Boom and Bust: Bird Stories for a Dry Country

boom & bust

Bird Stories for a Dry Country

Libby Robin • Robert Heinsohn • Leo Joseph [Editors]

boom & bust Bird Stories for a Dry Country

,IBBY2OBINs2OBERT(EINSOHNs,EO*OSEPH [Editors]

© in this edition CSIRO 2009 Copyright in the individual contributions is retained by the authors. All rights reserved. Except under the conditions described in the Australian Copyright Act 1968 and subsequent amendments, no part of this publication may be reproduced, stored in a retrieval system or transmitted in any form or by any means, electronic, mechanical, photocopying, recording, duplicating or otherwise, without the prior permission of the copyright owner. Contact CSIRO PUBLISHING for all permission requests. National Library of Australia Cataloguing-in-Publication entry Boom and bust : bird stories for a dry country / editors Libby Robin, Robert Heinsohn, Leo Joseph. 9780643096066 (hbk.) Includes index. Bibliography. Birds – Behavior – Australia. Birds – Effect of drought on – Australia. Nature – Effect of human beings on. Robin, Libby, 1956– Heinsohn, Robert. Joseph, Leo. 598.0994 Published by CSIRO PUBLISHING 150 Oxford Street (PO Box 1139) Collingwood VIC 3066 Australia Telephone: Local call: Fax: Email: Website:

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

Front cover: Melopsittacus undulatus. From John Gould’s The Birds of Australia. Original copy owned by the National Library of Australia. Call number RARE RBN ef F4773. Back cover: Masked woodswallows. From John Gould’s The Birds of Australia. Set in 10.5/16 Adobe ITC New Baskerville and Optima Edited by Janet Walker Cover and text design by James Kelly Typeset by Desktop Concepts Pty Ltd, Melbourne Printed in China by 1010 Printing International Ltd All illustrations are taken from John Gould’s The Birds of Australia, except for the illustration on page 147 which is reproduced courtesy of Frank Knight. The authors thank Andrew Isles for help sourcing the illustrations. 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.

FOREWORD It is with great pleasure that I recommend this book to you. It brings together some great writing by historians, ecologists, archaeologists, ornithologists and anthropologists. Each writer begins with the story of a bird and considers in different ways how it has adapted to its dry environment. People have changed the environment in Australia, dramatically – twice – through two great human invasions, as the environmental historian George Seddon said. The first people arrived some 55 000 years ago and spread throughout the land. Then there was the second invasion, by Europeans, in 1788. This quickly brought agricultural and industrial revolutions to a land that had been previously managed by fire for hunting. The environment changed irreversibly, and birds and other animals, plants and people have all had to adapt to live with these changes. These stories interweave the natural and cultural histories of the birds, and also consider the ways in which people have understood natural history in this place that so often breaks the rules developed by Western science for environments in the northern hemisphere. Australia has not ranked well in terms of its habitat and biotic losses since Europeans arrived. The situation is grave, but not all stories are about ‘bust’. Some bird species are well adapted to change. These stories of fauna, people and environment show how birds (and people) can adapt to change and variability in rainfall over shorter and longer timescales. iii

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The National Museum of Australia has had a long interest in Australia’s dry lands and deserts. It developed a major exhibition, Extremes, that compared the people and environments of deserts in Australia and other southern deserts in Chile, Argentina and Southern Africa. The book, 23 Degrees South: Archaeology and Environmental History of the Southern Deserts, edited by Mike Smith and Paul Hesse, is another important example of the Museum’s ongoing research in desert archaeology and environmental history. Dr Mike Smith, now a Senior Fellow in the Museum’s Centre for Historical Research, is one of Australia’s leading desert archaeologists, and also a key contributor to this book. Much of the work of the National Museum of Australia is about telling stories: through the objects in our collections, through our exhibitions and through our research and writing. The Museum has been an enthusiastic supporter of this book from its inception, because of the potential of such stories to helping us understand ourselves as well as our birds and the dry continent we call our home. At the National Museum of Australia we of course take a particular interest in all things Australian. But perhaps the most important duty of a good national museum is to be conscious of the role of Australia in the world. In a time of global climate change, when the whole planet is grappling with environmental variability and uncertainty, stories of people and environment in Australia can offer parables for a world where global climate change has added great variability and uncertainty to many environments. iv

F or e wor d

Boom and Bust: Bird Stories for a Dry Country will appeal to bird lovers, and also to all who are concerned about how people can adapt to changing environments everywhere. Craddock Morton Director National Museum of Australia Canberra August 2008

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CONTENTS

Foreword

iii

Contributors

ix

1

Introduction: boom and bust

1

Libby Robin and Mike Smith

2

The boom and bust desert world: a bird’s eye view

7

Libby Robin and Leo Joseph

3

Barcoo bantam: ‘It runs like hell!’

35

Graham Pizzey

4

Rain and grass: lessons in how to be a zebra finch

45

Steve Morton

5

Grey teal: survivors in a changing world

75

David Roshier

6

Australian pelican: flexible responses to uncertainty

95

Julian Reid

7

Night parrots: fugitives of the inland Penny Olsen

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8

Genyornis: last of the dromornithids

147

Mike Smith

9

Rainbirds: organising the country

185

Deborah Bird Rose

10

Woodswallows: a longer term, evolutionary view of boom and bust

205

Leo Joseph

11

White-winged choughs: the social consequences of boom and bust

223

Robert Heinsohn

12

Emu: national symbols and ecological limits

241

Libby Robin

Select bibliography

267

Index

285

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CONTRIBUTORS Robert Heinsohn is Associate Professor at the Fenner School of Environment and Society, Australian National University where his work focuses on the evolutionary ecology and conservation biology of birds. He has worked extensively on the behaviour of cooperatively breeding birds in Australia’s southeast woodlands, and on eclectus parrots and palm cockatoos on Cape York Peninsula and in New Guinea. Leo Joseph is Director of the Australian National Wildlife Collection at CSIRO Sustainable Ecosystems, Canberra. With an emphasis on the study of Australian birds in the research environment of a modern museum collection, his work integrates evolutionary history and present-day ecology. The aim of this work is to contribute to understanding of bird evolution. His roots are in natural history, especially in Australia’s arid and semi-arid country. Steve Morton is an animal ecologist who is especially interested in Australian deserts. Most of his career has been spent with CSIRO, first in Alice Springs and subsequently in Canberra. He is a member of CSIRO’s Executive and presently divides his time between Canberra, Melbourne and Alice Springs when possible.

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Penny Olsen is a research scientist based in the School of Botany and Zoology at the Australian National University. Her most recent book, Glimpses of Paradise: The Quest for the Beautiful Parrakeet, was published by the National Library of Australia in 2007 and won a Whitley Award in 2008. She edits Wingspan, the membership magazine of Birds Australia. Graham Pizzey AM (1930–2001) was one of Australia’s great bird observers, nature writers and conservationists. His series of Field Guides to Australian Birds began in 1965 with a commission from William Collins, the publisher of the Peterson guides to North America and elsewhere. The field guides continued to be revised until his death, the last being undertaken in partnership with CSIRO artist, Frank Knight. Pizzey was famous for a writing style that conveyed with immediacy the jizz (the appearance and demeanour) of every Australian species of bird. His descriptions of the ‘voice’ of the birds are invaluable in the field. Julian Reid is a biologist with expertise in avian and community ecology and a particular interest in Australian deserts. He divides his time between the Fenner School of Environment and Society, Australian National University and CSIRO Sustainable Ecosystems. Libby Robin is a historian of ideas at the Fenner School of Environment and Society, Australian National University and the Centre for Historical Research, National Museum of Aus-

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tralia. Her most recent book, How a Continent Created a Nation (UNSW Press), won the New South Wales Premier’s Award for Australian History in 2007. Deborah Bird Rose is Professor, Social Inclusion, at Macquarie University, where her work focuses on entwined social and ecological justice. She has carried out extensive research with Aboriginal people in Australia, and is currently working on a project on ‘love and extinction’. Her most recent solo book is Reports from a Wild Country: Ethics for Decolonisation (UNSW Press, 2004). David Roshier is a Senior Research Fellow with the Institute for Land, Water and Society at Charles Sturt University, Albury. He has broad interests in dispersal and movement ecology of birds, mostly in arid ecosystems. Currently, his research is focused on movement and migration of waterbirds in Australia and New Guinea. Mike Smith, an archaeologist, is a Senior Research Fellow in the Centre for Historical Research, National Museum of Australia. He has worked extensively across the Australian desert attempting to piece together the human and environmental histories of this fascinating region. In 2006 he was awarded the Rhys Jones Medal for Outstanding Contribution to Australian Archaeology.

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INTRODUCTION: BOOM AND BUST Libby Robin and Mike Smith

I love a sunburnt country, A land of sweeping plains, Of ragged mountain ranges, Of droughts and flooding rains. I love her far horizons, I love her jewel-sea, Her beauty and her terror The wide brown land for me!

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Core of my heart, my country! Her pitiless blue sky, When sick at heart, around us, We see the cattle die But then the grey clouds gather, And we can bless again The drumming of an army, The steady, soaking rain. Dorothea MacKellar (1904) – ‘My Country’ (verses 2 and 4)1

Not everyone shares Dorothea MacKellar’s delight in the contrasting beauty and terror of the ‘wide brown land’ of Australia. Droughts and flooding rains each bring their own difficulties, but the animals that have lived in this land over many years have developed impressive adaptations to cope with scarcity and plenty. The challenge for humans and animals alike in a land of environmental variability is uncertainty. When will those ‘grey clouds gather’ again? When will we ‘bless again / The drumming of an army,/ The steady, soaking rain’? The time between rains, ruled by ‘pitiless blue skies’, is a time for holding the nerve, of ecological stretch, and the irregular rains are at the core of a creative ecological pulse. ‘Boom and bust’ are the rhythms of Australia, so different from the regimented seasonality of northern Europe. It has been difficult for European Australians with the expectation of regular cyclical seasons to find a sense of permanence, to be at home in a place where drought and

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plenty stalk each other in unpredictable ways. In 1884 George Roberts, mailman on the Birdsville Track, disappeared. His packhorse was recovered but the mailman was not found. As 400 mm of rain had fallen in the district, people assumed that he had been drowned in the floods. But when his desiccated body was found on stony country near Haddon the following year, local people had to revise their opinion: he had died of thirst.2 This book uses a range of individual bird species as a lens for understanding environmental variability in Australia. It is more than a ‘natural history’; we have chosen stories that illustrate how natural systems play out under human-induced change and in some cases how humans have reacted to such events. Boom and bust is often a cyclical process for birds in the Australian environment. People and natural systems can both create booms and busts and human impacts can magnify or diminish natural patterns. People can interact with natural dynamics to constrain or amplify them in some way. Natural systems can also respond in unexpected ways. Sometimes change is irreversible. The story of biotic interactions with humans can only become more complicated as anthropogenic climate change alters rainfall patterns. Australia has not ranked well in terms of its habitat and biotic losses since Europeans arrived. It has the doubtful honour of leading the world in mammalian extinctions, with some 30% of its endemic mammal species extinct since 1788. It also has the highest number of threatened species on the

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planet. Conditions on this continent are already more variable and uncertain than in the rest of the world. In Australia, places with limited water – not just the desert country, but also the productive temperate south – are all predicted to receive less rainfall in the future.3 The situation is grave, but history suggests that not all stories will be of ‘bust’. The concept of boom and bust enables us to talk about the outcomes of environmental change for some bird species, and provides a key to exploring the idea of adaptation. If we do not understand the underlying natural rhythms of this continent, we will not be able to manage anthropogenic change. The evolutionary histories of Australian birds offer some insights into the natural dynamics of the continent. Europeans had to overcome expectations created by northern hemisphere conditions, particularly the idea that regular annual seasons are ‘normal’. In this context, the behaviour of birds adapted to Australian conditions has often been deemed ‘exceptional’ because it challenged established theories developed under European and Northern American conditions. A northern hemisphere perspective has sometimes made it difficult to observe the strategies of animals that live in landscapes ruled by variability and aseasonality. The difficulty of predicting whether any given year will be a boom or a bust year has become part of national stereotypes. Australia is the ‘El Niño continent’; a landscape ‘where creeks run dry or ten feet high’.4 But uncertainty is also increasingly a

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global phenomenon, and global climate change is already making living with uncertain seasons and extreme weather events more common, even in places where a well-ordered seasonality has been taken for granted in the past. In this book we draw on the natural history of Australia’s birds to explore the relations between fauna, people and environment in a continent where variability is normal and rainfall patterns do not follow regular seasons. It is a book that is aimed at Australian readers, but ideas about coping with boom and bust are also pertinent to other continents and increasingly relevant on the global scale.

Endnotes 1

MacKellar D (1904) My Country (poem). Republished with decorations and illustrations by JJ Hilder (1971) Angus & Robertson: Sydney.

2

This story is told in Tolcher HM (1986) Drought or Deluge: Man in the Cooper’s Creek Region. Melbourne University Press: Carlton South, p. 175.

3

Lindenmayer D and Burgman M (2005) Practical Conservation Biology. CSIRO Publishing: Melbourne, pp. 1–6.

4

This phrase appears in scientific as well as popular literature: Friedel MH, Foran BD and Stafford Smith DM (1990) Where the creeks run dry or ten feet high: pastoral management in arid Australia. In Australian Ecosystems: 200 Years of Utilisation, Degradation and Reconstruction. (Eds. DA Saunders, AJM Hopkins and RA How) pp. 185–94. Surrey Beatty and Sons: Chipping Norton, NSW.

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BOOM AND BUST DESERT

WORLD : A BIRD’S EYE VIEW Libby Robin and Leo Joseph

Picture the scene: mid-afternoon in the stony downs country of inland eastern Australia, somewhere near the border between South Australia and Queensland. It is a day so hot that all one can see are shimmering heat mirages rising from the ground. From nowhere, two four-wheel drive vehicles emerge. They stop at a dry creek bed framed by mineritchie, a small tree with bark that is made up of remarkably backward curving rough fibres of a burnt amber, an almost wine-red colour. The vehicles stop in the sparse shade the mineritchies

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provide and a tired-looking group of academics researching a book on boom and bust steps out of the vehicles. A few minutes later, after grumbling about the failed air conditioning units in their vehicles, one of them speaks in excited, hushed tones. He is an ornithologist who, despite the self-inflicted trauma of having qualified for an academic career in ornithology, still enjoys finding interesting birds. He draws everybody’s attention to a pair of Bourke’s parrots (Neopsephotus bourkii) sitting quietly in a nearby mineritchie. Thinking that his companions could not fail to be enthralled by the gem of information he is about to impart, he points out to them that Bourke’s parrot is one of the more remarkable Australian parrots and that recently emerging DNA sequence data is showing that its closest living evolutionary relatives, apart from a handful of local species, appear to live well beyond Australia. A behavioural ecologist in the group says that that’s all very interesting but is wondering where the nearest water is, knowing that these birds like to drink in the half-light of dawn and dusk. Just as he is pondering what part of their annual reproductive cycle they may be in, if indeed they have annual cycles out here, the anthropologist in the group, an authority on Indigenous linguistics, recalls that Bourke’s parrot is known to the Pitjantjatjara people further west as ‘Wilyurukuruku’. The historian in the group looks beyond the birds to a dilapidated stock-yard several hundred metres away. Aloud, she muses about when it was in use and what put it out of business. Meanwhile the archaeologist in the group is engrossed in the sharply angled

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stone flakes at his feet, clearly the result of human craftsmanship, but for what purpose? The palaeontologist in the group has turned to face the other direction, where she has spotted a geological outcrop in the middle distance and is feeling a flutter of hope that it might have just the right sediments she has been looking for to search for bones of that fascinating extinct bird she’s been working on. And so it goes. Just as different people can see wildly different things in one aspect of an Australian desert, so too the boom and bust phenomenon can awaken different interests in different people. In this book we will explore this generally and in this chapter we will explore it specifically in terms of desert birds. The history of ideas about Australian semi-arid country and deserts and the birds that live there provides an important background to this book. The bird stories we have chosen illustrate the booms and busts of life in the arid zone, but they have also contributed to the way we think about deserts themselves. In this chapter, we provide a brief overview of the relationship between ornithological ideas and our understanding of deserts. The first arid-zone bird ‘discoveries’ by Western science lacked the context of ongoing observation of desert country. As new agricultural and pastoral developments grew in semi-arid country after each of the World Wars, bird observers began to ask questions about how birds live in such country. In the post-war period, professional scientists turned their attention to desert country all over the world, and this brought the

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Australian desert into international focus. Since the 1970s, new ways of thinking about deserts have further shifted the critical questions of ornithology, and vice versa.

Arid-zone ornithology in the 19th century Ornithology in Australia had a strong start with the 19th century work of John Gould and his extraordinary collectors, particularly John Gilbert. Gould’s Birds of Australia, published in fascicles between 1840 and 1848, quickly became the definitive work on the subject, and remained so for the rest of the 19th century. It is still regarded as an extremely important reference in the history of ornithology. Gould, Gilbert and other collectors travelled widely, sometimes well beyond the limits of European settlement which in the 1830s and 40s hugged the coastlines closely. Their travels usually began in the major ports: Swan River (Perth and Fremantle), Port Essington and the Cobourg Peninsula in the north, Port Jackson (Sydney) and Hobart Town. Adelaide, Melbourne and Moreton Bay (Brisbane) were also developing as centres and their hinterlands provided birds for the collections. Charles Sturt (1795–1869) named the Darling River in 1828 and ventured well into the interior of the continent in 1844–1846, while John McDouall Stuart (1815–1866) led various expeditions into central Australia the late 1850s, culminating in a transcontinental crossing from Adelaide to the Timor Sea in 1862. Although intrepid explorers such as Stuart commented on the movements of birds when travelling in the interior, their

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expeditions were not focused on collecting birds for science. Of course, some birds whose populations do breed in central Australian deserts were collected closer to ports and coastal areas generally right from the first years of settlement in the 18th century. Emus, pelicans and other waterbirds such as the grey teal that feature in this book, were well-known in the Gould era. Gould also described species from specimens of individual birds now known to be desert and semi-arid specialists, most notably the night parrot, which Penny Olsen discusses in Chapter 7, the malleefowl (Leipoa ocellata) and the flock bronzewing (Phaps histrionica).1

New discoveries with new settlements: the early years of the 20th century As the semi-arid country fringing the interior was settled more intensively, particularly through soldier settlement schemes in the Western Australian wheat belt and the mallee country of South Australia and Victoria in the 1920s and later, arid-zone ornithology took its next steps. One of the bird mysteries solved during this period concerned the breeding patterns of the banded stilt (Cladorhynchus leucocephalus). By the end of the 19th century, ornithologists had regularly observed that breeding could occur over long periods in the tropics, but birds known from Australia’s temperate zones, including shorebirds like the banded stilt, were generally expected to be regular winter or spring breeders.2 Banded stilts look and behave like most other shorebirds, foraging in shallows of salty

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lakes and periodically dipping and digging their long bills into the sands for invertebrate food. They often co-occur with other shorebirds. For example, they were noted by George Keartland alongside avocets (red-necked avocet Recurvirostra novaehollandiae) and black-winged stilts (Himnatopus himantopus leucocephalus) on the 1896–97 Calvert Expedition to the Kimberley, in north-western Australia, although they were better known by ornithologists in places like Rottnest Island near Perth and the coastal flats of South Australia near Adelaide.3 The birds had been observed to disappear in large numbers, but no rookeries had been found, and their breeding areas remained unknown. At this time there was more known about birds breeding in the Arctic tundra at the mouth of the Lena River in Siberia than those in Australian deserts. There was speculation but little firm evidence about the actual breeding patterns of the banded stilt.4 Ludwig Glauert, palaeontologist and curator at the Western Australian Museum and one the state’s leading scientists, conducted a full zoological survey of Rottnest Island where the banded stilts were usually plentiful, yet sometimes absent altogether. He mused: ‘It seemed unlikely that the breeding places were outside Australia. Yet information concerning the ‘winter migration’ was of the vaguest possible character’. He was intrigued that such a well-known bird could ‘keep its [nesting] secret so successfully’.5 The wet winter of 1930 shed new light on the subject. A letter came to the museum from Mrs BE Cannon, a farmer

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and keen bird observer from inland south-western Australia reporting a huge rookery of breeding birds, ‘thousands of them on a sandy spit running out into Lake Grace’ near her home. The letter was accompanied by a blown egg and three photographs. The museum immediately sought more information, and on 4 September, a parcel of 23 unblown eggs followed, one of which proved to contain a chick that had died immediately before hatching. Glauert and his ornithological friend, the State entomologist, Clee Jenkins, realised they had the first solid evidence of where banded stilts breed, and that they could confirm that they were a bird of the Australian inland, not a Siberian migrant. The find was of particular interest to Ivan Carnaby, an experienced amateur egg collector, who kept close records of nesting in the Lake Grace area from 1928 for many years.6 He commented that before 1928, it was ‘a place where very little [had] been done in the way of ornithology’.7 Carnaby, an observant bush naturalist, was to continue to contribute important specimens and observations to arid-zone bird science throughout the 1950s and 1960s. Before 1930 was over, the idea that banded stilts bred in ‘winter’ was challenged by further finds at Lake Callabonna, north of Lake Frome in South Australia’s north-east. In late December, J Neil McGilp and Matt Morgan, long-time stalwarts of the South Australian Ornithological Association, heard reports from the manager of Moolawatana station that there was a rookery of nesting birds on a previously unvisited island in the lake. On 11 January 1931, they found ‘a densely

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packed mass of Banded Stilts’ – about 54 000 of them – which took to the air ‘with a whirr of wings, raising quite a dust as they did so, and revealing thousands of eggs on the bare ground’.8 There was no protection from the merciless heat (recorded at 104°F in the shade, = 40°C) or the hot wind, the only vegetation being ‘a few samphire bushes’. As McGilp and Morgan commented: It seems somewhat extraordinary that the second discovery should have so soon followed the first, after the nesting of these birds remaining a mystery for so many years.9

The banded stilt breeds ‘only in the arid inland at large shallow ephemeral salt lakes inundated by rain or floodwaters’.10 The irruption of the brine shrimp, its main food, is essential to a breeding event on this scale. The time of year and the weather are irrelevant. It is the boom, or pulse, of resources that makes breeding possible. McGilp and Morgan noted the ‘abundance of food’ in January 1931, the water swarming with small crustaceans 11

Professional ornithology and the Australian arid zone (1945 to 1974) International scientific interest in the Australian desert accelerated in the post-war era through various initiatives. Fulbright Scholarships encouraged North American researchers to

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travel and work in remote parts of Australia. Funded by these, Ruben Arthur Stirton and Richard H Tedford established the first tertiary sequences for the Australian desert, and were followed by specialist palaeo-ornithologists Alden Miller and Patricia Vickers Rich in the 1960s. Such palaeontological work was supported locally by David Ride, an Oxford-trained vertebrate palaeontologist, who was Director of the Western Australian Museum. Pat Vickers Rich subsequently moved to Australia and is now a world specialist in fossil non-passerine birds, especially the dromornithid group, of which the extinct Genyornis featured in this book is the best known example.12 Another major international program to build aridzone science in the 1950s was funded through the United Nations Educational, Scientific and Cultural Organization (UNESCO). It focused strongly on Israel and India, two new nations establishing themselves in the post-war years with significant desert country, and on the deserts of northern Africa, also politically charged in an era of pro-independence. UNESCO aimed to offer science that was above politics to combat ‘desertification’, a new and different enemy shared by diverse nations. There was a growing sense that deserts were not only ‘natural’ phenomena, but also often the product of humanity’s wasteful exploitation of soil. Deserts were expanding just at a time when the world needed more viable agricultural and pastoral land. This idea, a sort of cultural history of nature, was the theme of Paul Sears’ influential book, Deserts on the March, a treatise on the power of human cultures to

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destroy and remake nature.13 Sears focused on soil and land use in the United States at a time when it was reeling from the ecological tragedy of the ‘Dust Bowl’, but he included comparisons with China, India and Egypt. Australia also had a dust bowl tragedy in the 1930s, and Francis Ratcliffe (1904– 1970), the great writer and biologist, documented the destruction of pastoralism in desert places in Flying Fox and Drifting Sand.14 Both Sears and Ratcliffe were ecologists who had approached the issue of soil erosion as a natural science problem and found themselves writing about a major cultural issue. Each wrote prompted by a dust bowl crisis, but their books became much more widely read 20 years later in the post-war 1950s, when the idea of people changing nature became even more politically charged by the advent of the atomic age and the Cold War. In Australia, the Council for Scientific and Industrial Research (after 1949, the Commonwealth Scientific and Industrial Research Organisation, CSIRO) expanded to include a Wild Life Section. Francis Ratcliffe was appointed Officer-incharge. CSIRO also expanded its operations in Western Australia in this period and developed work in the arid centre around Alice Springs in the 1950s.15 Universities were also expanding. After 1948 it was possible to undertake doctoral research at Australian universities for the first time, beginning with the new Australian National University that was established specifically for this purpose. This increased the number of researchers working on Australian subjects, and enabled

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researchers to choose subjects demanding fieldwork remote from major cities. Often the first port of call for international researchers considering prospective research in Australia was CSIRO. One of the most internationally ‘networked’ scientists in Australia was ornithologist and fisheries expert, Dominic Serventy (1904–1988). He had worked in Sydney during the War years based at the fisheries division of the Council of Scientific and Industrial Research, but also maintained close contact with his home state, Western Australia, and headed the Perth office of the Wild Life Survey section when it was established in Perth in 1951.16 Animal problems in semi-arid rangelands country drove this development. Rabbits were the major concern across southern Australia but further north, pastoralists, especially in Western Australia, reported ongoing problems with native animals, including various kangaroos and emus.17 Serventy had been interested in birds since he was a young boy, but as a professional scientist (with a PhD from the University of Cambridge) he became interested in what ornithology could contribute to the wider biogeography of the arid zone. He worked with University of Western Australia geographer, Joe Gentilli, who was trying to gain a biogeographical understanding of Australia by developing an atlas that detailed the arid and semi-arid country on the edges of the Western Australian wheat belt. The correspondence between the two reveals that the movements and breeding patterns of birds were important tools for understanding the way the desert country worked,

