Bees of Australia: A Photographic Exploration

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BEES of AUST R ALIA

© James Dorey 2018 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. A catalogue record for this book is available from the National Library of Australia. Published by: CSIRO Publishing Locked Bag 10 Clayton South VIC 3169 Australia Telephone: +61 3 9545 8400 Email: [email protected] Website: www.publish.csiro.au Front cover: The face of a male Megachile apicata from Queensland. Collected by Olivia K. Davies. Title page: A female Lasioglossum sp. feeding from flowers in Victoria. Back cover: (left to right) side view of a female Leioproctus amabilis; dorsal view of a male Leioproctus amabilis; male Homolytictus dampieri. Photographs are by the author Set in 11/15 Adobe Garamond Pro Edited by Peter Storer Cover design by James Kelly Typeset by Desktop Concepts Pty Ltd, Melbourne Printed in China by 1010 Printing International Ltd. CSIRO Publishing publishes and distributes scientific, technical and health science books, magazines and journals from Australia to a worldwide audience and conducts these activities autonomously from the research activities of the Commonwealth Scientific and Industrial Research Organisation (CSIRO). The views expressed in this publication are those of the author(s) and do not necessarily represent those of, and should not be attributed to, the publisher or CSIRO. The copyright owner shall not be liable for technical or other errors or omissions contained herein. The reader/user accepts all risks and responsibility for losses, damages, costs and other consequences resulting directly or indirectly from using this information. Original print edition: The paper this book is printed on is in accordance with the standards of the Forest Stewardship Council ®. The FSC ® promotes environmentally responsible, socially beneficial and economically viable management of the world’s forests.

BEES of AUST R ALIA A P H OTO G R A P H I C E X P L O R AT I O N

JA M E S DOR EY

iv

Foreword Bees are viewed widely as beneficial. Humanity has long benefited from the domestication of a few species, most notably the European honeybee, Apis mellifera. These benefits include the production of honey and wax, as well as the pollination of crops and other plants. Their intricate behaviours, especially those associated with social existence, have presented stimulating models for evolutionary analysis, as well as impressive – often fanciful – templates for the imaginations of science fiction writers. Yet, few people appreciate the true variety and biological diversity of bees demonstrated so vividly in this book. Australia harbours numerous species of native bees, many of them inconspicuous to the casual observer, and most of them found nowhere else in the world. As with other groups of insects in Australia, early taxonomists allocated many newly discovered species to Northern Hemisphere genera with which they were familiar, and the high levels of endemism and independent evolution have only become recognised as our fauna is explored in more detail. The uniqueness of our bee fauna is itself a powerful motive for conservation, but the wellbeing of bees as predominant pollinators is also integral to the conservation of much of Australia’s flora. Bees can be affected by the loss of native flora and habitats and by the introduction of alien species, pesticides and other influences. Global declines of pollinators are a major concern and their wide ramifications are difficult to exaggerate. Tarlten Rayment commented in his A Cluster of Bees (1935, Endeavour Press, Sydney) – an enduring classic of Australia’s natural history literature – that bees are perhaps the most useful of all insect groups to humanity. Their wellbeing can be assured only through informed awareness, and any means that can increase this recognition is extremely welcome. Ways of attracting young people to wider interests in natural history (and redressing what is sometimes referred to as ‘the extinction of experience’) are extremely welcome. This book is a valuable contribution to that effort. James Dorey writes with knowledge and an infectious passion, and the wealth of information encompassed in his notes on each of the selection of bee species treated here will surely help to stimulate wider and enduring interest. The book is also a vehicle for displaying the author’s remarkable photographs: examples that demonstrate so impressively the beauty, wonder and structural variety of Australia’s bees. It also provides a useful ‘foothold’ for readers to start observing, collecting (with due regard to any permits needed), photographing and recording their findings on behaviour, abundance, which flowers are visited or ignored, and so on. It is easy to obtain original information – indeed, it is often difficult to find anything that has been recorded previously for many of our insect species! Novelties may include finding unusual species in a home garden, where bees can be encouraged by supplying suitable nectar sources

and nest sites. The general essays in this book focus on many topics related to this and are an important part of this volume. Some provide very practical advice on studying bees. One important lesson, perhaps a caution, is that, although the fine illustrations cover a representative selection of bees that may be encountered, a far larger number are not included. It is sure that any persistent observer will find additional species, some closely resembling those illustrated, but others clearly different. Some may never have been seen before, especially those from the more remote parts of Australia. Our ability to interpret bee diversity and patterns of species distribution and abundance depend on continuing documentation, and the availability of collected specimens for critical examination by specialists. Readers of this welcome book can easily become contributors to this endeavour. In short, bees are a significant focal group of insects. They are popular and appreciated, attractive in appearance and readily observed, as most are active by day – they are ideal for ‘citizen science’ projects or for individual interests. I hope that this book will foster these activities and lead to increased understanding of ‘how bees work’ in Australia. T. R. New Department of Ecology, Environment and Evolution, La Trobe University, Melbourne, Victoria 3086, Australia Email: [email protected]

Contents Foreword iv Acknowledgements xii Image captions

xiv

Introduction 1 New South Wales

4

Amegilla (Asaropoda) bombiformis

6

Megachile ustulata

8

Amegilla (Zonamegilla) asserta

10

Undescribed Leioproctus (Exleycolletes) sp.

12

Lasioglossum (Parasphecodes) lithuscum

14

Hylaeus (Macrohylaeus) alcyoneus

16

Lasioglossum (Chilalictus) lanarium

18

Leioproctus (Exleycolletes) cristatus

20

Exoneura sp.

22

Thyreus nitidulus

24

Why are bees important?

26

James Dorey

Queensland 30

viii

Austroplebeia australis

32

Palaeorhiza (Cnemidorhiza) disrupta

34

Megachile abdominale

36

Braunsapis sp.

38

Hylaeus (Euprosopoides) ruficeps

40

Euryglossina (Microdontura) mellea

42

Megachile apicata

44

Bees of Australia

Tetragonula carbonaria

46

Hylaeus (Gnathoprosopis) albonitens

48

Megachile aurifrons

50

Nomia (Hoplonomia) rubroviridis

52

Australian native bees as crop pollinators

54

Tobias J. Smith

Victoria 58 Homalictus punctatus

60

Leioproctus (Leioproctus) plumosus

62

Hylaeus (Gnathoprosopoides) philoleucus

64

Lasioglossum (Chilalictus) veronicae

66

Lasioglossum (Chilalictus) sp.

68

Amphylaeus (Amphylaeus) morosus

70

Pachyprosopis (Pachyprosopis) haematostoma

72

Nomia (Paulynomia) aurantifer

74

Euryglossina (Euryglossina) hypochroma

76

Apis mellifera

78

Threats to our bees

80

Matt Elmer

Western Australia

84

Amegilla (Notomegilla) chlorocyanea

86

Hylaeus (Euprosopis) husela

88

Homalictus dampieri

90

Xylocopa (Koptortosoma) parvula

92

Callohesma flavopicta

94

Austroplebeia essingtoni

96

Exoneura sp.

98

Hylaeus (Euprosopoides) obtusatus

100

Thyreus waroonensis

102

Social behaviour of bees

104

Tim Heard Contents

ix

South Australia

108

Brachyhesma houstoni

110

Homalictus urbanus

112

New Exoneura sp.

114

Hylaeus (Euprosopis) honestus

116

Lasioglossum (Callalictus) callomelittinum

118

Euryglossa adelaidae

120

Pachyprosopis (Pachyprosopula) kellyi

122

Brachyhesma sp.

124

Lipotriches (Austronomia) australica

126

Exoneura sp.

128

How to find native bees

130

James Dorey

Tasmania 134 Heterohesma clypeata

136

Hylaeus (Prosopisteron) perhumilis

138

Paracolletes (Paracolletes) crassipes

140

Leioproctus (Leioproctus) amabilis

142

Exoneura (Inquilina) sp.

144

Megachile (Eutricharaea) maculariformis

146

Hylaeus (Prosopisteron) quadratus

148

Bombus terrestris

150

How to attract native bees to your garden

152

Megan Halcroft

Northern Territory

x

156

Undescribed Amegilla (Asaropoda) sp.

158

Brachyhesma perlutea

160

Braunsapis sp.

162

Hylaeus (Rhodohylaeus) maiellus

164

Hylaeus (Euprosopis) elegans

166

Bees of Australia

Lasioglossum (Chilalictus) ochroma

168

Lipotriches (Austronomia) sp.

170

Meroglossa torrida

172

Xanthesma (Xanthesma) flava

174

Museums 176 Ctenocolletes smaragdinus

178

Megachile (Schizomegachile) monstrosa

180

Xylocopa (Koptortosoma) aruana

182

Palaeorhiza varicolor

184

Quasihesma gigantica

186

Lasioglossum (Chilalictus) hemichalceum

188

Amegilla (Asaropoda) dawsoni

190

Xylocopa (Lestis) aeratus

192

Hyleoides zonalis

194

The importance of museums

196

Michael Batley

Glossary 200 Appendix of species by family

203

Further reading

204

Index 205

Contents

xi

Acknowledgements There are many people that I must both acknowledge and thank profusely for helping me to make this book. Truly, it would never have been started, let alone finished, without the help of so many extremely generous people along the way. The first thanks must go to my ecology friends, Gergana Daskalova, Matt Elmer, Christina Elmer and Amelia Carlson, who encouraged me past the ‘I should do that’ phase into the ‘I can do that’ phase of the project. I would additionally like to thank Jeremy Whitehead, Tas Jouir and the many other friends who kept me going through the many hours, weeks and months of photo editing that were required to produce so many images of our native bees. I also need to thank my parents, not just for instilling in me a love of nature and encouraging me in any endeavour that I chose to undertake, but also for supporting me financially while I was driving around Australia for three and a half months collecting bees. On a similar note, I would like to thank Ron Dorey, my uncle, for giving me his was-to-be paddock basher van that took me around the country and that I named ‘Ron’. I would also like to thank him for the misunderstandings that I now suffer when anyone in my family asks me ‘How is Ron going?’ and the confusion that they suffer when I reply ‘He is in the shop’. I would also like to thank the collection managers, staff and academics at many Australian institutions who gave me their time to discuss my project face to face and show me around their collections. These lovely people included: Nikolai Tatarnic and Terry Houston at the Western Australian Museum; Mark Stevens and Peter Hudson at the South Australian Museum; Michael Schwarz at Flinders University; Ken Walker of Museums Victoria; Michael Batley at the Australian Museum; and Susan Wright and Geoff Thompson from the Queensland Museum. Michael Schwarz, Ken Walker and Terry Houston, in particular, took time out of their busy schedules to sit and impart their knowledge of bees to me, for which I am very grateful. Without the overwhelming support and encouragement of these people, I would never have passed the ‘I can do that’ phase of the project to actually completing the project, nor would I have a museums section to this book. Most of all, I need to thank all the contributors who gave up a substantial amount of time, energy and thought to help me make this book what it is. I could never have compiled the knowledge and expertise that is included in this book without the help of Michael Schwarz of Flinders University, Tim Heard of Sugarbag Bees, Tobias Smith of Bee Aware Brisbane, Megan Halcroft of Bees Business, Michael Batley from the Australian Museum, Matt Elmer of the University of Queensland and Remko Leijs of the South Australian Museum. It was always a goal of this book to bring together knowledge from a broad range of experts. Without the encouragement and dedication of these people, that would have been impossible. Contributions to species, subgenus, genus and state stories are listed below, while feature pages are credited in the text. In several cases, multiple authors contributed to a single description. Michael Schwarz: Amphylaeus (Amphylaeus) morosus, Braunsapis sp. (Qld); Braunsapis sp. (NT); Exoneura (Inquilina) sp., Exoneura sp. (SA); Homalictus punctatus, Hylaeus (Gnathoprosopoides) philoleucus, Hyleoides sp. xii

Bees of Australia

(Museums); Lasioglossum (Chilalictus) lanarium, Leioproctus (Exleycolletes) cristatus, Leioproctus (Leioproctus) amabilis, Lipotriches (Austronomia) australica, Lipotriches (Austronomia) sp., Meroglossa torrida; New Exoneura sp., Thyreus nitidulus, Thyreus waroonensis, Xanthesma (Xanthesma) flava, Xylocopa (Lestis) aeratus and Xylocopa (Koptortosoma) aruana. Michael also contributed to the state stories for South Australia, Victoria, Queensland and part of New South Wales. Michael Batley: Amegilla (Asaropoda) bombiformis, Amegilla (Notomegilla) chlorocyanea, Amegilla (Zonamegilla) asserta, Braunsapis sp. (Qld); Exoneura (Inquilina) sp., Exoneura sp. (WA); Hylaeus (Euprosopis) honestus, Hylaeus (Euprosopoides) obtusatus, Hylaeus (Euprosopoides) ruficeps, Hylaeus (Gnathoprosopis) albonitens, Hylaeus (Prosopisteron) quadratus, Hylaeus (Rhodohylaeus) maiellus, Lasioglossum (Chilalictus) lanarium, Lasioglossum (Chilalictus) sp. (Vic); Palaeorhiza varicolor, Paracolletes (Paracolletes) crassipes; Undescribed: Amegilla (Asaropoda) sp. and Megachile (Schizomegachile) monstrosa. Most identifications can also be attributed to Michael. Remko Leijs: Undescribed Amegilla (Asaropoda) sp. I would also like to thank Sarah Manning for proofreading and providing suggestions. Finally, I would like to thank the staff of CSIRO Publishing, who have not only given me the opportunity to publish, but helped me almost every step of the way. I would particularly like to thank Lauren Webb, who encouraged me, kept me on track and inspired me to get the book finished and include everything that should be included.

Acknowledgements

xiii

Image captions Pages iv–v: A female neon cuckoo bee roosting for the night on a twig in a northern New South Wales rainforest. Pages vi–vii: A tiny Euryglossinae sp. catching onto a stamen of a Melaleuca flower and appearing to wave ‘hello!’ Pages 4–5: The stunning cloud forests of the Gibraltar Ranges, New South Wales. Pages 26–27: A female Lasioglossum sp. feeding from flowers in Victoria. Pages 28–29: A female blue-banded bee (Amegilla (Notomegilla) chlorocyanea) coming in to feed from a flower in arid Western Australia. Pages 30–31: One of many long, straight and red roads running through dry eucalypt-dominated forests in outback Queensland. A great habitat for many bee species. Pages 54–55: A Tetragonula carbonaria queen and her court in their hive. Sealed and finished brood cells are evident next to their open and unfinished neighbours along the expanding front of the spiralling and tiered brood comb. Pages 56–57: Readying herself to fly, a Tetragonula carbonaria worker stands on the resinous lip of her nest entrance in a graveyard in Brisbane, Queensland. Pages 58–59: Lake Elizabeth in the Great Otway National Park. The lake was formed in the 1950s when a landslide dammed the valley and drowned the large trees that are now just slowly sinking giants. Lake Elizabeth now provides a home for many water-loving animals and plants, with platypus a common sight in the early mornings and evenings. Pages 80–81: Many sit-and-wait predators take advantage of flowers to find their next meal. That is just what this lynx spider has done, enjoying a Hylaeinae bee that has come to collect pollen and nectar for her young. While quite graphic, this is a natural threat to our native bees and one that should normally not threaten their existence. Pages 82–83: This European honeybee (Apis mellifera) is an introduced bee species which competes with native bees, birds, mammals and other insects for pollen, nectar and nesting places. Our positive relationship with these animals for crop pollination, honey, beeswax and more is at odds with the relationship that they have with our native flora and fauna. Pages 84–85: Western Australian boab trees dot the landscape around the Kimberley region, towering over the other scattered and otherwise low vegetation and grasslands; a testament to how successful their waterstoring strategy can be in this arid environment.

xiv

Bees of Australia

Pages 104–105: A single female Amphylaeus morosus found overwintering in her tree fern nest. This nest was collected as part of a study undertaken by Olivia K. Davies who works with Flinders University and the South Australian Museum. Pages 106–107: Worker Tetragonula carbonaria bees examine some damage caused during the splitting of a hive by Tim Heard in Queensland. Late stage larvae (right) and a pupa (left-most cell) can be seen exposed from the protective cells made by their siblings. While there will be a small cost to the colony from the split, overall they will benefit from a new hive and new opportunities. Pages 108–109: The gorgeous limestone cliff-faced coast along the Nullarbor plains at sunset. Pages 130–131: About 31 species of native bee (and one native wasp) captured in a three-day period on a single Melaleuca tree in suburban Brisbane, Queensland. Some individuals are males and females of the same species and some are possibly variants. Pages 132–133: A female Lasioglossum sp. pokes her head out of her ground nest in a cemetery in Brisbane, Queensland. Pages 134–135: A bridge over the incredibly reflective River Derwent in Tasmania. Pages 152–153: Stingless bees collecting pollen from a garden plant in suburban Brisbane, Queensland. Page 155: Native pea flowers planted in a native plant garden at the University of Queensland’s St. Lucia campus. These flowers seemed to be a favourite of native halictines. Pages 156–157: An enormous magnetic termite mound standing about 2.5 metres high in the tropical bush of the Cox Peninsula during the humid and hot wet season. Pages 176–177: A few of the specimens photographed in this book pinned with collection information written on 8 x 18 mm cards for future and current scientific studies. The undescribed Amegilla (Asaropoda) sp. featured on pages 158 and 159 is most prominent on the right-hand page. Pages 196–197: Three tiny micro-pinned Callohesma matthewsi all on the same piece of foam with the same collection information in the South Australian Museum. Pages 198–199: A literally gold-coated female Lasioglossum (Chilalictus) lineatum in the Victorian museum, which has been prepared for imaging with an electron microscope.

