydid nymphs are just as peculiar as the adults. The first instar nymph is fast-moving and long-legged relative to the adult. After a moult it becomes black with flecks of blue and white, and the intersegmental membranes become prominent and bright orange.

Genus Elephantodeta Brunner von Wattenwyl Hump-backed Katydids Elephantodeta currently includes two described species but there are a number of other undescribed ones as well. There is a related, undescribed genus with several species with which Elephantodeta could be confused. In

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Elephantodeta the fore coxa is armed with a spine, in the undescribed genus it is not. Hump-backed Katydids could be confused with Tinzeda. They are distinctive in having the hind femur armed with spines on the ventral surface and the pronotum with the lateral lobes as long as deep and with the ventral margin rounded or obtuse, not straight. The ovipositor is widest in the middle with the margins minutely serrate. Elephantodeta species live on or close to the ground in dry habitats. They are often numerous.

Elephantodeta nobilis (Walker) Noble Bush Katydid This is a spectacular species both physically and biologically. Nymphs are often bright yellow, resembling Acacia flowers. Adults are greenish-grey and resemble Acacia leaves.

Elephantodeta nobilis (nymph)

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Elephantodeta nobilis (female)

Elephantodeta sp. (male)

This is a nocturnal species with males flying from shrub to shrub seeking females. The calling song of the male is complex and lengthy. Females have an elongate, laterally flattened ovipositor which is probably used to insert their disk-like eggs into bark cracks.

Elephantodeta sp. Mystery Hump-backed Katydid This katydid is tentatively placed in Elephantodeta because it seems different from other species and also appears not to have a formal name. It

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seems incredible that such a spectacular katydid could have escaped the early taxonomists but such seems to be the case. It occurs in the dry interior of Western Australia.

Elephantodeta sp. 2. (This is another undescribed species that lives on acacias in central Australia.)

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Paracaedicia serrata (nymph)

Paracaedicia serrata (female)

Paracaedicia serrata (nymph) Photo: Jack Hasenpusch

Genus Paracaedicia Brunner von Wattenwyl Paracaedicia comprises 14 species from New Guinea, one of which is also known from north Queensland. The Queensland example is easily recognised.

Paracaedicia serrata Brunner von Wattenwyl Serrated Bush Katydid This species is so-called because of the ventral surface of the hind femur which bears closely spaced, short, stout teeth. This common species is easily recognised by its large size, hump-backed appearance and short, stout ovipositor. This rainforest species readily comes to lights and its loud ‘zip’like call can be heard at night from understorey vegetation.

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Most nymphs of Paracaedicia serrata are green but when they feed on the young leaves of various rainforest trees, they assume the reddish-brown or violet colour of the young leaves. Later when they move on to other plants, they become green.

Genus Tinzeda Walker Tinzedas Tinzeda is a genus of montane and arid Australia. In desert regions, during drought years, these katydids do not appear, but following nurturing rains, when the deserts bloom, they emerge and are quite common. Most species are sexually dimorphic. Males are fully winged and capable of flight and females of some species are flightless. When disturbed, males assume a defensive posture and display the surface of the abdomen which is often colourful. This is not unlike the display of the abdomen by the Mountain Katydid. Seven Tinzeda species are listed in the OSF but there are another 12 or so that remain undescribed. Users should be aware that these identifications are tentative and subject to change as revisionary studies are made. As presently understood, Tinzeda is characterised by the unarmed ventral surface of the hind femur, the lateral lobes of the pronotum being longer than deep, and with the ventral margin being nearly straight. The ovipositor gradually narrows from the base with its margins minutely serrate.

Tinzeda albosignata (Brunner von Wattenwyl) Stout Tinzeda This is a montane species found in the Kosciuszko Range in mid to late summer. It is often seen along roadsides in deep grass. The margin of the tegmen has a thin white stripe bordered on the outside by a much thinner brown stripe. There is a thin creamish-white stripe running for most of the length of the subcostal and radius veins. The surface of the pronotum bears a broad white stripe. The lower margin of the pronotum has a thin white margin. In the Kosciuszko Massif the white stripes on the tegmina are often dull and not as pronounced as they are in the Grampians.