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and also show how limited was the available information at the time. Gentilli wrote to Serventy in 1943: The idea of working out something about ornithological distribution is excellent, but when I asked the Australian Museum to let me know the supposed present distribution of mallee fowl and brush turkey, they could tell me nothing. I got the mallee fowl area rather well here from Mr Glauert, but he would not go outside Western Australia. So I think I shall have to give up the idea of showing the distribution of these very interesting species. The point is, that I am trying to start work on a much bigger undertaking, a sort of Atlas of Australia, for which I would enlist the collaboration of anybody willing to assist … This present work I am doing is only a textbook of Geography; but having seen how difficult it is to find any distributional maps, I have decided to give as much space as I can to maps of all kinds.18

The mysteries of the banded stilt’s breeding grounds had fascinated the interwar generation of ornithologists, but the 1950s generation turned to the breeding biology of the malleefowl, the only one of the distinctly Australasian megapode family to live in desert and semi-arid country. In this later, more rigorously scientific era of research, ‘southern’ bird groups held the key to understanding Gondwanan heritage. The malleefowl had long fascinated egg-collectors (oologists)

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and bird observers who knew that it incubated its eggs in a mound. But it took the systematic work of CSIRO scientist Harry Frith (1921–1982) to analyse the workings of the mound; how the male bird constructs them in sandy soils and then maintains them by adding and scraping away fermenting vegetation in such a way that the eggs remain at an almost constant temperature until they hatch. It also took systematic observation to confirm that the parental care of malleefowl was rather different from most other birds, with females taking no further part in rearing the young after laying the eggs, and neither parent having interactions with the young once they left the nest. Mound-digging behaviour demands light sandy soils, and although the birds were widely spread in Victoria, New South Wales, South Australia and Western Australia, they were generally found in mallee scrub, semi-arid country. In the 1950s, observers noted these birds in many new places, particularly on the edge of soldier settlement schemes, where timber was being cleared rapidly for new farms. In Victoria, a Lowan Sanctuary (nature reserve) was declared in mallee-broombush country at Kiata in 1955 in the semi-arid Wimmera-Mallee region in an early effort to protect the birds, which were seen to be threatened by agricultural and pastoral developments near the Little Desert.19 ‘Lowan’ was the Aboriginal name for the birds in this region; in Western Australia, their Aboriginal name was ‘Gnow’, and Serventy kept a ‘Gnow’ file for many years in the 1950s, as observant farmers and naturalists sent him notes expressing

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concern about their changing habits in the face of the rapid expansion of agriculture.20 Vermin Control Officer Corney, stationed at the Number 3 Vermin Fence (the Emu Fence) at Yalgoo in 1965 reported to his superiors that: I have noticed an increase in the Mallee Hen population this year. In places where I used to see an odd one or two, I now see so many as twenty or more and some in new places altogether. Galahs are another bird which is increasing in their thousands and are becoming a real major pest, not only in the farming country but all through the bush where they strip the trees and areas are going to die out from this.21

Serventy relied on a range of colleagues who knew that he was developing ideas about the semi-arid country and its birds.22 Official and semi-official reports were supplemented by the observations of CSIRO colleagues like John Calaby, who passed on observations from local observers like P Warren in Kalkannie: Mallee hens are coming back around Kalkannie. He has seen 4 birds in the past two years. He had never seen them before in his life. Last season, his brother was rolling scrub and found a mound with fresh eggs.23

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Harry Frith recognised that his scientific work on malleefowl was of international interest and he reported it at the International Ornithological Congress in Switzerland in 1954. He also later popularised his discoveries in a well-illustrated book.24 It was not easy for Frith to undertake serious ornithological work alongside his official CSIRO work on the orchard industry in Griffith in the New South Wales Riverina country. There was always a tension between doing such ‘pure science’ and reporting about the ‘applied science’ that CSIRO was expected to deliver. Francis Ratcliffe also had trouble getting his superiors to support the ‘pure science’ that was needed to understand the arid zone, although he was proud of the publications of Frith and others, and his own book Flying Fox and Drifting Sand was perhaps more influential in policy circles than in scientific ones. In 1955 Ratcliffe wrote personally to FG Nicholls, CSIRO Secretary of General Administration, appealing to him to reconsider a directive from head office that all research results had to be published by CSIRO.

25

He asked

Nicholls to grant him, as Officer-in-charge, ‘a certain amount of discretion’ about what might be regarded as ‘official’: Many of my staff publish, or want to publish a lot of unofficial stuff; and sometimes the line between the official and the unofficial is quite impossible to draw. Sometimes work which started as a purely hobby interest, done in a man’s spare time and at weekends, may develop into a piece of official research – the best example of this is

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Harry Frith’s interest in the Mallee Fowl, and the physics of its egg incubation … Dom Serventy presents an even more difficult case. As far as I can make out he spends his whole waking life being an ornithologist, at times acting and thinking as an officer of CSIRO and at times as a private individual.26

Serventy was always working at the frontiers of the possible, trying to make his nation-minded masters support ornithological work of serious international interest. In the mid-1950s, he maintained regular correspondence with AJ (Jock) Marshall (1911–1967), an expatriate Australian working as a physiologist at St Bartholomew’s Hospital in London, and American ecologist Don Farner (1915–1988) from the University of Washington. All three were developing ideas on how birds lived and bred in deserts. Serventy set up a folder on ‘Bird Irruptions’ that collected his and other people’s observations of irregular events from 1957 and 1958. The file includes his notes on some of the ideas about irruptions in the literature from other continents, and how they contrasted with the Australian cases.27 Marshall, a physiologist who had much experience of the breeding patterns of tropical birds in the New Hebrides, was interested in what stimulated the breeding cycle of birds from a hormonal perspective. He was keen to return to Australia after 15 years in Britain, to ‘retribalise’ his family. He undertook fieldwork in 1957 in association with Serventy,

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local ornithologist Ivan Carnaby, and the artist Russell Drysdale in the semi-arid and desert parts of Western Australia. His book Journey among Men (illustrated by Drysdale) popularised the natural history of the desert, and he also published scientific works on the physiology of desert breeding.28 In 1960 Marshall returned permanently to Australia, as Foundation Professor of Zoology at the new Monash University, where he established a department that focused strongly on Australian animals (particularly marsupials, but also birds). Don Farner was one of the non-Australian scientists developing theories about the behaviour of arid-zone species, particularly zebra finches (Taeniopygia guttata). These little birds appeared to be highly mobile and irruptive, especially after rain events. Farner kept an aviary to work systematically on their behaviour in the United States, but always sought opportunities to come to Australia to continue observations of birds in the wild as well. Klaus Immelmann, an ethologist from the University of Bielefeld, West Germany, spent a full year in Australia under a grant from the German Federal Research Organisation from 1959 until 1960, and made several other research trips to observe zebra finches. Serventy collected information on local zebra finch sightings and passed them on to both Farner and Immelmann. On his file, he had a note from SW Bowles, Fauna Warden for the Dowerin district, which reported 5000 black-tailed native-hens (Tribonyx ventralis) that ‘played havoc with local farmers’ Lucerne crops’ in the district on 12 December 1957 and ‘a small colony of about

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12 Zebra Finches’ in the Jennacubbine town-site nearby the next day. Even for black-tailed native-hens, this was a large group worth noting, but it is interesting that the appearance of such a small number of zebra finches was also mentioned by Bowles. The international interest in the bird probably stimulated a range of people to keep formal records of their movements and to pass these on to Serventy and others. As a better known species of a poorly understood biogeographical region, zebra finches were beginning to be seen as ‘indicator’ species in this period. The cheerful chatter of zebra finches signalled water in places where it was scarce. In 1961 Farner was selected by CSIRO to succeed Francis Ratcliffe as head of the Wild Life Survey Section, a choice that pleased Serventy who had persuaded Farner to apply for the job. However, Farner quickly realised that this would be a very full-time administrative job allowing little time for his zebra finch work. His home university made a counter offer of a new laboratory and aviaries for his finches in Seattle, very much bigger than his facilities in Pullman, and so he declined the Australian offer and moved to the University of Washington’s Seattle campus instead. CSIRO then appointed local scientist Harry Frith to the post. Frith, like Serventy, was well regarded internationally. In 1974, he was invited to organise the first International Ornithological Congress held outside the northern hemisphere since that meeting began in 1884. It took place in Canberra, close to the CSIRO facilities and strongly supported by CSIRO. It is a mark of the importance of desert

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ornithology in this period and the camaraderie amongst its practitioners that his fellow organisers included Serventy, Farner and Immelmann, the latter two being jointly responsible for a symposium on Physiological and Behavioural Adaptations to the Arid Zone.29

Rangelands and resilience science: new ideas for deserts By the 1970s, the need for more scientific definitions of deserts was becoming apparent. Dom Serventy commented in 1971: ‘A desert occurs wherever it is said to occur. A definition by acclamation!’30 Part of the difficulty was that deserts were both a natural and an anthropogenically induced phenomenon, and were often defined in terms of human use. Serventy’s working definition of a place where ‘dryness and heat are so excessive that normal agriculture is impossible in average years’ was typical, but it lacked precision and embedded Eurocentric expectations about ‘normal agriculture’. Even the idea that deserts had to be hot was changing by the end of the 1970s. JL Cloudsley-Thompson, editor of the Journal of Arid Environments which was established in 1978, called for articles about the ‘problems of the inhabitants of the world’s desert regions’ including ‘intensely … cold’ environments. 31 Rangelands science, which supported pastoral enterprises in semi-arid lands, grew strongly in the 1970s and 1980s and began to provide new, increasingly precise definitions of the ‘arid zone’. There was a growing awareness that dryness was

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not just about a lack of rain, but also about the degree of evaporation, so that country in tropical latitudes might still be ‘arid’ despite higher rainfall. The other question for people trying to develop enterprises in semi-arid areas was when the rain fell, and with what sort of reliability. Averages can be very misleading. In the 1970s, dry country in Western Australia could support the growth of wheat on 250 mm rainfall, because there was usually some winter rain for sowing and some spring rain for growing. In similar poor soils in north-western Victoria, an average rainfall of about 400 mm was not enough to grow wheat because the timing of the rainfall was less predictable. 32 Further out in the rangelands of the semi-arid zone, there are years where no rain falls at all. An ‘average rainfall’ is simply not indicative of what the country can or cannot grow. The Arid Zone Research Centre in Alice Springs defines ‘rangelands’ in Australia in terms of human use, but the arid zone itself in terms of rainfall, evaporation, land forms and vegetation. ‘Typical’ desert vegetation is something that is not necessarily annually or seasonally dictated, so this provides a somewhat independent measure of the uncertainty and variability of seasons in this country. Arid and semi-arid lands are: those remote and sparsely populated areas of inland Australia, defined by the presence of desert vegetation and land forms as well as by low rainfall. They are bound by median annual rainfalls of about 250 mm in the south

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but up to 800 mm in the north and about 500 mm in the east. Together with sub-tropical regions and the mountain high plains, they form the rangelands, where rainfall is too low or unpredictable or where terrain is too inhospitable for sustainable cropping or timber harvesting. The rangelands amount to 70% of Australia’s land surface, and by far the largest part is arid or semi arid.33

The growth of dynamic systems modelling and ‘resilience’ systems science since the early 1970s has influenced the way desert country and its flora and fauna are understood. The fundamental boom and bust idea that underpins international ecological understandings of desert environments came out of Australian rangelands science. In 1973, Israeli ecologist Imanuel Noy-Meir conceptualised the ‘pulse and reserve’ model, a term originally coined by Australian rangelands modeller, Mark Westoby.34 Ecosystems generally are driven or limited by a number of factors such as temperature, day length and so forth, but in the desert it is water. When you get some rain, you get a pulse of biological activity that gradually runs down. After the pulse of activity, the desert becomes quiet and the ‘clock’ stops ticking. As CS (Buzz) Holling, another systems scientist, put it in the same year as Noy-Meir wrote: Individuals die, populations disappear and species become extinct. That is one view of the world. But another view

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concentrates not so much on presence or absence, as upon the numbers of organisms and the degree of constancy of their numbers. These are two very different ways of viewing the behaviour of systems and the usefulness of the view depends on the properties of the systems concerned.35

The years since the 1970s have seen a growing scientific interest in the movements of populations of birds, and their evolutionary history. Individual sightings are still of interest, but the focus is more on how and why birds move into and out of arid country, as well as within it. How do they cope with its boom and bust conditions? Marshall and Serventy in the 1950s and ’60s concentrated on physiological triggers for birds breeding in arid regions, rejecting the idea that day length (seasons), for example, had major influence on the reproductive organs of many arid-zone species. By contrast later scientists such as Richard Kingsford, and including ornithologists like Dave Roshier and Julian Reid, whose work is represented in this collection, have closely documented massive breeding events and are using this work to understand the workings of desert river and ephemeral water systems in arid and semi-arid country.36 In 1990, botanist Mark Stafford Smith and zoologist Steve Morton, from their base in CSIRO at Alice Springs, wrote a classic paper that provided a conceptual framework that glued together the threads and themes that had been developing in previous decades and that we have mentioned above. They gave an overview of Australian desert systems and the drivers

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of ecological behaviours in an international context. While all arid regions have spatially and temporally variable rainfall, they noted that Australia’s particular problem was that temporal variability was higher than elsewhere. Most years have less than average rainfall, with just a few years of very high rainfall skewing that average.37 Small isolated rainfall events do little to stimulate plant growth, but big rains, especially those caused by cyclonic and monsoonal depressions, drive major and widespread effects on the biota, aided and abetted by the prevailing flatness of inland Australia. Their paper continues to be a key pillar in our understanding of Australian deserts and how boom and bust operates on systems within them.

A coda: a glimpse of future studies of birds in booming and busting deserts Eventually, our rag-tag group of academics researching their book on boom and bust, a little refreshed after stopping at the mineritchie-lined dry creek bed, fall back into their vehicles to slog out the next leg of their journey without air conditioning. The historian remarks that the spot where they just stopped encapsulated so much of what bust can mean in this environment. She expresses the hope of returning one day to witness first-hand a boom unleashed by inland Australia’s erratic and unpredictable rainfall. The behavioural ecologist remarks that waterfowl would likely appear very soon after such rains and the palaeontologist wonders aloud how birds like ducks can find water so quickly after rains. ‘Oh well’, the ornithologist

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chimes in, ‘it may be to do with infrasound, you know, the kind of long wave-length sound waves generated by thunderstorms.’ Silence. The kind of silence that voices loud scepticism. ‘Well, it’s been known since the 1970s that birds can detect infrasound,’ he continued. And so the conversation went. Clearly, just as solving the mystery of where banded stilts bred challenged us 100 years ago, so too today we still have plenty of scope to develop our understanding of the boom and bust phenomenon underlying the case studies discussed in this book. Boom and bust is a concept that can integrate diverse disciplines and technologies across orders of magnitude of time and space. It opens fields of study that yesteryear’s ornithologists who sought breeding colonies of banded stilts would not recognise. And to study those fields, we have tools available that those ornithologists could not have imagined. Fossils and subfossils yield signals of how populations in the past boomed or busted to extinction. Highly mobile birds like waterfowl are tracked over thousands of kilometres from satellites in orbit around the earth. That information is tied to what the sequence of the base pairs in their DNA tells us of their evolution and how natural selection has moulded their movement patterns. Isotopic methods can tell us of the diet of today’s species and even of extinct species. Studies not yet done of avian neuroanatomy will test whether waterfowl can detect infrasound. And unifying all of this is the idea that boom and bust is a concept that we humans need to understand if we are to read the lessons of

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the environment for our own survival. The stage is set for a boom in understanding.

Endnotes 1

Robin L (2001) The Flight of the Emu: A Hundred Years of Australian Ornithology. Melbourne University Press: Carlton; Gould J (1840–1848) Birds of Australia. The Author: London; Fisher CT (1992) The Importance of Early Victorian Natural Historians in the Discovery and Interpretation of the Australian Fauna, with Special Reference to John Gilbert. PhD thesis, University of Liverpool, UK; Lambourne M (1987) John Gould: Birdman. Osberton Productions: London.

2

Astheimer LB and Buttemer WA (2002) Changes in latitude, changes in attitude: a perspective on ecophysiological studies of Australian birds. Emu 102(1), 19–27, esp. p. 20.

3

See Jones J (1945) The banded stilt. Emu 45(1), 1–36, esp. p. 3.

4

Robin L (2005) Migrants and nomads: seasoning zoological knowledge in Australia. In A Change in the Weather: Climate and Culture in Australia. (Eds. T Sherratt, T Griffiths and L Robin) pp. 42–53. National Museum of Australia Press: Canberra; Glauert L and Jenkins CFH (1931) Notes on the banded stilt (Cladorhynchus leucocephalus) with a description of its eggs. Journal of the Royal Society of Western Australia XVII, 1–7.

5

Glauert L and Jenkins CFH (1931) p. 2. The museum’s 1930 collection of 19 successfully blown eggs and a chick was supplemented within a short time by two adult birds in full plumage, an immature bird, and a chick (nesting in down), sent by another Kukerin citizen, Mr W Broadwith.

6

Carnaby IC (1933) The birds of the Lake Grace District, WA. Emu 33(2), 103–4; Carnaby IC (1946) A further nesting record of the banded stilt. Emu 46, 156–58.

7

Carnaby IC (1933) p. 103.

8

Truran J (2000) The foundation period 1899–1933. In Birds, Birders and Birdwatching 1899–1999. (Eds. R. Collier et al.) pp. 18–56. SAOA: Adelaide; McGilp JN and Morgan AM (1931) The nesting of the banded stilt. The South Australian Ornithologist XI(2), 37–53, quote p. 40.

9

McGilp JN and Morgan AM (1931) p. 39.

10 Marchant S and Higgins P (Eds.) (1993) Handbook of Australian, New Zealand and Antarctic Birds. Vol. 2, p. 781.

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11 McGilp JN and Morgan AM (1931) p. 41. 12 Robin L (2001) pp. 198–99. 13 Sears P (1949) Deserts on the March. Routledge & Kegan Paul: London. 14 Ratcliffe F (1938) Flying Fox and Drifting Sand: The Adventures of a Biologist in Australia. Chatto and Windus: London (first Australian edition 1947); Robin L and Griffiths T (2007) Francis Noble Ratcliffe, 1904–1970. In New Dictionary of Scientific Biography. (Ed. N Koertge) pp. 207–11. Charles Scribner’s Sons: Farmington Hills, MI, USA. 15 R Birtles, unpublished ‘Summary of CSIRO Annual Reports 1926–1955’ (personal communication, 8 April 2008). 16 For example, he established a program of research into tuna in the Indian Ocean during the World War Two. Collis B (2002) Fields of Discovery: Australia’s CSIRO. Allen & Unwin: Sydney. See pp. 336–37. 17 Meeting 22 August 1945, University of Western Australia conducted by AEV Richardson (Chief Executive Officer, CSIR), Hedley Marston (Head of Animal Nutrition) and Dominic Serventy on the subject of pastoral pests, particularly in the north of the state. Pastoral interests were represented by ALB Lefroy, AM Stewart and EHB Lefroy and the Western Australian Department of Agriculture by GK Baron-Hay. CSIRO Wild Life Survey Section opened a branch in Perth in 1951, with Dominic Serventy as Officer-in-charge. 18 Letter from Gentilli to Dominic Serventy 16 January 1943 (Australian Archives Perth, Serventy Papers, Correspondence – Dr J. Gentilli). This letter was part of a correspondence that included discussions about offshore oceanic currents and fish distributions, as well as birds. It was not only lack of information but also uncertainty of employment in war-time Australia (for Gentilli, an immigrant of Italian background) that hampered this work. 19 Robin L (1998) Defending the Little Desert: The Rise of Ecological Consciousness in Australia. Melbourne University Press: Carlton, p. 8. 20 Serventy correspondence, Australian Archives (Perth). File Gnow. For example, a report from LJ McClennan of malleefowl unexpectedly sighted in the heavily forested area of Nannup 5 February 1953. 21 25 March 1965, AR Tomlinson to AJ Frazer – copy on DLS Gnow file. 22 They included senior Western Australian public servants, including the Chief Warden of Fauna (AJ Frazer) and the Chief Vermin Control Officer (AR Tomlinson), as well as AR (Bert) Main, who received notes

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from time to time at the Zoology Department of the University of Western Australia about the movements of birds and animals in remote corners of the state. 23 JHC typed note; Gnow file. 24 Frith HJ (1955) Incubation in the Mallee Fowl (Leipoa ocellata, Megapodiidae). In Proceedings of the 11th International Ornithological Congress, Basel, pp. 570–74; Frith HJ (1962) The Mallee Fowl: A Bird that Builds an Incubator. Angus & Robertson: Sydney. 25 Head Office Circular 55/42, ‘Publication of Research Results’, as cited in Ratcliffe to FG Nicholls, 16 August 1955, Ratcliffe–Serventy correspondence, Ratcliffe Collection, NLA. (Reference copy sent to Serventy.) 26 Ratcliffe to FG Nicholls, 16 August 1955, Ratcliffe–Serventy correspondence, Ratcliffe Collection, NLA, p.1. See also Robin (2001). 27 For example, Serventy’s own notes on the paper by Davis J and Williams L (1957) Irruptions of the Clark Nutcracker in California. Condor 59(3), 297–307, from p. 301, focused on the fact that the population seemed to build first, causing the shortage of food. (Notes by DLS, Irruptions file). By contrast, major breeding events of birds like the Banded Stilt followed the irruption of their food resource. 28 Marshall J (1960–61) Biology and Comparative Physiology of Birds. New York: Academic Press; Marshall J and Drysdale R (1962) Journey Among Men. Hodder & Stoughton: London. Marshall J (1998) Jock Marshall: One Armed Warrior. Melbourne: Bright Sparcs. http://www.asap.unimelb. edu.au/bsparcs/exhib/marshall/marshall.htm. 29 Robin L (2001) International ornithology comes to Australia. Historical Records of Australian Science 13(3), 233–54, esp. p. 240. Jock Marshall died in 1967; otherwise he would have undoubtedly also been involved in the conference organisation. Charles Sibley was also a key organiser at this conference; his work is discussed in detail in Leo Joseph, ‘Woodswallows’ (Chapter 10, this book). 30 Serventy DL (1971) Biology of desert birds. In Avian Biology (Eds. DS Farner and JR King) p. 292. Academic Press: New York. 31 Cloudsley-Thompson JL (1978) Editorials. Journal of Arid Environments 1(1), 1; and (1979) 2(1), 1. 32 Newman RJ (1959) Pasture improvement in the Little Desert. Journal of the Department of Agriculture, Victoria 57(1), 1–9.

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33 http://www.cazr.csiro.au/aridlands.htm. 34 Noy-Meir I (1973) Desert ecosystems: environment and producers. Annual Review of Ecology and Systematics 4, 25–51, see p. 30; Westoby M, Walker B and Noy-Meir I (1989) Opportunistic management for rangelands not at equilibrium. Journal of Range Management 42(4), 266–74. 35 Holling CS (1973) Resilience and stability of ecological systems. Annual Review of Ecology and Systematics 4, 1–23. 36 Kingsford R (2006) (Ed.) Ecology of Desert Rivers. Cambridge University Press: Cambridge UK. (See also Steve Morton’s reflective review of this book in Historical Records of Australian Science 18(2), 281–87); Dickman C, Lunney D and Burgin S (2007) (Eds.) Animals of Arid Australia: Out on their Own? RZSNSW: Mosman. 37 Stafford Smith DM and Morton SR (1990) A framework for the ecology of arid Australia. Journal of Arid Environments 18, 255–78.

34

BARCOO BANTAM: ‘IT RUNS LIKE HELL!’ Graham Pizzeyy1

From time to time I get a query about an unusual waterhen people have seen. It’s usually described as dark, with red legs – and a cocked tail that makes it look ‘like a little bantam’. Some say they have seen it far from water. Most agree that ‘it runs like hell’. Over the years I’ve had more technical enquiries, but there’s not the slightest doubt the bird in question in this case is that peculiarly Australian creation, the black-tailed native-hen (Tribonyx ventralis), a species of small rail. Over past decades, and during 1972–73 in particular, this remarkable bird came to wide attention over many parts of

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south-eastern Australia as a result of one of its great irregular invasions of our settled southern districts. Moving down from the drying inland early in 1972, blacktailed native-hens arrived in northern and central Victoria singly, then in hundreds, and then in thousands. They would stay in a district for days, weeks and in a few cases, months, then just as suddenly disappear. They appeared in a great arc from near Albury [Wodonga] in north-east Victoria to Nagambie, the outskirts of Melbourne and Geelong, and west to southern South Australia and even Kangaroo Island, where I believe they bred. Though the birds were a novelty to many who saw them, there was in fact nothing new about the influx: it’s been going on intermittently for tens of thousands of years, and is enshrined in the records of the pioneering era. In his great Handbook to the Birds of Australia, John Gould quoted his collector, John Gilbert, whose field journal reported that in the early 1840s, native-hens ‘visited the Swan River colony in myriads, treading down and destroying whole fields of corn in a single night’.2 ‘The natives’, wrote Gilbert, surprisingly, ‘not having seen them before, attributed their presence to the settlers, and called them the “White-man’s birds”: after the harvest was over they nearly all disappeared as suddenly as they arrived’. In South Australia, the experience was remarkably similar, no doubt brought about by the same climatic factors. From Adelaide, Captain Charles Sturt wrote that the native-hen:

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B a r c o o b a n t a m : ‘ I t r uns l ik e hel l ! ’

appeared suddenly … in 1840. It came from the north, fresh flights coming up and pushing on those which had preceded them. It was … evident that they had been unaccustomed to the sight of man, as they dropped in great numbers in the streets and gardens of Adelaide, and ran about like fowls … They took the entire possession of the creek near my house, and broke down and wholly destroyed about an acre and a quarter of wheat as if cattle had bedded on it. They made their first appearance in November, and left us in the beginning of March, gradually retiring northwards …3

Those irruptions may have been the first for many years in the two new colonies, but there have been many notable if less spectacular invasions since, at irregular intervals dictated by a combination of a run of wet seasons inland, which allow the bird’s numbers to increase exponentially, followed by severe dry conditions, which force them to stage the classic drought-time retreat of all Australian waterbirds and head for the coasts. The black-tailed native-hen is a smallish sturdy rail, about 40 cm long with a square folded, upright black ‘bantam’ tail. It is found only in Australia. In outback Queensland, it was (and still may be) known as the Barcoo Bantam. Brownish olive above and dark bluish grey below, with white streaks along the flanks, the underparts are blackish, the bill green with a reddish base, the eye yellow and rather piercing, the legs and feet bright red.