Image captions

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Introduction

People have long been attracted to bees. This is likely in part because humans have a long-standing practical history of honey collection and beekeeping. Or perhaps it is because the mostly docile bees appear with the coming of spring and herald the warmer, greener and more pleasant days to come. Bees have also long been the subject of study. The ancient Greek philosopher Aristotle even wrote about honeybees. Although Aristotle made some mistakes, such as suggesting that young bees were collected from flowers and calling the queen bee a king, he still contributed to our knowledge. Surely, some of what attracted people to bees then still attracts them now, but with the addition of their popularity across many types of media. They most often are presented in a positive light. However, without a certain level of intimacy it is difficult to develop a true fascination. My own fascination with bees was sparked because of a macro lens and an assignment to collect insects. During this assignment, I found and photographed 34 different species of native bee on a single tree in suburban Brisbane. Before then I had not thought much beyond the familiar honeybees and bumblebees, neither of which are Australian natives. That tree and those bees ignited my fascination. Through that lens (and a few more) I hope to instil and encourage that same fascination in you. To set the bees in a broader context, I want to pose a simple question: what are bees? Bees belong to an order of insects (Hymenoptera) that includes all bees, wasps and ants; bees are more closely related to wasps, a group that is often described with colourful language more than warm feelings. Bees, wasps and ants are further united in the group Aculeata by a common character among females: an egg-laying appendage (ovipositor) that has been modified into a sting. This is the reason that male bees cannot sting: they have no ovipositor. One of the major differences between bees and wasps is that bees rely on pollen as their main source of protein, rather than on other animals. In biology, though, there are always exceptions, and so there is a genus of stingless bees (Trigona) that collects carrion rather than pollen and stores this in pots like other stingless bees would honey. Other bee characters include branched, sometimes feather-like hairs and a hind basitarsus that is larger than the rest of the tarsal segments (see Fig. 1). Some other simple characters for telling bee groups apart can also be found on the face (see Fig. 2). There are over 1600 described and named bee species in Australia alone, with estimates suggesting a total of 2000 to 3000 species. Morphological differences between these species can be very small and frequently require a microscope to observe. This means that, although it is often possible to identify a bee to family and sometimes to genus in the field, identifying to species can be close to impossible, but not always. For further information on how to distinguish between native bee families, refer to Terry Houston’s Guide to Native Bees of Australia (CSIRO Publishing, 2018) and Charles Michener’s The Bees of the World, Second Edition (Johns Hopkins University Press, 2007). With so many different bee species, it is no surprise that many are yet to be described or even discovered by humans! This also means that very little is known about most species of Australian bees, leaving many Introduction

1

Fig. 1.  The major body segments of an insect head, thorax and abdomen, as well as some body parts referred to in the text of this book including the antenna, fore tarsus, rear tarsus and hind basitarsus. Shown on a male Hylaeus (Macrohylaeus) aclyoneus.

questions about each species to be answered, such as: Where do they live? What do they eat? Why did they become a species? When are they active? Could they help us? And, finally, who will resolve these answers? There is a lot of room for new work to be done and for new discoveries to be made when it comes to our Australian native bees. Most of the images in this book are of preserved specimens: bees that will be used in scientific studies and that have also been imaged and presented here for you. The bottom right corner of each species spread includes a to-scale silhouette of each sex or caste shown, giving a visual indication of how big – or small – each bee is. Most of the images are the result of a three and a half month road trip around Australia with a van that contained a very tiny, but very special, bee studio. The images in the Museums chapter at the end of the book were taken at some of our state museums (the Western Australian Museum, South Australian Museum, Australian Museum and Melbourne Museum). Information about species, subgenera and genera is often not available, and so the species included in the book are those that ‘science’ knows well enough to write something about. The book includes the bees that are found in Australia’s states and the Northern Territory, but you will notice that most species are found across multiple states and territories. The Australian Capital Territory has been included in New South Wales because of its small size. This is because bees care very little for the lines that we draw on maps (except where those 2

Bees of Australia

Fig. 2.  Features of the head referred to in the text of this book, including the ocellus, antenna, clypeus, mandible and compound eye.

lines mean a change in habitat). The distribution of many bees is instead governed by climate (e.g. tropical, subtropical, arid and temperate zones) and available habitat (e.g. food and nesting site availability). The bees are presented in this way so that you as readers can open the book to your own state, or those that you are interested in, and relate to those bees, as well as search for those species in your local area. On that note, this book is intended to be opened at any page and enjoyed at whatever depth of detail that you, the reader, wants. This book can also be enjoyed by people of any age or level of interest in bees. You may just want to look at the ‘pretty’ photos, or you may want to read all the accompanying text and feature pages. So please, pick the book up, enjoy the content and gain some more appreciation for our native bees and the hidden diversity of the insect world. I have tried to avoid technical jargon as far as possible, but a few specialised terms were inevitable. A glossary of such terms is provided at the back of the book. For the more involved reader, there is also an appendix of the bee species featured in this book, organised by family. Additionally, a list of further readings is provided to help you learn more about bees in Australia and worldwide. To see more bee and insect imagery feel free to check out my website at www.jamesdoreyphotography.com.au, and to read a small blog about my road trip around Australia collecting bees have a look at www.beelogblog.wordpress.com. Introduction

3

New South Wales National parks along the New South Wales coastline have a high level of bee diversity and are great places to go ‘bee hunting’. To the north of Sydney, many subcoastal parks are dotted with grass trees, and dead and dry flower stalks provide homes to many bees. The bright metallic green carpenter bees will burrow into the sides of these stalks and then excavate tunnels inside the stem. You can recognise their nests by a circular hole, about the size of a 5 cent coin. Ground-nesting bees are common in the sandy soil of places such as Ku-ring-gai Chase National Park. Wasps also nest in these sandy soils, but wasp nests have entrance tunnels that are irregular in shape and are usually sloped, while bee nests are always perfectly round and burrow straight into the ground, not at an angle. The many little coastal towns up and down the New South Wales coast are also great places to find halictine bees, particularly the tiny, colourful Homalictus and the slightly larger, but closely related, Lasioglossum. The parks and headlands along the coast often have a wealth of yellow daisies that these bees can visit in abundance. While bees are easy to spot against the bright yellow flowers, be wary of being fooled by hoverflies: another wonderful group of pollinators. The great gum forests of New South Wales, often regarded as important habitat for koalas, are also very important for bees, with individual flowering gums capable of attracting thousands of native bees. Many gums are too tall to spot these bees on, but finding smaller trees or those with lower limbs should reward you with a great bee-spotting experience. However, don’t forget to look up and see the super-highway of speeding insects around the flowers above your head. If you are lucky enough to own an insect net, a few sweeps across these flowers will give you an even better idea of what is relying on these trees, particularly the difficult to see green, yellow, orange and black euryglossine bees.

4

♂ ♂

♂ 6

Bees of Australia

♂

Amegilla (Asaropoda) bombiformis As the year warms up, suburban gardens along the coast from Brisbane to Sydney are enriched by these metallic orange beauties. It is easy to see why people call them ‘teddy bear’ bees. You may even find clusters of males roosting at night on twigs or grass stems. As the individuals get older, their colour may lose the metallic sheen and fade to pale yellow, but they are still a lovely sight as they search for their favourite flowers. Tubular flowers suit teddy bears nicely because they may contain nectar that other bees cannot reach.

New South Wales

7

♀

8

Bees of Australia

♀ ♀

Megachile ustulata Resin bees such as this M. ustulata are named so because of the materials that they use to create their nests. Unlike some other bees in the family Megachilidae, which make their nests using cut-up leaves and petals, these bees mix plant material and resin from trees to make nests for their young. Resin bees often make their nests in rock crevices or holes in wood made by wood-boring beetles (or bee-friendly people). Both nest site types can be found in many old houses, meaning that these bees will readily nest in old drill holes or gaps in brick or wooden houses. Resin bees will also nest in more opportunistic places, with nests reported in the folded canvas of a caravan’s awning and even some old oilskins. Megachile ustulata is a large species of bee, with the female a bit larger than a European honeybee; it is also a very striking bee, being dark black with a bright orange

abdomen. Although the scopa (a region of hairs under the abdomen) of this species is also a vivid orange colour, the lower abdomen may appear to be a different colour because these hairs are used to carry pollen: this individual is carrying yellow pollen. Footage by Tobias Smith of a mating pair of M. ustulata has shown some very interesting mating behaviour where the male quickly stroked the eyes and touched the antennae of the female. This is behaviour is speculated to be a way of calming females and to send ‘chemical communications’ to the female from odour glands on the male’s legs. Megachile ustulata can be differentiated from the common and large leafcutter bee, M. mystacea, by a lack of pale hairs on top of its head and its orange-coloured hair on its tarsi. Megachile ustulata has been found along the coast between Sydney and Brisbane, with one unusual capture reported in central Queensland.

New South Wales

9

♀

♀

10

Bees of Australia

♂

Amegilla (Zonamegilla) asserta Blue-banded bees are both beautiful and plentiful in suburban gardens. This species, with yellow face marks and a black streak down the hind leg of the female, is often seen in the cities along the eastern coast, particularly in herb gardens where many of the plants we grow have tubular flowers. Strange as it may seem, the number of different species has been unclear until very recently and the nesting habits of this species are almost unknown. The nests of one or two Amegilla species have been well studied and while we might assume that this species will behave similarly, we do not really know. Male Amegilla bees have a larger extent of pale markings on the face than do the females. They also spend more time

on the wing and less time on the flowers. If you see a bluebanded bee moving from flower to flower without touching any, then it is probably a male looking for a female. However, it is not so easy to tell one blue-banded species from another. If you are in Brisbane, Amegilla asserta is the one with yellow face marks and white, not orange, hair on the hind legs. In Sydney, the one with orange hair on the hind legs is not found, whereas in Melbourne A. asserta is the most common species. This is, of course, assuming that one of the less common species has not crept into town.

♀

♂

New South Wales

11

♀

♀

12

Bees of Australia

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Undescribed Leioproctus (Exleycolletes) sp. The taxonomy of many Australian bee groups is in a poor state, with countless species undescribed or described incorrectly. The taxonomic work is ongoing but often slow due to a lack of funding and expertise. This striking species of Leioproctus is one of those that are yet to be described. Even though specimens of this species do exist in museums, the

species has not yet been formally described and as such does not yet have a full scientific name. This male and female, of presumably the same species, were found in some numbers along the edge of a creek in the Northern Rivers region of New South Wales. This species has also been found in SouthEast Queensland.

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Lasioglossum (Parasphecodes) lithuscum Lasioglossum lithuscum is found in most of southeastern Australia, including Tasmania and, like many ground-nesting bees, has formed a relationship with mites. Many readers have likely heard of Varroa destructor, the mite that has been blamed for many honeybee declines overseas, but the relationship between bees and mites is not always a bad one. In fact, for many burrowing bee species such as L. lithuscum, mites play a very important role for the bees and their brood. The mites will feed on mould

growing inside the damp subterranean cells where bee larvae develop, protecting both the young bee and its food from fungal attack. For this reason, a female bee will shed some mites in each cell before sealing it up, allowing them to undergo their feeding and sexual stage. Once the bee is ready to leave the cell as an adult, the mites have another lift to their next home, continuing the symbiotic relationship between bee and mite. This is a possible explanation for the relatively large mites found on this individual.

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Hylaeus (Macrohylaeus) alcyoneus The stunning H. alcyoneus is quite large, ranging from 8.5 mm to 11 mm in length, and has a metallic blue abdomen (see Fig. 1). Male H. alcyoneus are usually larger than females, although not in the case of these two individuals. Larger males often have large spines under their abdomen; this individual has only smaller spines. The presence of spines and the larger males are believed to be an adaptation for territorial male–male competition. In Western Australia, this territory is likely to be a Banksia flower, but these bees will guard a wider range of flowers in the east, where banksias do not flower all year round. This has earned them their common name of ‘banksia bee’. These bees often make their nests in wood, and the individuals featured here were collected and reared from the dead stalk of a grass tree (Xanthorrhoea sp.) near Sydney, New South Wales. Ongoing genetic work by

Olivia Davies, which intends to resolve how bees of the subfamily Hylaeinae are related, has also uncovered some other very interesting biology of these bees. For example, the brood of this bee smell strongly of lemon myrtle, while their food provisions do not. Whether the smell is produced by the bee itself or extracted from the plant is unclear. The reason for this smell, however, might be as an antifungal compound or as an insect repellent. Yes, an insect using an insect repellent! Perhaps the strong smell of lemon myrtle confuses or deters brood parasitoids (such as Gasteruption wasps) and predators (such as ants), helping the young bees survive into adulthood. Hylaeus alcyoneus has been recorded in South-East Queensland, New South Wales, Victoria, South Australia and Tasmania, as well as in south-west Western Australia, but not in the vast expanse in between.

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Lasioglossum (Chilalictus) lanarium This is Australia’s largest Chilalictus bee, which is common in bushland and in many urban backyards in eastern and southern Australia, as well as south-west Western Australia. It is a communal species, where females cooperate in using a common nest but with each female rearing her own brood. It is an unusual species because females can move from one nest to another, provisioning cells in one nest, and then moving to another nest to do the same. This is probably a strategy to ‘spread the risk’ of rearing brood in just one nest, which might have a parasitoid wasp living in it and attacking all the bee larvae. Switching nests could be a strategy of ‘not putting all your eggs in one basket’. This

bee is also very aggressive and will quickly bite and sting any intruder that approaches the nest entrance – their stings can be very painful. Nests often occur in dense aggregations, and the nests are recognisable by a small rim or turret surrounding the entrance. When it was named in 1853, entomologists did not specify the derivation of the name, as they are now required to do, but it is safe to assume that the name came from lana, the Latin word for wool (because of the white hair on the abdomen). At the time, it did not matter that a couple of other species also had patches of white hair on the abdomen, but we now know that feature is not particularly distinctive.

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Leioproctus (Exleycolletes) cristatus This moderate-sized bee nests in bare, fine sandy soil, often in gardens and especially in coastal areas. Nests sometimes occur in dense aggregations and can be easily spotted by the ‘tumulus’ of freshly excavated soil around each nest entrance. Females are solitary and each nest comprises a single vertical shaft with short side tunnels radiating away from this with a cell at the end of each tunnel. Leioproctus is one of the largest bee genera in

Australia and belongs to the family Colletidae. For a long time the Colletidae was thought to be one of the most primitive bee families in the world, and colletids are most abundant and diverse in Australia. However, molecular studies now show that colletids are not as old as thought, but their diversity in Australia still dates back to the late stages of Gondwana, when Australia was connected to Antarctica.

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Exoneura sp. Some bees will form massive mating swarms, with thousands of bees all vying for the right to mate. This male Exoneura was found in one such swarm atop Mt Warning in northern New South Wales. The swarm was so dense that any swipe of a hand would leave you with two or three confused bees, which would immediately fly off again, not wanting to miss any mating opportunity. Exoneura species in Australia are mostly restricted to temperate and moist subtropical climates, and they are rarely found towards the tropics or arid zones of Australia.

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Thyreus nitidulus This handsome bee is actually a cleptoparasite of bluebanded bees in the genus Amegilla. It has shining metallic blue patches and bands on its thorax and abdomen and can be quickly recognised by these as it flies around. It will follow Amegilla females back to their nests and, when the host female leaves to collect food for her brood, Thyreus enters the nest and lays an egg in the brood cells. When the Thyreus larva hatches, it kills the Amegilla egg and then consumes the cell provisions. The species name nitidulus is Latin for ‘a little bit shiny’, but in fact this bee is very showy with its brilliant metallic blue markings.

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Why are bees important? James Dorey Australia’s bees are an amazingly diverse, visually stunning and important group of animals. Their ecological importance stems greatly from the ability to pollinate the flowering plants of the world. Pollination is a vital ecosystem service, critical for sexual reproduction in plants. Pollination is provided freely in the natural world by insects, birds, bats, lizards, possums, rodents and even lemurs. Although there are many animal pollinators, bees are considered to be the most significant pollinators of the lot, particularly for crop pollination. Crop pollination by the European honeybee alone has a commercial value of A$4–6 billion in Australia and A$240 billion worldwide, with 35% of the food that we produce and 75% of our food crops being reliant in some way on animal pollination. Bees also create wonderful and useful products such as honey, beeswax, pollen, propolis and more. Honey and beeswax production by honeybees in Australia is worth A$90 million annually. Although that is a lot of food and money, it is far from the end of the bee story. Pollination of crops and the making of bee products are the most commonly raised topics when discussing the importance of bees. Unfortunately, these issues are often raised only in terms of the European honeybee. Although it is a very economically important species, the European honeybee is actually an invasive species almost everywhere that it resides, frequently stealing nectar from native plants without pollinating them, competing with native pollinators for food and nest sites, and pollinating exotic weeds. The role of the honeybee in crop pollination and the making of bee products is also only the tip of the iceberg when it comes to how important bees are to us and life on Earth. With ~19 700 named species of bees across the globe, over 1600 of which are native to Australia, it should be clear that it is not just the honeybee that needs to be considered. Having co-evolved alongside flowering plants for over 100 million years, bees have become extremely important to the survival of many natural systems, as well as sustainable agriculture. Animals pollinate almost 90% of all flowering plants: that is over 308 000 plant species! Because bees are the primary pollinators in most ecosystems, it is difficult to overstate the importance of bees to the survival and success of many of these plants and the organisms that depend on them. One way to explain how important bees are in the natural world would be to imagine what might happen if they were to disappear tomorrow. Direct effects of pollinator loss or decline would result in the loss of pollination services for the plants in that ecosystem. Some flowering plant species rely on specialist pollinators that will pollinate only one or a few plant species. This is advantageous for the plant because it means that the pollinators must be loyal to them and are hence more likely to travel between two individuals of that same plant species, providing effective pollination. The obvious downside of such an arrangement is that both the pollinator and plant require each other to survive. Plants requiring a single species for pollination are particularly vulnerable to extinction if that pollinator were lost, and the bee would be vulnerable if the plant were lost. However, these isolated species interactions are rare and specialist relationships can also involve generalist plants and pollinators. Most flowering plants do not require a specialist pollinator but instead attract generalist pollinators. Although generalists may not be as efficient at pollinating as specialist pollinators, the risk of using such pollinators is much lower. The loss of one pollinator species will not be as much of a problem for these plants because other generalist pollinators will be able to pick up the slack or provide adequate pollination; such redundancy is typical of many plant– pollinator relationships. However, the problem can become catastrophic when more and more pollinators are removed from an ecosystem and the in-built redundancy starts to fall apart.