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Tinzeda albosignata (male)

Tinzeda albosignata (female with attached spermatophore)

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Tinzeda soror (female)

Tinzeda soror (Brunner von Wattenwyl) Kosciuszko Tinzeda This species is easily confused with the Stout Tinzeda because the two often occur together in the Australian Alps and Kosciuszko Range. There are subtle differences in the colour pattern and the female, which is longwinged and fully capable of flight. This species is often found in shrubbery.

Tinzeda minor (Tepper) Montane Tinzeda This is not a small species but has ‘minor’ colouration compared to the others. The ventral margin of the pronotum bears only a narrow white margin. The tegmen has a single white stripe along the leading edge but lacks the second stripe present on several of the other species. It is known from montane regions of Victoria and South Australia where it is found on the ground or in low shrubbery.

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Tinzeda minor (male)

Tinzeda minor (female)

Tinzeda sororoides (female)

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Tinzeda sororoides (male)

Tinzeda sororoides (Tepper) Southern Tinzeda This species is common in heath habitats in late spring and summer in southern Australia. The colours are subtle and there is a line of minute spots between the second white stripe and the dorsal margin of the tegmen.

Genus Torbia Walker Torbias Torbia is a widespread genus with at least one species that appears to occur from coast to coast. This may change if the species is studied in detail. The species range from small- to medium-sized katydids to large, robust species. They seem to be associated with eucalypts. The genus is easily recognised by the flat dorsal surface of the pronotum and the broad lateral lobe which is nearly as long as the dorsal length of the pronotum itself. The thoracic sterna are narrow and laterally compressed with the meso- and metathoracic lobes longer than broad. Torbia nymphs are the best known for their mimicry of ants, which they seem to resemble in both appearance and behaviour.

Torbia perficita Walker Giant Torbia This is the largest known species of the genus. It has a relatively broad range, occurring along the east coast from the tropics to Victoria. It is found

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in moist eucalypt and mixed woodland forests where it spends the day high in the trees. It is readily attracted to lights at night.

Torbia perficita (male)

Torbia perficita (female)

Apart from its large size, it is distinctive in having the veins of the tegmina strongly indicated both in structure and emphasised by colour. The posterior margin of the pronotum is well produced. There is a pinkishpurple mark on the thorax that is very prominent.

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Torbia viridissima (female)

Torbia viridissima (Brunner von Wattenwyl) Gumleaf Torbia This species is found from coast to coast in almost all habitats where eucalypts occur. It can be found at high elevations in the Australian Alps or in deserts in central Australia. Like T. perficita, there are marks on the thorax. In addition, the posterior margin of the lateral lobe of the pronotum is produced. The veins of the tegmina are not as well indicated as in that species. Torbia viridissima nymphs resemble ants and always attract attention. As they mature, the nymphs lose the ant-like appearance and use their green colouration as camouflage.

Torbia viridissima (nymph, first instar)

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Torbia viridissima (older nymph)

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Phasmodinae Stick Katydids

The Phasmodinae is endemic to Western Australia where the single genus with three species is relatively common in spring and summer. It is a member of the coastal sand plain heath community. All species are wingless in both sexes and show extraordinary convergence in body form with stick insects of the order Phasmatodea. The prognathous mouthparts, female ovipositor and hopping habit easily distinguish them promptly from stick insects. These katydids feed on flowers and can be of horticultural concern where intact native wildflowers are desired. Petals, anthers, stamens and pollen are eaten. Adults are found in the spring when there is an abundance of flowering vegetation. Phasmodines are nocturnal and can be easily spotted on heath flowers after dark. During the day they may take refuge deep in grasses or within shrubs. Eggs are laid in the ground. Males have the tibial auditory structure open, but faint on the anterior surface and open on the posterior surface. Females have only a faint trace of the tibial auditory tympanum on the posterior surface. All species are silent and produce no sound. Only a single genus is known.

Genus Phasmodes Westwood Stick Katydids The colour of most individuals is a mixture of green and brown. Mouthparts are obviously prognathous to facilitate feeding deep within flowers and for sipping nectar. They have been recorded as feeding on the flowers of Lambertia inermis, Anigozanthus manglesii and Lomandra, as well as grasses in flower such as spinifex (Triodia).