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Those legs are also sturdy and strong, as are the wings – for although it prefers to run, the native-hen can fly very swiftly, with rapid shallow wingbeats. But running is its forte: it can sprint on those strong red legs at impressive speed, well equal to that of a running horse, and by running in closepacked, spreading, shrinking, constantly changing companies, is often able to confuse and outwit predators. Unlike some other rails, it swims rather seldom, much preferring to potter along the margins of streams and swamps, or to roam well away from water, feeding mostly on green vegetation, seeds and some insects. But it nests near or over water, in dense low growth like lignum, usually in loose colonies. The nexus of water, green vegetation and seeds gives the clue to its way of life. In parts of the continent with regular rainfall and permanent swamps, related rails or gallinules like the dusky moorhen (Gallinula tenebrosa) and purple swamphen (Porphyrio porphyrio) do very well. Reliant on their home waters remaining stable, there has been no need for them to alter their basic shape and lifestyle. They find their food around the edge of swamps or while swimming, and if danger threatens, they retreat to the water. But for one of their kind to turn its back on these safe environments and take on the hazards of the rich but highly unpredictable environment of Australia’s inland was another evolutionary challenge altogether. To be successful the rail which tackled that had to be highly mobile and able to move great distances quickly when its temporary home waters dried

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up. It had to be able to respond to changing weather patterns and exploit new growth that results from the sudden heavy local thunderstorms that are a feature of our inland, and moreover to be able to breed at comparatively short notice in virtually any month of the year when such conditions occur. In the event, the bird clearly made that transition successfully. Those unpredictable rains, together with irregular but sometimes immense floods coming down the Channel Country from Queensland into the Lake Eyre basin, and similar flushes coming down the Darling, Lachlan, Murrumbidgee and Murray Rivers, spreading over whole landscapes of lignum-filled depressions, red gum and black box swamps – those have become the black-tailed native-hen’s reasons to be; they create its breeding and foraging strongholds. With the experience of ages it has hammered out an accommodation with those conditions and is able not only to breed irregularly when the waters arrive but even, like grey teal and others, somehow to ‘know’ when distant rain falls or rivers come down in flood. Then it miraculously packs up and travels, appearing on new waters within days or weeks of the event, as though this was the most natural thing in the world. Mostly native-hens travel on foot, taking refuge in numbers, camping by day along what waters they can find. They cross large open spaces by day or at night or find cover in stands of vegetation. But I suspect, when great distances are involved, they must also fly far at night, as other waterbirds do, to cross inhospitable regions in the cool of night and to avoid

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aerial predators, like falcons. On the move in big numbers, black-tailed native-hens provide one of the great wildlife spectacles that Australia has to offer. Police sergeant John Hobbs of Ivanhoe in far western New South Wales described such a mass movement of native-hens in 1972 and commented on their behaviour: On April 28, 1972 I was observing black-tailed native-hens at Bobuna Station, 15 miles north of Ivanhoe … A large sheet of open water had been formed a month earlier by heavy thunderstorms and, as at other similarly formed waters in the district, a huge invasion of native-hens had subsequently taken place. Some 4000 native-hens were feeding around the perimeter of the water in three big groups. As I walked round the water’s edge the birds moved ahead of me in solid mass, a moving carpet of black. Perhaps two or three took to the water … The remainder … headed ‘inland’, scurrying up the gentle slope to the open dry paddock beyond, across which most ran for up to half-a-mile from water … Unlike their relatives, … which head for the sanctuary of water when disturbed on land, the native-hens forget their gallinule ancestry, their watery birthplace, and rely on their running ability to carry them safely across dry land.

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John Hobbs went on to relate how later, when the mob was menaced by a grey falcon: … the native-hens scampered … away from the water across the drying land to the fringe of black box trees … Over 1000 birds were jammed against each other under each of the (three) protecting trees; not a pin, not a feather could have been pushed between them. The falcon made another run along the water’s edge, lifted over the fringe of trees and left.4

Confirming the timing of Hobbs’s comments, the well known Melbourne ornithologist Roy Wheeler, noted that in Victoria the 1972 irruption began in May and built up to a peak in October–November.5 The birds first appeared in mallee areas, sometimes far from water. Some were even reported living in north-west Victoria’s Big Desert, in desert banksia country – surely the last type of habitat one would expect. By spring most dams and swamps in western Victoria from Horsham to Hamilton and Edenhope had varying numbers of native-hens. The birds were reported as far east as Geelong and Ballarat; some reached Werribee [just west of Melbourne]. No breeding was reported. The end of this invasion in the summer of 1973 was described by the western district [pastoralist and] ornithologist, Claude Austin. Writing at Coleraine in western Victoria, he said:

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On February 4 I realised that the two [native-hens] that had been resident on one of my dams were not present. I then started ringing people whom I knew had been hosting them for some time. It is easy to say when you do see a bird but not easy to recall the last time you saw it. However, from what I can gather the black-tailed native-hen invasion finished on 2nd February, 1973. Nobody is sure that they saw one after that date … It is worth remembering that this date corresponded with the big rains in Central Australia and much of the north of the Continent.6

A similar situation obtained in South Australia, where the numbers were immense and some breeding took place. However, most had gone by the end of February 1973, the same month they left Victoria. In September 1974, Mr NW Schrader (also of Ivanhoe, in inland New South Wales) provided a neat postscript to the whole affair. Relating how when travelling by train west from Ivanhoe on 11 May 1973, Mr Schrader said he saw no nativehens. But on the return journey to Ivanhoe the next morning, after rain had fallen: As far as the eye could see there were moving waves of native-hens, running and zigzagging, rarely flying, away from the moving train … A rough count indicated that there were about 10 000 birds in sight of the track, over the length of the journey.7 42

B a r c o o b a n t a m : ‘ I t r uns l ik e hel l ! ’

The siren song of inland rain was drawing the blacktailed native-hens north again, as it always has, and always will, if we have the strength and resolve to protect their habitats and keep enough inland waters at least secure from exploitation for irrigation, cotton-growing, mining and uranium contamination.

Endnotes 1

This chapter was originally published as ‘Barcoo Bantams’ in Graham Pizzey (2000) Journey of a Lifetime: Selected Pieces by Australia’s Foremost Birdwatcher and Nature Writer. Angus & Robertson: Sydney, pp. 197–203, and is reproduced here with the kind permission of the Pizzey family and HarperCollins Publishers. Square brackets indicate editorial insertions in this version, which is reproduced in the original form apart from changes to the referencing style to conform to this book. Graham Martin Pizzey AM (1930–2001) was a well-known Victorian nature writer and conservationist, best known for his Field Guides to Australian Birds, the 1997 version being completed in partnership with illustrator Frank Knight. He travelled extensively throughout Australia observing birds, often with other leading ornithologists like Claude Austin, whom he quotes here.

2

Gilbert in Gould J (1965) Handbook to the Birds of Australia. The Author: London, pp. ii, 326.

3

Sturt’s 1840 diary, online at http://ebooks.adelaide.edu.au/s/sturt/ charles/s93n/part19.html ‘Birds: 126. TRIBONYX VENTRALIS, GOULD.—The black-tailed Tribonyx’.

4

Hobbs J (1973) Reaction to predators by the black-tailed native hen. Australian Birdwatcher 5(1), 29.

5

Wheeler R (1973) The black-tailed hen recent invasion. Geelong Naturalist 10(2), 45–50.

6

Austin C (1973) Black-tailed native hens. Bird Observer 497, 7–8. [The La Niña event of 1973–1975 was the greatest ‘wet’ event of the 20th century in Australia. At this time the normally dry Lake Eyre was filled to capacity and many of the inland floodplain areas were inundated.]

7

Schrader NW (1974) Invasion of black-tailed native-hens. Australian Bird Watcher 5(6), 234. 43

R AIN

AND GR ASS : LESSONS IN

HOW TO BE A ZEBR A FINCH Steve Morton

Two threads make up this story. One concerns that hardy speck of a bird, the zebra finch (Taeniopygia guttata), so aptly described as a ‘little Aussie battler’, a small and seemingly insignificant animal that nevertheless not only endures the toughest country in Australia but succeeds to the extent that it swarms in extraordinary abundance.1 The other involves the biologists who have studied these fascinating birds, beginning with the dedication of the great German ethologist, Klaus Immelmann, and including many of Australia’s most prominent ornithologists such as Jock Marshall, Harry Frith and

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Dom Serventy. In one way or another, each of the human characters in this story has been motivated by a desire to understand how it is that such a bird, one that must drink in order to survive on its diet of seeds, should be so successful in the face of a climate with one of the most variable rainfall patterns on earth. Each of these people has contributed significant parts to a complete picture of the zebra finch, while in passing also helping us to understand why Australian deserts look and feel as they do. But each, including myself, comes to the work’s end still facing some deep mysteries in this lively, diminutive creature.

The environmental setting for zebra finches The words ‘vast’ and ‘tough’ do not exaggerate the qualities of the Gibson Desert. This stretch of country in the centre of Western Australia has remained oblivious to the efforts of people of non-Indigenous origins to find uses for it. It is an arid land of thin stony soils with pale-yellow hummocks of spinifex, swathes of grey-green mulga, and big expanses of sand sheet with yet more spinifex. Even the optimistic and adventuring pastoralists of earlier times avoided such forbidding country, and cattle were never brought here. White people circled around it, and left it mostly empty of their aspirations. Even the name we place on the map, of poor Alfred Gibson, sprang out of his failure, misery and lonely death.2 Empty it is not, of course. At Patjarr, for example, Ngaanyatjarra people still live on their land, their home, as they have done for thousands of years.3 This settlement of some 20 build-

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ings sits on a low rise with spinifex, a tiny human community in an enormous landscape. Among the sparse acacia and grevillea bushes between the scattered houses fly groups of zebra finches, calling out to each other in their constant and characteristic refrain. The zebra finches are the most common birds in the settlement. They too see the Gibson Desert as home, and must have done so for almost countless generations. In their world-view there may be little here that could be described as harsh or difficult. Their songs sound cheerful and optimistic as they stir up in clusters from the puddles around the dripping taps, and steer off high in the morning air to seek out the richest patches of grass seed in the surrounding country. This scene is repeated daily at innumerable locations across inland Australia, at settlements, homesteads and stock waters, in creekbeds, soaks, and clay-pans, at rock-holes far distant from our vision. Of course it is possible to find places where zebra finches do not appear to be present. However, it is far more probable than not that, if one has the ability to wait, their carefree cries will eventually be heard, almost wherever it might be. Theirs is a story of persistence in an environmental setting that many other organisms find impossible. It tantalises us that these finches make their way so successfully in what most humans see as one of the most challenging landscapes on earth. The Ngaanyatjarra and their neighbours to the east, the Pitjantjatjara, know the zebra finch well; indeed, the bird is embedded in the Tjukurpa, the source of life and law.4 When

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CSIRO undertook a fauna survey of the Uluru-Kata Tjuta National Park in the late 1980s we were given a privileged glimpse of the ecological knowledge possessed by Pitjantjatjara people. 5 It struck me particularly how beautiful were the onomatopoeic names that these Western Desert peoples have for so many birds. The crested bellbird (Oreoica gutturalis) glories in its perfect designation of parnparnpalala. Neatly, the zebra finch is called nyii nyii, a fitting representation of the nasal call that it uses constantly for identity and contact. The common name of zebra finch provides its own logic and enjoyment but, whenever I think of the bird, in my head I hear nyii nyii. The depth of knowledge of natural history to be found among the Western Desert peoples is such that we in the scientific community may well have saved ourselves a lot of time by asking them earlier about the life-history of nyii nyii. Although this is not what happened, the story has its own fascination.

Klaus Immelmann and opportunistic breeding In May 1960 the eminent finch biologist, Klaus Immelmann, arrived in Alice Springs for a short visit as part of his year-long fieldwork in Australia. In November 1959 he had been in Kununurra, in the Kimberley region of Western Australia. There he had observed the response of zebra finches to the early storms of the approaching wet season, and documented the consequent increase in courtship, nest refurbishment and, eventually, nesting.6 Immelmann then traveled to central Aus-

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tralia, where he spent all of May. Happily, he was present early in the month when 22 mm of rain fell. Within hours, he noted, zebra finches were courting and, in a couple of cases, copulating. He also observed several other species, such as budgerigars and singing honeyeaters, gathering nesting materials the day after the rain. It is clear from his reporting of these events in English that his mind was made up. Rain itself must have a stimulating activity on gonadal development … Most of the birds of central Australia breed at any time of the year according to rainfall.7

Immelmann was an outstanding biologist yet, as Richard Zann has noted, most of his work was published in German and remains poorly quoted in the English-language literature. Zann’s own monograph on zebra finches was written to some extent ‘to make accessible to readers of English additional details of Immelmann’s fine fieldwork’.8 Zann pays due and abundant credit to Immelmann’s dedication, which began with the gift of a pair of zebra finches when the German was only 12 years old and continued for his entire scientific career.9 The domesticated zebra finch became a worldwide laboratory subject for behavioural research to a considerable extent because of Immelmann’s efforts. Subsequently, Richard Zann brought the zebra finch back to Australian science by devoting his career to working on it in the field. Both these men deserve our gratitude for the light that they have shed.

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Why did Immelmann so readily arrive at the conclusion that the zebra finch bred opportunistically? The answer is that this idea was in the Australian air, and no doubt Immelmann was deeply conversant with the writings of local biologists published throughout the 1950s. For example, the egg collector Ivan Carnaby produced a dramatic account of the reproductive response by numerous species of birds to rainfall in late summer, in the Gascoyne district east of Carnarvon in Western Australia.10 However, probably the most important impetus was created by the well-known ornithologist, Jock Marshall, who had undertaken his doctoral studies in Britain with the eminent zoologist John Baker, an expert on the breeding patterns of tropical birds.11 Marshall had been working throughout the decade to explore physiological control of gonadal activity, with a view to demonstrating that the males of many desert birds were capable of reacting rapidly to rainfall in order to prepare for a breeding event. He had good links with other active field workers such as Allen Keast, and in 1955–1956 Dominic Serventy joined him in Britain to study the breeding cycle of captive zebra finches.12 These scientists had the support of the influential Francis Ratcliffe, who in 1955 wrote to Serventy as follows: I have recently come to the conclusion that the conditions of our large continent with its unreliable rainfall put a high selective value on mobility and opportunism in relation to breeding … There must obviously be a high selective value in any mechanism that helps breeding to be triggered off

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by temporary and locally favourable conditions, while suppressing it at times and in places where the conditions are not favourable.13

By 1971, when Serventy published his magisterial overview of desert birds, opportunistic breeding appeared to have reached paradigmatic status. In writing of the ‘peculiar flexibility of the breeding pattern in Australia’, Serventy noted that the zebra finch and the budgerigar (Melopsittacus undulatus) ‘rank with the grey teal [Anas gracilis] as highly successful opportunistic breeders in arid areas’.14 Although Serventy found himself having to qualify the conclusion, he confirmed the accuracy of the notion of opportunism: There has been a tendency to claim … that birds in arid areas have broken free of the rigidity of regular breeding seasons … It appears, however, that all this is generally true only of the arid areas of Australia … In Africa … the breeding seasons … were rather more regular … Exceptionally heavy rains experienced in the Namib in September 1965 had very little effect on breeding activities … In Australia the response would have been massive … It would appear that the great flexibility in breeding, for which the arid areas of Australia are now noted, does not operate uniformly over the arid regions of that continent but characterises mainly the west and central parts … the truly arid parts.15

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Opportunistic breeding made sense, after all. As part of the adjustment by white Australians to the peculiarities of their new continental home, biologists were finally starting to voice pride in the country’s fauna and flora. In one prominent example, it became clear that part of the adaptive repertoire of marsupials was an ability to respond with flexible breeding to the variable climate of the continent, as was being revealed by studies of the reproduction of the big kangaroos.16 Subsequently, the notion spread that marsupials were not inferior mammals; rather, they were superbly adapted organisms.17 John Calaby, the wise historian of Australian zoology, put it this way: Once upon a time, about 30 years ago, when nearly all wisdom resided in the Northern Hemisphere, there were certain revealed truths about Australia … It was dogma … that Australia’s more distinctive vertebrates such as marsupials, were physiologically and ecologically inferior to their counterparts in the Northern Hemisphere … ‘Down under’ was a good place to banish such secondclass animals.18

In this new view, the continent was an exciting land of boom or bust with organisms cleverly adapted to sustaining themselves through thick and thin, and this truth was now exemplified by the birds of the Australian deserts. It may well have helped gain credibility for the idea that Immelmann, the

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great German biologist, was confirming an important Australian insight. My comments imply neither scepticism nor sarcasm. The insight into opportunistic breeding was partially true, and partially not. What emerges with the passage of time is a reproductive story in zebra finches that is more complex than was first thought, and far more interesting. I would hazard a guess that, 30 years hence, some of the thoughts expressed in this present paper may evoke charges of naïvety. Constant reinterpretation is the very nature of science, after all. In this spirit, events began to challenge the generality of the opportunistic breeding notion fairly quickly, as foreshadowed by Dom Serventy’s review. In December 1974, while undertaking my PhD studies on the reproductive pattern of an insectivorous marsupial, I met Gordon Maclean at the Fowlers Gap Research Station, north of Broken Hill. Maclean, a South African biologist interested in desert waterbirds, was visiting Australia on study leave. Both of us were wrestling with the fact that our arid study-sites were extremely difficult to gain access to because of the constant risk of becoming bogged. During 1974 Fowlers Gap received 630 mm, nearly three times its mean annual rainfall of 238 mm. It seemed that the creeks would never stop running, the saltbush plains never again dry out. It rained through the winter and continued on into the spring and summer. Maclean took advantage of the conditions to monitor breeding of as many bird species as he could find.

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When Maclean’s paper appeared describing the results of this important investigation I found it puzzling. The facts seemed evident: there was limited, if any, evidence of opportunistic breeding, despite the extraordinary nature of the seasons. Yet the paper seemed to struggle against stating this obvious result. Instead it read as if opportunistic breeding was an established fact, that it should have been occurring, and its apparent lack had to be explained away: As a result of unusually heavy rains since 1973, many bird species in north-western New South Wales became seasonal breeders, starting to nest in the spring of 1974. Opportunistic breeding was no longer released by rainfall … Even such a common and well-known opportunist as the zebra finch … started to breed late in the season and then only in small numbers.19

I was exposed first-hand to the paradoxical power of a paradigm to constrain thought while simultaneously purporting to offer a framework upon which it ought to be extended. That trip to Fowlers Gap brought about my involvement in this story. Along with Gordon Maclean, there was also Dick MacMillen on sabbatical leave from the University of California, Irvine. MacMillen was the pre-eminent physiological ecologist working with desert seed-eating rodents and birds.20 He has a generous and attractive personality, and in that short time together at Fowlers Gap we found much in common. By

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early 1977, I was working on granivorous mammals as a Postdoctoral Scholar in his laboratory in California, and on my return to Australia continued with studies of the seed-eating animals of our deserts. So I too was led to the zebra finch.

The breeding periodicity of zebra finches, beyond Immelmann Even at the time of Immelmann’s observations at Alice Springs in 1960, some data on the periodicity of breeding by zebra finches showed that an opportunistic pattern seemed unlikely to be universal. Harry Frith had been employed by CSIRO at Griffith, in the Riverina of New South Wales, in studies of orange trees in irrigated orchards. He found that most of the trees contained the nests of zebra finches and, after he transferred to the Wildlife Survey Section under Francis Ratcliffe in 1952, he was able to indulge his passion for birds by monitoring their breeding from 1953 to 1956.21 The results showed that the birds bred seasonally, with no reproduction in June or July.22 Another great Australian ornithologist, Jiro Kikkawa, had also become interested in breeding periodicity, and from 1961 to 1965 he examined zebra finches at Armidale in New South Wales.23 Here at the moist, cold end of the climatic envelope for the species, breeding was found to be distinctly seasonal although reproduction did occur in every month except June and July. In the next instalment, Stephen Davies worked at Mileura Station near Meekatharra in arid Western Australia, where the

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mean annual rainfall is about 200 mm. Davies sampled zebra finches specifically to test Serventy’s assertion that the bird was a highly successful example of an opportunistic breeder under arid conditions.24 Falls of rain in the winters of 1973 and 1974 failed to stimulate breeding. Davies concluded that at Mileura, rainfall alone was insufficient to initiate breeding, and that it took place in warm periods and especially in spring in years of effective summer rain when the birds’ food supply has germinated, established and seeded well. Subsequently, Zann re-analysed Davies’ data to reveal that zebra finches at Mileura have the ability to breed in any month and to breed continuously at low levels, while reproducing slightly more in spring and summer.25 The relationship between rain and breeding remained ultimately unclear, but obviously breeding may not be attempted for many months at a time. I admire Davies’ paper because it, for the first time, spoke vividly of the importance of understanding the resource on which the hatchlings of the zebra finches need to be fed. Until this time there seemed in the discussion to be a vacuum between rain and finch; perhaps there was a sort of ether that magically transformed moisture into young birds? The significance of this matter might seem evident enough to us now, but Davies had opened up a new domain by writing about grass seeds. Richard Zann was introduced to zebra finches by Jiro Kikkawa, and indeed assisted with the sampling of the Armidale birds. After he established himself at La Trobe University he began his long-standing field work on the species near

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Shepparton in northern Victoria. His studies throughout the 1980s revealed seasonal breeding from spring to autumn, with no reproduction occurring from May through August.26 He contrasted his findings to those from arid and northern populations by noting that rainfall did not stimulate breeding. At this point we have, therefore, a multitude of studies citing variations in breeding periodicity, ranging from marked seasonality through to poorly understood flexibility, the latter perhaps involving opportunistic breeding in response to rainfall. In trying to interpret this variation Zann, following Davies, spent much time trying to understand the connection of breeding to patterns of seed availability and nutrition of the young; and so we come to the grasses, where important lessons are to be found.

A profusion of grass seeds From the time of John Gould it has been known that zebra finches feed especially on grass seeds; the members of the family to which they belong, the Estrildidae, have long been called ‘grass-finches’. My work on seed-eating animals in the Australian deserts confirmed that avian granivores were more prominent here than in arid regions elsewhere in the world.27 When compared to several other deserts around the globe, Australia seems to possess few seed-eating rodents, and so I chose to examine the dietary ecology of the zebra finch, the smallest of our seed-eating birds and hence potentially the most rodent-like in size. Analysis of the crops from 97 zebra

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finches from two locations (Fowlers Gap Research Station, and Sandringham Station near Bedourie, Queensland) revealed an astonishing result.28 Every one of the 34 221 seeds that we counted came from a grass, and other material such as insects was vanishingly rare in the diet. Other analyses confirm the generality of this study.29 It is most impressive to consider how completely the zebra finch’s world revolves around grass. This may be due to the facility with which grass seeds can be handled: they are not firmly attached to their protective coat and are relatively easily husked, which zebra finches always do before swallowing; they are usually without sticky or hard protective layers; and they lack toxic chemicals.30 Most importantly, many grasses are prolific seeders and several species (including members of genera such as Enneapogon, Aristida, Panicum, Eragrostis and Eriachne) are particularly abundant in run-on areas of the landscape where they grow, mature and set seed quickly, even after small flushes of rain.31 We begin now to get a glimmer of understanding. A principal tool in the survival kit seems to be selection of a type of seed resource that can be quickly rejuvenated by small rainfalls. This resource stems from grasses that can generally be found in special places somewhere in a region, allowing for persistence of the birds in the drier times, thereby carrying them through to those periods when really big rains produce a flush of seeding grasses everywhere. The clever nyii nyii have found a way to avoid the worst features of an environment

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that superficially looks like it is dominated by a boom and bust dynamic: they have found a mechanism to smooth out the fluctuations. There is still a job to be done each day, of course. The daily metabolic requirement of the individual zebra finch is some 5 g of food, which translates to somewhere between 1000 and 6000 seeds (depending on their size). At first sight this number looks frightening, but the birds are adept at husking and consuming seeds when they are abundant. Perhaps the necessary volume of seeds can be accumulated in the crop within a couple of hours of foraging per day when conditions are suitable.32 Armed with this knowledge of the grass seed, perhaps we have a better chance of uncovering explanations for the wide divergence between populations in their breeding periodicity.