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The ultimate cost for any flowering plant species after the loss of pollination could be extinction. Extinction for large perennial flowering plant species that have lost their pollinator might be a slow and subtle process that may go unnoticed for many years. Other species may disappear in a flash or suffer a gradual population decline because of lessened seed set or seed quality from poor pollination. Flowering plants make up much of the physical structure in many ecosystems. This structure provides the conditions, shelter, nesting sites and habitat that many animals and plants need to survive. Flowering plants also provide food for many animals. A reduction of roots, stems, flowers, leaves, fruits or seeds produced by plants could set off a cascade of ecological effects. The many frugivores and herbivores that rely on these food supplies would be impacted by a loss of species producing these resources. A decrease in fruit and seed set alone would have a negative impact on the many bird, mammal and insect populations that use these resources. The cascade would then continue onto the many predators, parasites and parasitoids that prey upon these species and each other. There are many ecological and monetary reasons to preserve bee species and diversity. Perhaps the most often overlooked reason is the non-material benefits that people get from bees, and the wealth of biological history stored in the behaviour and genetics of each species. Without our wonderful diversity of bees, you would not be reading this book or enjoying these photos. You would not be able to go out in the garden or bush and see a multitude of bees conducting their important business. It is very unlikely that we will lose all our bees overnight. Bees are not our only pollinators and many flowering plants do not require animals for successful pollination. Yet, any pollinator species that is lost or whose population declines could be another chink in the armour that nature’s diversity and redundancy provides. Every chink could bring about unforeseen effects that might be another step towards the end of a population, a species, a crop or an ecosystem. Awareness and knowledge of the importance of bee diversity to humanity and the natural world is essential to the conservation of bees worldwide.

Queensland Like most Australian states, Queensland has a wide range of climates and habitats. Queensland is home to the Great Barrier Reef, stunning tropical islands, Australia’s northernmost point and a plethora of ecosystem types. In Queensland, you will find monsoonal rainforests to the north, persistent deserts to the west, dry grasslands to the south and lovely tropical and subtropical coastal regions to the east. Some of the most amazing habitats along Queensland’s eastern coast are the little pockets of remnant rainforest, which provide a view into Australia’s earlier wetter climate. Perhaps the most interesting Queensland bees occur in the wet tropical and subtropical rainforests. These include stingless bees, which are distantly related to honeybees. Stingless bees nest in cavities in tree trunks and their colonies can contain hundreds of workers. The stingless bees in Queensland are very small, ~5 mm in length, and are usually black. They are useful for pollination in horticultural crops such as avocados and macadamia nuts. They produce small quantities of honey, which is more sour than honey from the introduced European honeybee. Queensland is also home to many carpenter bees, which nest in dead rotting wood (such as branches of Banksia and Melaleuca trees). Carpenter bees are large, about twice the size of honeybees, and usually black with a yellow thorax, but some species in the Lestis group are coloured a bright metallic blue and green. Males can hold territories where they fight off other males and scent-mark leaves and stems, returning to these marked sites frequently where they produce sex pheromones that attract females. Queensland coastal dunes are often inhabited by small halictine bees in the genus Homalictus. They can be easily found feeding on the flowers of pigface (Carpobrotus sp.) along back dunes. They come in a range of colours, but the most beautiful species have red abdomens and bright metallic green or blue heads and thorax. Australian species of Homalictus are communal – they nest in underground tunnels and each nest can contain dozens of females, but all females lay their own eggs and rear their own brood – there are no queens or workers. One of the best places in Queensland to explore the nests of native bees is in the subcoastal regions from Maryborough south to the Gold Coast. Much of this area contains open heathy meadows where grass trees (Xanthorrhoea) are common. Once the flower spikes of Xanthorrhoea have died and become dry, they can snap off and many different native bees will dig nesting burrows in the remaining stalks. The bee nests can be recognised as perfectly circular and clean holes in the broken surface at the top of the dead stalk.

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Austroplebeia australis This eusocial bee can be found in the Northern Territory, Queensland and New South Wales. It has the most southern distribution of any Austroplebeia species and hives of this species are commonly kept in parts of Australia. The colouring of these bees varies markedly from central Australia to the central coast of New South Wales, Cape York in Queensland and the Kimberley in Western Australia. These changes occur between

populations across the continent. Perhaps it was these variations that led to A. australis being given five separate names, the four others being: A. percincta, A. cockerelli, A. ornata and A. websteri. But the oldest name is the one that must stick, so this little beauty remains as A. australis. When it comes to taxonomy, there is no prize for coming second (or third, fourth and fifth) and these other names are no longer used for this species.

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Palaeorhiza (Cnemidorhiza) disrupta Palaeorhiza bees are often large compared with many others in the subfamily Hylaeinae, which contains many charismatic Australian bees, such as the masked bees. They have also undergone major diversification in New Guinea, with every subgenus resident there. They extend as far west as the Lesser Sundas and as far south as Port Macquarie. With ~150 species in the genus, many being described from

a single specimen, it is likely that there are many more to be found. Unlike most other hylaeine bees, representatives of Palaeorhiza (Cnemidorhiza) have been found nesting in the ground instead of in wood. Bees in this subgenus are usually metallic with brilliant colouration. Palaeorhiza (Cnemidorhiza) disrupta is found along most of Queensland’s eastern coastline.

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Megachile abdominale Male Megachile species can often have unusual modifications to their bodies. Megachile abdominale is one such species, where males have very long fore tarsi (front feet). Males of Megachile ramulipes, a related bee, also has similarly long fore tarsi but otherwise looks quite different, lacking a striking orange abdomen. An easy way to tell

a male from a female is that Australian Megachile males have a downwardly curved abdomen tip, whereas a female’s abdomen generally points straight back. This is presumably an adaptation to get around a female’s abdomen to mate. Megachile abdominale has been recorded only in the greater South-East Queensland region.

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Braunsapis sp. Bees in the genus Braunsapis are very like those of Exoneura but always have a black abdomen; unlike the typical red abdomen of most Exoneura. A big difference between these two groups is that Braunsapis evolved in Africa and then spread to Australia via tropical Asia, whereas Exoneura evolved in southern Australia and is adapted to cool and wet climates. This difference in their geographic histories is reflected by their distributions: Exoneura lives in cool temperate forests and rarely occurs north of southern Queensland, while Braunsapis is most common in the tropics and rarely extends below Sydney on the east coast. Interestingly, both Braunsapis and Exoneura have their own social parasites: bee species that are closely related to them, and look very similar, but which surreptitiously enter their nests and lay eggs that are reared by the host females.

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Hylaeus (Euprosopoides) ruficeps If you see this bee in eastern Australia, you will have no difficulty recognising the species by the female’s red head, which (as you may already have noticed) is what the Latin name means: ‘red-headed’. There are two colour forms in which the pale marks on the face and

back are either yellow or white. However, in Western Australia things are more difficult: the head is dark and, although males can be distinguished from H. obtusatus by their face patterns, the differences between females of the two species are much more subtle.

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Euryglossina (Microdontura) mellea The only described species in the subgenus Microdontura is E. mellea, which has only four recorded collection localities from Queensland and Victoria, although these sites do span almost the entire east coast. It is entirely likely that there are more species in this subgenus, but the last revision of the Euryglossina was completed in 1968! Even back then, the author noted that her collections often included new species of Euryglossina, making it possible that new species are sitting in collections just waiting to be described.

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Megachile apicata In many species of Megachile, the males can have some very strange modifications to their body. Quite often these can involve the broadening or lengthening of the front legs: modifications that might be used during mating and for species recognition. Perhaps it should come as no surprise that males within a single species can also have somewhat diverse morphology. The large, flattened antennal tips

of this specimen looks like a fantastic character to work out the species; they were, however, a red herring. Both the colouration and antennae appear to vary within the species. The best characters to tell that this was indeed M. apicata resided on its front and rear ends: small ridges on end of the abdomen and its paleyellow mandibles. You can find M. apicata around most of mainland Australia.

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Tetragonula carbonaria Tetragonula carbonaria is probably the most commonly kept and cultivated of Australia’s native bees. Nests of T. carbonaria can be identified by their amazing single or multiple spiral-shaped brood comb. Virgin stingless bee queens will undertake mating flights before breeding and, once their flights are completed, their abdomens will swell in size, leaving them unable to fly. A queen will then lay eggs for the rest of her life, never leaving her nest. Males will often form large congregations, waiting for a virgin queen to fly by. When the males mate, their genitals will lock into place

and be left within the queen, which must then seek the help of her workers to have them removed: this is likely a strategy to prevent other males from mating with the queen during that flight. The worker bees are sterile females that build and provision the hive, while monitoring the queen to ensure that she is doing her job … and doing it well. Tetragonula carbonaria, although predominantly a subtropical species, extends its range further south than any other Australian stingless bee, making it almost to the Victorian border along the eastern coast.

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Hylaeus (Gnathoprosopis) albonitens A beautiful masked bee. The members of this subgenus are all small and they most frequently visit eucalypts and other flowers in the Myrtaceae family. Two species are metallic blue, but the other five are black with yellow on the face, collar and legs. It is always worth looking closely at flowering eucalypts for these small gems. Hylaeus albonitens can be found from the tropical north of Western Australia, the Northern Territory and Queensland, as well as down the eastern coast to just beyond the New South Wales border. As is the case with many bees, the males have a paler face than the females.

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Megachile aurifrons Megachile aurifrons is a large and very striking bee: the eyes of the female are red and only surpassed in beauty by those of the smaller male. This bee has a massive range, which spans most of mainland Australia. This was made very apparent to me when I found one individual in Dotswood, Queensland, and then later, another at Arrowsmith, Western Australia: two sites just about as far apart from each other as you can get without taking a swim. So, unless you are living in Tasmania, do

not give up hope of seeing this lovely bee in the flesh. In general, megachilid bees can be recognised by their rather stout, bullet-shaped, body form, and this species can be further distinguished by its golden/orange face (in Latin aurifrons means golden face) and white bands on its abdomen. This species nests in burrows that have been created in dead wood by other insects such as wood-boring beetles, and it is a common occupant of ‘bee hotels’ in urban gardens.

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Nomia (Hoplonomia) rubroviridis Nomia bees can often be recognised by the colourful bands on their abdomen, although some species lack bands. The colour of the bands can vary greatly, making it a poor species character. Two male N. rubroviridis that were caught in Dotswood, Queensland, at the same time displayed very different coloured bands: one yellow and the other blue and green. Nomia rubroviridis is found across the top of Australia and down the east coast to Brisbane.

You might be forgiven for thinking that a Nomia was a blue-banded bee (Amegilla sp.), but an easy way to tell them apart is to check their faces because Nomia lacks the striking yellow facial markings of many blue-banded bees. Nesting biology is known for a Nomia (Hoplonomia) species from Japan, H. punctulata, where bees form communal nesting burrows with several females in occupancy.

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Australian native bees as crop pollinators Tobias J. Smith In Australia, we grow over 50 crops that benefit from, or are dependent on, pollination by insects. These include a range of tree crops, other fruit and vegetable crops, seed crops and textile crops, to name a few. Like most countries around the world, Australia focuses heavily on the European honeybee, Apis mellifera, for the pollination of crops pollinated biotically (by animals). Farmers may pay for the services of commercial managed honeybee hives or benefit from the free pollination services of other nearby colonies (feral or managed), or both. Although honeybees are an inarguable pillar of our agricultural system, we over-rely on this single species, and we should aim to diversify our crop pollinator communities, for multiple reasons. Honeybees are vulnerable to a range of ailments, and in some parts of the world have experienced declines. In addition, these bees are an introduced species in Australia, and there have been documented negative environmental impacts both here and overseas. Having different types of pollinators within crops is beneficial for several reasons. First, diversity can add beneficial back up in the event of the decline of a single species. In addition, in many cases, having two or more pollinator species in a crop may result in higher rates of pollination than with any of the species on their own. Honeybees are incredibly important and versatile; however, it is becoming increasingly understood that the collective contribution of other insect pollinators to global crop pollination is actually greater than the contribution from honeybees alone. The role of Australian native bees in crop pollination is under studied. Although over 1600 species have been described, most species probably make little to no contribution to crop pollination. There are, however, a few species and groups that have great potential as crop pollinators, but fewer than 20 species have actually been documented pollinating crops. The best understood are two stingless bee species: Tetragonula carbonaria and Austroplebeia australis. Tetragonula carbonaria has been recorded as a pollinator of macadamia, blueberry, longan, mango and, with limited success, capsicum. Austroplebeia australis has been recorded pollinating celery and, with limited success, also capsicum and carrot. In addition to these two stingless bees, several other native bee species have been documented pollinating crops, including: two Amegilla and two Xylocopa species (tomato); seven Megachile species (lucerne); one Lipotriches species (lucerne); and one Lasioglossum species (white clover). Several other species have been recorded as occasional crop visitors, including Lasioglossum, Braunsapis, Exoneura, Homalictus and Hylaeus species. It is highly likely that some of these species are pollinators of some crops, and that there are also other unrealised native bee crop pollinators. Further, as some bee-keen gardeners might know, Lipotriches and Lasioglossum species are common visitors to tomato, and several Megachile species are common visitors to cucurbits. In most cases, the use of native bee species as crop pollinators in Australia is not aimed at replacing honeybees, but rather, at complementing them. In some cases, though, native bee species do offer pollination benefits that honeybees do not. For example, some native species in the genera Amegilla, Xylocopa, Lipotriches, Lasioglossum and Hylaeus can perform ‘buzz pollination’. Buzz pollinating bees can vibrate their bodies while clinging to flowers to shake pollen free. The flowers of crops including tomato, capsicum, eggplant, chilli and blueberry can have increased rates of pollination if they are visited by a buzz pollinating bee than if they are visited by a non-buzz pollinating bee species, because of the way their pollen is held within the flower. One of the hurdles in using Australia native bees as pollinators of crops is ensuring they occur in high enough numbers within crops to have a meaningful impact on overall pollination. Therefore, diversifying insect crop pollinators using native bee species can be undertaken using two different pathways: (1) using managed native bee species; and (2) through protecting, promoting and using wild bees in agricultural landscapes.

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Managed native bee species

Australia has only a small number of native managed bee species, all of which are stingless bees, and only one of which is used as a crop pollinator on a commercial scale: Tetragonula carbonaria. Being a highly eusocial bee, T. carbonaria lives in colonies of thousands of individuals and is easily kept in artificial hive boxes. The cultivation of T. carbonaria is well established and a community of beekeepers exists. The number of hives available to agriculture is slowly increasing as the number of commercial beekeepers providing hives to agriculture (renting or selling) grows. Tetragonula carbonaria is primarily used as a managed pollinator of macadamia and blueberry, but is also used in avocado and occasionally in several other crops. It has long been established that T. carbonaria is a frequent visitor to macadamia, and an effective pollinator, but recent research has gone one step further and demonstrated that T. carbonaria is the most efficient insect pollinator of macadamia. In blueberry, recent research has shown that T. carbonaria is an effective pollinator. With several recent funding investments in research focused on T. carbonaria as a managed crop pollinator, it is likely that we will soon hear more about the potential usefulness of this species as a managed pollinator in other subtropical crops. Promoting wild bees in agricultural landscapes

Native bee pollination in crops can also be delivered as an ecosystem service, provisioned by ecosystems in agricultural landscapes. In fact, the majority of native bee pollination of crops in Australia is probably derived from wild bees. For wild bees to persist in agricultural landscapes, they require access to a few key resources: diverse sources of nectar and pollen; nesting sites; and nesting materials. As such, protecting remnant vegetation and restoring areas of natural vegetation is key to supporting bees and other pollinators in agricultural landscapes. In addition to protecting natural vegetation, farmers can provide supplementary forage and nest sites for bees within and around crops. Such measures can include planting complementary nectar and pollen sources, such as diverse windbreaks or cover crops, with year-round flowering in mind. Native bees can also be supported with the provision of nesting sites. These may be as simple as artificial wooden nest blocks for solitary above-ground nesting species. However, as more is understood about the life cycles and crop pollination potential of other bee species, there may be other forms of nest ‘management’ that may be used. For example, the installation of carpenter bee nests, or the preparation of appropriate soil patches for the ‘transplanting’ of ground-nesting bee nests, both of which are undertaken in other parts of the world. Lastly, but importantly, wild bees, like other insects, are vulnerable to insecticides. Therefore, with any use of insecticides in agricultural landscapes, wild bees, their nest sites and their forage must be carefully considered and protected. The future

The pollination of Australia’s biotically pollinated crops is currently dominated by the use of managed European ­honeybees, and the free services of wild pollinators in agricultural landscapes. Although there is a small but growing industry of managed stingless bees as crop pollinators, there are multiple pathways in which we can further use native bees in crop pollination. These include supporting and protecting bees in agricultural landscapes, and actively encouraging them through the provision of suitable nesting sites. As research continues into the pollination systems of various crops in Australia, it is likely that we will find an even greater potential for contributions from native bee species. Native bees are very likely one of our greatest partners in delivering better pollination of crops in Australia, and it is safe to assume that their full potential is still far from realised.

Victoria Victoria has a wide range of habitat types and a correspondingly wide range in bee diversity. The northwestern regions contain typical arid-zone bees such as minute euryglossines and large, hairy colletines. The cool montane forests of the Great Dividing Range are home to many allodapine species. These are easy to find in the Dandenong Ranges close to Melbourne: simply look for small and perfectly round entrance holes in the ends of old tree fern fronds that have fallen to the ground. Another bee common in the Dandenong Ranges is the large black and yellow colletid bee Amphylaeus morosus. These also build tunnels in dead tree fern fronds, but they are much larger than allodapines and have a powerful sting. Amphylaeus morosus is the only species in the family Colletidae that is known to be social, but colonies rarely contain more than three females. Halictine bees are also common in Victoria, especially in open forests where they often nest in bare soil along roadsides or pathways. They can also be common in urban backyards and their small entrance holes are surrounded by a small turret of loose soil that has been excavated from the tunnels below. Some of the Victoria halictines are especially interesting because they have large flightless males with enormous sickleshaped mandibles. These males remain in their nests and fight to the death with each other, and the survivors mate with females as they emerge from their underground cells.

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Homalictus punctatus Homalictus is a widespread genus of small, often metallic, bees that nest communally in soil. They range all the way from coastal dunes to inland deserts and up to alpine areas. They are one of the most common native bees in urban gardens where they can nest in lawn and bare soil, but they often escape attention because of their small size. Each nest has a single entrance but the burrow branches after this, leading to a

maze of tunnels with brood cells leading off from these. It appears that all Australian Lasioglossum and Homalictus species are communal, lacking queens and workers. This is very unlike halictine bees in other continents, which are usually solitary or eusocial, and prompted the question of the ‘Australian enigma’ – why is communal nesting ubiquitous in Australian halictine bees, but rare elsewhere?