Phasmodes ranatriformis Westwood 182

Kings Park Stick Katydid This species has the broadest range of any known member of the genus. It occurs from the vicinity of Mt Ragged, along the coast and inland in appropriate heath habitats to Cockleshell Gully, north of Perth, Western Australia. At the latter locality it is sympatric with P. jeeba and some 35 km north-west it occurs with P. nungeroo. The colour of this species is uniformly green and only rarely straw-brown. The male cercus is strongly incurved and without a prominent internal flange. The female subgenital plate has an apical notch.

A Phasmodes species feeding.

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Phasmodes ranatriformis (male)

Phasmodes ranatriformis (male)

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Phasmodes nungeroo (male)

Phasmodes nungeroo Rentz Kalbarri Stick Katydid This species occurs along the coast from Kalbarri south to the Eneabba area, Western Australia. Inland populations have been discovered at Coorow, Western Australia. It is either green or grey-brown with a streaky appearance, especially in the southern portion of its range. Green and brown forms predominate in the Kalbarri region. This species seems to prefer tall and reedy grasses as well as Spinifex (Triodia). It has been observed feeding on grass seeds as well as pollen. P. nungeroo is distinguished from all other species by the following combination of characters: grey or greybrown colour, elongate male cercus with a less prominent internal flange and the female with an apical notch at the tip of the subgenital plate.

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Key to subfamilies of Australian katydids

Key to subfamilies 1a Head strongly projecting (prognathous); body form ‘stick-insectlike’ 2 1b Head not projecting; body form variable but not stick-like 3 2a Both sexes wingless; body form very stick-like and slender (Stick Katydids) Phasmodinae 2b Body form less stick-like; both sexes winged or female wingless (Pollen & Nectar-feeding Katydids) Zaprochilinae 3a Head rounded, not usually slanted or frontally flattened; fore tibia in section square, especially apically (Fig. 1); prosternum unarmed; ovipositor variable in size and shape but usually sickle-like (Figs. 2,3) (Bush Katydids) Phaneropterinae 3b Without above combination of characters 4 4a Pronotum large, hind margin acute, lateral margins dentate (Fig. 4); males lacking typical stridulatory region of tegmina (Hooded Katydids) Phyllophorinae 4b Pronotum not as above; males with stridulatory region on dorsal surface of tegmen 5 5a Antennal sockets strongly rimmed (Fig. 5), especially on internal margins; thoracic auditory spiracle small (Fig. 6), inconspicuous, not concealed by pronotum (True Katydids) Pseudophyllinae

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5b Antennal sockets not strongly rimmed (Fig. 7); thoracic auditory spiracle large, elongate, often partially concealed by pronotum (Fig. 8) 6

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6a Tibial auditory tympanum usually open (Fig. 1); if closed on one or both sides, then slit is directed laterally and the opening is broad and the margins curving (Fig. 9); prosternum armed with a pair of spike-like processes (as in Fig. 13) (Long-legged Katydids) Mecopodinae 6b Tibial auditory structure either open or closed, the slit distinctly directed dorsally relative to tibia (Fig. 11); if open then the opening of nearly uniform width (Fig. 12); prosternum either armed or unarmed. 7 7a Prosternum unarmed; tibial auditory structure usually open (Fig. 12); small, delicate, highly vagile arboreal and epiphyllic species, greenish or greenish-yellow (Leaf Marauding Katydids) Meconematinae 7b Prosternum armed or unarmed; other characters not as above 8 8a Prosternum unarmed; greenish, striped, brachypterous species with the tegmina mostly concealed under the pronotum; tibial auditory structure slit-like on both sides; size minute, males of some species 5–8 mm in total body length (Micro Katydids) Microtettigoniinae 8b Without above combination of characters (see Figs. 13, 14) 9

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9a Fore tibia bearing a single apical spur on posterior margin of dorsal surface (Fig. 15); fastigium of vertex as broad as width of first antennal segment to half its width (Figs 16, 17); hind tarsus with a plantula (Fig. 18) which, in most species, is at least half the length of the basal segment (Shield-backed Katydids) Tettigoniinae 9b Not possessing at least two of the above characters 10

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10a Hind tibia without apical spurs on dorsal surface; posterior portion of lateral lobe of pronotum produced or not produced (Sluggish Katydids) Austrosaginae 10b Hind tibia with at least an external apical spur on dorsal surface, if not, then posterior portion of lateral lobe of pronotum produced 11 11a Frons vertical; hind tibia not bearing an internal apical spur on dorsal surface (Fig. 20); if not, then posterior lobe of pronotum produced (Fig. 19); meso- and metasternum not spiniform; sexually dimorphic, males winged (one species male micropterous), females wingless (Balloon-winged Katydids) Tympanophorinae 11b Without above combination of characters 12