Seven long years Richard Zann came to spend sabbatical leave with me at CSIRO’s laboratory in Alice Springs in 1986. He turned up during one of the extended dry spells that characterise the inland, but good fortune delivered him 170 mm of rain in late June within a short period of his arrival (the mean annual rainfall is 263 mm). His observations confirmed that, prior to the rain, no breeding had occurred for many months. But he was especially struck by the fact that no breeding behaviour – no courtship, no nest-building – was stimulated by the massive June rain. One of Immelmann’s greatest admirers saw with his

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own eyes that reconciliation of such diverse observations on rain and finches was required. Together with his American colleague, Nancy Burley, who had joined this effort bringing a behavioural perspective, Zann began sampling free-flying zebra finches near the laboratory at a walk-in trap baited with commercial bird-seed. Young birds were allocated to four age categories based upon the colour of the bill and irides, and these categories used to estimate laying date. This method works until the young birds reach about three months of age, after which time they attain adult features. When he was preparing to return to Melbourne, Zann convinced me that it would be a relatively simple matter to continue the sampling in order to obtain the type of extended record necessary to resolve the long-standing questions about periodicity of breeding. The hero of the resulting seven-year sampling effort was Kevin Jones. Despite many other demands made on his time in assisting me with different studies across the southern Northern Territory, until December 1992 Jones took monthly samples whenever he could manage it, gaps notwithstanding. Breeding occurred in every month of the year, with a peak in October and the lowest point in April.33 Heavier rainfall led to more intense breeding episodes. Once a breeding episode began, surges of reproduction were stimulated by follow-up rains. There was a lag between the onset of the first rains and the onset of breeding of one to two months in summer and two to three months in winter, a variation clearly designed to

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ensure that hatching coincides with the first availability of ripening grass seeds. It appears that the timing of seed set is complexly related to the prevailing temperatures immediately after rain and the amount and timing of previous falls of rain; it may also be affected by the species composition of the grasses. In short, we concluded that zebra finches track these complex factors and breed opportunistically so as to time the hatching of their young with the availability of fresh supplies of half-ripe seed. We believe we can now, finally, fully interpret Immelmann’s observations of rainfall stimulating breeding behaviour in May 1960. The rainfall figures for the preceding six months suggest that a breeding episode may well have begun in November 1959, peaked in February or March 1960, and tailed off through April. The rainfall witnessed by our predecessor was probably a stimulus for a small resurgence in reproduction rather than the impetus for an entirely new episode. Furthermore, the results from Alice Springs provide context for interpreting prior studies of the timing of breeding in zebra finches.34 The annual patterns across Australia are coupled with underlying seasonal pulses of plant production. Like almost all avian species, zebra finches in the south and east breed principally in spring and summer. Those in the inland breed irregularly, and it seems probable that, because of temperature gradients, the further north one proceeds the less likely it is that such breeding will be suppressed in winter if moisture allows it. Underlying this irregularity in the arid

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zone, nevertheless, is a consistent trend over periods of years towards stronger breeding in spring months. Overall, zebra finches everywhere have the potential for very long periods of breeding to an unusual degree compared to most other Australian birds. Conversely, they may also fail to breed for extended periods. Immelmann, Marshall, Serventy and their colleagues were fundamentally correct in concluding that the zebra finch was an unusual opportunistic breeder, and that occupation of the highly uncertain environment of inland Australia could be enhanced by such habits. The subsequent work has swept away a couple of myths about the nature of that opportunism, and replaced them with knowledge of some unexpectedly complex relationships among finches, rain and grass.

Is opportunistic breeding widespread? Some authors reacted against the generalisation, for example in Serventy’s overview, that birds of the Australian deserts exhibit opportunistic breeding to a greater degree than on other continents – even with his qualification that it is characteristic only of the ‘truly arid parts’ (whatever that might mean).35 For example, some 20 years later Hugh Ford concluded that the Australian environment was rather more ‘equable’ than other authors had been inclined to admit; by equable he meant lacking strong seasonality: This equable environment and rather restrained breeding is in marked contrast to the picture that is often portrayed

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of Australia as a land of extremes to which birds respond in an opportunistic pattern … Opportunistic breeding, especially after rain, has probably been over-emphasised.36

Tellingly, Ford went on to comment on an underlying reason why his fellow scientists may potentially have indulged in such exaggeration: ‘In central Australia it is perhaps the ornithologists as much as the birds that are opportunistic’.37 Ford was suggesting that too many conclusions had been extrapolated from scientific observations made during excursions into the inland rather than from the long-term studies necessary to sort signal from noise, a point of view for which, given my involvement in the zebra finch story, I naturally have strong sympathies. Furthermore, Ford has some logic on his side because most birds are insectivores, and the abundance of insects may be directly depressed by the low temperatures that characterise winter across much of the arid zone, regardless of the timing of rainfall. Let us not swing too far the other way, however. Given the substantial coastal bias in knowledge of life-histories of Australian birds, Ford’s conclusions about equability could themselves be exaggerated.38 Those with substantial experience in Australia’s arid landscapes are likely to emphasise a different equilibrium between the two extremes of opportunism and mild, equable seasonality. Richard Schodde is one such ornithologist, and the conclusions in his review of 1982 seem to me to balance around the correct point:

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There is no doubt that the incidence of opportunistic breeding is proportionately much higher in the Australian arid zone than in peripheral mesic regions … Furthermore, it is correlated with nomadism … Its converse, that sedentary species breed seasonally, is equally true … Winter sets the stage for spring breeding ... followed in spring by an increase in plant growth and an associated increase in the abundance and activity of insects. No matter how weakly expressed, this is the one regular cycle in the desert to which birds can gear their reproductive rhythms; increasing day-length and perhaps temperature are, in turn, its only consistent clues. These factors, rather than rain or food itself, thus appear to be the primary zeitgebers triggering breeding for most birds of the Australian arid zone, particularly the more sedentary species.39

In closing, then, the complexities of the patterns exhibited by the zebra finch, a species far better known than most, lead us to be cautious in assuming too much about opportunism in other species in the absence of convincing information.40

Desert life can be harsh Patjarr Creek emerges from the Clutterbuck Hills some 10 km north-west of the settlement of Patjarr. It is a small watercourse, but where it leaves the low hills it has enough energy to cut a channel a couple of metres deep. We arrived there about 9 am

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from Minna Minna, the coolibah claypan 4 km away which the creek floods occasionally before dissipating among the sand hummocks. A pool some 5 m2, and at its deepest about 40 cm, remains in the lowest part of the gully. It is late October, and warm, and the nyii nyii need this water. The mulgas around the soak are packed with them, each thin branch pulsating with small bodies, and the air is vibrating not only with their nasal voices but with the rush of their massed wings as they rise out of the bushes in mobs of hundreds to whirr down to the pool. There are so many of them that my powers of estimation are overcome. Are there five thousand, maybe as many as eight thousand? It is a startling and enthralling vision of abundance, incontrovertible evidence of a hugely successful combination of life-history and behaviour in the heart of the Australian desert. Within minutes of our arrival a dark shape swoops in. An Australian goshawk sprays birds from the pool as it flashes across the creek before coming to rest as a silhouette at the centre of the densest mulga on the bank. The nyii nyii set up an even greater racket than before, but they have to abandon drinking. After a few moments I put the predator up by jumping into the bed of the creek and clambering out the other side; he flies off up the stony slope, where I see his larger mate awaiting him. Within a further minute the nyii nyii are back at the water, pouring in living, feathered waves out of the mulga shrubs, over the lip of the bank, and down to the precious moisture.

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Earlier I emphasised the adaptive fit of the zebra finch to its country, but recognition of this point needs to be balanced with an appreciation of the knife-edge of survival on which each bird must perch. Life can be snuffed out in a sharp instant by the talons of a raptor. Eggs are destroyed, one clutch after another, by other birds and by ubiquitous goannas.41 Death from heat exhaustion is always a possibility.42 As if all this were not enough, consider the task of finding water and abundant grass seed in an arid landscape, every single day. Despite all that we have discovered about the zebra finch, its ability to rise to this challenge still leaves me astonished, feeling that I have not yet penetrated to the mysterious core of the success of these tiny birds. How do they do it? In the laboratory zebra finches can persist without drinking water indefinitely, through mobilisation of water from metabolism of the carbohydrate in seeds.43 In the wild, though, especially when conditions are warm to hot, drinking water is almost certainly required every day.44 I have sat in camp debating endlessly with my colleagues how the nyii nyii seen nesting far out on the waterless sand-plain could possibly be locating a soak and flying back and forwards the necessary 50 km every day. What may account for their water-locating abilities? It may well be possible that the birds regularly fly 50 km or more in their daily lives. In order to do so, though, they must possess outstanding navigational abilities, because some of the waters that they gain access to are minute and inconspicuous in a sea of sand, stone and bush. Of course it is possible

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that we biologists severely underestimate the availability of water. Contrasts of the cartography of waters from Indigenous and non-Indigenous knowledge-sources can reveal dramatic differences; whitefellas may simply be unaware of waters well known to Indigenous people.45 In turn, nyii nyii may know of many more waters than we do, may see far more than the apparently featureless vastness that sometimes overwhelms us. We do not know the likelihood of each of these possibilities. Assume, though, that the bird is secure in its knowledge of water. Now it needs to monitor the even more obscure pattern of rich patches of the special grass seeds that it must harvest at the rate of thousands a day. It must create another map in its head of all such places, ranked in priority for foraging. It must constantly be on the alert for signs that the patch being used is dropping below an energetically secure threshold, and be ready to abandon it in favour of the next option. Zann’s work shows remarkably high mobility by banded birds, such that a particular colony experiences both pronounced immigration and emigration.46 Perhaps the birds are sampling different patches on different days, in order constantly through testing to maximise foraging success. Not even one mistake can be afforded, however; if both food and water are not forthcoming, rapid death awaits. In this country of long dry spells, the birds must be ready not just to abandon a locale but perhaps even an entire region. We have only the vaguest of ideas about the range of movements, unfortunately, and so understanding of how zebra

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finches manage this extreme spatial uncertainty in food supply still eludes us. Some observers have suggested that mass death can occur during extended droughts, and in some places, such as the Gibson Desert, the species can be classified as ‘irruptive’.47 If the birds choose a bad option for movement as seeds and water decline – by waiting too long or, like the wretched Alfred Gibson, by travelling in the wrong direction – such outcomes must occur occasionally. There is little doubt in my mind that sometimes the nyii nyii do indeed go bust. But the quality of the spatial perception that the birds possess should not be underrated too hastily. When I consider the mystery that still lies at the core of the animal’s successful persistence, I visualise a sophisticated neuronal network busily pulsing in its brain. This unseen network achieves the most extraordinary results. It somehow calculates the probable time-course by which rainfall will be translated into half-ripe grass seed for hatchlings; it must do so by integrating a record of likely soil moisture prior to the most recent rainfall with current and probable future temperatures in order to estimate seeding by the grasses. While this longer term analysis is occurring, the brain is also processing a dynamic geographical map in daily time. It is calculating the spatial pattern of waters in relation to rich patches of grass seeds, again undertaking sophisticated integration to judge when previously sampled sites are expected to come into top condition as their grass seeds fall to the ground in peak abundance. A map is being constructed by which the most efficient pathways of daily

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movement among water and seed are decided. The brain’s data files are also being updated daily in order to prepare decision rules for abandoning the region altogether. When that happens, it seems not beyond belief that the neuronal network accesses an archival map of previously inhabited regions that the bird can fall back upon with some expectation of survival. Because of these considerations, I remain in awe of the little nyii nyii.

Rain and grass Perhaps it is too easy to verge on anthropomorphism with a bird that gives such a sense of vivacity. The zebra finch seems so full of life, its voice so cheerful, that one can be seduced into thinking it capable of meeting any challenge the uncertain desert environment might throw at it. For it is true that inland Australia is subject to tremendous fluctuations in supply of the two critical factors, rain and grass: those conditions can boom, and they can bust. But the bird’s capabilities go a long way towards smoothing out the impacts of those fluctuations on reproduction and survival. Its breeding is undeniably remarkable in its flexibility. Numerous unanswered questions remain about the navigational behaviours that are used to minimise the hazards of such uncertain supplies of seed and water, questions that will remain mysterious until some clever scientist places tracking devices on zebra finches and analyses their undoubtedly sophisticated movements.48 My guess is that, when this information comes in, we will be astonished yet

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again at the panache with which the nyii nyii succeed in dealing with country that seems to us in our ignorance to be so inimical to occupation.49

Endnotes 1

Zann RA (1996) The Zebra Finch: A Synthesis of Field and Laboratory Studies. Oxford University Press: Oxford, p. vi.

2

Giles E (1889) Australia Twice Traversed. Sampson, Low, Marston, Searle and Rivington: London.

3

Patjarr is 170 km north of Warburton, Western Australia, at 24°28 S. 126°17 E.

4

Cane S (2002) Pila Nguru: The Spinifex People. Fremantle Arts Centre Press: Perth, pp. 137–39.

5

Baker L, Woenne-Green S and the Mutitjulu Community (1993) Anangu knowledge of vertebrates and the environment. In Uluru Fauna: the Distribution and Abundance of Vertebrate Fauna of Uluru (Ayers Rock-Mount Olga) National Park, N.T. Kowari. (Eds. JRW Reid, JA Kerle and SR Morton) pp. 79–132. Australian National Parks and Wildlife Service: Canberra.

6

Immelmann K (1963) Tierische Jahresperiodik in ökologischer Sicht: ein Beitrag zum Zeitgeberproblem, unter besonder Berücksichtigung der Brutund Mauserzeiten australischer Vögel. Zoologische Jahrbücher Abteilung für Systematik Okologie und Geographie der Tiere 91, 91–200.

7

Immelmann K (1963) Drought Adaptations in Australian Desert Birds. Proceedings of the XIII International Ornithological Congress, pp. 649–57; esp. p. 650 and p. 655.

8

Zann RA (1996) p. v.

9

Zann RA (1996) p. xv.

10 Carnaby IC (1954) Nesting seasons of Western Australian birds. Western Australian Naturalist 4, 149–56. 11 Astheimer LB and Buttemer WA (2002) Changes in latitude, changes in attitude: a perspective on ecophysiological studies of Australian birds. Emu 102, 19–27. 12 Robin L (2001) The Flight of the Emu: A Hundred Years of Australian Ornithology. Melbourne University Press: Melbourne, p. 189; Keast JA and Marshall AJ (1954) the influence of drought and rainfall on reproduction in 70

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Australian desert birds. Proceedings of the Zoological Society of London 124, 493–99; Serventy DL and Marshall AJ (1957) Breeding periodicity in Western Australian birds: with an account of unseasonal nesting in 1953 and 1955. Emu 57, 99–126; Marshall AJ and Serventy DL (1958) The internal rhythm of reproduction of xerophilous birds under conditions of illumination and darkness. Journal of Experimental Biology 35, 666–70. 13 Robin L (2001) pp. 189–90. 14 Serventy DL (1971) Biology of desert birds. In Avian Biology. (Eds. DS Farner, JR King and KC Parkes) pp. 287–339, esp. p. 307 and p. 310. Academic Press: New York. 15 Serventy DL (1971) pp. 306–7 and p. 309. 16 Frith HJ and Calaby JH (1969) Kangaroos. FW Cheshire: Melbourne. 17 Parker P (1977) An ecological comparison of marsupial and placental patterns of reproduction. In The Biology of Marsupials. (Eds. B Stonehouse and D Gilmore) pp. 273–86. University Park Press: Baltimore. 18 Calaby J (1984) Foreword. In Vertebrate Zoogeography and Evolution in Australasia: Animals in Space and Time. (Eds. M Archer and G Clayton). Hesperian Press: Sydney. 19 Maclean GL (1976) Rainfall and avian breeding seasons in northwestern New South Wales in spring and summer 1974–75. Emu 76, 139–42, esp. p. 139 and p. 142. 20 MacMillen RE (1990) Water economy of granivorous birds: a predictive model. Condor 92, 379–92. 21 Tyndale-Biscoe CH, Calaby JH and Davies SJJF (1995) Harold James Frith 1921–1982. Historical Records of Australian Science 10, 247–63. 22 Frith HJ and Tilt RA (1959) Breeding of the zebra finch in the Murrumbidgee Irrigation Area, New South Wales. Emu 59, 289–95. 23 Kikkawa J (1980) Seasonality of nesting by zebra finches at Armidale, NSW. Emu 80, 13–20. 24 Davies SJJF (1977) The timing of breeding of the zebra finch Taeniopygia guttata at Mileura, Western Australia. Ibis 119, 369–72. 25 Zann RA (1996) pp. 123–124. 26 Zann RA (1996), pp. v–vi; Zann RA and Straw B (1984) Feeding ecology and breeding of zebra finches in farmland in northern Victoria. Australian Wildlife Research 11, 533–52; Zann RA (1994) Reproduction in a zebra finch colony in south-eastern Australia: the significance of monogamy, precocial breeding and multiple broods in a highly mobile species. Emu 94, 285–99. 71

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27 Morton SR (1985) Granivory in arid regions: comparison of Australia with North and South America. Ecology 66, 1859–66. 28 Morton SR and Davies PH (1983) The diet of the zebra finch (Poephila guttata), and an examination of granivory in birds of the Australian arid zone. Australian Journal of Ecology 8, 235–43. 29 Davies SJJF (1977); Zann RA and Straw B (1984). 30 Soobramoney S and Perrin MR (2007) The effect of bill structure on seed selection and handling ability of five species of granivorous birds. Emu 107, 169–76. 31 Davies SJJF (1986) A biology of the desert fringe. Journal of the Royal Society of Western Australia 68, 37–50; Stafford Smith DM and Morton SR (1990) A framework for the ecology of arid Australia. Journal of Arid Environments 18, 255–78. 32 Zann RA and Straw B (1984). 33 Zann RA, Morton SR, Jones KR and Burley NT (1995) The timing of breeding by zebra finches in relation to rainfall in Central Australia. Emu 95, 208–22. 34 Zann RA (1996) pp. 125–29. 35 Serventy DL (1971). 36 Ford (1989) Ecology of Birds: an Australian Perspective. Surrey Beatty & Sons: Sydney, p. 110 and p. 232. 37 Ford (1989) p. 234. 38 Clarke MF (1997) A review of studies of the breeding biology of Australian birds from 1986–95: biases and consequences. Emu 97, 283–89. 39 Schodde R (1982) Origin, adaptation and evolution of birds in arid Australia. In Evolution of the Flora and Fauna of Arid Australia. (Eds. WR Barker and PJM Greenslade) pp. 191–224, esp. pp. 199–200. Peacock Publications: Adelaide. 40 Astheimer LB and Buttemer WA (2002). 41 Zann RA (1996) p. 76. 42 Zann RA (1996) p. 70. 43 MacMillen RE (1990). 44 Zann RA (1996) pp. 62–64.

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45 Donaldson T (2002) ‘What they call that in the Whites?’: Ngiyampaa and other placenames in a New South Wales Ngurrampaa. In The Land is a Map: Placenames of Indigenous Origin in Australia. (Eds. L Hercus, F Hodges and J Simpson) pp. 207–38. Pandanus Books: Canberra; Yu S (2002) Ngapa Kunangkul (living water): an indigenous view of groundwater. In Country: Visions of Land and People in Western Australia. (Eds. A Gaynor, M Trinca and A Haebich) pp. 33–55. Western Australian Museum and Lotteries Commission of WA: Perth; Walsh F (2008) To Hunt and to Hold: Land Use Systems and Perceptions of Martu and their Relevance to Karlamilyi National Park Management, Great Sandy Desert, Western Australia. PhD thesis, University of Western Australia, Perth. 46 Zann RA (1996) pp. 151–55. 47 Zann RA (1996) pp. 139–41; MacGillivray DWK (1929) Through a drought-stricken land. Emu 29, 113–29; Burbidge AA and Fuller PJ (2007) Gibson Desert birds: responses to drought and plenty. Emu 107, 126–34. 48 Roshier DA and Reid JRW (2003) On animal distributions in dynamic landscapes. Ecography 26, 539–44. 49 The chapter is dedicated to the memory of Kevin Jones, who died well before his time. Kevin was my Technical Officer for 10 happy years. His native scepticism about the antics of scientists was always overtaken by his extraordinary work ethic, and I honour him here for both those qualities. Richard Zann deepened my interest in Zebra Finches by stimulating the Alice Springs study; furthermore, his monograph informs every part of this paper. I am indebted to Margaret Friedel and the staff of CSIRO’s Alice Springs laboratory for hosting me as a Visiting Scientist, to Tara Leckey for her hospitality at Patjarr while the first draft was being prepared, to Fiona Walsh for advice on Indigenous matters, and to Faye Alexander, Libby Robin and Tom Griffiths for their encouragement.

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GREY

TEAL: SURVIVORS IN A

CHANGING WORLD David Roshier

The incongruity of waterbirds occupying one of the driest landscapes on Earth has captured the imagination of explorers, travellers and scholars since the early 19th century when the explorer Charles Sturt observed large numbers of waterfowl and pelicans flying inland.1 Despite evidence from other explorers to the contrary, Sturt concluded that the only explanation of his observations was the existence of an ‘inland sea’ and in 1844 he set out overland with a boat to find it. Sturt failed to find an inland sea, in part because his European perspective limited his understanding of the nature of the

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movements, motivations and constraints on grey teal and other waterbirds that occupy the arid interior of the continent. In more recent times, the grey teal (Anas gracilis) has become the exemplar ‘avian nomad’ and scholarly writers still draw on the scant detail we have of its movements to compare and contrast patterns of movement and migration globally.2 The grey teal is indeed extraordinary in its ability to find temporary waters wherever they occur. While crossing the Strzelecki Desert, I have seen pairs of birds on shallow pools by the road just hours after a storm momentarily dampened the usually dry desert. Away from the large aggregations of waterfowl that we occasionally observe on the more persistent desert lakes, this is often the circumstance in which we see grey teal – a handful of individuals a long way from any known permanent water. While other waterbirds use these temporary pools, more often than not it is a grey teal or two that surprises us simply by being there.

Getting about the desert To any earthbound traveller in inland Australia, the overwhelming impression is one of unending dry scrub and desert. However, pick the right date and view the landscape from above and a surprising amount of water may be evident on claypans between the dunes or in wide shallow lakes that cover many square kilometres. This surface water may have resulted from a passing storm during the infrequent summers when monsoonal lows penetrate deep into the inland, or the result

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of long, slow floods that drift in pulses down dryland rivers to fill lakes and billabongs many hundreds of kilometres away, like those that occur along the floodplain of Cooper Creek and the Diamantina River in the Lake Eyre Basin. 3 The frequent but irregular floods that redistribute water from the tropical north down the river systems of the Lake Eyre Basin into the desert regions of south-west Queensland and northern South Australia is the boom that sustains many Australian waterbird populations.4 The waters of once dry but now flooded lakes produce a pulse of productivity that at its peak seems inexhaustible, and many waterbirds take the opportunity to breed, sometimes producing more than one clutch if the favourable conditions persist long enough. For a wetland-dependent species, finding mostly temporary resources widely dispersed in time and space on a vast arid continent appears a daunting task. Moreover, arriving in time to take advantage of the peak of food abundance to maximise the likelihood that breeding will be successful seems miraculous, particularly given that the various food resources that each species prefers are not necessarily available at the same stage of the wetting and drying cycle. The first species to arrive in numbers on the desert lakes after a flood are often hardhead (Aythya australis), followed soon after by grey teal and others. With time the fish-eating pelicans, cormorants, egrets and herons arrive as fish numbers increase, as do a rich mix of ducks and shorebirds to exploit the now abundant food resources.