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Leioproctus (Leioproctus) plumosus These amazing metallic green and blue solitary ground-nesting bees have been found along much of Australia’s coastline. I have found them in Western Australia, Victoria and Tasmania. Although these widespread wonders are solitary, they still nest in aggregations, with anywhere from one to three generations a year. There are some benefits to nesting in aggregations, such as making it easier to find a mate, but there are also some serious drawbacks. One such drawback is that an aggregation will create a hotspot for parasites, predators and parasitoids. Indeed L. plumosus

has been found to fall victim to a common and species-rich family of parasitoid wasp: the Ichneumonidae. Two species of ichneumon wasps have been observed patrolling nesting sites for likely hosts in which to lay their eggs. Once a female wasp finds a likely nest, she will lay her own eggs in the nest and her young will consume the developing bee larvae. When the next generation of bees is ready to emerge, so too will the wasp, ready to continue the cycle of parasitism. Parasitism may be gruesome but it is also an amazing and ecologically significant process.

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Hylaeus (Gnathoprosopoides) philoleucus This subgenus contains only two species, H. philoleucus and H. bituberculatus, both of which are small bees with characteristic short and blunt mandibles. Hylaeus philoleucus has a generally more northern distribution than its sister species, ranging from Victoria to north Queensland. These sister species can be distinguished by the white markings on H. philoleucus and the yellow markings on H. bituberculatus. The name philoleucus loosely means ‘lover of white’,

because philo means ‘lover of’ and leucus means ‘white’. This is likely because of the striking white patterns on the faces of the male and female bees. This specimen, collected in Melbourne, is the southernmost specimen recorded. The next southernmost specimens were collected just over the border from New South Wales, near Echuca, and two females on Kangaroo Island, South Australia, a haven for bees in a highly human-disturbed state.

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Lasioglossum (Chilalictus) veronicae The genus Lasioglossum is closely related to Homalictus, with the relationship between the two being difficult to discern. The attractive metallic greens on the head and thorax of L. veronicae, as well as its small size (5 mm), make it appear Homalictus-like. The differences between the two genera are not that great, with Lasioglossum having simpler hairs than the feather-like hairs of Homalictus and different hair formations on their rear legs. Both characters require close examination under a microscope. There is, however, a

shortcut when looking at male Lasioglossum (which can often be distinguished from females by their long antennae): almost all male Lasioglossum have a yellow or white marking on their clypeus, the integument just above their mouth, while most Homalictus males do not. This forms a cute, pale ‘upper lip’ which is fantastic for bee identification! The Homalictus-like Lasioglossum veronicae can be found confusing taxonomists and enthusiasts in south-west Western Australia, South Australia and Victoria.

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Lasioglossum (Chilalictus) sp. This may be one of the most important groups of native pollinators in southern parts of Australia. Members of the group appear in large numbers in museum collections and are known to visit flowers in at least 30 different families. They nest in soil, with several females taking turns at guarding the entrance to a shared vertical shaft. Each female constructs brood cells branching off the main tunnel. There are roughly 50 Australian species with a similar overall appearance but a range of sizes.

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Amphylaeus (Amphylaeus) morosus This bee is one of the most remarkable bees in Australia. It belongs to the huge worldwide family Colletidae, but it is the only species in this entire family that is known to be social. It can be identified by its large body size (about the size of a honeybee) and golden yellow markings on the thorax and its face. It digs nest tunnels into soft dead stems and branches, such as fallen tree fern fronds and the dead flower scapes of Xanthorrhoea grass trees. After females have excavated these tunnels, they line them with a silvery secretion that is licked on by their mouthparts, and this

helps to keep the nest waterproofed. Sometimes two or three females can share a nest, but they do not form hierarchies (such as queen or worker castes) and each female is reproductive. Females have a bright yellow diamond marking on their black face, which makes it easy to identify them as they sit in their nest entrances waiting to forage. Recent studies by Olivia Davies have shown that this bee has a remarkable genetic system involving a puzzling lack of variation in their mitochondria (the intracellular bodies that regulate metabolism).

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Pachyprosopis (Pachyprosopis) haematostoma Pachyprosopis haematostoma is the only member of its genus with metallic colouration. It is a brilliant blue and orange, making it a very eye-catching bee. This bee used to go by two names, P. haematostoma and P. aurantipes. This was until the genus was revised by Elizabeth Exley (who is sometimes referred to as ‘the grandmother of Australian bee research’) who considered them to be the same species: a common story for many Australian bees. The two original species names were based on work in 1915 by Theodore Dru Alison Cockerell, an English/American zoologist who did much of the early work on Australian bees and who described the males and females of this species as different because,

as with many Pachyprosopis, the males and females look very different from one another. Cockerell did suggest that the males and females could have been of the same species, remarking on his labels that ‘if this is correct, it is very remarkable, as they differ extremely in colour and general appearance’. Pachyprosopis haematostoma has been found nesting in vacated beetle burrows in a tree stump, with some indication that they were also actively burrowing and not just riding on the coat tails of the beetle larvae’s hard work. These bees can be found along much of the east coast, as far south as Adelaide and Melbourne, and as far west as south-west Western Australia.

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Nomia (Paulynomia) aurantifer Found only in Australia, the subgenus Paulynomia has only two species, which extend up the east coast from Melbourne to Sydney and north to Queensland. The actual characters for Paulynomia can be difficult to observe, sometimes requiring a microscope and sometimes even the dissection of male genitalia. But when compared side-by-side, the two species, N. aurantifer and N. swainsoniae, are not too difficult to tell apart: the former having bright orange stripes and the latter more yellow stripes. Existing collection records suggest that N. aurantifer is associated with rainforests (this individual, for example, was caught foraging on a rainforest

vine on the edge of a rainforest), while N. swainsoniae is associated with sclerophyll forests, where it is commonly found visiting pea flowers. One of the major theories behind how one species can become two is called the ‘taxon cycle model’, where a species can expand its niche into a new habitat, which might eventually lead to reproductive or geographic isolation within the species resulting in speciation. This is possibly how Nomia (Paulynomia) now has two species. If you see either species, you could take a photo and post it to BowerBird: a citizen science website that can be a useful way for scientists to collect data from everyday people.

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Euryglossina (Euryglossina) hypochroma Bees from the genus Euryglossina are commonly found across much of Australia, with an apparent exception of most arid areas of central and western Australia. They are recognisable by an inward sloping of their clypeus (the section of integument just above their mandibles); for such small bees, this is sadly difficult to observe without the aid of a microscope. Interestingly, almost all male bees have 13 segments to their antennae, which make it easy to sex specimens. However, a few Euryglossina have only 12: an evolutionary change certain to confuse students of entomology.

Euryglossina can be found in their thousands around the essential oil-producing group of plants Myrtaceae, particularly members of the genera Eucalyptus, Corymbia, Melaleuca, Leptospermum, Tristania, Syncarpia and Syzygium. Some Euryglossina species have been found nesting in timber. For example, E. hypochroma was found nesting in a wood pile in Adelaide, South Australia, while E. xanthogena and E. philoxantha have been found on separate occasions to be nesting in beetle-infested timber.

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Apis mellifera The European honeybee is perhaps the most recognisable bee in the world. It has been transported to almost every corner of the Earth and is prized for its pollinating abilities, production of honey, propolis, beeswax and more. People often think that wild colonies of honeybees are ‘native’ bees, but they are not. They are in fact escaped ‘feral’ colonies that are adapting to Australian ecosystems. The recent invasion and establishment of Apis cerana (the Asian

honeybee) near Cairns means that A. mellifera is no longer the only introduced honeybee species in Australia. The European honeybee can be recognised by its moderate to large size (larger and hairier than A. cerana) and often black and gold-striped abdomen. European honeybees can, however, be anywhere from a bright golden orange to almost black. The species can be found across all of Australia, although it becomes less common towards the tropics.

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Threats to our bees Matt Elmer Bees are one of the most important groups of insects on the planet. Native bees are the primary pollinators of natural ecosystems and are important pollinators of many agricultural crops. They can even be more efficient at pollinating some crops than European honeybees (Apis mellifera). There has been growing evidence, however, that pollinators are declining around the globe, which could have disastrous consequences for ecosystems and agricultural productivity. The biggest threats to native bees are habitat loss, exotic species, climate change and pesticides, and the combination of these stressors acting together. Habitat loss

The loss of natural habitat is the biggest driver of decline in native bees, and all biodiversity around the world. As the human population has grown and expanded, vast amounts of natural habitat have been cleared for practices such as logging, urban expansion and agriculture. This destruction reduces the diversity, abundance and availability of floral resources and nesting sites that are needed to support wild bee diversity and maintain adequate population sizes. Because of habitat loss, the remaining vegetation can often be fragmented into smaller, separate patches. This fragmentation can lead to altered community structures, reduced connectivity and gene flow, and further decrease the diversity and abundance of native plants and bees. Although it depends on the ecosystem and type of disturbance, bees that are rare, social, specialist pollinators or above-ground nesters can be particularly threatened by the disturbance of their natural habitat. In Australia, almost 40% of native forest has been cleared since European colonisation, with clearing still continuing today. It is important to conserve the natural habitat that we have left, because it is crucial for supporting native bee communities that pollinate our ecosystems and crops throughout Australia. Exotic species

Exotic, non-native species are another major threat to native bees. The main exotic bee species around the world are European honeybees, which can now be found on every continent except Antarctica, and bumblebees (Bombus species). Exotic bees can compete with native species indirectly, by using large amounts of floral and nesting resources in a landscape, or directly, by species interactions. In Australia, honeybees have been observed directly displacing native bees from flowers. When comparing similar habitats, native bees can be over three times more abundant in regions where honeybees are absent. Exotic bees can also bring and spread threatening diseases and parasites. Honeybees and bumblebees can host many pathogens and parasites that can be spread throughout their introduced range to related, and sometimes unrelated, species. The spread of disease and parasites is particularly associated with the decline of native and managed bumblebees. The biggest threat posed by exotic bees, however, is their contribution to the pollination and spread of invasive plant species, which themselves threaten native bees. Invasive plants can contribute to habitat degradation, outcompete native plants and disrupt plant–pollinator mutualisms. Exotic bees can often visit exotic plant species more frequently than native bees and pollinate them more effectively. The close association of honeybees with the spread and seed set of lantana (Lantana camara) in Australia is a prime example of this. Australia also has several other exotic bee species. Of particular note is the buff-tailed bumblebee (Bombus terrestris) in Tasmania, a species that can effectively pollinate several threatening weed species, such as tree lupin (Lupinus arboreus). Climate change

Climate change is an enormous immediate and future threat facing species all around the world. Bees are expected to be significantly affected, from their individual biology to their interactions within ecosystems and plant–pollinator

mutualisms. As the average temperature increases around the globe, species will be threatened with extinction if they are unable to shift their ranges to stay within their temperature limits or adapt to the changes. In general, range shifts are expected to be to higher latitudes and altitudes. Bees will be particularly threatened if the plants they rely on for food and nesting are unable to shift their range or adapt in similar ways. Species already living at high latitudes and altitudes are especially at risk and, given the rate of global warming, adaption to elevated temperatures will be unlikely in many cases. Climate change can also advance the emergence dates of bees and flowers, which could be similarly problematic if the plants that bees depend on do not advance their flowering times at a similar rate. Increased temperatures can also increase competition, by leading to range expansions of bees adapted to higher temperatures, the formation of novel communities and facilitating the establishment of some exotic species. Climate change is also expected to alter rainfall patterns and increase the frequency and severity of extreme weather events such as drought, heatwaves, hurricanes, floods and bushfires. In Australia, floods can already decimate clusters of ground-nesting bees, and bushfires can destroy vast amounts of native woodland. Taking measures to mitigate the effects of climate change, particularly reducing greenhouse gas emissions, is necessary to minimise these risks. Pesticides

Another threat associated with the decline of wild and managed bees is pesticide use. Although other agrochemicals and pesticides can contribute to bee declines, insecticides are the most threatening. Insecticides are important for controlling disease-carrying insects, such as mosquitoes and agricultural pests, where appropriate insecticide application can protect a significant proportion of a crop’s yield. The problem is that many insecticides are applied unnecessarily and are ‘broad spectrum’, meaning that they kill pest and beneficial insects such as pollinators (and natural enemies of pest species). Bees can be exposed to insecticides by direct contact, ingestion of contaminated floral resources, handling of contaminated nesting materials and from insecticide sprays drifting into nesting or foraging areas. Insecticides can also result in many damaging sub-lethal effects. Neonicotinoid insecticides, for example, are particularly associated with bee declines and can seriously impair reproduction, development, foraging efficiency, immune function, learning and memory. Although much of this research has been performed on honeybees, many wild bees show similar or increased sensitivity to insecticides and are likely affected in similar ways. In Australia, broad-spectrum pesticides are incredibly cheap and commonly used. Many pesticides that are banned in Europe and elsewhere are still permitted in Australia, such as organophosphates, which are neurotoxic and a risk to human health. Reducing the impact of insecticides requires responsible application and the use of integrated pest management strategies. This includes spraying only when determined necessary by surveys of pest levels, spraying at times when bees are not active, and conserving natural enemies and nearby native vegetation. Multiple stressors

These threats often occur together, resulting in far more harmful effects. Multiple stressors are most threatening when they interact synergistically, meaning that the combined effect of two stressors can be many times greater than the sum of their individual effects. For instance, climate change is expected to greatly exacerbate the effects of habitat loss. This can lead to a greater reduction in floral resources that can increase competition and enhance the decline of bees already competing with exotic species such as honeybees. Reductions in floral diversity can also induce dietary stress, which can increase the virulence of parasites and pathogens. The severity of disease and parasites can also be enhanced by various pesticides. Some relatively harmless fungicides can also increase the toxicity of harmful pesticides up to 1000 times. One bee species in Australia that could be particularly threatened by multiple stressors is the native eusocial bee Exoneurella tridentata. Grazing by exotic species such as goats, rabbits and sheep has largely prevented the growth of new western myall (Acacia papyrocarpa) trees, which E. tridentata predominantly use for nesting. These

trees have long maturation times and rely heavily on inundating rainfall for widespread germination, which could be affected by changes in temperature and rainfall patterns associated with climate change. Already with a small population size and high competition for nest sites, the interaction between exotic species, habitat loss, and climate change could pose a serious threat to E. tridentata in the coming years. Native bees are a diverse and valuable group of insects. To conserve them we need to: reduce habitat loss; conserve native vegetation; reduce greenhouse gas emissions; use pesticides responsibly; follow integrated pest management strategies; and uphold strict quarantine protocols to restrict the arrival of exotic plants and animals. To maintain our beautiful ecosystems and ensure food production into the future, we must protect our native bees.

Western Australia Western Australia accounts for around a third of Australia’s landmass. This large state has much more to offer than its well known and vast deserts such as the Great Sandy Desert, the Gibson Desert and the Great Victoria Desert. The northern end of the state is an extensive tropical area, with high summer rainfall. While the state’s south has Mediterranean-like conditions, with most rain falling during the winter months, much of the state experiences maximum temperatures of over 40°C every year, while many inland areas can fall below freezing at night. This means that bees must be able to cope with large fluctuations in temperature. Furthermore, the huge expanse of Western Australia’s ecosystems means that the state’s bee fauna is highly diverse. The south-western region harbours many cool-adapted bee groups that are otherwise only found in eastern Australia, such as allodapine bees. Most of these bees nest in the dead flower stalks of kangaroo paw and they can be very abundant all the way from Margaret River down to Denmark. Moving further inland, the allodapine bee Exoneurella tridentata is more common, nesting in old beetle burrows in western myall and bullock bush trees. E. tridentata is a fascinating bee with distinct queen and worker castes where the queens are nearly twice as large as workers. Western Australia also has a strong fauna of stenotritid bees, including one of Australia’s prettiest bees, Ctenocolletes smaragdinus: a large brilliant metallic-coloured bee that is covered in sparse, pale hairs. Stenotritid bees occur only in Australia and are likely to have evolved before Australian broke away from Gondwana, at a time when Australia was still linked to South America via Antarctica. Another group of very interesting bees in the south-west of Western Australia is the genus Paracolletes. These large bees nest in the ground and construct unusual brood cells that have elongated and hooked necks. The food provisions in brood cells are nearly all nectar, with only a small amount of pollen; the nectar ferments with natural yeasts and the larvae consume the fermentation products. Interestingly, a very similar strategy occurs in diphaglossine bees from South America, which are related to Paracolletes, and suggests an ancient connection between Australia and South America. One of Western Australia’s prominent tourist attractions is also a prominent bee attraction! Every year a large part of the state explodes in the brilliant colours of wildflower blooms. Western Australia has over 12 000 species of wildflower, more than half of which are endemic (found nowhere else). Most of the plant diversity can be found in the south-west corner, where the Mediterranean conditions cultivate the biodiversity hotspot.

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Amegilla (Notomegilla) chlorocyanea Anywhere south of the Tropic of Capricorn, from coast to sandy desert, you are likely to see this lovely bee. Blue-banded bees are quite difficult to tell apart, but this species is the exception. You may have to look closely, but if you see a metallic blue or green colour on any of the legs then it is

Amegilla chlorocyanea. With practice, you may even learn to recognise the small notches in the last couple of bands on the abdomen. Watch out for colour variations. The blue-green bands can fade to almost white, and about one in 10 specimens has distinctly orange bands.

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Hylaeus (Euprosopis) husela Hylaeus husela is the type species for the subgenus Euprosopis, meaning that when someone wants to examine the characters that define the subgenus they use H. husela as the standard. If the subgenus were ever split into two, the original name (Euprosopis) must stay with the part containing H. husela. The subgenus Euprosopis is the most abundant of

the Australian Hylaeinae and members chiefly forage on flowers of the family Myrtaceae. This species can be found across much of Australia’s tropical north from Western Australia to Queensland. It is also very similar in appearance to the common and beautiful H. elegans, but H. husela is characterised by two striking yellow stripes running down its back.

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Homalictus dampieri Some bees will aggregate to roost at night. People will often find roosting aggregations of blue-banded bees (Amegilla sp.) or species such as Lipotriches australica. Frequently these are the male bees forming a ‘bachelor pad’ for the night. This little Homalictus dampieri was found forming one such pad, blackening the

bottom of a few leaves high overhead where it would brave the night with the other males before looking for a mate in the morning. These bees can be found from inland of Exmouth in Western Australia, north along the coast of the Northern Territory and Queensland and all the way to northern New South Wales.