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12a Fore tibia usually bearing five to seven long, moveable, outwardly bowed, opposing spines, the longest of which in many species is as long as or longer than the combined lengths of the first two tarsal segments; fastigium of vertex (Fig. 21) narrow, strongly laterally compressed, its greatest width less than that of first antennal segment in most species and projected above same and usually sulcate 13 12b Fore tibia of most species with spines not unusually lengthened and not as long as the combined lengths of the first two tarsal segments. Fastigium of vertex variable in width (Figs. 22, 23) and not sulcate Conocephalinae (see p. 84 for key to tribes) 13a Male subgenital plate constricted in the middle; styles very elongate, bowed outwards; Hexacentrus sexually dimorphic for tegmina shape; ovipositor ensiform (Hexacentrus) or falcate (Alison) (Fierce Predatory Katydids) Hexacentrinae 13b Male subgenital plate normal, scoop-shaped, not constricted in the middle; styles short, stout, usually parallel (Predatory Katydids) Listroscelidinae

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Glossary apical at or near the apex (end) of any structure aposematic colouration warning colouration apterous completely wingless body length length from the top of the head to the end of the abdomen, excluding wings and cerci brachypterous with tegmina and wings shorter than the abdomen but overlapping or touching each other dorsally carina a ridge (plural: carinae) cercus a paired structure, one on each side of end of the abdomen at the base of supra-anal plate and paraprocts, variably sized and shaped, and often species distinctive cline a gradual or nearly continuous change of a characteristic over a geographical region clypeus facial sclerite between the frons and the labrum coastal sand plain used here generally for the region just north of Kalbarri, Western Australia, south to the Cape Arid region concolorous of the same colour coxa basal segment of the leg, by which the leg is attached to the body diapause temporary stoppage of development dimorphism difference in appearance of individuals of the same species, particularly between males and females endemic restricted to a given geographical region

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epiphallus the titillator, the usually sclerotised portion of the male phallus episternum the small, triangular lobe protruding under the lateral lobe of the pronotum facultatively parthenogenetic eggs developing with or without fertilisation falcate sickle-shaped fastigium of vertex anterior part of the vertex, projecting anteriorly between the eyes femur the third and usually the stoutest part of the leg from which the tibia is attached frons anterior portion of the face that merges above with the fastigium of the vertex below with the clypeus on sides with eyes and facial carinae Gondwanan pertaining to the southern super-continent consisting of continental blocks of South America, Africa, Madagascar, India, Antarctica and Australia Holarctic the faunal region combining the Nearctic and Palaearctic regions intercercal between the cerci interocular between the eyes labrum upper lip of the mouthpart lateral at or from the side lobiform shaped like a lobe macropterous with tegmina and wings fully developed, exceeding or at least reaching the end of the abdomen, and overlapping dorsally mesal pertaining to the middle

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mesosternum ventral portion of the mesothorax mesothorax the second thoracic segment metanotum dorsal surface of the third thoracic segment mesopleura pleuron of the mesothorax mesopterous in between short- and long-winged; wings not reaching the end of the abdomen. metathorax the third thoracic segment micropterous with greatly reduced tegmina and wings; tegmina may overlap micropyles a pore or pores on the surface of the egg through which a spermatozoon can enter morphs individual variations mulga either Acacia aneura or a vegetation type dominated by that species Nearctic the ecozone covering most of North America including Greenland and the highlands of Mexico nocturnal active at night nymph the juvenile stage of hemimetabolous insects occiput posterior part of the head merging anteriorly with the vertex and usually not having a definite line of separation ocelli three small simple eyes, one in the middle of frontal ridge and two near the sides of the compound eyes omnivore eating many kinds of foods orthopteran of or pertaining to the order Orthoptera; often used broadly to include also cockroaches, mantids and stick insects