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The first evidence that grey teal were highly mobile across the continent and that the arid interior was no barrier to movement came in the 1950s. Harry Frith of CSIRO banded hundreds of waterfowl, including grey teal, at several sites across the continent. Relative to the numbers of birds released, recoveries were few but enough to demonstrate impressive feats of migration over thousands of kilometres; birds banded in south-eastern Australia flew as far as Western Australia and New Guinea.5 Then in the 1980s and 90s scientists became increasingly aware of large concentrations of waterbirds numbering in the tens of thousands on remote desert wetlands, prompting questions as to the origin of the birds and the frequency with which individuals ventured into the arid interior or even whether individuals remained there all or most of their life.6 Recently, satellite tracking has enabled us to follow individuals as they move about the continent, often at high speed. From September 2003, Martin Asmus and I fitted 23 grey teal with transmitters and tracked them for up to 879 days from sites in the Strzelecki Desert and the agricultural landscapes of the Riverina in southern New South Wales.7 By late 2003 drought was biting hard and waterbirds of any kind were difficult to find in significant numbers anywhere in inland Australia. In each region, we needed to find upwards of several hundred teal feeding in the one locale to have any chance of trapping the dozen birds required for the study. The Coongie system in northern South Australia still had water in its southern lakes but few birds; they’d either moved or

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perished. With a tip-off from Richard Kingsford following his annual aerial survey of waterbirds in eastern Australia, we located approximately 40 000 ducks – mostly grey teal and pink-eared duck (Malacorhynchus membranaceus) – on Lake Hope in the Strzelecki Desert in October 2003. This remote lake had filled from Cooper Creek in March 2000 to a depth of several metres, but by late 2003 little water remained in the rapidly drying lakebed. It was clear that birds would soon have to move and the fate of the nine birds released with small solar-powered transmitters provide some interesting insights into the behaviour of these desert dwelling ducks. By late October, the waters of Lake Hope were heavily fouled with bird droppings and little remained of the abundant zooplankton and invertebrates that proliferate following inundation and provide a rich food source for waterfowl and others. One tagged bird departed Lake Hope on 2 November 2003 for Andracunie Swamp, 104 km to the north across the Strzelecki Desert, returning to Lake Hope five days later having apparently not found what it was seeking. By late November, Lake Hope was virtually dry and the first escapee departed on 14 November 2003, while the remaining eight tagged birds departed over three days from 22 November 2003. Despite the almost synchronous departure of birds, their subsequent movements over the next months and years differed markedly. One hapless, perhaps inexperienced, individual departed Lake Hope for the dunefields of the Strzelecki Desert on 23 November 2003 before returning nine days later

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to the now dry Lake Hope. Finding nothing to sustain it, it departed the same day to the north-west seeking the shallow temporary wetlands that infrequently form between the sand dunes after rain. This bird died four days later having finally run out of options in the now dry Strzelecki Desert. By contrast we tracked another of the escapees from Lake Hope for another 850 days. This bird remained in the region for nearly 10 months on shallow temporary wetlands to the west before moving south on a journey that took it through pastoral lands east of the Flinders Ranges, west to the far shore of Lake Torrens, south to Port Augusta, the Clare Valley and eventually arriving at the coastal wetlands of the Coorong in the southeast of South Australia in March 2005. Still restless and seeking better opportunities to feed or breed, it departed for the low hills of the Barrier Range around Fowlers Gap in western New South Wales, 600 km to the north-east, following heavy rain in July 2005. This bird reached the area in a single flight and stayed for four weeks before returning to the Coorong, where it stayed until we lost track of it in March 2006. The fate of the other birds released at Lake Hope was equally mixed. Another quickly perished in the Strzelecki Desert to the south of Lake Blanche, while the remainder were tracked for the next year or more moving along and between the floodplains of the Diamantina River and Cooper Creek. Each appeared to follow an individual path with few sharing the same destination or fate. In December 2003 one bird was tracked to the house tank of Mulka Station on the Birdsville track. I was interested in whether this bird was travelling alone 80

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or in a flock, so I rang the station manager, Gary Overton, and asked him to report back on the number of teal on the tank. He duly responded that there were two, presumably our bird and its travelling mate. The travels of an isolated pair of teal across the desert seeking opportunities to breed contrasts starkly with the vast numbers of waterfowl observed in the migration flyways of the northern hemisphere each spring and autumn, all heading in the same direction with approximately the same plan for breeding success. In January 2004, heavy rains in northern Australia resulted in a moderate flood inundating the upper reaches of Cooper Creek. It takes weeks for the flood waters to reach the middle reaches to the east and west of Innamincka and two birds headed north to meet the flood waters on the floodplain to the south of Windorah in south-west Queensland. The timing of one of these flights in relation to rainfall and movement of floodwaters was extraordinary. Floodwaters arrived near Windorah in mid January following rains that commenced on 12 January 2004 and were at their heaviest locally in the 24 hours after 9 am on 15 January 2004 when 62 mm fell. One tagged bird departed a permanent waterhole on Cooper Creek 360 km to the south-west near Innamincka sometime after 7 pm on the same day and arrived near Windorah on the morning of 18 January 2004, within 24 hours of the arrival of flood waters. Somewhat surprisingly this teal returned to near its original location on Cooper Creek two months later about six days after the floodwaters reached that location and approximately one day after the flood peak, demonstrating an 81

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extraordinary understanding or sense of where and when water was in different parts of the landscape. Another bird flew north to meet the flood waters south of Windorah from a ground tank among the sand dunes near Orientos Homestead in the south-west corner of Queensland. Once the flood passed, this bird continued north and east before being taken by a bird of prey in woodlands 100 km west of Quilpie. We know the fate of this bird as I recovered the mangled and holed transmitter with part of the talon of the bird of prey still embedded in the side of the unit. We repaired this transmitter and fitted it to a teal at Lake Toontoowaranie in the Coongie system in February 2005. In an extraordinary flight, this bird flew 982 km north across the Simpson Desert in little more than one day to the Barkly Tablelands in the Northern Territory via the flooded Eyre Creek in south-west Queensland. Clearly the desert is no barrier to the movements of this wetland-dependent species. While the distances between isolated desert wetlands appear not to be a significant constraint on movement of grey teal, it is notable that most extended legs of long flights were undertaken at night, with the bird departing soon after sundown and stopping by the early hours of the morning. Nighttime travel is common among long-distance avian migrants and may be to reduce risk of predation or to reduce energetic costs. Ducks are energetic fliers with high wing loadings and must keep flapping to stay aloft. Like all birds, ducks have a higher body temperature than similar sized mammals.

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Evidence from related European species of waterfowl suggests that they become overheated during periods of extended exercise when the air temperature is above 15ºC and lose large amounts of water to maintain body temperature.8 Such a low ambient temperature as a limit to extended flight of grey teal seems unlikely, given its general tolerance for the desert environment. However, examination of the available weather data suggests otherwise. Weather stations with high altitude wind and temperature data are few across inland Australia but examination of the records from the nearest to a flight path show that ambient temperatures in the study area tend to drop by about 5°C per 1000 m of altitude and night-time temperatures can be 20°C or more less than day-time maxima. Matching the timing of flight to the weather records showed that ambient temperature was near or below 15°C at 1000 m, or likely to be so given the air temperature at ground level at the time. While we don’t know the altitude teal fly when moving about the continent, waterfowl have been observed at much higher altitudes than 1000 m and it remains a possibility that the physiology and tolerance of elevated ambient temperatures during flight in grey teal are similar to their northerntemperate counterparts, even though they inhabit a hot, dry continent with a vastly different climate.

What drives the movements of grey teal? Grey teal are wide-ranging and dispersive across most of the continent. They utilise a broad range of aquatic habitats in

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their travels but most favour large, often temporary, shallow inland wetlands that provide an abundance of food sources for a limited period.9 Frith’s banding studies and the recent tracking studies have shown that some individuals undertake movements of hundreds, sometimes thousands, of kilometres. As with long-distance seasonal migrants, these irregular movements are thought to be undertaken to exploit resources for breeding or survival.10 But what cues start these journeys? In contrast to northern-temperate environments, where changes in day length and temperature trigger hormonal changes in many species that act as cues for migration to breeding habitats, the cues for movement in a highly variable desert landscape are less obvious. Hormonal control of movement and migration is less likely to facilitate exploitation of temporary wetlands that are so highly variable in both time and spatial extent. Apart from the monsoon that deluges the Top End most summers, few places in inland Australia provide reliable food resources for waterbirds at a similar time each year. Moreover, the few permanent wetlands often don’t provide the abundant food resources that come with the floodwaters that pulse through our dryland rivers. It has been suggested that waterbirds in these environments find temporary wetlands by visual cues, low frequency sound and/or temperature and pressure gradients that develop around passing weather systems.11 Smell has also been suggested as a means for waterbirds to find highly productive wetlands on a dry continent.12 While visual cues are certainly important at some scale, the

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cues used to find temporary wetlands far beyond the horizon can only be speculated upon. However, tracking of individual teal gives us some clues. Most grey teal that we tracked apparently responded to distant cues such as rainfall and flooding at some stage during the study.13 The precise nature of the cue or cues used to find distant wetlands is less clear, although some flights were in response to thunderstorm activity hundreds of kilometres away. Thunderstorms generate low frequency sound, infrasound that can be detected over many hundreds of kilometres by birds and others to navigate.14 More difficult to explain are the flights over hundreds of kilometres to intercept floodwaters long after the weather signal that produced the flooding has dissipated. Intuitively we know that grey teal must be able to find distant wetlands or else they would perish on a mostly arid continent. A clue to how these birds interact with the landscape comes from long flights over hundreds of kilometres into regions that at the time had little surface water, settling on small persistent water sources in regions that rarely had significant areas of surface water. These apparent prospecting flights have been described as ‘ranging behaviour’, or a search for a place to feed or breed that should stop when the resources being sought are encountered.15 This is consistent with our general understanding of what constitutes nomadic movement behaviour. The movements of individuals across regions of diverse and numerous wetland resources to settle on distant wetlands, bypassing many wetlands encountered along the way and

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continuing in response to no apparent environment cue runs counter to the generalisation that ranging behaviour is a suitable description for all long-distance flights of grey teal. For example, one bird in the Riverina flew 286 km overnight across extensive areas of irrigated rice crops to settle on a sewage treatment works at Swan Hill for 17 days before returning to Barrenbox Swamp near Griffith. One can only assume that this individual bird had been to the treatment works at Swan Hill previously. Species that interact with temporary resources at broad scales may use periods of high resource abundance to locate particular resources for future use.16 Spatial memory is known to be highly developed in some bird species and is one of a number of orientation and navigation processes used to return to known resources.17 The longest observed flight in our studies of grey teal was that of a bird released at Fivebough Swamp near Leeton in southern New South Wales. After making its way to the agricultural districts around Moree in northern New South Wales, which had recently experienced heavy summer rain and flooding, this bird flew 600 km west into the Paroo River region that was still in the grip of the drought, and returned to Moree via Cunnamulla in southern Queensland over the next week or so. The rate of movement during the early phase of this 1268 km flight was rapid, with the bird travelling 502 km in the first 6.6 hours, with an average groundspeed of 99.9 km/h for half that period.18 This flight was also impressive as a feat of navigation as the bird returned to the

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same group of wetlands it had been previously. This bird continued its travels around the Murray-Darling Basin, covering more than 2500 km in the first 90 days, returning to Fivebough Swamp near Leeton in the Riverina before making another sojourn out to the Paroo River. In total, this bird travelled over 4800 km in the year we were able to track it. So why explore? One explanation is that knowledge of the distribution of resources at broad scales can be gained during periods of high resource availability at reduced energetic cost or risk of predation compared to periods of low resource availability. For teal, wet periods when food is plentiful enable them to improve their condition and energy reserves and undertake long flights into regions where the availability of food is less certain. As food resources decline, there is likely to be greater fitness and probability of survival for those individuals whose movements are guided by prior knowledge and move directly to known resources. This has been observed in other wetlanddependent species that range over large areas seeking shortlived prey that are most abundant when once dry wetlands are first inundated.19 In contrast to the wanderings of some, one bird released near Leeton in November 2004 flew south in December to recently flooded farmland near Gunbower on the Murray River and stayed for the next 393 days. This bird only occasionally moved away from a small cluster of wetlands alongside an irrigation canal and we began to wonder whether this bird was fully fit or whether there was some other feature that made

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this spot so attractive. In June 2005 we spent a day in the area and eventually observed the bird on a large billabong. It was as surprised to see us as we it and flew strongly back in the direction of its favoured canal and wetlands. The canal that was its preferred haunt was heavily wooded along the banks with dead timber and reed beds breaking the water’s surface. Previously it had been a creek that flowed intermittently but now delivered water to the farmer’s fields guided by gates and banks. We called into the house for a chat and were informed that unlike others thereabouts they didn’t allow any shooting on the property. Clearly one nomadic teal had found food and security and had few reasons to move!

Understanding movement and migration The movements of Australia’s waterbirds are often described as ‘erratic’ or ‘unpredictable’ and nomadic movement patterns are usually classified separately to those observed in seasonal migrants, the assumption being that waterfowl occupying a desert landscape are different in some unspecified way.20 However there is little evidence to support this notion apart from the fact that the direction and timing of movement is not consistent and does not conform to a seasonal timetable. Bird migration has fascinated humans for centuries and the ecology and physiology of migration is now well understood for many species.21 Most of what we know and understand of bird migration comes from the study of long-distance seasonal migrants in northern-temperate regions. Lengthen-

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ing spring days prompt millions of birds to move towards higher latitudes where a pulse of food production increases the likelihood of breeding success. In ornithology, this has become the prevailing model for understanding long-distance movement of birds, although we are increasingly aware that even in northern-temperate regions the model is not universal in its applicability to migrants or biomes.22 Are grey teal nomads or migrants? Some authors define migration broadly to encompass all movement between occupied sites or home ranges.23 Using this definition, five basic characteristics of migration common to all taxa have been identified: persistent movement between distant sites; directional movement; non-responsiveness to arresting stimuli when migrating; distinct behaviour for arrival and departure; and energy allocation to support movement. It is likely that all or most of these characters are true of so-called nomadic species and the primary difference between seasonal migrants and wide-ranging nomads is that the breeding and non-breeding ranges are disjunct in the former and overlapping in the latter. In ornithology ‘migration’ has had a more restricted definition that refers to the periodic (usually bi-annual, seasonally driven) movement to and from regular breeding and non-breeding grounds; it implies a strong tendency towards breeding site philopatry, that is returning to the same place each year to breed. Considering birds only, a recent comparative analysis of migratory behaviour identified the behavioural and physiological characteristics common to all migrants.24 These were

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precise timing of seasonal physiological events; an ability to find the way over large distances; endurance performance (extended fasting and intense exercise); ability to prosper in contrasting environments (different food, competitors, predators and parasites); increased predation risk and potential for overheating during flight; tracking or predicting of food resources; seasonal time pressures, and some degree of variation or unpredictability of resource distribution. None of these are exclusive to long-distance seasonal migrants, and as we track teal and others it is becoming increasing clear that many of the above characteristics and constraints also apply to nomadic species. In particular, waterfowl utilising temporary wetland habitats in arid environments for breeding are time constrained as evaporation and prey dynamics combine to limit the time for which prey items are available to feed young. Evidence of fat deposition in a nomadic species comes from the white-faced whistling-duck of southern Africa.25 Storage of fat prior to arrival at breeding sites reduces ‘reproductive uncertainty’ associated with using time-limited resources. Females that arrive with body stores have a headstart in the race to accumulate sufficient energy reserves to initiate a clutch of eggs. This is a strategy used by many long-distance seasonal migrants that breed in the Arctic, another environment where food to feed young is only available for a few weeks. So, is there something extraordinary about our desert ducks, and the grey teal in particular, that enables them to survive the rigours of the Australian desert and distinguishes them

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from their northern-temperate counterparts? I don’t think so, but some might disagree. Members of the Anas genus are remarkably similar in their ecology and breeding biology, even though realised patterns of movement contrast markedly.26 Many aspects of migration are thought to be under endogenous control, so it has been suggested that most facets of migratory behaviour have a direct genetic basis and have evolved and reevolved many times in birds.27 However, there appears to be no consistent expression of migratory traits within a population, species or any other phylogenetic grouping, which would be expected if migratory traits had a direct genetic basis.28 Equally likely is the possibility that some phylogenetic groupings, such as dabbling ducks, are not significantly constrained by changing patterns of resource distribution – even at the extremes of variability experienced in Australia. Thus, patterns of movement in Anas populations (including both seasonal migration and nomadism) may be part of a broader behavioural strategy that has a genetic basis but whose expression is dependent on both the realised pattern of resource distribution through time and the energetic constraints of moving between breeding, non-breeding or migratory staging sites.29 If so, our understanding of movement and migration is too simplistic as we have tended to concentrate on the outcome rather than the means by which birds find the resources they need to survive and reproduce. In a changing world, understanding those means will be critical to understanding which species are vulnerable to global climate change and those that are likely to continue to

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prosper. In this context, Australia has an important place in the development of a broader understanding of patterns of movement and migration as all strategies are evident in a fauna that constantly deals with the extremes of crushing drought and infrequent flooding rains. Some depart the continent each autumn for more productive or less extreme environments, while others such as the grey teal simply deal with the local extremes of boom and bust.30

Endnotes 1

Stokes E (1986) To the Inland Sea. Charles Sturt’s Expedition 1844–45. Hutchison: Hawthorn, p. 17.

2

Dingle H (2006) Animal migration: is there a common migratory syndrome? Journal of Ornithology 147, 212–20.

3

Roshier D, Whetton P, Allan R and Robertson A (2001) Distribution and persistence of temporary wetland habitats in arid Australia in relation to climate. Austral Ecology 26, 371–84.

4

Kingsford R, Curtin A and Porter J (1999) Water flows on Cooper Creek in arid Australia determine ‘boom’ and ‘bust’ periods for waterbirds. Biological Conservation 88, 231–48.

5

Frith H (1957) Breeding and movements of wild ducks in inland New South Wales. CSIRO Wildlife Research 2, 19–31; and Frith H (1959) The ecology of wild ducks in inland New South Wales. II. Movements. CSIRO Wildlife Research 4, 108–130; and Frith H (1962) Movements of the grey teal, Anas gibberifrons Muller (Anatidae). CSIRO Wildlife Research 7, 50–70; Frith H (2002) Waterfowl in Australia (2nd edition). Angus & Robertson: Sydney.

6

Kingsford R (1995) Occurrence of high concentrations of waterbirds in arid Australia. Journal of Arid Environments 29, 421–25; Roshier D, Robertson A and Kingsford R (2002) Responses of waterbirds to flooding in an arid region of Australia and implications for conservation. Biological Conservation 106, 399–411.

7

Roshier D, Klomp N and Asmus M (2006) Movements of a nomadic waterfowl, grey teal Anas gracilis, across inland Australia – results from satellite telemetry spanning 15 months. Ardea 94, 461–75; Roshier D, 92

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Asmus M and Klaassen M (2008) What drives long-distance movements in nomadic grey teal Anas gracilis in Australia? Ibis 150(3), 474–84. 8

Engel S et al. (2006) Exhaled air temperature as a function of ambient temperature in flying and resting ducks. Journal of Comparative Physiology B – Biochemical, Systems and Environmental Physiology 176, 527–34.

9

Marchant S and Higgins P (1990) (Eds.) Handbook of Australian, New Zealand and Antarctic Birds. Volume 1 Ratites to Ducks. Oxford University Press: Melbourne, pp. 1266–81.

10 Kingsford R and Norman I (2002) Australian waterbirds: products of the continent’s ecology. Emu 102, 1–23. 11 Simmons RE, Barnard P and Jamieson IG (1998) What precipitates influxes of wetland birds to ephemeral pans in arid landscapes? Observations from Namibia. Ostrich 70, 145–48. 12 See Roshier D et al. (2006). Also see Wallraff HG (2004) Avian olfactory navigation: its empirical foundation and conceptual state. Animal Behaviour 67, 189–204. 13 See Roshier D et al. (2008). 14 Hagstrum JT (2000) Infrasound and the avian navigational map. Journal of Experimental Biology 203, 1103–11. 15 Dingle H (1996) Migration: The Biology of Life on the Move. Oxford University Press: New York. 16 Bennetts RE and Kitchens WM (2000) Factors influencing movement probabilities of a nomadic food specialist: proximate foraging benefits or ultimate gains from exploration? Oikos 91, 459–67. 17 The ability of birds to navigate using spatial references was most unusually demonstrated in pigeons that used the motorways of Europe to navigate home. Lipp HP et al. (2004) Pigeon homing along highways and exits. Current Biology 14, 1239–49. 18 See Roshier et al. (2008). 19 See Bennetts RE and Kitchens WM (2000). 20 Ford H (1989) Ecology of Birds. Surrey Beatty & Sons: Sydney; Griffioen PA and Clarke MF (2002) Large-scale bird-movement patterns evident in eastern Australian atlas data. Emu 102, 99–125. 21 Alerstam T (1990) Bird Migration. Cambridge University Press: Cambridge; Berthold P, Gwinner E and Sonnenschein E (2003) Avian Migration. Springer: Berlin; Greenberg R and Marra P (2005) Birds of Two Worlds: The Ecology and Evolution of Migration. John Hopkins: Washington DC. 93

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22 Newton I (2006) Advances in the study of irruptive migration. Ardea 94, 433–60. 23 Dingle H (2006); Dingle H and Drake VA (2007) What is migration? Bioscience 57, 113–21. 24 Piersma T et al. (2005) Is there a ‘migratory syndrome’ common to all migrant birds? Annals of New York Academy of Science 1046, 1–12. 25 Petrie S and Rogers K (2004) Nutrient-reserve dynamics of semiaridbreeding white-faced whistling ducks: a north-temperate contrast. Canadian Journal of Zoology 82, 1082–90. 26 Briggs S (1992) Movement patterns and breeding characteristics of arid zone ducks. Corella 16, 15–22. 27 Alerstam T, Hedenström A and Åkesson S (2003) Long-distance migration: evolution and determinants. Oikos 103, 247–60; Pulido F (2007) The genetics and evolution of avian migration. Bioscience 57, 165–74. 28 Coppack T and Both C (2002) Predicting life-cycle adaptation of migratory birds to global climate change. Ardea 90, 369–78; Piersma T et al. (2005); Pulido F and Widmer M (2005) Are long-distance migrants constrained in their evolutionary response to environmental change? Causes of variation in the timing of autumn migration in a blackcap (S-Atricapilla) and two garden warbler (Sylvia borin) populations. Annals of the New York Academy of Sciences 1046, 228–41. 29 Roshier D, Doerr V and Doerr E (2008) Animal movement in dynamic landscapes: interaction between behavioural strategies and resource distributions. Oecologia 156, 465–77. 30 Acknowledgements: None of this work would have been possible without the enthusiasm and good sense of Martin Asmus. Martin and I shared many weeks by remote desert wetlands or gathered around a computer to view the latest data from the satellites and I thank him for his many efforts that contributed to the ideas expressed in this essay. I would also like to thank Mike Schultz, Mark Wilson, Julian Reid, Richard Kingsford, Ross McDonnell, Marcel Klaassen, Craig Poynter, Erik Doerr and Veronica Doerr who contributed to the body of work behind this essay by sharing their knowledge and skills. Finally, thanks to the property owners and residents of rural and remote Australia that responded positively to our requests for access and shared their knowledge.

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PELICAN : FLEXIBLE

RESPONSES TO UNCERTAINTY Julian Reid

While studying the ecology of the Coongie Lakes in far northeast South Australia in the late 1980s and early 1990s, I would sometimes lie back in awe and watch the spectacle of one or two thousand pelicans take off from one lake, flapping initially and increasingly soaring upward in spirals, until at a great height one bird would take the lead, break out of the thermal that was carrying it aloft and head for a distant lake, with successive birds peeling off one by one to follow its lead, while the take-off from the lakes’ surface by small skeins continued. It could take half an hour for a large flock to take flight in this

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staged manner, climb the thermals and follow the lines of departing birds to their new feeding grounds. This was a middle-of-the-day phenomenon, often late morning, when the thermals had begun to develop. Another memory I have is of hot, humid, uncomfortably still nights, camped with my colleagues on the shore of a lake, struggling to sleep, listening to the night sounds of crickets and birds, when a sudden clapping roar of sound would rush across the lake’s surface, as a mob of pelicans flew off from a distant roost or nesting island to feed. Similarly I recall the clear, sharp ‘shshshshsh’ breaking the stillness as a group of pelicans alighted unseen from the night sky, skimming onto the water’s surface next to our camp. Being so large, on the rare occasions when pelicans congregate inland in huge flocks, they must consume staggering amounts of fish, and generally they cannot avoid making their presence known to humans in their midst. The Australian pelican (Pelecanus conspicillatus) has mythological significance for Aboriginal people. It was harvested for food by the Yandruwandha people in the Coongie Lakes, has long been persecuted and had nesting colonies destroyed by European fisher folk as perceived competitors for their fish stocks. It learns quickly to capitalise on human and other novel sources of food (friendly anglers, municipal garbage dumps), and is a common sight around wharves and harbours in coastal Australian towns, particularly fishing centres.1 Amongst the heaviest flying birds in the world, pelicans as a group evolved from marine ancestors, but six of the seven

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species including the Australian pelican spend most of their time on rivers, lakes, inland seas and coastal waters.2 Male Australian pelicans can weigh in excess of 10 kg, but 8 kg is a more usual upper weight.3 Renowned for and recognised instantly by their massive bill and pouch, pelicans mainly eat fish, and they nest colonially. They soar spectacularly on thermals, occasionally to great heights (3000 m or more), and they display precise orchestration of movement when feeding cooperatively or flying in V formation. The population of pelicans in Australia is thought to number 300 000–500 000.4 There is a sharp contrast in many respects between the sight of a few pelicans scattered along the shore of coastal towns and that of tens of thousands of birds massed at breeding colonies on inland Australian floodwaters. As well as the obvious visual contrast, the dispersed small groups of coastal birds are an everyday and ‘everywhere’ occurrence, whereas the massive inland breeding aggregations are infrequent and episodic, in both time and space.5 Viewed from one perspective, that of the location of a massive inland colony, 30 000–50 000 pairs in the four recorded examples, namely Lake Cawndilla (Menindee Lakes), Lake Eyre South, Lake Goolangirie (Coongie Lakes), and Lake Machattie (Georgina River), this is classic boom-bust.6 One cannot imagine a more arid, harsh and unproductive ecosystem than the salt crust surface of Lake Eyre South in most years, yet in the first six months of 1990 following its flooding from local rains a year earlier, 100 000 pelicans successfully fledged as many as 90 000

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chicks.7 However, the same species forms small breeding colonies of tens to hundreds, rarely a few thousand, on an annual basis on many small islands in coastal and sub-coastal Australia. This is not the hallmark of a boom-bust species. It displays characteristics of both boom-bust and ‘regular-annual’ reproductive strategies.8 Behavioural flexibility and adaptability to new environments and resources (reservoirs, other unnatural impoundments, artificial islands, human garbage, exotic fish as prey) would seem to be defining features of the Australian pelican, as much as its anatomical features and colonial breeding that are prescribed by its membership of the pelican family.9 In this chapter I explore the theme of ‘plasticity’ as evidenced by the Australian pelican’s behaviour and responses to resource variability and environmental novelty. The interactions between humans and pelicans provide a springboard for the exploration of this theme. I then focus on the episodic aggregations of tens of thousands of birds in huge colonies, as these events exemplify the boom-bust nature of Australian dryland rivers and lakes, and because they introduce one of the great unsolved natural history mysteries: how do pelicans know when to migrate into the interior, presumably largely from all parts of the continent’s periphery, and how do they know precisely where to congregate? By contrast, the trigger for waterbirds deserting the interior subsequently is readily apparent: drying of wetlands and drought force birds to evacuate if they are to survive.