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Xylocopa (Koptortosoma) parvula The great carpenter bees of the subgenus Koptortosoma, such as this X. parvula, are among the biggest bees in Australia. Together with their size and black and gold colouration, they are relatively easy to identify. This species can be found across the top end of Australia in Western Australia, the Northern Territory and Queensland, living in open

forests, shrub lands and even agricultural or urban environments. These charismatic northerners like to nest in dead branches, making linear nests of up to seven cells. Xylocopa parvula has been recorded visiting the flowers of many plants, including those of Eucalyptus, Solanum, Grevillea and several native pea plants.

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Callohesma flavopicta The beautiful yellow, black and orange Callohesma bees can be found in every Australian state and territory, but are in their highest numbers in Australia’s southern and eastern regions. Described as a single genus, recent genetic analyses on the group suggest that Callohesma and Euhesma, two closely related genera, are in fact paraphyletic and not monophyletic. This means that,

although species within the genera are genetically distinct from one another, the genera themselves are not. Some Callohesma species appear to be more closely related to Euhesma than to other members of their own genus. The inclusion of genetic information can help to confirm the members of bee genera but, as in this case, it can also do the opposite.

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Austroplebeia essingtoni Named after Port Essington in the Northern Territory, where it was first collected, this little eusocial stingless bee’s range extends from the top of the Northern Territory almost to Exmouth in Western Australia. In such hot and often dry parts of Australia, these bees have been found to survive in places where the annual rainfall is as low as 300 mm! Nests are usually built in tree hollows, but A. essingtoni seems to be the only species of Austroplebeia to also nest in wall cavities or the crevices of cliffs.

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Exoneura sp. Females of Exoneura species are readily recognised by the wedge-shaped abdomen that they share with females in the genera Braunsapis and Exoneurella. They nest within tunnels in stems and twigs, and the female whose turn it is to guard the entrance uses the flattened end of her abdomen to block the entrance hole. Unlike other Australian bees, members of all three genera feed their larvae as they grow instead of sealing them up with enough food for the whole of their

development. Interestingly, their social behaviour is intermediate between the solitary behaviour of most bees and the highly social behaviour of the small number of species in Tetragonula and Austroplebeia. One of the surprising things about Exoneura is that, although they are common on the coast and adjacent ranges of southern Australia and we know a great deal about their social behaviour, the number of species and their correct names are poorly understood.

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Hylaeus (Euprosopoides) obtusatus Occasionally nature takes pity on bee watchers and gives us species that are (relatively) easy to identify. Hylaeus obtusatus and Hylaeus rotundiceps are two such species. Both are masked bees with a yellow patch on the back, and the males have face markings that are quite distinctive with, just to help, yellow for the H. obtusatus and cream for the H. rotundiceps. The

upper face of the females has coarse sculpturing consisting of large round depressions (technically ‘punctures’) that can be seen with the naked eye or a simple magnifier. Although the females of both species look rather alike, H. obtusatus is found only in Western Australia and H. rotundiceps found only along the east Australian coast and in Tasmania.

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Thyreus waroonensis This is a genus of cleptoparasitic bees with striking patches of metallic blue setae (hairs) on their black integument. They parasitise blue-banded bees (genus Amegilla) by surreptitiously entering the host nests, making a small hole in the host brood cells and depositing their egg inside the cell. Amegilla are very fast flying bees and can zigzag suddenly and quickly when moving between

flowers. However, Thyreus is even better and can accurately follow Amegilla, staying ~15 cm behind its intended host where it cannot be seen and following it back to the host’s nest. It will then wait until Amegilla embarks on another foraging trip so it can enter the nest to lay its own egg. The young T. waroonensis larva has enlarged mandibles and kills the host egg and then consumes the cell provisions.

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Social behaviour of bees Tim Heard Most bee species are solitary. This is their primitive state; the first bee species were solitary but later some species began to cooperate with each other and evolved social behaviour. The members of these cooperating social colonies are usually, but not always, closely related to each other. A crucial aspect of social behaviour is that members of a colony are able to instantly recognise each other. They guard against entry of not only pests, predators and parasites but also others of their own species. Here we explore the social life of bees, starting with solitary bees and then societies of increasing complexity. In the case of solitary bee species, an adult female working alone finds a site and constructs her nest. She mates with a male to fertilise her eggs. But, as is the case of all bees, the males do not contribute to raising young at all; the females do everything. The nests are usually built in a protected position such as in soil or in hollow plant stems. Nests can be built in an existing cavity, as is the case for the leafcutter bees, or may be created by digging into the substrate. Carpenter bees bore into wood, and many halictid bees excavate tunnels into soil. The nest may consists of one or many cells. The female collects food to provision each cell, combining pollen and nectar to form the larval food, either as a solid mass or a semi-fluid paste. She then deposits an egg on the provisions and closes the cell. She repeats that process as many times as she can before she dies. Her eggs hatch and the resulting larvae consume the food, grow, pupate and then emerge as adults. Generations do not overlap; the nesting female dies before her offspring reach adulthood. Social behaviour occurs in other insect groups and has evolved independently many times, even within bees. Because social behaviour evolved several times independently, there is a rich and fascinating diversity of social life. The most basic kind of social behaviour is nest aggregation, as exemplified by female blue-banded bees. In this case the bees are effectively solitary, each building her own nest, but preferring to build those nests close to others. The next step in the hierarchy of social complexity is communal or egalitarian social behaviour, in which nest entrances are shared but each female builds and provisions her own cells within the nest. The advantage of a shared nest entrance is improved defence, because one female is usually present to deter intruders. Communal behaviour seems to be the typical state in Australian halictids. Look for ground-nesting Lasioglossum bees in your area, even in your backyard. You may find nest entrances shared by two or more females. An example of nest sharing from a different group is Amphylaeus morosus, a hylaeine bee from the Colletidae family that nests communally in hollow stems. Progressively more complex social behaviours have evolved, such as cooperative brood care, in which females work together to rear their young. This strategy is used by some of the Australian reed bees, members of the Exoneura genus (allodapine bees from the subfamily Xylocopinae). Females of these bees share a nest, cooperate in maintaining the nest and foraging and share egg laying more or less equally. No castes are obvious. A notable behaviour of the allodapine bees is that they progressively provision the larvae by feeding them one meal at a time. Solitary bees, and most social species, mass provision the larvae with a single load of food. Another oddity of allodapine bees is that their nests are not divided into cells, but the larvae share a common space. The next evolutionary advance is reproductive division of labour, in which some individuals dominate reproduction while others lay fewer eggs or are sterile, and do more work than the reproductive nest mates. For example, the Australian allodapine Exoneura robusta shows reproductive differentiation: each colony contains one or two reproductive queens and the remaining nest mates have small ovaries. An important factor in social behaviour is the overlap of generations: that is, a female lives long enough to meet her offspring as adults (solitary bees usually die before their offspring emerge as adults). An example of this temporary

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association between females of two generations has been reported in the large carpenter bees (Apidae, Xylocopini). Adult bees of Xylocopa (Lestis) aeratus hibernate communally, with both sexes in the nests. In spring, females may start nesting solitarily while some form associations with either their adult offspring or a formerly reproductive guarding female. The nest structure of the common Xylocopa (Koptortosoma) aruana suggests that the species may also show social behaviour. When the overlap of generations meets a caste system, we witness the evolution of eusociality, one of the most extraordinary innovations in the history of life. Eusociality has evolved also in some wasps and in all the ant and termite species alive on the planet today. Eusocial behaviour in bees may be primitive or advanced. An example of primitively eusocial species are the bumblebees and some halictid bees. The species form annual nests that are founded each year by a single female working alone. She rears a cohort of daughters that serve as her workers and the foundress stays at home to concentrate on laying eggs. The most advanced highly eusocial (truly social) behaviour has evolved in the Apidae family, with two of its tribes: the stingless bees (Meliponini) and the honeybees (Apini). In these highly eusocial bees, the queen caste totally dominates reproduction, the queen is morphologically distinct from the worker caste, there is no solitary phase in the life cycle of colonies, and colonies are perennial (they live for many years). They are also the only two groups of bees that make and store honey. Another twist to the diversity of social behaviour is that it may be obligate or facultative. In the highly eusocial stingless bees and honeybees, it is obligate because caste is inflexibly set before adulthood. But in some bee groups the individuals can change within their adult life and so are not consigned permanently to specific roles; in this case, social behaviour is facultative. In apid groups, all species are highly eusocial with little diversity in the expression of that social behaviour. By contrast, the bee family that shows the highest abundance in social diversity is the Halictidae. The halictids include species that are solitary, communal, semi-social and eusocial. In addition, eusociality in halictids is evolutionarily labile; that is, with the passage of long periods of time, a eusocial species can revert to solitary nesting, I dedicate this box to Michael Schwarz from Flinders University who has spent a lifetime investigating the natural history and evolution of bees and, with his students, has made many wonderful contributions.

South Australia Although South Australia is a big state, most of its ecosystems are arid or semi-arid. Wetter and cooler regions are restricted to the south-eastern corner and the Mount Lofty Ranges. In these areas, we can find some bee groups that are more typical of cooler climates, such as allodapine bees. In the Mount Lofty Ranges, allodapine bees can be common in introduced plants with soft stems, such as blackberry, roses and even grape vines. Kangaroo Island is especially interesting for native bees because it is now the only place in South Australia where we can find the spectacular green carpenter bee, Xylocopa (Lestis) aeratus. This bee occurs along the southern coastal regions of Australia, from New South Wales to Queensland, and in the early 1900s was recorded from South Australian localities such as Aldinga Beach and Naracoorte, but is now restricted to Kangaroo Island. The extensive arid and semi-arid regions of South Australia are dominated by colletid bees, which mostly nest in bare soil or sand. Many of these bees are tiny, especially euryglossine bees, which are usually yellow or light brown in colour and can have body lengths that get down to ~3 mm or less. The central and northern arid zones of South Australia are also host to species of the bee family Stenotritidae. This ancient family only occurs in Australia, and is the only bee family to be restricted to a single continent. Stenotritids are large bees that nest in the soil, and in arid regions of South Australia they can be found feeding on eucalypts surrounding dry watercourses.

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Brachyhesma houstoni Elizabeth Exley named B. houstoni after Dr Terry Houston, who collected the original specimens back in 1964. In the original paper she wrote, ‘It is a pleasure to dedicate this species to Mr. T. Houston who collected the specimens’. There is an international code of zoological nomenclature that guides authors in the way that they can name species. Species, for

example, can be named after friends (or even enemies), but it is frowned upon to name them after yourself. Brachyhesma houstoni can be found in South Australia and south-west Western Australia. Females can be recognised by the brown colouration on top of their head and thorax, while males require closer examination of long hairs on their hind legs.

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Homalictus urbanus Homalictus urbanus is one of the more common bee species found in Australia and has been found in every state on the mainland. The species has over 3500 sightings reported on the Atlas of Living Australia Site! This beautiful little bee is most commonly found in October and November, yet sightings have been made all year round.

I have found this bee in South Australia, Queensland and New South Wales, visiting Melaleuca, Eucalyptus and Asteraceae flowers. This species seems to be quite a generalist pollinator, which may well contribute to its ability to be active through much of the year, taking advantage of many different plant species when and where they flower.

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New Exoneura sp. This newly discovered species of Exoneura from Blinman in the Flinders Ranges is very unusual. The genus Exoneura is almost completely restricted to cool temperate forests and wet coastal regions, so finding a new species in the harsh arid landscape around Blinman presents a real puzzle. It must have some very unusual adaptations to be able to live in such a harsh region. The male shown here has large bulbous eyes that are generally used to detect flying females, and the long hairs on the face and

thorax are probably used to disseminate volatile sex pheromones that attract receptive females. But the hind legs are very unusual for a male because of their abundant and long setae. In female bees these are called ‘scopae’ and are used for collecting pollen to feed larvae in the nest. That this new species has such setae in males seems very peculiar. This new species reinforces the notion that there are a great many surprises waiting for us as we delve deeper into the Australian bee fauna.

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Hylaeus (Euprosopis) honestus This attractive masked bee is common in southern parts of the continent. It is closely related to Hylaeus elegans but does not have the red abdomen and does not appear to inhabit the drier areas that the latter species includes in its range. The species is found in South Australia, and extends along the coast to southern Western Australia, Victoria, New South Wales and South-East Queensland, as well as south to Tasmania.

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Lasioglossum (Callalictus) callomelittinum Very few details are known about L. callomelittinum or its subgenus, Callalictus. Members of this subgenus have partly or completely red or yellow thoraxes, making them visually striking bees. Lasioglossum callomelittinum can be found across much of

Australia’s south-east, extending from Adelaide to Brisbane with a specimen also being recorded from near Hobart. This individual was found foraging in coastal heath along the South Australian coast, south of Adelaide.

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Euryglossa adelaidae In the Euryglossa genus, as is the case for much of the subfamily Euryglossinae, the males and females bear little resemblance to one another: a fact that, as you can imagine, might make associating sexes to a single species very difficult without collecting them from a single nest. Some Euryglossa bees have been found nesting in the ground, with loose soil covering

their nest entrance. Bees in this group are often black or black and red and have characteristic clubbed hairs, although the latter cannot be seen with the naked eye. Despite being named E. adelaidae, Adelaide seems to be near the western edge of its distribution, which extends through Victoria and up the eastern coast.

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Pachyprosopis (Pachyprosopula) kellyi Males of the subgenus Pachyprosopula often have extensive yellow markings compared with the females and look quite different. This means that associations between males and females are made based on collection similarities, a far from certain method of telling if specimens are from the same species. Nest collections can help solve this problem, in the absence of closely related parasitic species, but nesting biology is known only

from two species in the genus and none from this subgenus. Pachyprosopis kellyi can be found in all eastern states, including Tasmania, and is found in its highest densities in Victoria. Females can be identified by an inverted ‘U’ on the top of their head, reaching half way down to their antennae. Identification of males can be more difficult, sometimes requiring the dissection of their genitalia.

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Brachyhesma sp. The Brachyhesma genus is one of my favourites, partly because this is a stunning group of bees with very pleasant colouration. The only issue with observing these spectacular bees in the wild is that they are so small and often visit the flowers high up in Eucalyptus trees, where their pale colouration blends in well with the pale flowers of the

tree. For this reason, whenever I collect a Brachyhesma species that I have never photographed before, I am always very excited to see how they look up close. Brachyhesma are thought to be ground-nesting bees. The genus is prevalent in arid regions of Australia, with 41 named species. They live in every state and mainland territory except for Tasmania.

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Lipotriches (Austronomia) australica This pretty bee is about the size of a honeybee and is a buzz pollinator. Some plants, such as tomatoes and the native hibbertias, have their pollen inside tube-like ‘poricidal’ anthers. Buzz pollinators vibrate these anthers with their abdomens, allowing the pollen grains to be released and pollination to occur. Lipotriches

australica is another communal Australian bee, with up to three females sharing a nest, but each female constructs and provisions her own brood cells. They often nest in dense aggregations, sometimes with many tens of thousands of nests per hectare when conditions are right.

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Exoneura sp. Exoneura is a genus of Australian bees that have small female colony sizes but very complex social behaviour. However, the males are interesting for quite separate reasons. In many species, the males have very long and dense hairs covering the front of their face and their thorax. These hairs (called setae) help disseminate volatile sex pheromones that attract females. Sometimes males will aggregate in sunny but protected

spaces in the forest, probably because their combined scents will attract more females. The behaviour of males coming together in order to better attract receptive females is called ‘lekking’ and occurs in many animal species, ranging from flies through to birds. Exoneura males also have enlarged compound eyes that allow them to better recognise females who are attracted into their leks.

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How to find native bees James Dorey I am often asked, ‘How do you find so many bees?’ This is far from a silly question, because if you are reading this book you are probably very interested in finding some amazing pollinators in the flesh (or perhaps in the exoskeleton). ‘Where should I look?’ is often the first question that people will ask, but ‘where’ is not the only factor to consider when looking for bees. The first thing that you likely need to do is loosen your preconceptions of what a bee might look like. This is because if it is not a European honeybee or bumblebee many people will not recognise it as a bee. For example, the world’s smallest bee is only 1.8 mm long and is from Cape York, Queensland (in the genus Quasihesma), while the world’s largest bee measured in at 39 mm long with a wingspan of 63 mm and was found in Indonesia (Megachile pluto). You will also find bees in a myriad of colours, far from the ‘standard’ orange European honeybee or black and yellow bumblebee. Looking through this book will put you in good stead to expect the unexpected! Many bee species are also restricted by time of year, time of day and the weather. During the colder months of the year, it is unlikely that you’ll find a great diversity of bees. I often find nothing more than the occasional European honeybee taking advantage of a warmer day. It stands to reason then that, until the sun is up and the air temperature is a bit more favourable, even during the warmer months of the year you are unlikely to find many bees. You might have the most success searching in between the cool of morning and the intense heat of noon: somewhere around 10 a.m., perhaps an hour earlier in the heat of Australia’s tropical north and an hour later in the temperate south and Tasmania. Bees will feed whenever they can, when they need to and when the flowers are most profitable to visit. This means that, weather permitting, they can still be found throughout much of the day. I might suggest that you avoid searching in the rain, intense heat or cold (although you may learn a thing or two about which bee species do not mind such unfavourable conditions). Now that your mind is ready to find the unexpected, and you know what seasons and weather to search, you can begin to think about the ‘where’. It may come as no surprise to many that the best place to start looking is where bees go to feed and collect food for their young or find a mate: flowering plants. These plants can be anything from the tallest gums to the smallest herbs and shrubs. Bees will search out either the best or their favoured food source. It is important to keep in mind that bees often prefer certain groups of plants, and sometimes even individual plants. This may be because a plant is flowering more, has more available floral resources or even just because it is in the sun while another is not! Thus, the best times of year to search for bees will likely depend on what time plants are flowering in your area. Native plants generally attract the most native bees, while some weeds and non-native garden plants attract no pollinators or merely introduced species such as the European honeybee or bumblebee, although there are exceptions. Such plants are not useful in your search or for native bees. Some of the best plants that I have encountered to find a diversity of bee species are the native melaleucas and eucalypts. Yet, if you look around enough on the street, around your house or in the bush you will begin to see a pattern of what flowering plants the bees near you like to visit. So, keep your eyes open for trees, herbs, shrubs, vines or any other flowering plant and you might be surprised what brings in the pollinators year after year. Bees do not spend all their time hanging around flowering plants and getting a feed. They still need to find a place to make a nest or roost for the night, which means nesting sites are another place to find bees. It is always very exciting to find bees where they nest, but it is not always easy. Most of the nests that I have found are those in the ground, often consisting of several small holes shared semi-socially by a few bees (often with one on guard just inside the

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entrance). So always keep your eyes open for little flying insects over a small section of mostly uncovered ground. Once you see these insects, watch where they land to see if they are bees and where their little entrance holes are. These nesting aggregations can be found almost anywhere: I have found aggregations in many places, ranging from a sandy unpaved sidewalk to the dirt in a local graveyard (a local haven for animals and plants). Other bees such as carpenter bees, masked bees and reed bees dig their nests out of branches, twigs or the fallen stalks of grass tree flowers. These can be more difficult to find, but may be discovered by searching for the almost perfectly round entrance hole and waiting to see the occupants come or go from the nest. Finally, the males of some bees, such as blue-banded bees, neon cuckoo bees and teddy bear bees, come in to roost at night on twigs or grass stalks. These roosting sites can host one to hundreds of bees. Roosts can be used night after night and in some cases across generations and years. To find them, you’ll need to search before they wake up, in the early evening when they are coming in to roost or (my personal favourite) at night with a torch! For many species, almost nothing is known of their biology, so be prepared to observe something new and exciting. Perhaps the most important thing of all to do is to keep your eyes peeled and to search for the tiny creatures with which you share the world.