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orthopteroids insects of the orders Orthoptera, Mantodea, Phasmida, Blattodea ovipositor the egg-laying structure protruding from the abdomen of the female Palaearctic one of eight ecozones dividing the earth’s surface; the Palaearctic is the largest including Europe, Asia, north of the Himalayas, northern Africa and the northern and central portions of the Arabian Peninsula paraprocts a pair of lateral lobes representing parts of the eleventh tergite and located on the sides of the anus partly below or alongside the supra-anal plate parthenogenesis egg development without fertilisation plantula a pair of pads at the base of the first segment of the hind tarsus; variously modified but characteristic of some groups such as the Tettigoniinae where it is developed as a stabilising prop when living in tall grasses or as an adjunct to jumping prognathous with strongly projecting mouthparts pronotal carinae ridges positioned on the surface of the pronotum. pronotum the dorsal shield of the thorax pronotum, lateral lobes the downward-projecting lateral portions of the pronotum pronotum, metazoan part of pronotum posterior to the posterior groove or sulcus prosternum the narrow region between the first pair of legs prothorax the first segment of the thorax proximal the part of an appendage which is nearest to the body, as opposed to distal

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pseudocercus extensions of the paraprocts spermatheca the receptacle of the sperm spermatophore a sticky package containing sperm that is transferred from the male to the female during copulation spermatophylax the highly nutritious portion of the sperm package provided by the male during mating and eaten by the female spines of hind tibia two rows of rigid spines on the sides of the tibia spiniform spine-like spiracle an external opening of the tracheal system spurs larger than spines and articulated, often curved apically, usually one or two pairs at the apex of the hind tibia sternum, sternites the ventral part of the thorax; usage, pro-, meso- and metasternum stridulation the process of producing sound by a file and scraper method stridulatory file the sound-producing structure on the underside of the male left tegmen in Tettigoniidae subgenital plate the last, scoop-shaped segment of the underside of the abdomen sulcus a groove, suture or furrow supra-anal plate the last abdominal tergite covering the anus from above sympatric the occurrence of two or more populations or species in the same area synonym one of two or more scientific names of the same rank to denote the same taxon tarsus the five-segmented distal part of the leg

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tibia the part of the leg between the femur and the tarsus titillators processes at the end of the aedeagus, often above the dorsal phallic lobes tribe subdivision of a subfamily containing one or more genera, the name of which ends in ‘ini’ trochanter a small sclerite between the coxa and the femur truncate squared off at the end unsclerotised membranous, not hardened like the exoskeleton ventriloquial having the ability to produce a call as if coming from some place other than the source vertex the upper part of the head merging in front with the fastigium of vertex and behind with the occiput. wallum an ecosystem in south-east Queensland and northern New South Wales characterised by a floristically rich component on deep, acidic, sandy soils, often dominated by Wallum Banksia, Banksia aemula; a high water table may create swamps from time to time

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References Bailey W J (1979). A review of Australian Copiphorini (Orthoptera: Tettigoniidae: Conocephalinae). Australian Journal of Zoology 27, 1015– 1049. Bailey WJ and Rentz DCF (Eds.) (1990). The Tettigoniidae: Biology, Systematics and Evolution. Crawford House, Bathurst & Springer-Verlag, London. Brunner von Wattenwyl, C (1891). Additimenta zur Monographie der Phaneropteriden. Verhandlungen Zoologisch-botanischen Gesellschaft 412, 1–196. Eades DC and Otte D (2009). Orthoptera Species File Online. Version 2.0/3.3 http://Orthoptera.SpeciesFile.org Gwynne DT (2001). Katydids and Bush-crickets: Reproductive Behavior and Evolution of the Tettigoniidae. Cornell University Press, Ithaca, NY. Gwynne DT, Yeoh P and Schratral A (1988). The singing insects of King’s Park and Perth gardens. Western Australian Naturalist 17, 25–81. Ingrisch S (1998). Monograph of the Oriental Agraeciini (Insecta, Ensifera, Tettigoniidae): Taxonomic revision, phylogeny, biogeography, stridulation and development. Courier Forschungsinstitut Senckenberg 206, 1–391. McCoy F (1886). Natural History of Victoria. Prodromus of the Zoology of Victoria, or figures and descriptions of the living species of all classes of the Victorian indigenous Animals XII, 1–375.