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Pelican biology and resource availability in inland Australia For such an obvious regularly seen species, there are surprising gaps in scientific knowledge of the Australian pelican’s basic biology. A captive bird was reported to be 60 years old, yet age structures of natural populations are not known, nor are typical longevities in the wild.10 The minimum age of first breeding was assumed to be three to four years based on circumstantial evidence and from information on related species, although recent research has found three-year-olds breeding.11 Nor is the maximum age of breeding known, but the species appears to be long-lived. Breeding pairs are thought to be monogamous but only for the duration of a single breeding season; pairs reform at the onset of each breeding event, with several males vying for the attention of an unpaired female.12 Both sexes acquire prominent courtship colours for a limited period and the colours fade quickly once brooding begins. Large colonies grow sequentially over an extended breeding season, and dates of first and last eggs laid may span several months.13 The usual clutch size is two and both parents brood the eggs and feed chicks. The incubation period is 32–35 days, and if two eggs hatch the second hatches two to three days after the first.14 This hatching asynchrony is correlated with substantial rates of siblicide where the older and larger chick kills its weaker sibling.15 Chicks leave the nest after 10 days and join creches; they fledge at about 80–90 days, and are thereafter independent.

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Since prospecting birds typically spend a month or more settling on and inspecting the eventual colony site, a five month period would appear to be the absolute minimum for successful breeding.16 Colony sites needs to be secure from predators for a lengthy period, and for inland breeding locations this may translate to a minimum period of four months of inundation around the colony. This may partly account for the infrequent intervals at which large pelican colonies form in inland Australia where in most years rainfall is insufficient to generate sufficient floodwaters (extent and persistence) for a secure breeding event. Prey abundance and availability may well be a more potent limiting factor. Compared with other deserts globally, rainfall in arid and semi-arid Australia is highly variable in its amount from year to year and to a degree unpredictable in its season.17 Skewed by the very large events in occasional years, mean annual rainfall is relatively high for deserts generally. Consequently, the presence and persistence of wetlands are greater than first imagined. David Roshier and colleagues modelled wetland availability across arid Australia over a 108 year period, and concluded there was only one 12 month period without some wetland persistence.18 However, only certain wetlands provide suitable habitat for fish-eating predators. Much of inland Australia, particularly the central-Western Desert region, is unsuitable for pelicans, as fish populations in these areas of uncoordinated drainage are, if present at all, sparse and usually confined to small species and a few

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localised wetlands. Furthermore, and as anticipated earlier in this chapter only some flooding events will be of a magnitude to produce sufficient fish biomass to sustain a large breeding event by pelicans.19 Spatio-temporal variability of rainfall is amplified by coordinated catchment and runoff processes that define the major river systems of inland Australia, such that across a 30 year period from 1973 to 2003, the annual discharge of Cooper Creek at Innamincka in north-east South Australia varied across almost three orders of magnitude (50–15 000 GL), an order of magnitude greater than the annual rainfall variations (35–950 mm) in the same period.20 There are other climatic, hydrological and biological processes that conspire to amplify or reinforce the significance of the largest flood events. Jim Puckridge and colleagues examined the Southern Oscillation Index, rainfall data, Cooper Creek flow volumes at Currareeva, Queensland, and fish responses to environmental fluctuations in the Coongie Lakes.21 They drew the following conclusions: large flood events are clustered such that usually one large flood is followed or preceded by another large event within 12 months; this partly arises from the clustering of La Niña (high rainfall) episodes; flood clustering is also entrained by catchment processes such that a second, smaller flow volume will flood a disproportionately larger area of floodplain and downstream lakes (‘flood memory’); waters in the Coongie Lakes became successively warmer and more transparent, hence more productive, over a cluster of three large floods in

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1989–1991; recruitment and biomass of five native fish species, including the bony herring or ‘bony bream’ (Nematalosa erebi), a major prey item of the Australian pelican, boomed in the Coongie Lakes over the three-year flood event, such that fish abundance increased at a greater rate than lake volumes. A huge pelican colony (30 000–50 000 pairs) formed on an island in one of these lakes towards the end of this flood cluster.22 A big colonial fish predator like the Australian pelican is dependent on extensive river, floodplain and lake systems with the capacity to produce massive fish populations (booms), such as Cooper Creek and other rivers in the Lake Eyre Basin. Other river systems that may well sustain huge pelican colonies periodically are the north-western rivers in the Murray-Darling Basin (Darling, Paroo), the Bulloo, the Barkly Lakes system (Northern Territory), and the lower Sturt Creek (Lake Gregory, Western Australia).

Interactions with humans An Aboriginal informant from the Innamincka region recounted how his grandparents’ generation of Yandruwandha people understood the breeding biology of pelicans in the Coongie Lakes: This is how they used to catch pelicans on the lakes. When the young ones are getting big, they build a big yard on the bank [shore] of the lake. Then they go into the water and they herd the young birds back, the whole lot of

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them. If a little one gets tired it turns around and heads back to the island while they hunt [head, herd] the rest of them, the biggest ones, on and put them in the yard. Then for two or maybe three weeks they camp there, living on the birds, while the birds feed on fish that their parents bring them and give them in the yard. They don’t have to feed them, the parents do that. While they are eating some the rest are getting fat. The biggest ones, that are nearly ready to fly, they kill and cook and eat. When they finish them off they go down again and muster some more.23

These Aboriginal people obviously knew about the creching behaviour of pelicans, that the parent birds would find the location of the ‘new creche’ and continue to feed their chicks, and also that breeding in a pelican colony is often staggered with new birds commencing to breed after the first birds’ eggs have hatched, such that there could be repeat human visits to the nesting island to gather new birds for rearing and fattening. The Coorong is a long narrow estuarine wetland at the mouth of the ailing Murray River along the south-east coast of South Australia. It is one of the six icon sites recognised by the Living Murray strategy, and boasted ‘vast hordes’ of pelicans in the 19th century.24 The connection between the Australian pelican and the Coorong achieved moderate fame worldwide through the popular children’s book, Storm Boy, and the successful film of the same title released in 1976, in

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which real pelicans were used to represent the role of ‘Mr Percival’, an orphaned pelican chick raised after its parents were killed by sportsmen.25 David Paton, an ecologist with the University of Adelaide, began ecosystem studies there nearly 30 years ago. The aquatic birdlife of the Coorong has changed considerably over this period. It has generally diminished, and the failure of pelicans to breed for three years in succession at a site on which they were thought to have bred on an annual basis since European settlement sparked much consternation and media activity in late 2005.26 The Coorong fishing community had started to destroy the pelican colony by at least 1870, and its demise appears almost complete through the combined actions of drought, the cumulative impacts of upstream river regulation and over-extraction of the Murray’s water, the irony being that these same forces and social currents had led to the collapse of the Coorong commercial fishery.27 However, both the Coorong and its pelicans are more resilient than first appeared. As the prolonged drought in the new millennium continued to bite, the spring of 2006 witnessed a large colony of pelican back at their usual haunts, islets near Policemans Point, with 1500 chicks counted.28 Paton explained this unpredicted event as a drought response, that pelicans had probably gathered at the Coorong as other sites around the country dried up, and warned that without urgent resumption of River Murray flows into the Coorong the ecosystem might well go past its tipping point beyond which recovery could be difficult.

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An example of the adaptability of the pelican to anthropogenic-driven environmental change is the establishment in the past four decades of a breeding colony on an artificial island at Outer Harbor, Adelaide’s main shipping port. Greg Johnston of the Adelaide Zoo and Flinders University began studying this colony at Pelican Point in 1990 and reports that they have attempted to breed annually since, although failing completely in 2007.29 The island was only built in 1978 and pelicans began to nest there by 1985. A feature of this population was that some of the nesting birds regularly foraged at the nearby Wingfield garbage facility. That the pelicans continued to nest successfully after the closure of the Wingfield facility led Johnston to conclude that the importance to the colony of food scavenged from the dump was exaggerated.30 Similarly, pelicans commenced to breed on Penguin Island and other small islets offshore at Perth in Western Australia in 1998, and have bred there annually since.31 Clearly the Australian pelican is an opportunist in its choice of nesting sites and a highly generalised ‘predator’. As well as the legendary (apocryphal?) stories about eating small yapping dogs, waste food, birds and exotic fish are definitely on their menu.32 Notwithstanding the flexible and opportunist feeding and nesting behaviours exhibited by the Australian pelican, there is concern for its continued survival. Johnston was quoted as saying breeding sites in Victoria have been reduced from 10 to two in recent years, in response to drought and increasing disturbances at colonies.33 Most south-eastern

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Australian populations of waterbirds are in serious decline due to similar pressures and due also to excessive water extraction from the Murray-Darling Basin. 34

The energetics of a huge breeding colony While the energetics of nesting Australian pelicans have not been studied, equations describing the field metabolic rate of a suite of marine fish-eating birds of warm waters, including relatives of pelicans, have been derived.35 Applying this equation to an average 5 kg pelican yields an approximate daily energy requirement of 2.2 megajoules (MJ). If we consider the best documented of the massive pelican colonies in inland Australia, that of 50 000 pairs raising 90 000 chicks on Lake Eyre South in the first six months of 1990, and accept five months as the minimum period in which pelicans can settle, court, nest and raise chicks to independence, then it is a simple matter to multiply the numbers: 100 000 individuals × 150 days × 2.2 MJ = 33 million MJ of energy required by the nesting adults.36 Using the same conversion factor for tonnage of fish to energy requirements – approximately 200 tonnes of fish per million megajoules of seabird energy expenditure we derive a conservative estimate of 6600 tonnes of fish consumed. This figure equates to a daily consumption of 0.4–0.5 kg of fish for each nesting adult.37 How does an estimate of 6600 tonnes compare with measured abundances of fish in terminal Lake Eyre Basin wetlands? Lake Hope is a large freshwater lake on the lower Cooper, about a tenth the size of Lake Eyre South, and is within flying 10 6

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range (200 km) of pelicans nesting on Lake Eyre. A commercial fishery operated at Lake Hope over two years between 1992 and 1994, and harvested 309 tonnes of adult golden perch or callop (Macquaria species), and this off-take was estimated to be 4% of the total callop biomass in Lake Hope, approximately 7700 tonnes. 38 Callop is the largest fish in the Lake Eyre Basin, growing to a maximum size of about 3 kg. Two other species of fish, Lake Eyre hardyhead (Craterocephalus eyresii) and bony herring, dominate numerically the fish communities in Lake Eyre when it fills. It was estimated that 40 million carcasses of these two species lined the drying shores of Lake Eyre in 1975 during a major fish kill caused by rising salinities.39 Without knowing the size class distribution of these fish or their relative proportions, one can only speculate at the biomass this fish kill represented, presumably at a stage of flood recession long after peak production, but a moderately large bony herring weighs 200 g, and so it is feasible that Lake Eyre is capable of the production of many thousands of tonnes of fish during a major flood. In fact, as salinities rose above the lethal limits for fish in Lake Eyre South in mid-1990 in the final stages of the huge breeding event, adult pelicans were flying to Lake Hope to feed and catch fish for their young.

The great unsolved puzzle It is well documented that pelicans become scarcer in their usual coastal haunts when the big floods occur in central Australia, and that after the floodwaters recede birds then 10 7

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disperse widely.40 How could 100 000 pelicans coordinate their movements, presumably from all parts of the continent, and know both when and where it is the time and place to gather? The regular annual migration of numerous bird species around the world seems quite explicable by comparison, drawing on combinations of instinct, predictable seasonal cues such as daylength variation, and reinforcement borne of repeated traverses along the same route, for older birds at least. The mystery of the mechanisms underpinning these great inland movements for the purpose of breeding in the wake of major floods is not confined to pelicans but pertains to a wide range of nomadic Australian birds that respond in a broadly similar manner.41 There are important distinctions, however, between pelicans and many other waterbirds, in that fish-filled floodwaters are restricted in their distribution spatially (compared to the more generally distributed production of aquatic vegetation, plankton and invertebrates) and the production of sufficiently high densities of fish in the middle and lower reaches of large inland rivers can occur several months after the rains given the long hydrological lags in these systems. Ducks and many other waterbirds (for example, the rail, blacktailed native-hen) are renowned for arriving at locations in arid Australia immediately after heavy drought-breaking rains.42 By contrast, pelicans may need to delay their movement until fish have had time to reproduce and expand their populations.43 Another issue concerns the mode of flight in different waterbirds. Whereas ducks, rails, grebes and shorebirds use

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direct flapping (self-powered) flight to undertake large-scale movements, pelicans soar in the main with the occasional flap, and so may need suitable atmospheric conditions for long-distance flights.

The scout model John Read suggested that coastal groups of pelican send out scouts periodically to assess conditions inland and, if favourable conditions are found, that the scouts return and communicate the information to their peers, and then a larger group sets off to colonise these ephemeral feeding grounds.44 This process would be repeated across many such groups around the country until a large number of birds amass in the inland flooded region. Once birds are in the region it is easy to understand that they could explore the extent of the floodwaters, find the best feeding grounds, assess whether and when conditions (including their own body condition) are appropriate for breeding, and then choose a suitable site or sites for colony formation. Read contended that the soaring flight of pelicans expends little energy, thus making it feasible for scouts to fly the great distances required – a three-way trip – to source floodwaters, back to collect their group members, and then return. I have doubts about two elements of this theory. First, that the scouts would display such altruism (there does not appear to be any fitness advantage, that is, increased survival and reproductive success relative to other group members), and second, whether directed soaring flight

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is that metabolically inexpensive or even feasible, since the scouts would have to be able to move the vast distances in opposite directions within a short period of time. There is an even simpler model than that proposed by Read, a null model which needs to be disproved before more elaborate models should be considered. This simple model has similar assumptions to Read’s about the ease of transcontinental flight, that pelicans can indeed travel vast distances virtually at their whim and in any direction. Under this ‘periodic random movement’ model the majority of pelicans in all parts of the country periodically and randomly move varying distances, searching for better feeding (and other survival) opportunities than they currently face, and that they settle where they find better conditions (or return to known locations where they predict that they can at least survive). Under this model, provided the movements are sufficiently frequent and large, it could be expected that a substantial proportion of the pelican population would gradually gravitate to the areas of highest productivity, such as inland floodwaters. There is no evidence, however, that the majority of pelicans regularly and randomly move around Australia in this way. On the contrary, they seem to stay out of arid Australia during the customary dry periods and extended droughts. Besides, I suspect that transcontinental flight is actually quite energetically expensive, involves considerable risks of mortality, and is probably not possible at all times of the year due to prevailing wind directions and other atmospheric conditions such as thermal

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formation. Rather, I believe that the migrations of waterbirds into the inland after major rains and floods are purposeful, directional and occur in response to detected environmental cues. I have no idea what those cues are. There is no evidence that pelicans have a highly developed sense of smell or advanced levels of communication.45 It is difficult to envisage that visual cues could be used over distances greater than the order of 50–100 km if we accept 3000 m as the maximum altitude a soaring pelican can attain.

Communication needed for flocks of birds to coordinate a group departure The fact that only a proportion of the Australian pelican population undertakes the great movements inland from coastal regions in any particular event raises several intriguing issues, such as whether there could be a genetic component to the behaviour (as is commonly assumed for migratory birds).46 It also begs the question whether some form of communication is needed when different birds in a discrete coastal population choose to move or stay, given that they are all exposed to the environmental cues that trigger the migration by some. My speculation on this issue is informed by the first example I described about flock behaviour in the Coongie Lakes: a single bird out of a flock of 1000–2000 is the first to take flight from one lake, and (virtually) all others follow the lead bird’s example. I suspect that such ‘follow the leader’ behaviour is ingrained in flock-living birds generally, and that relatively few

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subtle signals serve to distinguish a short ‘comfort’ flight, say, from other take-off and flight manoeuvres that might signify a longer distance intention. Also, just as the lead position in V formation flight is often rotated to give all birds the slipstreaming benefits, I suspect there are many leaders in pelican groups and that these leaders will generally be the older, experienced birds, so it is in the interests of the younger individuals to observe the leaders and follow their example. Accordingly I conclude that elaborate forms of communication are not required, provided my basic model of flock behaviour is plausible. There appear to be no great disadvantages associated with leadership, unlike the altruistic costs that accrue to scout behaviour – so what if 10, 50 or 2000 other pelicans follow you to an adjacent lake 20 km away or to the Lake Eyre Basin wetlands some 1000 km distant? The leaders and the followers will share similar benefits that accrue from cooperative feeding and other flock behaviour at the new location. I will make one more assertion for which I have no evidence, but nor is it essential to the model outlined above. I suspect that it is again older, experienced birds that can detect and interpret the ‘far-off flood cues’ more expertly than young birds, and that having already experienced at least one great inland flood event, they are better able to navigate to the same or similar locations (see Chapter 5). They probably store the equivalent of maps in their brain; knowledge which is compiled and added to every time they follow new flight paths or explore new wetland regions once arrived. Forged in the

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evolutionary crucible where the next error may be fatal, we could expect all highly mobile and visually oriented vertebrates to have evolved sophisticated, ‘map-like’ navigation aids and processing skills. The great mystery that remains then is the identity of the physical, detectable cues, associated with torrential rainfall events or ensuing great floods, which at least some birds can sense at distances of many hundred to several thousand kilometres.

Conclusions Pelicans are long-lived birds. They cannot breed until they are in their fourth year. La Niña driven clusters of large floods in Cooper Creek persist for two to three years at most, and have a return time of four to six years.47 Hence we can rule out the possibility that newly produced generations of pelicans contribute, in situ, to the occasional massive colonies that form during the major inland flood events; there is insufficient time for the progeny to mature to breeding age before the floods have receded. It is probable that most birds, progeny and adults alike, have to vacate these arid and semi-arid river systems before the next major flood event returns, although because of the spatio-temporal variability in rainfall across inland Australia there will be some opportunities for waterbirds to move from one internal drainage basin to another.48 Fish-eating species like the pelican will have fewer such opportunities than other waterbirds, and so they tend to redistribute

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themselves around the periphery of the continent in the periods between major inland flood events. Pelicans show a high degree of flexibility in their responses to environmental uncertainty and the provisioning of novel resources; they certainly do not adopt an ‘all or none’ breeding strategy. Just as some birds remain in their coastal breeding colonies when the occasional massive colony forms in the interior, not all pelicans within the flooded interior nest in huge colonies. Roger Jaensch and I found several small colonies of pelicans scattered across the Channel Country during the 2000–2001 flood event when huge colonies formed in successive years on Lake Machattie.49 Also, small numbers breed in the interior on a semi-regular basis, capitalising on smaller flood and localised, heavy runoff events. Accordingly while the rain and flood-filled Lake Eyre Basin wetlands can certainly be described as boom-bust ecosystems, the pelican does not fit the bill of a boom-bust species; its reproductive rate is simply too low. Rather, it is an opportunistic user of boom-bust ecosystems on a ‘fly in, fly out’ basis. I have demonstrated the extraordinary productivity of Lake Eyre Basin wetlands in the boom periods, on the basis of the tonnages of fish required to sustain the enormous pelican colonies described here. These booms can cause biological reverberations throughout the continent (and beyond), in the following years and decades as dispersing birds forced to leave the drying interior colonise new areas. 50 Indeed the remarkably wet period of 1974–1976 seems to have led directly to a per-

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manent change in status of Australian pelicans in New South Wales coastal districts, the species having only become common and generally distributed there since the mid 1970s.51 I presented a simple, untested conceptual model to describe how 100 000+ pelicans from scattered parts of the continent’s periphery might coordinate a mass (if fragmented) migration into the interior after major flooding rains. Key elements of the model include: 1) the acquisition of knowledge, skills and experience through a long life; 2) these old experienced birds detect rain or flood cues from afar and do so expertly though probably imprecisely; and 3) simple if subtle behaviours may suffice to alert groups of birds to follow when lead birds in the flock signal their intention to take long-distance flights, and that these leaders will be the older birds in the flock that have travelled between coast and interior on several occasions. Much intriguing biology is yet to be learnt about this most familiar but little-studied bird, and the mystery of the mechanisms, by which the Australian pelican senses and navigates to far-off wetland filling events in the vast interior, remains. 52

Endnotes 1

Kerwin B and Breen G (1986) The way it was [Yandruwandha story (Innamincka dialect)]. In This is What Happened: Historical Narratives by Aborigines. (Eds. LA Hercus and P Sutton) pp. 37–38. Australian Institute of Aboriginal Studies: Canberra; Deborah Rose, personal communication, 12 November 2007. On fish stocks see Chapman FRH (1963) The pelican in South Australia with special reference to the Coorong Islands. South Australian Ornithologist 24, 6–14; Eckert J (1965) Pelicans breeding near Milang. South Australian Ornithologist 24, 36–37.

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2

Marchant S and Higgins PJ (1990) (Eds.) Handbook of Australian, New Zealand and Antarctic Birds, Volume 1B, Australian Pelicans to Ducks. Oxford University Press: Melbourne, p. 739.

3

Marchant S and Higgins PJ (1990) p. 746.

4

Jaensch RP (2003) ‘Wetlands to be included in a National Audit of Waterbird Populations as Determined from the 2003 Australian Population Estimates’. Unpublished report to the Commonwealth Department of the Environment and Heritage: Canberra, p. 11, see Table 1.

5

Remarkably, this may not have even been the case generally, as at least before the mid 1970s pelicans were thought to be uncommon in most coastal parts of New South Wales; in the mid to late 1970s dispersing birds – presumed to have originated from major breeding events during the great filling of Lake Eyre (1974–1976) and other inland lakes – first appeared in numbers, settled, and have remained ever since, e.g. Marchant S and Higgins PJ (1990) p. 741; and McAllan IAW, personal communication, 19 April 2008.

6

Lake Cawndilla, Menindee Lakes: Macgillivray W (1922) The nesting of the Australian pelican Pelecanus conspicillatus. Emu 22, 162–74; Chapman FRH (1963) p. 9; Lake Eyre South: Waterman MH and Read JL (1992) Breeding success of the Australian pelican Pelecanus conspicillatus on Lake Eyre South in 1990. Corella 16, 123–26; Read JL (2003) Red Sand, Green Heart: Ecological Adventures in the Outback. Lothian: Port Melbourne, pp. 64–76; Lake Goolangirie, Coongie Lakes: Reid J (1992) ‘Terrestrial monitoring of Coongie after flood, an assessment of the effects of flooding on the terrestrial biota in the Coongie Lakes District’. Final Report to the Reserves Advisory Committee of the National Parks and Wildlife Service of South Australia: Adelaide, p. 28; Lake Machattie, Georgina River: Costelloe JF, Hudson PJ, Pritchard JC, Puckridge JT and Reid JRW (2004) ‘ARIDFLO scientific report: environmental flow requirements of arid zone rivers with particular reference to the Lake Eyre Drainage Basin’. School of Earth and Environmental Sciences, University of Adelaide: Adelaide, p. 492. See also Kingsford RT, Curtin AL and Porter J (1999) Water flows on Cooper Creek in arid Australia determine ‘boom’ and ‘bust’ periods for waterbirds. Biological Conservation 88, 231–48; Kingsford RT and Norman FI (2002) Australian waterbirds – products of the continent’s ecology. Emu 102, 231–48.

7

Waterman MH and Read JL (1992) p. 126.

8

Marchant S and Higgins PJ (1990) pp. 740–41.

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9

Vestjens WJM (1977) Breeding behaviour and ecology of the Australian pelican Pelecanus conspicillatus, in New South Wales. Australian Wildlife Research 4, 37–58; Marchant S and Higgins PJ (1990) p. 742.