Tasmania Tasmania is one of Australia’s most naturally stunning states, with more than 400 protected areas and many World Heritage sites making up over 20% of the state. Home to rainforests, eucalypt forests, savanna woodlands, coastal heaths and much more, but, of course, all with a temperate twist. The uninhabited southwestern corner is one of three great temperate wilderness areas left in the Southern Hemisphere. With the state’s wonderful habitats and biodiversity, it is no wonder that tourism is the second biggest contributor to the Tasmania’s economy. Tasmania has been separated from mainland Australia for ~12 000 years. This separation began with the rising sea levels associated with the end of the last glacial maximum (ice age), which isolated much of the plant and animal life from mainland Australia. As a result, most groups of Australian native bees can be found in Tasmania, although there are some notable absentees. Sadly, Australia’s favourite group of eusocial, honey-producing bees is not present – the stingless bees (the genera Austroplebeia and Tetragonula). However, this is not unsurprising, because these are mostly restricted to the more northern, warmer parts of Australia. Tasmania also lacks the big and beautiful carpenter bees (Xylocopa spp.). Despite these two missing genera, there are many bees to find in Tasmania. Halictines such as Lasioglossum and Homalictus are commonly found foraging on many plants, from tiny weeds to the large gums over the entire state. Allodapines – reed bees – can be found even in the highly disturbed forest plantations that are common in parts of Tasmania. The relatively large red and black Exoneura species can nest in the pithy stems of many small herbaceous plants and weeds, making them excellent survivors in both disturbed and undisturbed areas. Tasmania’s isolation and temperate climate may protect it from some mainland introduced pest species, but it also provides some protection against pests reaching mainland Australia. The large earth bumblebee (Bombus terrestris) is an invasive bee that was introduced to Tasmania, with the first reported sighting in 1992 in Hobart. The bumblebee has now spread to cover the entire island, becoming one of the most prominent bee species there. Commonly used to pollinate greenhouse tomatoes, the bumblebee has also been found to compete with native bees and birds, including the critically endangered swift parrot. It also pollinates and contributes to the spread of some invasive weed species.

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Heterohesma clypeata The genus Heterohesma has only two species in it: H. weiri and H. clypeata. Neither of these species seems to be particularly common or to have a particularly large range. In fact, until recently the only available sightings of H. clypeata were from around Sydney and the Blue Mountains. This individual then might well be the first found in Tasmania! Yet, it is in good company as one of the few collections of H. weiri is also in Tasmania. The rarity of sightings for this species highlights the fact that there isn’t a huge amount of collection

information for many rare insect species, particularly those that aren’t as large and charismatic as the Richmond birdwing butterfly, for example. Without such information, it is impossible to tell if a species might be at risk of disappearing, naturally uncommon or just difficult to find. But it also means that if you live in Tasmania and would like to see a wonderful bee that is otherwise unrecorded in the state such as Megachile aurifrons, there might still be hope for you yet. Ta s m a n i a

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Hylaeus (Prosopisteron) perhumilis Females from the subfamily Hylaeinae do not carry pollen on hairy scopae like many other groups of bees. Instead they swallow the pollen, using stiff brush-like hairs on the fore legs to sweep pollen from flower anthers into their mouths. The subgenus Prosopisteron is polyphyletic, meaning that some of the species in it do not share a common ancestor. Hence, it does not represent a true related group. Hylaeus

perhumilis itself is also very closely related to two Hylaeus bee species from New Zealand, suggesting a recent dispersal to New Zealand and speciation event. Hylaeus perhumilis lives in Tasmania, as well as along the coast from Brisbane south to Adelaide and Perth. It was also collected in South Africa between 1930 and 1950 as an introduced species, but has since not been collected and is likely no longer present.

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Paracolletes (Paracolletes) crassipes Paracolletes crassipes have been found foraging on Melaleuca and other Myrtaceae, with this individual and several others found foraging on lemon myrtle (Backhousia citriodora). This species can be found in Tasmania, along much of Australia’s east coast, as well as in south-west Western Australia. Paracolletes crassipes is about the length of a honeybee, but fatter and the abdomen is dark brown with a faint metallic sheen. Paracolletes was used as the type genus of the tribe Paracolletini, which is why the name had to be based on ‘paracollet-’. The tribe contained all the Australian genera in the subfamily Colletinae plus some of the genera

found in South America. Recent genetic work suggests that these genera formed a group that had arisen from a common ancestor and the authors of the study concluded that the tribe should be raised to the level of subfamily. Unfortunately, Paracolletes did not really belong in this new subfamily and so the subfamily’s name could not be Paracolletinae, but had to be the oldest available name, Neopasiphaeinae. The researchers could not decide where to put poor old Paracolletes, which was left out of all the proposed subfamilies. These suggestions will have a major effect on future studies of the genera involved.

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Leioproctus (Leioproctus) amabilis Leioproctus is one of the largest genera of bees in Australia, with a large number of species that are found in all kinds of habitats. They all live solitary lives in underground nests. They are very handsome bees and can often be quite large, with the largest about the size of a honeybee. Females of most species can be recognised by a tuft of hairs at the end of their abdomen and many have metallic tints of blue, red or bronze colouration. This genus of bees has close affinities to some bee groups in South America and dates back to the times of

Gondwana when Australia was still connected to South America via Antarctica, and polar climates were warm enough to allow forests to flourish in regions that are now covered with thick ice. Leioproctus amabilis lives in much of eastern Australia, including Tasmania, and can also be found around Adelaide and in south-western Western Australia. Individuals of this species can vary in colouration, with one individual boasting a stunning purple abdomen while another looks almost entirely golden.

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Exoneura (Inquilina) sp. Colonies of Exoneura bees consist of a relatively small number of females, but their social behaviour is very complex. This is not surprising because they have had more than 30 million years of living in social groups. Colonies can be egalitarian or have dominance hierarchies, but how this works out depends on the relatedness between females, which have a remarkable ability to assess their kinship to other females. In some ways they are like primates: social groups might be small, but interactions within each group can be very complex. Their complex living arrangements

have opened the door to a special kind of social parasitism. Females in the closely related genus Inquilina have evolved to exploit social living in Exoneura, and are able to kill Exoneura eggs and replace them with their own. Exoneura then rear the parasite’s brood as their own. Inquilina females spend their entire life in their host nests, relying on the hosts for both pollen and nectar. Indeed, Inquilina females have lost the special pollen-collecting hairs on their legs and their mouthparts are greatly reduced, because all their food is provided by the hosts.

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Megachile (Eutricharaea) maculariformis One of the most common and easily identifiable characters of many male megachiles is their enlarged and modified fore tarsi (front feet), but only half of the Megachile species exhibit this feature. These often unusual and amazing structures, such as those on this male M. maculariformis, are believed to be used during mating as a species-recognition tool. During courtship,

the male will rub his fore tarsi over the female’s eyes. Presumably, if they were not the same species the female would cease courtship and they would both resume their tasks and never speak of it again. Megachile maculariformis is common in Tasmania and along Australia’s east coast, and the range may extend along the south coast to southwest Western Australia.

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Hylaeus (Prosopisteron) quadratus This is one of the many masked bee species that are black with a patch of yellow on the back. The face marks of the male are reasonably distinctive, but females are harder to identify. Found mostly on Eucalyptus and other myrtaceous flowers, and usually within 100 km of the coast, the range of this species includes Tasmania and regions from Queensland to south-western Australia. However, it is not as abundant as some other species of Hylaeus.

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Bombus terrestris Bombus terrestris, or the large earth bumblebee, is now a common sight in Tasmania. In fact, being such a large, loud and admittedly cute bee they are hard to miss! It originates from the Northern Hemisphere but was introduced to New Zealand last century to help with the pollination of red clover, but is also used in crop pollination. Although it is certainly a Tasmanian resident now, this bumblebee was first recorded in

Tasmania in the early 1990s. Whether this introduction from New Zealand was intentional or accidental is unclear. Although the bumblebee is quite good at pollinating some agricultural plants such as tomatoes, it is also very good at pollinating noxious European weeds, competing with native bee species for resources and stealing from native plants, making it an unwelcome guest in the Australian bush.

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How to attract native bees to your garden Megan Halcroft Food resources

Suburbia, with its well-watered and tended gardens, has great potential to support bees and other pollinating insects. But how do we know what plants are right to help feed the bees? Flowering plants are the most important resource you can provide for bee health. Easy-care grasses and non-flowering trees, such as conifers, are of no use to bees and simply create a flowerless, foodless landscape. Flowers produce nectar, which contains minerals, vitamins and rich sugars, and provides adult bees with the energy they need to mate, forage and reproduce. When nectar is mixed with pollen by the female bee, this food is high in energy and protein and enables the bee grub to grow and develop into a healthy adult. Thus the cycle continues … When designing a pollinator habitat garden, it is important to remember the resources that are attractive to bees. Pollen is a good source of protein for bees and produces odours that attract bees to the plant. For these reasons, flowers that produce pollen with high-quality protein are often more attractive to bees. Plants such as Banksia, Callistemon, Melaleuca and Eremophila have both high-quality pollen-protein and abundant sugar-rich nectar. Many native gum trees produce good pollen-protein, and the large bowl-shaped nectaries attract the world’s most diverse collection of short-tongued bee species. Many of our native plants, such as Leptospermum, Xanthorrhoea and Macadamia, have only low to moderate quality pollen-protein, but the nectar content is rich and abundant. On the other hand, if you include some Acacia species, which have no floral nectaries but provide high-quality pollen-protein, your garden will offer a balanced floral resource for native bees. Many of our flowering plants are uniquely Australian, and native bees (which have co-evolved alongside them) have developed characteristic morphological structures to help them collect and transport nectar and pollen back to the nest. Long-tongued bees, such as blue-banded, teddy bear and carpenter bees, often prefer tubular-shaped flowers, such as Correa, Prostanthera and Scaevola. Female bees in the Megachilidae family, such as leafcutter bees and resin bees, have a characteristic thick scopa (region of hairs) under the abdomen to store and transport pollen. Megachilid bees often prefer pea flowers and are some of the best pollinators of these flowers. The bee alights on the flower’s keel, which hides the anthers and pollen, and as she pushes into the flower to collect nectar, the pollen is presented directly to her scopa. Once she has finished drinking the nectar, she scrabbles the pollen into her scopa and heads back to her nest. Some of the preferred pea flowers include Pultenaea, Dillwynia and Hardenbergia. Many of our small, ground-nesting bees, such as Lasioglossum and Lipotriches, forage on shallow flowers such as daisies. However, some seem to favour native flowers that require buzz pollination, such as Dianella, Tetratheca, Hibbertia and native lilies. The flowers in this group have poricidal anthers. Pollen is hidden inside the elongated anther, and is only released when a buzz-pollinating bee holds onto the anther and vibrates her thoracic muscles. This sonication shakes the pollen grains out of the slits or pore of the anther, thus enabling pollination of the flower, and freeflowing pollen grains are attracted to the bee’s branched, electrostatic hairs. There are ~100 species of native bee that are ‘specialist’ bees. These bees have evolved so closely with particular genera of plants that the young adults will emerge from the nest at the same time as their mutualistic partner begins to flower. Such bees include the female Persoonia bee, Leioproctus incanescens, which has special hooks on her forelegs to rake pollen from the anthers of Persoonia flower petals, and a polished face, to push down to the nectary.

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Most of our native bees are generalist foragers and will collect resources from a variety of flowers, including exotic ornamentals and weeds. If possible, allow some of the weeds in your lawn to flower, because these are often the first floral resource to appear in spring, providing much needed food. Nesting resources

Approximately 30% of bee species nest in pre-existing cavities. These naturally occur in the form of hollow plant stems, cracks and crevices in tree bark or rock walls, abandoned borer (wood-boring beetle or moth larvae) holes in branches and tree trunks. Our predilection for tidy gardens removes much of this potential habitat. Any sign of borer damage to trees usually results in the removal of the tree, and thus hundreds of potential nesting places for cavitynesting bees. While it is understandable that people desire tidy or hazard-free homes, it is quite simple to adapt our behaviour to help the bees. As you tidy and prune your garden shrubs, look to see if the stems are hollow, or have a pithy centre. Cut these into 20–30 cm lengths and bundle them up with a couple of pieces of thin wire. Leave some length to the wire and use this to secure the bundles to branches of trees or shrubs in the garden. Reed bees (Exoneura, Exoneurella or Braunsapis species) and masked bees (hylaeine bees) are often attracted to small cavities in hollow stems. Find a small area in the corner of your garden to stack tree limbs with deep bark ridges or rolled bark. If you have a dead tree stump or log in your garden, drill a few holes in it, to encourage resin bees to nest there. By providing a variety of hole sizes, between 3 mm and 8 mm in diameter, you will attract a wide range of species. The smaller bees prefer smaller holes, while larger bees need larger holes. You can also provide artificial habitat for some of our cavity-nesting bees. Small ‘bee hotels’ placed around the garden can look attractive or quirky. It is recommended that, rather than creating one large hotel, several small nests will better mimic nature and there is less risk of attracting predators and parasites. Most of our native bees are ground-nesting insects, burrowing up to half a metre deep into the soil. Heavily mulched soil is great for preserving soil moisture but it makes it very difficult, if not impossible, for the small (5–12 mm) bees to get to the soil surface to start nesting. Try to leave an area of un-mulched ground for these bees. By providing a variety of floral resources, which will flower throughout the foraging season (late winter to late autumn in temperate climates), bees and other pollinating insects will have access to food to rear their young. This, combined with a ready source of nesting substrate, will ensure that existing populations of bees are supported and can increase over time.

Northern Territory The Northern Territory is home to Australia’s Red Centre and parts of its tropical north. Both these regions can undergo some amazing transformations with the coming and going of the life-giving monsoonal rains. It is hard to pinpoint exactly where the Red Centre becomes the tropical north, particularly when the rains of the wet season turn red to green and cracked dry earth into vast lagoons that reflect the blue sky above. The Red Centre is vast and arid, occasionally bursting into greens and colourful flowers that partially cover the red soils when rains roll over the landscape. The tropical north is both incredibly stunning and in the wet season absolutely sweltering. The rains green the country, providing more flowers and resources for the Australian native bees. Seemingly isolated from much of the country and not possessing the status of ‘state’, the rugged and harsh Northern Territory may often be forgotten by many people, but it has not been forgotten by the bees. Like parts of Queensland, the Northern Territory has a relatively high diversity of meliponines – the stingless bees (this is relative to the rest of Australia’s states, because we have only two of the 50 Meliponinae world-wide genera: Austroplebeia and Tetragonula). Although both genera are mostly tropical, Austroplebeia extends its range much further south into the Northern Territory than Tetragonula, existing in places drier than is usual for the group. These wonderful little eusocial bees are our only native honey-producing bees, making small honeypots with flavours vastly different from those of the European honeybee. The tropical north is in fact one of the few places in Australia that the introduced European honeybee does not thrive. Most of the sampling done for bees in the Northern Territory concentrates around Alice Springs and Darwin, with a line of samples that suspiciously follows the Stuart Highway. A lack of sampling in many other parts of the Territory suggests that there might be much room for discovery!

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Undescribed Amegilla (Asaropoda) sp. Currently this species does not have a formal name, but it will be described in a revision of all Australian Asaropoda bees by the Amegilla expert Remko Leijs, at the South Australian Museum. It is very similar in appearance to Amegilla (Asaropoda) rhodoscymna, the red singer bee, which is found mostly along the Queensland coast. Unlike A. rhodoscymna, this species has

been found only in the tropical north of the Northern Territory, with all specimens but this one having been found in patches of monsoonal rainforest around Arnhem Land. This specimen was found very close to the Northern Territory’s border with Western Australia, flying along one of the impressive rocky plateaus that are common in the area.

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Brachyhesma perlutea Being bright yellow and green, you might think this bee would be easy to spot. However, measuring only 2.5 mm long, it is in fact one of the smallest and most difficult to find species of bee that I have yet to encounter. These little beauties have been found to nest in hard, rocky soil, digging down over 10 cm to make the cells for their young under the earth. This may not

seem terribly deep, but it is 40 times its body length. If I wanted to dig down 40 times my own length I’d need to go 72 m – a feat that I would not be interested in attempting! Previously, B. perlutea has only been recorded in Western Australia and South Australia, making this finding near Tennant Creek an interesting one.

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Braunsapis sp. The genus Braunsapis originally evolved in Africa and then spread to Asia and from there to Australia. These very long distance ‘Indian Ocean Rim’ dispersal routes can take many millions of years to traverse, with many new species evolving along the way. Braunsapis is an allodapine genus (like Exoneura) and is social, living in colonies of up to about half a dozen females. It does not have true queens and workers; instead it has ‘reproductive

queues’ where older females lay eggs and provide food for them. When those females become old they are replaced by the next female in the queue. Some Braunsapis species are social parasites that enter the nests of other Braunsapis species, laying their own eggs that are then tended by the host species. These socially parasitic species have reduced mouthparts and pollen-collecting hairs, because they rely on their hosts for food collection.

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Hylaeus (Rhodohylaeus) maiellus The subgenus Rhodohylaeus is a large group of hylaeine bees that have extensive red markings (rhodo in Greek means rose or red). This species also has bright yellow patches on the face, legs and thorax, and so are very striking bees. But,

despite being common and attractive, very little is known about their nesting biology. Some Rhodohylaeus species have been found nesting in old beetle burrows in dead wood, but apart from that, little else is known.