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Nasrecki P and Otte D (1999). An illustrated catalog of Orthoptera, Vol. 1, Tettigonioidea (CD-ROM). Orthopterists’ Society at the Academy of Natural Sciences of Philadelphia, Publications on Orthoptera Diversity, Philadelphia. Nickle DA and Naskrecki P (1997). Recent developments in the systematics of Tettigoniidae and Gryllidae. In The Bionomics of Grasshoppers, Katydids and Their Kin. (Eds. SK Gangwere, MC Muralirangan and M Muralirangan) pp. 41–58. CAB International, Wallingford, UK. Pitkin L (1980). A revision of the Pacific species of the genus Conocephalus Thunberg (Orthoptera: Tettigoniidae). Bulletin of the British Museum (Natural History) 41, 315–455. Redtenbacher J (1891). Monographie der Conocephaliden. Verhandlungen Zoologisch-botanischen Gesellschaft 41, 315–562. Rentz DCF (1985 ). A monograph of the Tettigoniidae of Australia. Vol.1: The Tettigoniinae. With an appendix by DH Colless. CSIRO Australia/Brill, Melbourne. Rentz DCF (1993a). A Monograph of the Tettigoniidae of Australia. Vol. 2: The Phasmodinae, Zaprochilinae and Austrosaginae. With appendices by N Ueshima and DH Colless. CSIRO Australia, Melbourne. Rentz DCF (1993b). Tettigoniidae of Australia. Disk 1. The Austrosaginae, Zaprochilinae and Phasmodinae. Audio CD. CSIRO Australia, Melbourne. Rentz DCF (1993c). Orthopteroid insects in threatened habitats in Australia. In Perspectives of Insect Conservation. (Eds. KJ Gaston, TR New and MJ Samways) Ch. 7, pp. 125–138. Intercept, Hampshire. Rentz DCF (1996a). Grasshopper Country: Australia’s Abundant Orthopteroid Insects. New South Wales University Press, Sydney.

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Rentz DCF (1996b). Grasshopper Country: Sounds of Australian Orthopteroid Insects. Audio CD. CSIRO Publishing, Melbourne. Rentz DCF (2001). A Monograph of the Tettigoniidae of Australia. Volume 3. The Listroscelidinae, Tympanophorinae, Meconematinae and Microtettigoniinae. With Chapters by DH Colless and N Ueshima. CSIRO Publishing, Melbourne. Rentz DCF and Colless D (1990) A classifi cation of the shield-backed katydids (Tettigoniinae) of the world. In The Tettigoniidae: Biology, Systematics and Evolution. (Eds. WL Bailey and DCF Rentz). Crawford House Press, Bathurst; Springer, Berlin. Rentz DCF, Su YN and Ueshima, N (2006). Studies in Australian Tettigoniidae: The Mecopodine Katydids Part 2 (Orthoptera: Tettigoniidae; Mecopodinae; Sexavaini) Queensland Palm Katydid. Transactions of the American Entomological Society, 132, 229–241. Rentz DCF, Su YN and Ueshima N (2008). Studies in Australian Tettigoniidae: New Phaneropterine katydids from Queensland rainforests. Zootaxa 1964, 1–39. Riley CV (1874). ‘Katydids. Report of the noxious, beneficial, and other insects of the state of Missouri’. pp. 150–169. State Entomologist Office, Missouri. Webber BL, Rentz DCF, Ueshima N and Woodrow IE (2003). Leucopodoptera eumundii, a new genus and species of katydid from the tropical rainforests of north Queensland, Australia (Orthoptera: Tettigoniidae; Phaneropterinae; Holochlorini). Journal of Orthoptera Research 12 (1), 79–88.

Orthoptera food mix 6 Parts of muesli 2 parts of flake type tropical fish food

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1 part of grass seed a small grained grass free from pesticides!) 1 part of pollen available from health food stores)

Pampel’s fixative 2-4 Parts of glacial acetic acid 15 parts of 95% ethanol 30 parts of distilled water 6 parts of formalin formaldehyde in water)