10 Nelson B (2005) Pelecaniformes: Pelicans, Cormorants and their Relatives. Oxford University Press: Oxford, p. 270. 11 Vestjens WJM (1977) p. 39; Harrison P (1985) Seabirds, An Identification Guide, 2nd edition. Croom Helm: London, p. 283; Marchant S and Higgins PJ (1990) p. 738; Johnston G, unpublished data, personal communication, 26 March 2008. 12 Vestjens WJM (1977) p. 45. 13 Waterman MH and Read JL (1992); Reid J (1992) pp. 27–28. 14 Vestjens WJM (1977) p. 37. 15 Nelson B (2005). 16 This requirement of a one month period for ‘settling’ on the colony site may not apply to desert breeding birds, where the time constraints may be so acute (Roger Jaensch, personal communications); Vestjens WJM (1977) p. 45. 17 Stafford Smith DM and Morton SR (1990) A framework for the ecology of arid Australia. Journal of Arid Environments 18, 255–78. 18 Roshier DA et al. (2001) Distribution and persistence of temporary wetland habitats in arid Australia in relation to climate. Austral Ecology 26, 371–84. 19 Kingsford RT and Porter JL (1993) Waterbirds of Lake Eyre, Australia. Biological Conservation 65, 141–51. 20 Unpublished data, courtesy of Bureau of Meteorology and South Australian Department of Water, Land and Biodiversity Conservation. 21 Puckridge JT, Walker KF and Costelloe JF (2000) Hydrological persistence and the ecology of dryland rivers. Regulated Rivers: Research and Management 16, 385–402. 22 Reid J (1992) p. 28. 23 Kerwin B and Breen G (1986) pp. 37–38. 24 The Living Murray, Icon Sites, MDBC webpage <www.thelivingmurray. mdbc.gov.au/iconsites>; Parker SA et al. (1979) An Annotated Checklist of

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the Birds of South Australia, Part One: Emus to Spoonbills. South Australian Ornithological Association: Adelaide, p. 24. 25 Colin Thiele (1963) Storm Boy. Rigby: Adelaide. 26 Paton DC (2002) Troubled waters for the Coorong. Nature Australia 27, 60–69; Paton DC (2005) Monitoring of Biotic Systems in the Coorong Region, 2004–2005. University of Adelaide: Adelaide. 27 Walker KF, Sheldon F and Puckridge JT (1995) A perspective on dryland river ecosystems. Regulated Rivers: Research and Management 11, 85–104; Chapman FRH (1963) pp. 8, 11. 28 Australian Broadcasting Commission ‘7.30 Report’ [transcript of program 10 January 2007]. 29 Johnston G, unpublished data, personal communication, 26 March 2008; Andrew Faulkner, The Australian, 28 January 2008. 30 Johnston G, unpublished data, personal communication, 26 March 2008. 31 Rippey E, Rippey JJ and Dunlop N (2002) Management of indigenous and alien Malvaceae on islands near Perth, Western Australia. In Turning the Tide: The Eradication of Invasive Species. (Eds. Veitch CR and Clout MN) pp. 254–59. Occasional Paper of the IUCN Species Survival Commission, No. 27, Gland: Switzerland and Cambridge: UK; Elizabeth Rippey, personal communication, 22 April 2008. 32 See also Vestjens WJM (1977) pp. 38 and 54, for accounts of nesting on human structures and for the majority of fish consumed being species introduced to the Murray-Darling Basin. 33 Chapman FRH (1963) p. 8, writing in the 1960s, described the ‘marked decline’ in pelican abundance in southern South Australia; Andrew Faulkner, The Australian, 28 January 2008; Martin O’Brien of the Victorian Department of Sustainability and Environment confirms the recent demise of coastal colonies in Victoria (personal communication, 6 May 2008). 34 Kingsford RT and Norman FI (2002) p. 50. 35 Birt-Friesen VL et al. (1989) Activity-specific metabolic rates of freeliving northern gannets and other seabirds. Ecology 70, 357–67, esp. Table 4, p. 363. 36 Waterman MH and Read JL (1992) p. 126; Read JL (2003) pp. 64–76. 37 Roby DD et al. (2003) Quantifying the effect of predators on endangered species using a bioenergetics approach: Caspian terns and juve1 18

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nile salmonids in the Columba River Estuary. Canadian Journal of Zoology 81, 250–65. 38 Turner DB (1994) An Investigation of a Fish Kill and the Catch History of the Mulka Fishery, North-East Desert Region, South Australia. Aquasearch South Australia: Adelaide. 39 Ruello NV (1976) Observations of some massive fish kills in Lake Eyre. Australian Journal of Marine and Freshwater Research 27, 667–72. 40 Parker et al. (1979) p. 25; Marchant S and Higgins PJ (1990) p. 741; Read JL (2003) pp. 77–83. 41 Reid JRW, Badman FJ and Parker SA (1990) Birds. In The Natural History of the North East Deserts. (Eds. MJ Tyler et al.) pp. 169–82. The Royal Society of South Australia: Adelaide; Kingsford RT and Norman FI (2002) p. 51; Costelloe JF et al. (2004) p. 74. 42 Marchant S and Higgins PJ (1993) Handbook of Australian, New Zealand and Antarctic Birds. Vol 1B and Volume 2, Raptors to Lapwings. Oxford University Press: Melbourne. See also Chapters 5 and 3 in this volume. 43 Note use of the word ‘may’. This has not been verified as yet. Pelicans may respond directly to the flood-triggering rains, fly into the region, and follow the passage of floodwaters and wait until it is appropriate to breed; however, it is unlikely most birds follow this strategy, see Kingsford et al. (1999), p. 242. 44 Read JL (2003) pp. 77–83. 45 Nelson B (2005); McKilligan raised a similar suite of issues (prevailing winds, thermal formation, the experience of old birds) in relation to the long-distance movements of the straw-necked ibis, see McKilligan NG (1975) Breeding and movements of the straw-necked ibis in Australia. Emu 75, 199–212. 46 Pulido F (2007) The genetics and evolution of avian migration. Bioscience 57, 165–74. 47 Puckridge JT et al. (2000) p. 390; Kingsford RT et al. (1999) p. 231. 48 Roshier DA et al. (2001) Continental-scale interactions with temporary resources may explain the paradox of large populations of desert waterbirds in Australia. Landscape Ecology 16, 547–56. 49 Costelloe et al. (2004) p. 492. 50 Marchant S and Higgins PJ (1990) p. 741. 51 McAllan IAW, personal communication, 19 April 2008. 1 19

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52 This essay is a tribute to my dear friend, the late Jim Puckridge, with whom I shared numerous experiences on field trips to the Lake Eyre Basin, and from whom I gained invaluable diverse insights. His passion for the protection and conservation of Cooper Creek, the Coongie Lakes and wider Lake Eyre Basin wetlands has inspired many. I thank the following for sharing their knowledge with me: Justin Costelloe, Jake Gillen, Roger Jaensch, Greg Johnston, Leo Joseph, Steve Morton, Martin O’Brien, David Paton and David Roshier.

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PARROTS : FUGITIVES OF THE INLAND Penny Olsen

Rainfall drives most natural cycles in Australia, the driest continent on earth, particularly in the three-quarters of the mainland that is arid and semi-arid. In this perverse inland environment, as ecologist Michael Letnic has written, ‘boom means bust, rain means fire’.1 Rare flooding rains bring water, plants bloom and seed and the animals that feed on them rush to breed. But inevitably the green flush bakes to red, fires may flourish, and animals must dig in, move or perish. Charles Darwin was not the first to notice cycles of boom and bust in nature, but he realised that they provided a mechanism for his pet theory, explained in the Origin of Species:

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if there be, owing to the high geometrical powers of increase of each species, at some age, season, or year, a severe struggle for life, and … if variations useful to any organic being do occur, assuredly individuals thus characterized will have the best chance of being preserved in the struggle for life … This principle of preservation, I have called, for the sake of brevity, Natural Selection.2

In the arid inland, this struggle is writ large for residents and visitors alike. Its denizens have had to adapt to both the extremes and to the unpredictability. Extreme conditions require extreme responses, some obvious, others obscure. For over a century and a half, the desert’s and the nation’s most obscure bird has held many of the secrets of its survival close. In all that time, the night parrot (Pezoporus occidentalis), that ‘fat budgie’ with its raft of evocative aliases – the porcupine parrot, spinifex parrot, midnight cockatoo and solitaire – has eluded all except one man, who held most of its secrets until his watery end. Like all the inland’s inhabitants, the night parrot must capitalise on the good times and survive the bad. Does it have a boom and bust lifecycle, like the budgerigar (Melopsittacus undulatus), zebra finch (Taeniopygia guttata), letter-winged kite (Elanus scriptus) and black-tailed native-hen (Tribonyx ventralis), which have explosive breeding, rapidly multiplying and bursting from the desert as it dries again, or does it take a more conservative approach? When conditions are severe, as

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they are many more years than not, does it stay put and tough it out, or, like the emu (Dromaius novaehollandiae), travel extraordinary distances, guided by some mysterious force, to where food plants are seeding?3 Frederick Andrews, employed as a taxidermist and collector for the South Australian Museum early in the 1860s and 1870s, was the European who knew the parrot best, for most if not all other encounters with the secretive species have been serendipitous. Andrews noted that the parrots were not seen when seeding was poor: they come and go according to the nature of the season. When the early season is wet the porcupine grass flourishes and bears large quantities of seed, on which many birds feed; but if, on the contrary, the season is a dry one the grass does not seed, and no birds are to be seen.4

Andrews reported what he saw and did not speculate on the mobility of the parrot, but many have since. 5 Unfortunately for those attempting to understand the nature of the night parrot, his words can be taken to mean that the birds were too few to be seen in poor years, or that they left and returned when conditions had improved. The star-struck parrot was a short-lived sensation when, as plunder of Empire, an unfortunate individual was sent on a sea voyage to London’s Zoological Gardens. ‘The most extraordinary circumstance connected with this bird is, that it is

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nocturnal!’ marvelled its donor. ‘It lives in the rocky caves of the ranges, and comes out at night to feed’.6 Received in November 1867, just two months later it ‘took what appeared to be a fit of sneezing, and died within a day’.7 Dissection revealed the congested lungs of a victim of pneumonia. Little wonder, given that during the day it ‘remained motionless on its tuft of grass’ with not, apparently, even a cave for warmth or shelter from the British winter. Its dissector found that the ‘body was so fat and oily’ that it was difficult to remove the skin intact. Inspired by John Gould’s remarks on the delicate flavour of its congener the ground parrot – ‘equalling if not surpassing, that of the quail’ – he nonetheless declined to taste the bird, but remarked that its ‘delicacy of appearance’ indicated that it would also be pleasant eating.8 Gould managed to see the bird soon after it arrived at the gardens. He was anxious to observe it live, having first described the species for science just six years before from a skin collected in Western Australia, sent direct from Perth.9 Of the specimen skin he wrote: short and dumpy [with several] remarkable differences [from its closest relative the ground parrot:] the head is disproportionately large, the mandibles short and robust, the nostrils high and round, the tarsi and toes short and delicate, and the nails unusually diminutive … the wings large and long, while the tail is very short.10

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Gould was always keen to be the first to describe new species, but had been in possession of this specimen for some years. Even though he had long left the peculiar parrot undescribed, he thought it sufficiently different to deserve its own new generic appellation, Geopsittacus, to separate it from Pezoporus, the ground parrot. He must have had a few reservations though, when he added ‘ornithologists can adopt it or not as they please’.11 Time has shown that they have not. Still, when Gould saw the captive and learned that, as he had suspected, it was a creature of the night, he felt justified: it has never been seen to perch, but moves over the floor of its domicile in a series of jumps, much like those of a Sparrow … Mr Bartlett informs me that, like all other nocturnes, it becomes much more wakeful and active at night, nibbling its tuft of grass, watercress, millet, and canary-seed like a Rabbit. As yet it has not been heard to utter any sound, except a faint whistle.12

On better acquaintance, Mr. Bartlett found that ‘its voice was a double note, harsh and loud’.13 When he made the first description of the night parrot, Gould did not know where in the huge colony of Western Australia his specimen was obtained other than that it came from the interior.14 Nearly 80 years later it was traced to the Mt Farmer district and the 1854 Austin expedition to the area where the Gascoyne River rises.15 Assistant Surveyor Robert

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Austin and party started out from Northam in July 1854. They were dogged by disaster. In August, several horses ate poisonous native plants and died in agony, forcing Austin to discard much equipment and press on with limited mounts. In September, one of the men, Charles Farmer, accidentally shot himself and died in agony eight days later. As temperatures rose, the meagre surface water evaporated and the men had to resort to drinking their own urine and that of their remaining horses before retreating coastward in November. Among the small collection of skins they retained was ‘a beautiful ground parrot (pezoporus [sic] formosus)’16, shot at a native well amongst granite ranges a few days after Farmer’s unfortunate accident. The name was repeated in the appendix on the natural history specimens, written by WA Sanford, and the two misidentifications may well have misled Gould, who never explained his delay in describing it. As eventually became apparent, the parrot had eluded recognition by science for yet another decade, and Gould had also missed another highly significant specimen collected during a previous but equally miserable inland venture. Whereas Gould finally described the night parrot in 1861 from the bird that was collected in 1854, the species was first collected in 1845, during the Sturt expedition. Like Austin, explorer and public servant Charles Sturt was defeated by desert and returned with a night parrot that lay overlooked for years.17 Searching for a way to salvage a stuttering career, at 49 years old, with failing eyesight and deeply in debt, Sturt set

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off in August 1844 with 15 men, six drays, a boat and 200 sheep. He was driven by a conviction that at the heart of Australia there was an inland sea and the hope that he would achieve the honour of discovering it. His party’s send-off from Adelaide was splendid, a public breakfast, making Sturt feel ‘thankful that I have been permitted to witness the happiness of thousands’.18 By December they were short of water but moved further north into the Grey Range, where they were fortunate to find permanent water at Depot Glen on Preservation Creek. The summer heat trapped them there, surrounded by inhospitable country. Scurvy claimed Sturt’s second-in-command and later plagued Sturt himself. Finally, in July, heavy rain released them. Sturt moved his party in a north-westerly direction and made a push towards the centre of the continent to discover sand dunes, but not those of the seaside. They had struck the Simpson Desert at its most unforgiving. The skeletal Sturt and his exhausted party made their way back to Adelaide. Sturt was bitterly disappointed, believing he had found only desolation: gloomy and burning deserts over which I wandered during more than thirteen months … I wish not to hide from my readers the disappointment, if such a word can express the feeling, with which I turned my back upon the centre of Australia, after having so nearly gained … an achievement I was not permitted to accomplish.

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His career, he feared, was ‘probably terminated forever’.19 But, by the time of Sturt’s return in January 1846, a new administration more sympathetic to Sturt and his achievements managed the colony’s affairs and his status improved somewhat, though he might have been more fortunate. In May 1847 he took leave and arrived in London just too late to receive personally the gold medal of the Royal Geographical Society. After his return to Adelaide he was appointed Colonial Secretary but his eyesight forced early retirement. In 1869 he died, still in financial difficulty and before the formalities were completed for his knighthood.20 During Sturt’s period of leave in London, he completed an account of the expedition. He described how in October 1845, north of Cooper Creek, South Australia, one of his party, John McDouall Stuart, later to achieve fame from his own inland explorations, shot a ‘beautiful ground parrot’ on a [salt] plain covered with samphire, salsolae and mesembrianthemum, which gave it the appearance of a sea marsh.21 He also spent time with his old acquaintance, John Gould, with whom he wrote the appendix to his narrative. In this the bird is identified as ‘the ground parrot’, which was ‘only twice seen in the interior, but on both occasions in the same scrubby and salty country it is known to frequent in New South Wales and other places’. The pair described it as having ‘dark green plumage mottled with black’ and ‘a patch of dull red over the bill’.22 The specimen is still in the collection of Gould’s benefactor Lord Derby, and now in the National Museums, Liverpool.23 It is definitely a night parrot that, unlike the ground parrot, 128

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does not have a red frontal band. It seems that Sturt and Gould wrote the account without the bird before them. Indeed, a letter from Lord Derby to Gould suggests that the specimen may have gone astray for a while along with others intended for Gould.24 Sturt had evidently sent off two packages of specimen skins and Gould received the wrong one. While working with Gould, Sturt had noticed the error. A trade was eventually made but the uniqueness of the parrot evidently still escaped notice. Another South Australian, London Zoo’s expatriated parrot, had been captured on a sheep station at Nonning Run leased by Mr Charles Ryan in the newly settled Gawler Ranges west of Spencer Gulf. This was the find that probably sent Frederick Andrews to the ranges in search of more specimens of the highly sought-after rarity.25 In March 1873, the Zoological Gardens received a second bird.26 It was quite likely captured by Andrews and by July it too had expired. These two captives, a male and female, separated by death and six years, were the only night parrots known to have been shipped out of Australia live. Their skins, now in the British Museum, are among the scant 24 night parrots known to be in collections around the world.27 Andrews, in his role as taxidermist for the South Australian Museum, almost certainly collected between 17 and 19 of them, most likely at the Director’s urging because of their value for trades with other museums. Between 1871 and 1883, Andrews spent several periods around Nonning. It seems that he collected the parrot near 1 29

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waterholes in the sandhills on the northern side of the low, spinifex-clad ranges, where rain falls erratically and the arid zone begins. Presumably, he rode the steamer which made a twice weekly run from Port Adelaide to Port Augusta, and from there made his way west 109 km to his base at Nonning. He may well have gone with the mail run from the port to Yardea just west of Nonning, which began in 1867 was said to be the fastest in Australia.28 Although most of his specimens were from the Gawler Ranges region, Andrews also had success elsewhere. As he reported, despite the previous drought, he ‘shot some specimens at Cooper Creek in 1875, when out collecting for the late Mr JW Lewis in his exploration of the country about Lake Eyre’.29 However, the number of specimens is not recorded in the account of the expedition and the register of the South Australian Museum lists only one, presumably that now in Museum Victoria.30 Hence, about 20 specimen skins originated in South Australia, two in Western Australia and two in Queensland. Despite this bias, and only by a quirk of history, the bird was named from Austin’s Western Australia specimen. In 1883 Andrews’ sole publication on the subject of his unique expertise appeared.31 In it he explained that ‘numerous mistakes were made concerning [the night parrot’s] habits and economy which I have endeavoured to correct by many years of study and observation’. He recorded something of the parrot’s roosting and breeding habits:

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During the day this bird lies concealed in the inside of a tussock or bunch of porcupine grass (Triodia), the inside being pulled out and a snug retreat formed for its protection. Here, also, its rough nest is formed, and four white eggs laid.

At Lake Eyre, however, Andrews observed that: ‘They were in that district observed to conceal themselves during the day in thick patches of shrubby samphire, on the salt flats bordering on the creeks,’ and on the lake itself. But Andrews left much unrecorded. One of the secrets to his astonishing success was revealed in a letter to the Director of the South Australian Museum in which he wrote that at Murnia Rockholes (about 8.5 km from Moonaree Homestead): ‘next moon is the time at which I have always obtained night parrots there’.32 Another was the parrot’s apparent dependence on waterpoints: When the dark shades of evening have fairly set in it comes out to feed, but generally flies direct to the nearest water, which is often at a considerable distance from the nest; in some instances I have known them to fly a distance of four or five miles. After drinking and shaking themselves up a little they fly off to feed on the seeds of the porcupine grass, returning to water two or three time during the night.33

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It seems that Andrews spent spend extended periods in wait near water, whereas most others collected as they travelled. He worked by the light of the new moon, when most collectors would have been tucked up in their swag. About a year after presenting his paper to the Royal Society of South Australia, the blue-eyed, grey-bearded Andrews was dead, aged about 60. On 19 October 1884, when he did not return to his host’s home near Mt Jagged, south of Adelaide, a search was mounted. He was found drowned in a waterhole. The inquest determined that he ‘came to his death by falling into a waterhole while suffering the effects of sunstroke’.34 Since he had no relatives in the colony, he was buried locally, at Willunga. Andrews’ background is a mystery. He was said to have begun with ‘slight knowledge of natural history’ when he was hired in his late thirties to work for the museum.35 One source also claims that he had played the ophecleide in the band of the Coldstream Guards when that regiment was at Dublin in Ireland, but there is no mention of him in regimental records.36 A report of the Board of Governors wrote of the loss to the museum, his long service of 25 years and more, and his ‘extensive acquaintance, from actual observation, with … birds’ and concluded that: ‘of late years his heath had been failing, and it was not probable that he could have much longer borne the fatigue and exposure which a collector must necessarily undergo’.37

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Andrews had apparently been unwell since sailing from Port Adelaide to the Aru Islands, in the Arafura Sea, with Adelaide natural history collector Samuel White. White’s boat departed in April 1880, dropping White’s family off in Sydney, and collecting intermittently as they made their way up the east coast. On Queensland’s Barnard Islands fatigue set in and Andrews was ailing and ‘the work from all appearances seemed too much for him’.38 White noted: ‘I am sorry now that I brought Andrews. None but very hardy men are fit for this work, some of the crew are complaining’. On approach to the Aru Islands, White continued to complain: both my taxidermists are laid up with bad feet, considering the little work they have done on the few islands we touched at, coming up the coast, I am afraid they will not stand the hard work in the tropical islands we are bound for. 39

The lush forest must have seemed world away from the parched Gawler Ranges and in the Aru Islands, Andrews was twice lost. But he displayed some of his old resilience and resourcefulness. Lost overnight, he was found in a hollow tree and none the worse for wear, having eaten some large green frogs ‘which he described as being of excellent flavour’.40 Andrews’ employer White proved dogged, but his ornithological endeavours were beset by bad luck. This expedition was no exception. He was possibly too demanding and fell out with

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his captain and crew. With hands festering from the arsenic soap used to preserve specimen skins, and intending to continue the expedition later, he returned to Sydney, but within days a ‘chill’ ended his life.41 White had left Andrews on Thursday Island and returned to Sydney with his other taxidermist, James Cockerell. James was the son of John Thomas Cockerell. Amongst the collection of bird skins sold to a London buyer by Cockerell Senior and donated to the British Museum in mid 1881 was one of the few night parrots unlinked by its label or place of collection to Andrews. Father and son were competent collectors and ornithologists, but ‘creating mischief appeared to be a family failing’.40 Sold as a unique collection containing many as yet unnamed species, ‘the celebrated Cockerell Collection’ turned out to be nothing of the sort. There was ‘not a single new species to be described’. Worse, ‘the data on the skins have been found to be incorrect’.42 In 1880, just before the sale, Cockerell had withdrawn some of the skins from his home museum, Brisbane, following a dispute and accused the museum of maladministration over the disappearance of several specimens, which became the subject of an inquiry by a Senate Select Committee. While waiting for White’s ship to be built in 1879, Cockerell junior was employed to collect in Western Australia. Yet he is not known to have travelled into country where he might have collected a night parrot. A specimen of the night parrot

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was recorded as present among the collections in the Australian Museum at the time of White and Cockerell’s return to Sydney in 1880 but subsequently went missing.43 Was this to become the Cockerell specimen? Despite his best efforts, Andrews’ source of night parrots had dried up by 1883, but several other collectors went further into the desert hoping to find the bird. Lamed by a childhood accident, George Keartland nonetheless served stoically on two difficult explorations of Central Australia: the Horn (1894) and Calvert (1896) expeditions. From the Horn expedition he returned with many specimens, but empty-handed as far as the night parrot was concerned. During the ill-fated Calvert expedition to north-west Australia, he had to abandon over 300 bird-skins, but he saw a night parrot ‘disturbed by Mr. Wells’ camel in passing through some spinifex on a stony rise on which scattered mulga trees were rather too numerous’.44 He regretted that he was unable to shoot it. He also observed that ‘the feathers of these birds were often found in the nests of various small birds or amongst the spinifex, where they had no doubt fallen victims to the hungry dingoes’. On the earlier expedition Keartland had been shown other victims left by predators. He famously reported that at Alice Springs telegraph station during the summer of 1892, ‘many were brought in by cats … and … I was shown … portions of skins, wings and tails’ that covered several picture frames.45 Keartland was able to discern from the remains that the bright green of the adults was replaced by brown in the young. Apparently the remains were

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sufficiently intact for him to note that some had been moulting from one plumage to the next. He further observed that, ‘As they frequent the dense porcupine grass, in which they hide during the day, a good dog is necessary to find them’.46 By then the night parrot was vanishing and hard to find anywhere, dog or not. Keartland’s observations pointed to cats and dingoes as the cause, but there was more to it. As Baldwin Spencer, leader of the 1894 Horn expedition, often regarded as the first scientific expedition to the central deserts of Australia, articulated, the Eyrean fauna was adapted to flood and famine: To this irregular alternation of seasons, and to a great diurnal variation in temperature every animal and plant must become accustomed if it is to survive.47

Still, pastoralists were seduced when they saw the country following heavy rain ‘covered with a most luxuriant vegetation’, leading them to imagine it to be its normal condition, and … send out large numbers of stock to graze … and when the inevitable drought occurs … find their stock dying by hundreds of thousands for want of water.48

But it was more than the optimism of individuals – governments required it. In South Australia, for example, to prevent speculation the 1857 Land Regulations for the wastelands of 136

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the Crown, as less temperate parts of the State were known, had to be stocked with 100 sheep or 16 cattle equivalents per square mile (19 per square kilometre).49 By 1862 the whole of the Gawler Ranges was leased to graziers. Around that time, some 90 000 sheep were shorn at Nonning Run near what was to become Andrews’ night parrot beat. Initially shepherds were employed, but by the late 1870s most were replaced by fences. The savage 1863–1867 drought made it clear that the aridity, unreliable water and watering large numbers of stock from a few water points made the businesses unsustainable. The scale of overstocking (and consequent damage to the land) is demonstrated by several adjustments downwards to a minimum of 20 sheep in 1904 and five in 1939. The land was denuded and eroded, waterholes clogged and trampled. Rabbits took over in the 1870s and they and the livestock attracted greater numbers of dingoes, and so it went on, leaving little space for a parrot dependent on groundcover, good seedings of native plants and water. The 1897 Royal Commission, held towards the end of the next major drought in the State, found that the pastoral industry was severely depressed. It was a scenario typical of the history of much of rural Australia and still pertinent today, as marginal rural industries prosper in the good seasons then fold in the bad. The often untold casualties are the native plants and animals. Virtually all of the night parrot’s range had been turned to pastoral use when, in the early decades of the 20th century, several capable ornithologists, including the son of Samuel 137

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White, Samuel Albert White, made a concerted effort to rediscover the species. Their searches of the Gawler Ranges and other of the parrot’s known or suspected haunts proved futile.50 By 1915, prominent ornithologist Archibald James Campbell was calling the bird ‘missing’. 51 His friend, Alfred Walker, a 25-year resident of Innamincka, not far from where Stuart, on the Sturt expedition, netted his night parrot, and familiar with the parrot reported that according to ‘both whites and blacks’ it had ‘entirely disappeared’, and Campbell concluded that ‘evidently this Parrot has been exterminated. Few skins remain of this remarkable species, while there is not an egg in any collection’. But the parrot still clung to life, concealed by the great interior. In 1912, the year that Samuel Albert White was searching the Gawler Ranges, Martin A Bourgoin, an employee of the Western Australian Public Works Department, was hunting bronzewings in the spinifex-clad hills near Nichol Spring. 52 Mistaking them for pigeons as they rose, he shot one of four parrots. The strange bird caused much discussion among the men in the camp, some maintaining it was a ground parrot and others a night parrot. Bourgoin recognised it correctly and subsequently described its immature plumage, not at that time known. Bullocky, an Aboriginal, knew it too: ‘That one all same rabbit, live along burrow, drink and feed about at night’. They made it into a specimen skin but it was poorly preserved and deteriorated; by the 1930s the skin’s whereabouts could not be traced and the whole incident became yet another fleeting encounter between man and night parrot. 138