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Hylaeus (Euprosopis) elegans This little beauty I have heard called ‘one of the most common bees in Australia’. Certainly, true to this statement, I found quite a few of them! They are found over most of the country except in the tropical north and isolated Tasmania. Although they visit many flowers, they seem to

prefer Eucalyptus and Melaleuca flowers. The four individuals that I recorded were all found feeding on gums (Eucalyptus). Their stunning red, yellow and black markings of both the male and female made certain that finding one would always bring a smile to my face.

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Lasioglossum (Chilalictus) ochroma Found mostly in Australia’s arid zones, L. ochroma is recognisable by its unusually pale ochre colouration. Interestingly, this bee has some crepuscular characteristics (crepuscular animals are those that are mostly active at dusk and dawn). The pale pigmentation and large ocelli (the three

simple eyes on top of its head) indicate that it might be adapted to lower light conditions. However, this individual was caught at high noon, and though the bee was possibly inactive and hiding among the flowers or foliage, this is not quite suggestive of a crepuscular animal.

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Lipotriches (Austronomia) sp. Lipotriches is a group of ‘chunky’ bees found across Australia. They are only slightly smaller than honeybees, sometimes with green or blue metallic tints to their bodies. They are buzz pollinators and this is important for a variety of native plants that require specialised insect pollinators that can vibrate flowers at high frequencies, allowing pollen to be released from tube-like anthers. Australian native plants that require buzz pollinators include the genera Hibbertia, Solanum and Dianella. Austronomia bees build underground nests and prefer sandy

soils. The nest is a single burrow that leads to an open chamber containing multiple brood cells. Austronomia species are weakly social and two to three females can share a single nest, but without any queen or worker castes. This kind of egalitarian social behaviour is common in Australian bees, but is rare in other regions of the world. Understanding why Austronomia have evolved this kind of behaviour will help us understand social evolution more broadly. But, as with most Australian native bees, we need more information on their biology.

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Meroglossa torrida Meroglossa is a genus of ~20 species, ranging throughout Australia. They can be large bees and often have attractive yellow and red-brown markings on the face and thorax. Their nesting biology is largely unknown. Although Meroglossa has been treated as a genus, new genetic evidence suggests that it should be included within the large genus Hylaeus.

Hylaeus first evolved in Australia, where it is highly abundant and diverse, managing to disperse out of Australia only once but then succeeding in spreading to all the continents except Antarctica, including remote islands such as Madagascar, New Zealand and Hawaii – in that sense Hylaeus is a real Australian ‘success story’!

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Xanthesma (Xanthesma) flava Xanthesma are small, mostly yellow bees in the colletid subfamily Euryglossinae, and are common in arid and semi-arid regions of Australia. Unlike most other bees, they do not carry pollen on specialised hairs on their legs or abdomens, but instead swallow it and then regurgitate pollen into their brood cells. They nest in soil and have branching tunnels leading to brood cells. In at least one species, several females can share a nest, so they may be communal like Australian halictine bees. Detailed observations of their nesting biology

are likely to yield exciting data that could help us understand how social behaviour first evolved. The Euryglossinae are completely restricted to Australia, New Guinea and New Zealand, but their closest relatives belong to the genus Scrapter, which occurs only in southern Africa. Somehow or another, the common ancestor to these two bee groups managed to disperse from Australia to Africa, creating a real puzzle for understanding how ancient bees could disperse between widely separated continents.

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Museums Museums are centres of learning, discovery and wonder. Each one houses its own incredible displays to entice the public in and bring them on a journey through time and human knowledge. The importance of museums as the custodians of our art, culture, history and scientific knowledge is difficult to understate. Behind the large entry doors and incredible displays at our state museums, there is much more going on than you might think. Away from the public eye, museums hold rooms of invaluable collections, maintained and studied by dedicated researchers. The collections house the artefacts and remains of civilisations and cultures lost to time, and those that still thrive. They are also home to collections of natural history, with animals, plants, fossils and more being kept, curated and studied. Researchers in each museum explore their area of expertise and share this knowledge with the scientific world and the public. Some collections are never exhibited to the public but are available only for scientific research. This chapter features some of these specimens. Bees from the large insect cabinets and drawers of the South Australian, Victorian and Western Australian museums have been photographed so that, even if they never make it to an exhibition, they can at least be admired here. And they give a taste of the wonder and importance of museum natural history collections.

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Ctenocolletes smaragdinus Ctenocolletes smaragdinus has been found only in south-west Western Australia. It appears that the species has only one generation every year, with adults of C. smaragdinus emerging from early September with the coming of spring. During this time, males will patrol areas with flowering plants, making circuits in search of likely mates. Some males were even observed hovering in one spot near food plants, turning one way then another and then making intermittent sprints to other parts of the plant. Males were also often seen chasing one another, apparently trying to keep the best mating territory for themselves. In one observed case, a male tackled a female that was feeding on nectar to the ground where the pair struggled before flying off, with the male gripping onto the female’s back. Ctenocolletes smaragdinus can be distinguished from other Ctenocolletes by its brilliant metallic green colouration.

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Megachile (Schizomegachile) monstrosa This monster truly deserves its Latin name, monstrosa, being one of Australia’s longest bees. Only the large carpenter bees and Dawson’s bee can match its 20 mm length (though the Indonesian species Megachile pluto can be twice as long). The best place to see M. monstrosa is on flowers of Eucalyptus or related trees in hot dry areas, but be careful not to confuse it with M. semiluctuosa, which is also black and white, frequents similar areas and is rather more common. The latter species is only ~15 mm long and has more white hair, but the unambiguous differences are apparent only on close inspection. Megachile monstrosa has been found in Western Australia, Queensland and New South Wales; however, available collection records indicate that the eastern and western populations do not meet.

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Xylocopa (Koptortosoma) aruana There are several species of Koptortosoma in Australia. They are probably the most easily recognisable of all the Australian native bees. They are large black bees, usually with a bright yellow thorax, and are two to three times the size of honeybees. They build tunnels in dead branches and their large body size gives them the ability to excavate through reasonably hard wood. In some countries, species of the same subgenus can become pests as a result of their tunnelling through timber frames in houses. Females can occasionally nest in small social groups, but without queen or worker castes. Males, such as this bee, will hold territories that they patrol to drive off other rival males, often returning to a central point in their territory where they will scent-mark leaves and branches with pheromones that smell like citrus. Xylocopa aruana lives along the east and north coast from New South Wales to Western Australia and even extends as far north as Papua New Guinea.

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Palaeorhiza varicolor Bees of the genus Palaeorhiza are often very striking, with metallic colours and plentiful yellow markings. Some species, however, can be almost entirely red or black. They can be found along Australia’s east coast and as far south as Melbourne, but are also a relatively widespread group of tropical bees. Although the subfamily Hylaeinae originated from Australia, this genus appears to have recently diverged and diversified in New Guinea and subsequently re-colonised Australia. Palaeorhiza varicolor itself has been found along Queensland’s eastern coast, with this individual originating from near Cairns. The colour of the integument of Palaeorhiza bees is the result of optical interference, much in the same way that oil slicks on wet roads can produce rainbow effects. The bee’s integument has a series of interleaved layers with different optical properties. If the distance between the layers changes then the colours will be altered. This can cause nightmares for taxonomists when specimens stored in liquid and those that are stored dry can change colour, making a species seem variable in colour, when it may not be.

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Quasihesma gigantica Quasihesma is the genus that holds Australia’s smallest bees, many of which are minuscule: just 2 mm long! Quasihesma gigantica is found only in the Northern Territory and was named in 1974 by Elizabeth Exley, who has described well over 200 bee species and educated and encouraged many of Australia’s important pollinator researchers. Clearly, she also had something of a sense of humour to dub a 3.5 mm long bee ‘gigantica’. However, the bee may deserve the name, being almost twice the length of many others in the genus Quasihesma. Elizabeth collected this individual in 1973 and, being a ‘paratype’, this specimen was used in the original classification of the species back in 1974.

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Lasioglossum (Chilalictus) hemichalceum This is a small bee with a black head and thorax and red or black abdomen. Lasioglossum hemichalceum is one of the most interesting bees in Australia because of its ‘communal’ social behaviour. Each underground nest can contain dozens of females and multiple tunnels, but all tunnels share a single entrance at the soil surface. Each female constructs and provisions her own brood cells, but all females help defend the nest by spending some time guarding the nest entrance. However, there are two even more remarkable features of this bee. Even though females do not cooperate in provisioning brood, they exchange food with each other in a behaviour called trophallaxis. It is likely that this helps the colony as a whole to survive by ensuring that each female has enough food to contribute to colony defence. The second remarkable feature is that many, but not all, of the sons that females produce have enormous mandibles and heads but tiny wings and so are unable to fly. Other ‘normal-appearing’ males can fly and search for mates outside of the nest. This system is quite different from that seen in other eusocial bee species, where sexual dimorphism and caste systems are restricted to the females of the colony. So why on earth would a species have such distinct castes among their males? They use these mandibles to fight with other males in the nest, with the losing males being killed. The surviving males then mate with newly emerging females in the colony. This bee species therefore shows behaviours that are both highly cooperative (females sharing food and nest defence), as well as highly aggressive (males fighting to the death with the winner mating with females). A further explanation might be that, because of size of big-headed male heads coupled with their small antennae and wings, these males might be useful in nest defence. Their big heads house large muscles attached to powerful mandibles that could be used to deter predators and parasitoids from attacking the nest. Found across much of Australia, except for the tropical parts of Western Australia, the Northern Territory and Queensland, this interesting species is quite common but to find a big headed male you might have to look a little lower than you normally would.

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Amegilla (Asaropoda) dawsoni Dawson’s bee is the largest Anthophorini bee (the tribe of bees that contains blue-banded and teddy bear bees) in Australia. These bees form very impressive nesting sites, with anywhere between two and 10 000 nests in a single area! Females make the nests, digging into the ground and provisioning cells. Larger males fight with each other to hold territories around these nest sites, whereas smaller males patrol forage plants in search of virgin females with which to mate. Native to parts of Western Australia, these bees appear to breed only from mid-July to mid-September, with young developing and then entering a state of diapause (suspended development) for a year or more.

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Xylocopa (Lestis) aeratus This bee builds nests in dead Xanthorrhoea flower scapes or in the dead, rotting trunks and branches of Melaleuca trees. It is another communal bee, with up to four females sharing a single nest, with each female building and provisioning her own brood cells. However, nest sharing is not always peaceful and dominant females can ‘bully’ their subordinate nest mates to get food from them. There are only two species of Lestis, ranging from New Guinea down to Kangaroo Island in South Australia. This is a buzz-pollinating bee and studies have shown that it could be useful for helping to increase tomato yields in greenhouse crops. In the south, L. aeratus used to occur in western Victoria and mainland South Australia, though that southern distribution has now reduced to just Kangaroo Island.

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Hyleoides zonalis Hyleoides is a genus of striking hylaeine bees. They have bands of red or orange across their body, which is warning colouration, indicating that they can have dangerous stings. Hylaeine bees do not have a region of specialised hairs (the scopa) on their legs for carrying pollen; instead they swallow pollen and then regurgitate it when back in their nests. The lack of a scopa makes their legs look thin and waspish and, combined with their warning colouration and darkened forewings that mimic the dark folded wings of wasps, they are easily mistaken for wasps. The evolution of red and black warning colouration by multiple species is called Müllerian mimicry. If a predator such as a bird is stung by a yellow and black wasp, it is more likely to avoid attacking other insects with the same kind of colouration.

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The importance of museums Michael Batley In 2015, two eminent scientists named a new species of bee-fly from Africa in a leading taxonomic journal. The publication re-ignited a vigorous debate among taxonomists via a flurry of publications with multiple authors. One had 123 signatories and others had as many as 79 authors. Eventually, in 2017, the editor of one journal announced no further articles on the topic would be accepted, pointing out that the topic was not new and, anyway, the rules had been clarified. So, what caused all the fuss? The bee-fly was large, black and yellow and an excellent mimic of an African carpenter bee. Its describers were able to take 19 good photographs and to capture a specimen. But it escaped! From the photographs, it was possible to work out that it belonged to a genus containing rare species and, as experienced dipterists (experts on flies), the describers knew that it was unlike any known species. They argued that in these special circumstances the rules for naming new species, contained in the International Code of Zoological Nomenclature, allowed them to create a new name without depositing a specimen in ‘an institution that maintains a research collection, with proper facilities for preserving them and making them accessible for study’. The strength of the reaction to this suggestion demonstrated how important the idea of type specimens is to taxonomists. Simply stated, what would happen if there turned out to be two similar bee-fly species that could be distinguished only by small features invisible in the photographs? Which one would carry the published name? Even the dipterists that were involved and the few people who suggested other possible exceptions to the rule emphasised that deposition of type specimens should occur if at all possible. Examples of the need for such a rule occur regularly in the experience of all taxonomists. Such a case arose when I and two colleagues began looking at bee species in the genus Amegilla and the British Museum generously allowed us to examine type specimens from their collection. Many of you will be familiar with the blue-banded bees and teddy bear bees often seen in suburban gardens. Both are in the genus Amegilla, but are usually placed in separate subgenera. The colour and presence (or absence) of bands make them easy to separate, but there are less obvious differences in the shape of the face and other parts of the body. In 1854, Frederick Smith of the British Museum described the Australian teddy-bear bee for the first time and called it Amegilla bombiformis, but added a note saying that he had another specimen that was a bit smaller and had a patch of black hair on the hind leg. Size and even hair colour may vary within species, so he decided to call the smaller specimen variety α (alpha) of A. bombiformis. When the English-American bee expert Theodore Cockerell looked at the same specimen in 1904, he thought the size and colour differences were sufficient to treat the smaller specimen as a new species, which he called Amegilla alpha. (Actually, he called it Saropoda alpha, but that genus became part of Amegilla in 1956.) To our surprise, when we looked at the same specimen, it was immediately obvious that it belonged in the banded subgenus and was not a ‘teddy-bear’ species at all. The shape of the face and the underside of the abdomen were unmistakeable. The reason that this was not apparent in 1904 was not due to technological changes, but simply because there are now many more specimens available and hence patterns are easier to discern. Because the earlier authors were unaware of these patterns, their published descriptions contained no clues that would have led us to this conclusion had there been no type specimen. We were able to confirm our conclusion with the aid of molecular evidence, but simply seeing the holotype was enough. Similar cases were encountered many times before we reached the conclusion that quite a lot of Amegilla species had been given more than one name. A list of 43 names was reduced to 10 genuinely different species, and only four

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new species were added. Without safe storage of type specimens in museum collections we would still have a long list of unnecessary names and little idea of what they meant. So, the reason for the vigorous reaction to the bee-fly publication was that taxonomists are exposed, almost daily, to the importance of type specimens and many have experienced long waits and difficult expeditions in order to collect rare species. There are, however, special cases (but not ones that are likely to apply to insects). The endangered Galápagos pink land iguana, featured in David Attenborough’s television series about the islands, was named in 2009 without a preserved type specimen. The holotype was micro-chipped and released. This is the sort of exception to the rule about type specimens that the International Commission on Zoological Nomenclature indicated might be allowed in Declaration 45, published in 2017 to clarify the section of the code that gave rise to the bee-fly controversy. Looking back over the episode, one cannot help noticing how carefully the participants prepared their cases and the thoroughness with which they were prepared to re-examine the rules of taxonomy. But there was an underlying concern that the debate might be misunderstood outside their profession and that all type specimens could be replaced with photographs. Perhaps it is best to let one group of participants speak in their own words. ‘A transformation of the nature and purpose of natural history museums seems to be underway in the mind of some administrators and decision makers: from centres of knowledge development and provision of the past and present to entertainment centres largely devoid of scientific background.’ – Dalton Amorim et al. (2016) Timeless standards for species delimitation. Zootaxa 4137 (1): 121–128.