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Appendix 1: Keeping katydids alive and preservation of specimens Provide a source of water for any katydids that you want to keep alive. Rolled dental cotton, available from dental supply houses, can be propped into a small vial or elongate, cylindrical jar, such as a pill bottle. Water will be absorbed into the cotton and lapped off the cotton by the katydid. Cereal mites also find the water an ideal breeding site so check regularly and change the cotton if necessary. Most species will accept a simple ‘Orthoptera food mix’ (see page 205). Only the few obligate predators will reject it. Use only a small amount of food because cereal mites will eventually find it and breed in excessive numbers and foul the jars and kill moulting katydids. Food should be changed regularly and the jars washed thoroughly from time to time to reduce problems caused by the mites. The main component of the mixture is natural (not toasted) muesli. If the bits are too large, the mixture should be slightly crushed by hand or for a short time in a blender. You want the particles to be small enough for small nymphs to be able to deal with them. The mixture should be kept dry and ideally stored in a refrigerator to prevent infestation from cereal mites. If it does eventually become infested, the mixture should be discarded. Feed only a small amount. Less than half a teaspoon should be scattered on the bottom of the container or provided in a small feeding dish (a bottle cap is often useful). Dry dog food is theoretically useful but it may be infested with mites, cereal moths or beetles. Special foods mixed specifically for crickets that are used by fishermen may also be used but quality control may not be very high and the mixtures may be contaminated. Best to mix it yourself.

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Kill the specimen by freezing or in a killing bottle. After removing the body contents of a katydid you want to preserve for your collection, dust into the body cavity a small amount of dry stuffing powder made from mixing one part of talcum powder with one part of boric acid powder. This is a safe mix and the boric acid powder can be obtained from a chemist. The talcum helps to dry the specimens quickly and the boric acid powder tends to reduce bacterial action. The key to good colour preservation is rapid drying of the specimens. Placing them in an oven will ‘cook’ them and discolour them drastically. Treated as above and left on a setting board in a dry room will preserve most of the colour.

Preservatives and fixatives Preservation of specimens in liquid is important if you want to conveniently examine legs and genitalic structures without destroying them. Placing the specimens in 75–80 per cent ethyl alcohol (ethanol) provides very acceptable results. Better preservation can be gained by fixing the specimens first in Pampel’s Fixative (see page 205) and then transferring them to 75 per cent ethanol for permanent storage. You may encounter some difficulty in obtaining some of these chemicals. A high school or university chemistry department may assist.

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Appendix 2: Special interest groups and entomological supplies There are a number of organisations that can be of help to people who want to learn more about Orthoptera and insects in general. The Orthopterists’ Society is an international organisation of more than 300 members that was founded in 1978. People with an interest in systematics, biology, behaviour, ecology, biogeography, genetics, morphology and control belong to this society. Meetings are held every four years at venues around the world. The Journal of Orthoptera Research is published annually and contains articles on a variety of subjects in the fields noted above. Metaleptea is the society’s newsletter and contains news of interest to members including book reviews, notices, reports and short notes on orthopteroid ecology, systematics and orthopteroid biology. Website: http://140.247.119.138/OS_Homepage/ provides current news and recent books as well as a bulletin board and members database. The Orthoptera Species Files (OSF) (http://orthoptera.speciesfile.org/HomePage.aspx) is a catalogue of all Orthoptera species and is an essential tool for taxonomists. It is kept up to date on a regular basis and should be consulted by all those who need to determine the current status of any scientific name. There are a number of entomological societies within Australia that cater to professional and amateurs alike. Papers and notes of interest to students of Orthoptera appear from time to time. The Australian Entomological Society is the largest organisation of Australian entomologists. The Society has two publications. The Journal of the Australian Entomological Society publishes articles on a quarterly basis in a number of different fields. Myrmecia is the newsletter of the society and is issued quarterly. It publishes executive reports, minutes, regional news, notices of events and short notes of interest. Other society activities include annual meetings in various capital cities, awards to students and professionals and competitions.

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For additional information on the Australian Entomological Society check the website: http://www.agric.nsw.gov.au/Hort/ascu/myrmecia/society.htm New South Wales, Queensland and Victoria have regional entomological societies and produce publications and hold meetings. The Society for Insect Studies is an entomological society in New South Wales for amateurs and professionals. Membership is open to any person with an interest in entomology. It operates in the Sydney area and holds meetings every other month in the Australian Museum. Website: http://miller.emu.id.au/pmiller/books/stickinsects/group/insect.studies/index.html The Australian Entomologist is a quarterly journal published by the Entomological Society of Queensland. This is an illustrated magazine devoted to entomology in the Australian region, including New Zealand, Papua New Guinea and the south Pacific islands. Website: www.esq.org.au/ Australian Entomological Supplies has provided entomology supplies, equipment and books to both the entomology professional and amateur entomologist for over 35 years. Website: www.entosupplies.com.au/

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