Glossary Abdomen – the third functional region of an insect’s body behind the thorax. The abdomen includes the propodeum, which is fused to the thorax to form the mesosoma in bees, wasps and ants (see Fig. 1, p. 2). Aggregation – a group of bee burrows in the ground. Each burrow can be inhabited socially (by multiple bees) or solitarily (by one bee). Some aggregations are believed to be caused by limited nest sites, but some bees seem to prefer nesting near other bees, making it a social phenomenon. Allodapini (adj. allodapine) – a tribe of bees in the subfamily Xylocopini (family: Apidae) that nest in hollow or pithy stems or in other cavities. Allodapines are unusual because they raise their brood without the use of cells. Antennae – two long sensory organs that protrude from the head of an insect (see Fig. 2, p. 3). Arid – a dry climate. Biotic pollination – pollination of plants by animal vectors. Brood – bee larvae. Brood comb – the beehive cells where a queen lays an egg (eusocial bees). Buzz pollination (or sonication) – a method used by many bees, including blue-banded bees and carpenter bees, to get pollen out of a plant’s anthers (pollen-containing organ of a flower). Buzz-pollinating bees can vibrate their flight muscles or head very fast, causing the pollen to dislodge from the anther. Caste – specialised individuals within a bee species that carry out different tasks. For example, in eusocial bees the queen and worker bees are two different castes that carry out different tasks (laying eggs and tending the hive respectively). Cleptoparasite – a species that lays its eggs in the nest of another, leaving the host species to raise the young, such as a cuckoo bird (or bee). Clypeus – the region above the mandibles on bees (see Fig. 2, p. 3). Colletidae (adj. colletid) – a family of morphologically and species diverse bees that nest in soil, rotting wood, or pre-existing holes in stems, wood or even volcanic rocks. All colletid species are solitary, with the notable exception of the Australian Amphylaeus morosus. Compound eye – an eye that is made up of many smaller visual units, found on insects and crustaceans (see Fig. 2, p. 3). Crepuscular – describes organisms that are mostly active during twilight. Diapause – a period of halted development in insects or other animals, often during unfavourable conditions (for bees, this is often winter). Ecosystem – a community formed through the interaction of both the biotic (living) and abiotic (non-living) components of an environment. Euryglossinae (adj. euryglossine) – an Australian subfamily of Colletidae with small to minute species that nest in the ground and rotting wood. Eusociality – is defined by species that have cooperative care of brood, overlapping generations, and have a division of labour with both reproductive and non-reproductive individuals. Honeybees and stingless bees are eusocial. Exotic species – a species that is not native to a certain area. Often these species are intentionally or accidentally moved outside their native range by humans. 200

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Facultative – ‘optional’. For example, a facultatively social species would be one that can either live socially or solitarily, such as the stem-nesting Australian bee species Amphylaeus morosus. Fore tarsi – the last group of segments on the front legs of insects (see Fig. 1, p. 2). Gondwana – a supercontinent that broke up 180 million years ago and included present-day South America, Africa, Arabia, India, Madagascar, Australia and Antarctica. Halictidae (adj. halictid, halictine) – a family of bees with members that largely burrow in soil (sometimes in rotting wood). Commonly called ‘sweat bees’, halictids are the second most species diverse group of bees in Australia, after the colletids. Hylaeine – a subfamily of Colletidae with species that have short and usually sparse hair that can make them resemble sphecid wasps. Hylaeinae has a worldwide distribution but is most diverse in the Australian region with all genera except for Hylaeus restricted to Australia, New Guinea, New Zealand and neighbouring islands. Integument – the tough protective outside layer of insects that forms their exoskeleton. Larvae – developing bee young before pupating and becoming an adult. Lekking – males aggregating together to better attract females. Males then compete with one another for a mate. Mandibles – a pair of appendages near an insect’s mouth that are often used for cutting, crushing or grasping. Some uses include: eating, nest construction and fighting (see Fig. 2, p. 3). Monophyletic – a group of organisms that have evolved from a common ancestor (cf. Paraphyletic). Müllerian mimicry – when one or more species mimics another to warn off potential predators. These species can be distasteful or, in the case of bees, have a powerful defence such as a sting. Once a predator experiences the defence of one of these species, it learns to avoid them all. Mutualism – when two species have a relationship that is mutually beneficial to both. Nest – a place where bee young are reared that is made by either the mother or worker bees. Niche – the ecological place that a species fills or could fill. It takes into account both the abiotic and biotic factors that a species prefers in its environment. Obligate – ‘by necessity’. For example, an obligately social bee species needs to exhibit social behaviour to survive and reproduce (i.e. worker bees are sterile and cannot reproduce, while the fertile males and females require the workers to provide food and shelter for them). Ocellus – a simple eye, with a single lens. Bees have three of these on top of their heads (see Fig. 2, p. 3). Paraphyletic – a taxonomic group that is descended from a common ancestor, but does not contain all the descendants within that group. Bees, for example, form a monophyletic group within the superfamily Apoidea, which leaves the rest of the superfamily, the sphecoid wasps, as a paraphyletic group called the Spheciformes. Parasite – an organism that lives on or within another organism (a host) and uses that host’s resources to fuel its own growth and reproduction. Typically, parasites do not kill their host. Parasitoid – an insect that develops as a parasite but eventually kills its host. Paratype – one of the specimens used to originally describe a group and on which the original descriptions were based (cf. type specimen). Pathogen – an infectious agent that can cause a disease, such as bacterium, virus, protozoa, fungus etc. Pheromone – a chemical cue that is meant to elicit a response from members of the same species (conspecifics). For example, when some social bees, such as honeybees, sting or bite they might release a pheromone to stimulate an aggressive response in conspecifics. Glossary

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Polyphyletic – a group of organisms that do not have a common ancestor. Members of this group should not be placed within the same taxon. Primitive eusociality – species in which a colony is started by a single female that constructs and provisions a nest; when her daughters emerge there is division of labour between the foundress (or queen) and the workers. Workers and queens look morphologically similar but differ in behaviour and physiology. These colonies are usually ephemeral, unlike the more permanent eusocial colonies. Propolis – a mixture of bee saliva, beeswax and exuded plant materials (e.g. sap) used to seal gaps in a hive. Pupa – a life stage of an insect where it undergoes metamorphosis into an adult. Pupate – when a larva becomes a pupa. Range shift – a change in geographic distribution as a natural response of species and populations to changes in climate. Revision – a review of the species classifications or taxonomic ranks within a group of organisms. Often revisions are done when new material and knowledge become available. Scopa – a region of hairs (setae) on a bee that is used to carry pollen. The scopa can be under the abdomen or on the rear legs of a bee. Semi-social bees – a colony of bees with multiple females, often sisters, that show division of labour. These colonies will have a principal egg-layer and one or more foragers. These bees might be considered primitively eusocial. Setae – rigid hair-like structures, particularly of an invertebrate. Sexual dimorphism – morphological differences between the sexes of a species, meaning that males and females look different. Social parasitism – a parasitising species relies upon a host species to provision for their young in a mixed-species colony or nest. Solitary bees – solitary bees are those in which a single female will construct, provision and lay eggs in her own nest with no help from other bees, and usually dies before her offspring mature. Speciation – the formation of a new species, distinct from its ancestors or closely related species. Sting – when a bee injects venom into another animal with her modified ovipositor. This differs from a bite, which does not inject venom in bees and uses the mandibles. Taxonomic rank – the relative rank of taxonomic groupings (e.g. kingdom, phylum, class, order, family, genus, species). Genus, subgenus and species-level names are all shown in italics throughout the book, whereas higher taxonomic ranks are not (e.g. Lasioglossum (Chilalictus) ochroma is in italics, but its family Halictidae is not). Thorax – the middle segment of a bee, between the head and the abdomen (see Fig. 1, p. 2). Trophallaxis – an exchange of regurgitated liquids between social insects and their larvae or other adults. Tumulus – a loose mound of soil or sand around the entrance of a nest resulting from its excavation. Type specimen (or holotype) – the primary specimen on which a published name is based. If it were to be found that there were two similar species, the holotype specimen would be the reference to be used to decide which species should carry the previously published name. Virulence – the ability of a pathogen to cause disease or damage a host.

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Appendix of species by family Species

Page

Apidae

Species

Page

Hylaeus (Prosopisteron) quadratus

148

Amegilla (Asaropoda) bombiformis

6

Hylaeus (Rhodohylaeus) maiellus

164

Amegilla (Asaropoda) dawsoni

190

Hyleoides zonalis

194

Amegilla (Notomegilla) chlorocyanea

86

Leioproctus (Exleycolletes) cristatus

20

Amegilla (Zonamegilla) asserta

10

Leioproctus (Leioproctus) amabilis

142

Undescribed Amegilla (Asaropoda) sp.

158

Leioproctus (Leioproctus) plumosus

62

Apis mellifera

78

Undescribed Leioproctus (Exleycolletes) sp.

12

Austroplebeia australis

32

Meroglossa torrida

172

Austroplebeia essingtoni

96

Pachyprosopis (Pachyprosopis) haematostoma

72

Bombus terrestris

150

Pachyprosopis (Pachyprosopula) kellyi

122

Braunsapis sp. (Qld and NT)

38, 162

Palaeorhiza (Cnemidorhiza) disrupta

34

Exoneura (Inquilina) sp.

144

Palaeorhiza varicolor

184

Exoneura sp. (NSW, WA and SA)

22, 98, 128

Paracolletes (Paracolletes) crassipes

140

New Exoneura sp.

114

Quasihesma gigantica

186

Tetragonula carbonaria

46

Xanthesma (Xanthesma) flava

174

Thyreus nitidulus

24

Halictidae

Thyreus waroonensis

102

Homalictus dampieri

90

Xylocopa (Koptortosoma) aruana

182

Homalictus punctatus

60

Xylocopa (Koptortosoma) parvula

92

Homalictus urbanus

112

Xylocopa (Lestis) aeratus

192

Lasioglossum (Callalictus) callomelittinum

118

Colletidae

Lasioglossum (Chilalictus) hemichalceum

188

Amphylaeus (Amphylaeus) morosus

70

Lasioglossum (Chilalictus) lanarium

18

Brachyhesma houstoni

110

Lasioglossum (Chilalictus) ochroma

168

Brachyhesma perlutea

160

Lasioglossum (Chilalictus) sp.

68

Brachyhesma sp.

124

Lasioglossum (Chilalictus) veronicae

66

Callohesma flavopicta

94

Lasioglossum (Parasphecodes) lithuscum

14

Euryglossa adelaidae

120

Lipotriches (Austronomia) australica

126

Euryglossina (Euryglossina) hypochroma

76

Lipotriches (Austronomia) sp.

170

Euryglossina (Microdontura) mellea

42

Nomia (Hoplonomia) rubroviridis

52

Heterohesma clypeata

136

Nomia (Paulynomia) aurantifer

74

Hylaeus (Euprosopis) elegans

166

Megachilidae

Hylaeus (Euprosopis) honestus

116

Megachile abdominale

36

Hylaeus (Euprosopis) husela

88

Megachile apicata

44

Hylaeus (Euprosopoides) obtusatus

100

Megachile aurifrons

50

Hylaeus (Euprosopoides) ruficeps

40

Megachile (Eutricharaea) maculariformis

146

Hylaeus (Gnathoprosopis) albonitens

48

Megachile ustulata

8

Hylaeus (Gnathoprosopoides) philoleucus

64

Megachile (Schizomegachile) monstrosa

180

Hylaeus (Macrohylaeus) alcyoneus

16

Stenotritidae

Hylaeus (Prosopisteron) perhumilis

138

Ctenocolletes smaragdinus

178 203

Further reading Heard T (2015) The Australian Native Bee Book: Keeping Stingless Bee Hives for Pets, Pollination and Sugarbag Honey. Sugarbag Bees, Brisbane. This multi-prize winning and best-selling book by the ex-CSIRO scientist Tim Heard provides a brilliant general description of native bees, with a focus on Australia’s native stingless bees. Full of wonderful illustrations and images, this is a great book to purchase if you want to learn more about stingless bees and how to keep them. Houston T (2018) A Guide to Native Bees of Australia. CSIRO Publishing, Melbourne. A Guide to Native Bees of Australia provides a detailed introduction to the estimated 2000 species of Australian bees. Illustrated with stunning photographs, it describes the form and function of bees, their life-cycle stages, nest architecture, sociality and relationships with plants. It also contains systematic accounts of the five families and 58 genera of Australian bees. Photomicrographs of morphological characters and identification keys allow identification of bees to genus level. Michener CD (2007) The Bees of the World. 2nd edn. The John Hopkins University Press, Baltimore, MD. Written by Charles (‘Mich’) Michener, this book covers the diversity, biology, ecology, evolution and behaviour of bees worldwide. Mich was a towering researcher of bees worldwide, universally loved and respected by his students and colleagues, and this magnum opus is a truly monumental work, over 900 pages in length. His book covers all aspects of bees, but especially focuses on taxonomy and so might be more useful for more technically minded readers. NSW DPI (2016) Australian Native Bees: A Practical Handbook. NSW Department of Primary Industries, Orange, NSW. Like the book you are reading now, the Australian Native Bees book brings together expertise from many of Australia’s leading native bee researchers. It covers general bee biology and ecology, some basic bee identification, how to keep and encourage the broad diversity of native bees, including the solitary ones, and more. Rayment T (1935) A Cluster of Bees. Endeavour Press, Sydney. Tarlton Rayment was a prolific, and also very eccentric, amateur bee researcher working from his home in a bayside suburb of Melbourne up until the late 1950s. He described hundreds of Australian native bees, along with wasps and other insects. He was regarded as an Australian version of entomologist Jean-Henri Fabre and he lovingly detailed the behaviour and nesting biology of bees. Many of his accounts of bee biology were combined with his whimsical and philosophical observations on nature and the human condition. This is a very rare book, but most Australian state libraries will have a copy. It provides a unique insight into a long-bygone era and is a true gem. 204

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Index Page numbers in bold refer to main species account. abdomen  2, 9, 17, 152, 175, 196 aggregation  19, 21, 63, 91, 104, 127, 132 Allodapini  58, 84, 104, 108, 134, 163 Amegilla (Asaropoda) bombiformis  6–7, 196 dawsoni  181, 190–1 undescribed sp.  158–9 Amegilla (Notomegilla) chlorocyanea  86–7, see also bluebanded bee Amegilla (Zonamegilla) asserta  10–11, see also blue-banded bee Amphylaeus (Amphylaeus) morosus 58, 70–1, 104 antennae  2, 3, 9, 45, 54, 67, 77 Apis mellifera  1, 15, 26, 30, 54, 56, 78–9, 81–2, 106, 130, 156, see also honeybee arid zones  3, 23, 58, 77, 108, 115, 125, 156, 169, 175 semi  108, 175 Austroplebia  134, 156, see also stingless bee; sugarbag bee australis  32–3, 54 essingtoni  96–7 banksia bee  16–17 biotic pollination  54, 56 blue-banded bee  10–11, 53, 86–7, 91, 103, 104, 132, 152, 191, 196 Bombus terrestris  1, 81, 106, 130, 134, 150–1, see also bumblebee Brachyhesma houstoni  110–11 perlutea  160–1 sp.  124–5 Braunsapis  38–9, 54, 99, 154, 162–3 brood  15, 17, 19, 25, 30, 47, 145, 189 care 104 cells  25, 61, 69, 84, 103, 127, 171, 175, 189, 193 comb 47 bumblebee  1, 81, 106, 130, 134, 150–1 buzz pollination  54, 127, 152, 171, 193

Callohesma flavopicta  94–5 carpenter bee  4, 30, 56, 92–3, 104, 106, 108, 132, 134, 152, 181, 182–3, 192–3, 196 caste  2, 71, 84, 104, 106, 171, 183, 189 system 106 cleptoparasite  25, 103 Colletidae  21, 58, 71, 104, 108, 175 crepuscular 169 Ctenocolletes smaragdinus  84, 178–9 cuckoo bee  102–3, 132 neon  24–5 diapause 191 ecosystem  26, 28, 30, 56, 79, 81, 83, 84, 108 service  26, 56 Euryglossa adelaidae  120–1 Euryglossina (Euryglossina) hypochroma  76–7 Euryglossina (Microdontura) mellea  42–3 Euryglossinea  4, 58, 108, 121, 175 eusocial  33, 56, 61, 82, 97, 106, 189 Exoneura  22–3, 39, 54, 98–9, 104, 114–15, 128–9, 134, 154, 163 Exoneura (Inquilina)  144–5 exotic species  26, 81–3, 154 facultative social behaviour  106 Gondwana  21, 84, 143 Halictidae  104, 106 Heterohesma clypeata  136–7 Homalictus  4, 30, 54, 67, 134 dampieri  90–1 punctatus  60–1 urbanus  112–13 honeybee  1, 15, 26, 30, 54, 56, 78–9, 81–2, 106, 130, 156 Hylaeine  104, 154, 165, 195, see also masked bee Hylaeus (Euprosopis) elegans  89, 117, 166–7 205

honestus  116–17 husela  88–9 Hylaeus (Euprosopoides) obtusatus 41, 100–1 ruficeps  40–1 Hylaeus (Gnathoprosopis) albonitens  48–9 Hylaeus (Gnathoprosopoides) philoleucus  64–5 Hylaeus (Macrohylaeus) alcyoneus  16–17, see also banksia bee Hylaeus (Prosopisteron) perhumilis  138–9 quadratus  148–9 Hylaeus (Rhodohylaeus) maiellus  164–5 Hyleoides zonalis  194–5 integument  77, 103, 185 larvae  15, 19, 63, 73, 84, 99, 103, 104, 115, 154 Lasioglossum (Callalictus) callomelittinum  118–19 Lasioglossum (Chilalictus) hemichalceum  188–9 lanarium  18–19 ochroma  168–9 sp.  68–9 veronicae  66–7 Lasioglossum (Parasphecodes) lithuscum  14–15 leafcutter bee  9, 36–7, 44–5, 50–1, 104, 152 Leioproctus (Exleycolletes) cristatus  20–1 undescribed sp.  12–13 Leioproctus (Leioproctus) amabilis  142–3 plumosus  62–3 lekking 129 Lipotriches (Austronomia) australica 91, 126–7 sp.  170–1 masked bee  132, 154, see also Amphylaeus, Hylaeus, Hyleoides, Meroglossa and Palaeorhiza species Megachile  54, 130, 181 abdominale  36–7 apicata  44–5 aurifrons  50–1, 137 ustulate  8–9

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Megachile (Eutricharaea) maculariformis  146–7 Megachile (Schizomegachile) monstrosa  180–1 Megachilidae  9, 51, 152 Meroglossa torrida  172–3 monophyletic 95 Müllerian mimicry  195 mutualism  81–2, 152 nest  3, 4, 9, 17, 19, 26, 28, 30, 47, 56, 82, 104, 106, 154, 189, 191 niche 75 Nomia (Hoplonomia) rubroviridis  52–3 Nomia (Paulynomia) aurantifer  74–5 obligate social behaviour  106 Pachyprosopis (Pachyprosopis) haematostoma  72–3 Pachyprosopis (Pachyprosopula) kellyi  122–3 Palaeorhiza (Cnemidorhiza) disrupta  34–5 Palaeorhiza varicolor  184–5 Paracolletes (Paracolletes) crassipes  140–1 paraphyletic 95 parasite  28, 63, 81, 82, 104, 154 social  39, 123, 145, 163 parasitoid  17, 19, 28, 63, 189 paratype 187 pathogen 81–2 pheromone  30, 115, 129, 183 polyphyletic 139 propolis  26, 79 pupate 104 Quasihesma  130 gigantica  186–7 range shift  82 reed bee see Braunsapis and Exoneura species resin bee  8–9 revision  43, 159 scopa  9, 115, 139, 152, 195 semi-social  106, 130 sexual dimorphism  189 speciation  75, 139 Stenotritidae  84, 108, 178–9

sting  1, 19, 58 stingless bee  1, 30, 32–3, 46–7, 54, 56, 96–7, 106, 134, 156, see also sugarbag bee sugarbag bee  1, 30, 32–3, 46–7, 54, 56, 96–7, 106, 134, 156, see also stingless bee sweat bee see Lasioglossum sp.

waroonensis  102–3 trophallaxis 189 tumulus 21 type specimen  196, 198

teddy bear bee  6–7, 132, 152, 191, 196 Tetragonula  99, 134, 156, see also stingless bee; sugarbag bee carbonaria  46–7, 54, 56 thorax  2, 25, 30, 67, 71, 111, 115, 119, 129, 165, 173, 183, 189 Thyreus nitidulus  24–5

Xanthesma (Xanthesma) flava  174–5 Xylocopa  54, 106, 108, 134 Xylocopa (Koptortosoma) see carpenter bee aruana  182–3 parvula  92–3, 106 Xylocopa (Lestis) aerates  106, 108, 192–3, see also carpenter bee

virulence 82

Index

207