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Working inland on the Canning stock route, Bourgoin recorded other close encounters in his diary. He saw several night parrot pairs and family groups between 1920 and 1935: again at Nichol Spring, and at Bolger’s Soak, Pinyerinya Pool, Windich Spring and Ned’s Creek. The latter sighting involved eight birds – two adults and six young in the mouth of a cave. He described their call when coming and going to water as ‘a long drawn-out mournful whistle’. 53 Andrews had earlier reported that the name given to this bird by the Aborigines was an unparrot-like ‘Myrrlumbing’: from the supposed resemblance of their whistling note to the sound of that word. They have also a very peculiar croaking note of alarm whilst at the water, which much resembles the load croak of a frog.54

The combined meagre knowledge of the bird, gathered over 80 years ago, has been plumbed by parrot seekers and poets alike. Seekers hope to filter a night parrot from the night sounds of the desert and glean other clues to its whereabouts. Poets often find inspiration in parrots and the nocturnal parrot just happens to be the antithesis of the bold, bright, noisy stereotype.55 John Kinsella questions the night parrot’s very existence: If at all, then flitting obscurely the rims of water tanks, the outlands of spotlights and filaments of powerlines … 139

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in brief nocturnal flight, with long drawn out mournful whistle.56

For Dorothy Porter, in a chapter headed ‘Chance Meeting’, it represents an interior voice: Myrrlumbing! just an extinct desert call a ghost singing harsh, loud excited57

Perhaps all are drawn by the ephemeral qualities of the peculiar parrot with the strange cry, and the challenge and chance of its refuge, the desert. It was a chance conversation on a train that led able natural history collector FB Lawson Whitlock to be asked to venture to Central Australia in search of the parrot. A commuting grazier had revealed in conversation that he had flushed what appeared to be a night parrot when moving cattle near the Palmer River. Whitlock’s sponsor, well known oologist and specimen collector Henry Luke White (no relation to the Samuel White and his son) had got wind of the report and was keen for him to go. The collector ‘hesitated somewhat, as I always consider it more or less of a gamble to explore these dry interior regions, where the chances for and against a favourable season lean to the adverse’.58

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Still, the expedition was fully financed, so in March 1922 Whitlock began his ‘longest and most arduous’ undertaking for his employer.59 Starting by train to Adelaide and Oodnadatta and then by belligerent camel, he headed inland to Hermannsburg Mission. After some months spent searching the area, in July 1923 he was summoned to the Mission to discover that some local children had fired the spinifex, flushing a parrot that dived into an isolated clump, from which they had caught and cooked it, feathers and all. Inspecting the site, Whitlock found bird prints and, after the ‘natives’ uprooted several Triodia, ‘a small lair, not unlike the nest of the western ground parrot’.60 They brushed away the prints and returned the next day to find fresh footprints but no mate to the unfortunate bird. That was as tantalisingly close as Whitlock came to the fugitive species. As it transpired, Whitlock himself was something of a fugitive. When he settled in Perth in 1901, aged about 40, Whitlock was already an established ornithologist and had published several books on birds.61 He had been an accountant and involved in insurance before emigrating to Australia ‘for reasons of health, to seek a more equable climate’.62 Almost immediately, he began work as a collector and for nearly thirty years travelled widely, mainly in remote parts of Western Australia and under harsh climatic conditions. But health may well have been the least of his troubles. Amongst the criminal cases in the British National Archives at Kew are extradition orders dated November 1897 for Frederick Bulstrode Whitlock. His

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sentence was ‘Extradition from Australia’ and he is described as a ‘fugitive offender’.63 Whatever his crime, he recovered from his fall from grace (and health) and lived a long and successful life as a collector and field naturalist, dying at age 93. In 1923, Whitlock returned from his month-long trip to the Centre in the firm belief that ‘the bird still exists in small numbers, especially in country not stocked with cattle or horses’.64 Clearly, Frederick Andrews was lucky that he went to the Gawler Ranges when he did – in the very early days of pastoralism. By the 1880s, as grazing concerns went bust, he was no longer finding night parrots. There is no doubt that he knew how to secure them, when other talented collectors did not. Since the days of Whitlock and Bourgoin (whose specimen was viewed by ornithologist Dudley Le Souëf), the night parrot has made just two confirmed appearances. In 1990, then again in 2006, it turned itself in. Both times the body of a dead parrot was happened upon alongside a road stretching across a gibber plain by someone with enough knowledge to recognise its significance.65 The record shows that the night parrot has gone bust, but not yet turned to dust. A few fugitives remain, protected by the desert. We continue to wonder whether its population ever underwent violent cycles of dearth and plenty. The brief, tiny boom in its collectability in the Gawler Ranges between 1867 and 1880 has been taken as evidence of a final flush (from a local breeding boom or from movement from the inland) associated with a few wets years about that period. But chance

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may well have placed its collector there just at the point before human impact hit hard. Chance has played a major role in all encounters with the parrot and in the fluctuating fortunes of those who have gone in search of it. Just as it rules the denizens of the desert, and is embodied in the elusive nature of the night parrot, chance also drives boom and bust in the tides of men.

Endnotes 1

Letnic M (2003) In the desert, boom means bust, rain means fire. Australian Geographic 69, 85.

2

Darwin C (1859) Origin of Species, Chapter 4. See http://www.literature. org/authors/darwin-charles/the-origin-of-species/index.html.

3

See Chapters 3 and 4, on black-tailed native-hen and zebra finch respectively, and Chapter 12 on the emu.

4

Andrews FW (1883) Notes on the night parrot (Geopsittacus occidentalis). Transactions of the Royal Society of South Australia 6, 29–30, quote p. 29.

5

Wilson H (1937) Notes on the night parrot, with reference to recent occurrences. Emu 37, 79–87, esp. p. 84; Higgins PJ (1999) (Ed.) Handbook of Australian New Zealand and Antarctic Birds, Vol. 4: Parrots to Dollarbird. Oxford University Press: Melbourne, p. 609.

6

Sclater PL Untitled [Mr PL Sclater read notes upon some recent remarkable additions to the Society’s Menagerie]. Proceedings of the Royal Zoological Society of London 28 November 1867 & 1868, 890–91.

7

Murie J (1868) On the nocturnal ground-parrakeet (Geopsittacus occidentalis, Gould). Proceedings of the Zoological Society of London 27 February 1868, 158–65, quote p. 159.

8

Murie J (1869).

9

Gould J (1861) On a new genus and species of parrakeet from Western Australia. Proceedings of the Royal Zoological Society of London 1861, 100–01.

10 Gould J (1867) The Birds of Australia: Supplement. The Author: London, Part 4, opp. Pl. 66.

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11 Gould J (1865) Handbook to the Birds of the Australia. The Author: London, p. 88. 12 Gould J (1867) opp. Pl. 66. 13 Murie J (1869). 14 Gould J (1867) opp. Pl. 66. 15 Wilson H (1937) p. 81. 16 Austin R (1855) Journal of Assistant-Surveyor R. Austin [commanding an expedition sent by the government to explore the interior of Western Australia, north and east of the settled districts, for extensive tracts of fertile land available for pastoral and agricultural purposes, supposed to exist in the vicinity of a large sheet of inland water, called ‘Cow-cowing[’] and upon the Gascoigne River, flowing into the north arm of Shark’s Bay, which the settlers, feeling the want of more extensive pastures for their rapidly increasing stock, memorialized and aided the government to form and equip this party to examine and report upon.] E. Stirling Government Printer: Perth, p. 46. 17 Cleland JB (1937) The history of ornithology in South Australia. Emu 36, 197–221. 18 Sturt C (1848) Narrative of an Expedition into Central Australia, [performed under the authority of Her Majesty’s Government during the years 1844,5, and 6. Together with a notice of the province of South Australia in 1847]. T & W Boone: London, vol. 1, pp. 38–39. 19 Sturt C (1848) p. 35. 20 Gibney HJ (1967) ‘Sturt, Charles (1795–1869)’. Australian Dictionary of Biography, Vol. 2, Melbourne University Press: Melbourne, pp. 495–98. 21 Sturt C (1848) p. 35. 22 Sturt C (1848) Vol. 2, Appendix p. 41. 23 Forshaw JM, Fullagar PJ and Harris JI (1975) Specimens of the night parrot in museums throughout the world. Emu 76, 120–26. 24 Sauer G (2001) John Gould the Bird Man; Correspondence. Vol. 4 – 1846 through 1851. Maurizio Martino: Connecticut, pp. 215–16 [letter of c. 3 March 1848]. 25 Paton JB (1975) Birds of the Gawler Ranges, South Australia. South Australian Ornithologist 26, 180–92. 26 Sclater PL (1873) Untitled [Additions to the Society’s menagerie during the months of March and April 1873]. Proceedings of the Royal Zoological Society of London 9 May 1873, 1875, 433–34. 14 4

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27 Forshaw JM et al. (1975). 28 D’Arcy Webling D (1995) (Ed.) The Journals of Alfred Charles Webling: Narrating Experiences and Descriptions of Early South Australia, Particularly the Port Augusta Region of Spencer Gulf, and the Escape Cliffs Settlement, Adelaide River, in the Northern Territory, from Voyages Aboard HM Schooner Beatrice during 1862–1866. The Editor, 1995. 29 Andrews FW (1883). 30 Lewis JW (1876) Journal of Mr. Lewis’s Lake Eyre Expedition, 1874–5. South Australian Parliamentary Paper, 1875, no. 19, 1876, pp. 40–410; Waterhouse FG (1875) Specimens from Lake Eyre. The Register [South Australia], vol. XL, no. 8, 968, 1875, 7; Forshaw JM et al. (1975). 31 Andrews FW (1883) Notes on the night parrot (Geopsittacus occidentalis). Transactions of the Royal Society of South Australia 6, 29–30. 32 Forshaw JM et al. (1975) p. 120. See also Joseph L (1983) Further notes on birds of the Gawler Ranges. South Australian Ornithologist 29, 46–54. 33 Andrews FW (1883) p. 29. 34 Whittell HM (1954) The Literature of Australian Birds. Paterson Brokensha: Perth, p. 14. 35 Cleland JB (1937) p. 299. 36 Whittell HM (1954) p. 14. 37 Cleland JB (1937) p. 299. 38 White SA (1920) The Life of Samuel White: Soldier, Naturalist, Sailor. WK Thomas & Co: Adelaide, p. 44. 39 White SA (1920) p. 54. 40 White SA (1920) p. 61. 41 White SA (1920) p. 61. 42 Whittell HM (1940) Frederick Lawson Whitlock. Emu 39(4), 279–85. 43 Forshaw JM et al. (1975) p. 120. 44 North AJ (1897) List of birds collected by the Calvert Exploring Expedition in Western Australia. Transactions of the Royal Society of South Australia 22, 125–92. 45 North AJ (1896) Aves. In Report on the Work of the Horn Scientific Expedition to Central Australia. II. Zoology. (Ed. B Spencer) pp. 53–111, quote on pp. 107–08. Dulau and Co.: London. 14 5

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46 North AJ (1896) p. 108. 47 Spencer B (1896) Narrative and summary of results. In Report on the Work of the Horn Scientific Expedition to Central Australia. I. (Ed. B Spencer) pp. 1–199, quote on pp. 9–10. Melville, Mullen and Slade: Melbourne. 48 Spencer B (1896) p. vii. 49 Gawler Ranges Soil Conservation Board (2004) Gawler Ranges Soil Conservation District Plan. Gawler Ranges Soil Conservation Board. 50 White SA (1913) Field ornithology in South Australia: in the Mallee. Emu 12, 179–85; White SA (1913a) Field ornithology in South Australia: the Gawler Ranges. Emu 13, 16–32; White SA (1912) Field ornithology in South Australia: Port Augusta District. Emu 12, 122–30. 51 Campbell AJ (1915) Missing birds. Emu 14, 167. 52 Wilson H (1937). 53 Wilson H (1937) pp. 83–84. 54 Andrews FW (1883). 55 Kinsella J (2005) Parrotology: on the necessity of parrots in poetry. Australian Book Review October 2005. 56 Kinsella J (1989) Night Parrots. Fremantle Arts Centre Press: Fremantle. 57 Porter D (1984) The Night Parrot. Black Lightning Press: Wentworth Falls. 58 Whitlock FL (1924) Journey to Central Australia in search of the night parrot. Emu 23, 248–281, quote p. 248. 59 Whitlock FL (1924). 60 Whitlock FL (1924) p. 263. 61 Whittell HM (1954). 62 Whittell HM (1954) p. 280. 63 United Kingdom Home Office Records Catalogue Item HO 144/514/ X67243. 64 Whitlock FL (1924) p. 264. 65 Boles WE, Longmore NW and Thompson MC (1994) A recent specimen of the night parrot Geopsittacus occidentalis. Emu 94, 37–40; Boles WE, Longmore NW and Thompson MC (1991) The fly-by-night parrot. Australian Natural History 23, 688–95; Stafford A (2007) Night parrot find suppressed. Wingspan 17(2), 12–13. 14 6

GENYORNIS:

LAST OF THE

DROMORNITHIDS Mike Smith

For such a large bird, taller than a man and weighing in at 275 kg, Genyornis newtonii is surprisingly difficult to find.1 Only one articulated near-complete skeleton of this extinct Australian bird has ever been found and that was excavated at Lake Callabonna by EC Stirling and AHC Zietz in 1893.2 However, Genyornis eggshell is abundant in parts of the arid zone, where it provides a rare archive of environmental and biogeographic information on a crucial time period, around 40–60 000 years ago. At this time, global climate change put Australian ecosystems under pressure, fragmenting woodland and riverine habitats along the desert margins, as the continent swung

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from the boom of the well-watered conditions of the last interglacial (Stage 5.5) into the bust of the lower temperatures and enhanced aridity of the last full glacial (Stage 2).3 At a critical moment in this cycle, people arrived (probably between 50 000 and 60 000 years ago),4 adding a new layer of opportunistic predation to Australian ecosystems. In the history of Genyornis – and its extinction 40–50 000 years ago – I suspect we have something to learn about the ecological resilience of Australia, though much of the detail is still sketchy. From this perspective, the boom and bust that Genyornis speaks to is not the annual or inter-annual climatic variability of ecologists but the longer cycles of Quaternary climate change of the earth scientists.

Getting to know a dromornithid Until the 1990s, Genyornis was considered to be a ratite, like an emu, cassowary or ostrich.5 The ratites, it was thought, were a Gondwanan lineage of large ground-dwelling birds, with the flat sternum characteristic of flightless birds (who do not need the extra muscle attachments that a keeled chest provides) and with a primitive, reptilian palate. So perhaps it is not surprising that reconstructions, like those by Henry Galiano in 1974 and 1979 (Figure 8.1) and Frank Knight in 1985 (see start of this chapter) made Genyornis look like an emu Dromaius novaehollandiae, both in body form and in the long filamentlike feathers that are so characteristic of Dromaius.6 For Genyornis, the problem was that the palate (the key feature in

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this phylogenetic scheme) was either not visible or poorly preserved on the few fragments of fossil crania available. In the early 1990s all this changed, and it was the fossil evidence of much earlier dromornithids (the family that includes Genyornis) that provided the key to a new phylogeny for Genyornis. By then, Peter Murray and Patricia Vickers Rich had recovered enough fossil material of Miocene dromornithids, to show that the ancestors of Genyornis had a hinged rather than solid palate. This drove a reassessment of the evolutionary position of Genyornis, which like all the dromornithids was now placed in a family more closely related to anseriform birds (ducks, geese and swans).7 Scientific perceptions of the biology and ecology of this bird shifted commensurately. Rather than a stocky emu, the latest reconstructions of Genyornis now show a gigantic goose-like bird (see Figure 8.1).8

Palaeoecology Although many of the dromornithids occupied sub tropical woodlands, Genyornis was clearly adapted to the arid interior, as this species had a wide range in south-eastern and central Australia. Its ecology, however, appears to have been quite different to that of the emu (see Table 8.1).9 Genyornis was not only a slightly taller bird, but also a much heavier bird than the emu: more than five times as heavy on conservative bodymass estimates.10 Nor was it much of a runner. With its heavy build and stocky legs it is unlikely it could have run more than 15–20 km per hour (compared to 40–50 km per hour recorded

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Figure 8.1 The changing image of Genyornis newtoni, reconstructed as a stocky emu-like ratite in 1979 (left) (after H. Galiano in Rich (1979)) and as a gigantic goose-like anseriform bird in 2004 (right) (after P. Murray in Murray and Rich (2004)). These are shown on either side of the skeleton of this fossil bird (after Rich (1979)).

for emus). For Stirling and Zietz in 1900, the ‘strange disproportion’ between the size and robustness of the upper leg bones compared to the tarsometatarsus and feet, suggested ‘a slow sluggish habit’.11 The presence of nasal salt glands indicates that Genyornis could tolerate saline waters in arid zone rivers and playas, and eat saline plants (halophytes) without ill

Table 8.1

Comparison of Genyornis and Dromaius.74 Genyornis

Dromaius

Height m

2.15

1.90

Body mass kg

275

50

Running speed km/h

15–20

50

Clutch size

<5

5–15

Egg size mm

125 x 155

90 x 134

Egg weight g

1300

610

150

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effect.12 Its height suggests it browsed on fruits and seed pods, but the location of the fossil finds suggest that samphire and chenopods might also have been important in its diet. Like the emu and ostrich, it might also have been omnivorous, opportunistically eating invertebrates like caterpillars and grasshoppers, or even small reptiles and other animals. And like these birds, it also swallowed small stones (gizzard stones) to mechanically aid digestion, a trait that is absent in birds that are mainly carnivorous. But, unlike the emu, Genyornis seems to have been restricted to riparian zones along lakes, arid rivers and floodplains (where its eggshell is found). The field evidence is clear on this point at least: Genyornis laid clutches of 3–4 large eggs (at 1.3 kg, twice as large as emu eggs) in hollows scraped in sandy ground near rivers, salt flats, or ephemeral lakes. 13 Less is known about its population ecology, but such a large-bodied bird must have needed to forage over a large area to meet its dietary needs. If so, its population density cannot have been high, and if Genyornis was gregarious then any groups of birds must have been widely spaced.14 Without the long-range migratory capability of the emu, Genyornis was probably tethered to corridors of its favoured habitat, perhaps moving up and down channel systems and floodplains as seasonal conditions allowed. Certainly, this is what the distribution of fossil remains suggests. Genyornis had a wide range in those part of the arid interior that are interlaced with river channels (Figure 8.2) – in western New South Wales, the Lake

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Figure 8.2 The distribution of Genyornis newtoni, based on fossil bone (Horton, 1984; Rich, 1985) and eggshell (Williams, 1981; Miller et al. 1999). (Illustration by Clive Hilliker)

Eyre Basin, the Murray-Darling Basin, and in south-western Queensland.15 It appears to have been especially abundant in semi-arid areas between Port Augusta and Lake Torrens, and extended south along coastal salt flats into more temperate regions and into the maze of peat swamplands and old beach ridges that make up the Naracoorte-Coonawarra region of South Australia. There may have been an isolated population

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in south-western Western Australia and there are isolated finds of Genyornis eggshell at Ningaloo on the west coast and at Lake Lewis in Central Australia.16 The latter almost certainly represents a limited extension of range following the corridor of the Finke River, during good years. This distribution of Genyornis corresponds to the modern distribution of chenopod shrubland, with the exception of the Nullarbor Plain.17 This association also seems to hold true over time: at Cuddie Springs, Genyornis was present when the region supported a chenopod shrubland, but disappears when chenopods are replaced by grassland and open Eucalypt woodland.18 At Lake Callabonna, the fossil-bearing muds show that the lake margins were chenopod shrubland around 75 000 years ago when Genyornis lived there.19 Ideas about Genyornis inevitably turn on a comparison with the emu (Dromaius novaehollandiae): the stories of these birds are intertwined. The emu is lanky, arid-adapted and wide-ranging, able to swing out across several hundred kilometres of open desert.20 In contrast, the emerging picture of Genyornis is of a stocky slower-moving bird, ‘graviportal’ rather than ‘cursorial’, tethered to fertile patches near rivers, coasts and lakes. Vulnerability to climatic change, or to interdecadal variability in climate, is greatest where a species is restricted to a fragmented habitat of resource rich ‘islands’ with only a limited capacity to migrate between patches. Predation pressure is at its most devastating where a species is geographically circumscribed in some way. If we think of

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continental extinction as the end point of a series of smaller local extinctions, then this suite of circumstances set up this improbable bird for disaster. The fact that the range of Genyornis overlapped with a focal region for early human settlement – the Murray-Darling Basin – only heightened the likelihood that this would occur. For the story of how this unfolded we need to turn to some remarkable research on emu and Genyornis eggshell.

Eggshell narratives Avian eggshell, including that of the emu and Genyornis, is a bio-ceramic made up of about 95% calcium carbonate built around a flexible structural framework of keratin, a fibrous protein.21 Both of these components have been exploited to reconstruct the history of Genyornis: s 4HE ORGANIC COMPONENT THE AMINO ACIDS IN KERATIN has been the key to building an extinction chronology for this bird; s 3TABLE ISOTOPES IN THE INORGANIC CALCIUM CARBONATE component, in emu and Genyornis eggshell, provide a record of the changing diet of these birds and of a palaeoclimate in the arid interior of Australia. Both lines of research reflect major developments in Quaternary science in the 1990s, in geochronology and in stable isotope research, fuelled in part by the availability of a new generation of scientific instruments capable of analysing milligram-sized samples. 154

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The stage was set in 1981 when Dominic Williams, a young palaeontologist at Flinders University, identified Genyornis eggshell at Dempsey’s Lake near Port Augusta.22 He found three types of fossil eggshell eroding from an old dune. Two of these were readily identified as emu or duck. The third, a thick smooth-surfaced shell, with a distinctive pattern of pores, was unlike any living species.23 Williams examined the size and ultrastructure of this eggshell. By a process of elimination, he concluded that it must belong to Genyornis newtoni.24 Williams died prematurely in 1984 and it was Gifford Miller, a palaeoclimatologist at the Institute of Arctic and Alpine Research, University of Colorado, who picked up his research on fossil eggshell.25

A date for extinction The first task was to establish a chronology for Genyornis. The problem was that most of the fossil eggshell is beyond the effective range of radiocarbon dating. Even the most recent Genyornis eggshell gave 14C ages (ranging from 40 680 BP to >44 400 BP) that probably underestimated the true age of the eggshell. 26 Miller, however, had come to Australia, looking to apply an alternative dating technique, amino acid racemisation (AAR), that he had successfully used to date ostrich eggshell in Africa.27 The great advantage of this technique is that: it requires only milligram samples of eggshell; it can be run cheaply and efficiently on a high-performance liquid chromatograph (HPLC), and can give ages back to at least 140 000 years. 155

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Miller promptly analysed more than 700 samples of Genyornis eggshell, as well as more than 450 pieces of emu eggshell as a control series.28 This was a remarkable research effort, providing the largest statistical sample dating any Australian megafaunal species. It showed that Genyornis disappeared from the Lake Eyre region about 50 000±5000 years ago, and also from the Murray-Darling Basin and Lake Frome areas at about the same time. This ‘extinction window’ should be broadened to 42 000–58 000 years ago to take into account uncertainties with the dating methods. But even this is a remarkable picture of a bird that was relatively common in arid and semi-arid Australia prior to 50 000 years ago, and gone after 40 000 years ago. Of course, not everyone agreed that this was a record of ‘continental’ extinction. At Cuddie Springs, near Brewarrina, there is a bone bed that contains the remains of Genyornis that appear to have survived until 30 000–35 000 years ago. Cuddie Springs is a site with its own dating problems, but taking this at face value suggests that Genyornis underwent a sharp contraction of range beginning 50 000 years ago and that extinction was a regional process taking many millennia, rather than an ‘event horizon’.29 Although Miller’s study included several regions, only the Lake Eyre series was adequately calibrated against other dating methods, leaving just enough uncertainly to cloud the issue of whether extinction was instantaneous or took place at different times in different regions. The eggshell dates suggest that extinction occurred

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a little earlier at Lake Eyre and Lake Frome in the arid zone (50 000 years ago), than on the semi-arid margins in the Murray-Darling and Port Augusta areas (48 000 years ago) or in temperate regions such as at Wood Point near Port Pirie (43 000–46 000 years ago).30

‘An instantaneous signal of diet’31 Eggshell is not only useful for dating; it also provides a window on the diet of birds like emu and Genyornis. This second line of eggshell research was developed by Beverley Johnson, a postdoctoral researcher at the University of Washington, working with the long series of eggshell samples already assembled by Miller.32 Australian plant species use one of two different photosynthetic pathways: C3 or C4. This affects how they manage CO2 during photosynthesis and how much of the heavier isotope of carbon (13C versus 12C) they build into their tissues. This characteristic 13C/12C ratio is passed along the food chain to animals that consume the plants (or eat the insects that consume the plants). All trees, most shrubs, and grasses growing in temperate regions are C3 and have

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C values of –20‰ to –35‰

(per mill). Tropical grasses and chenopods like Atriplex (saltbush) and Halosarcia (samphire), able to grow in hot seasonally dry conditions, use the C4 pathway, and typically have 13C values of –9‰ to –16‰.33 In the early 1980s, PW Hattersley at the Australian National University identified the climatic determinants of C3 and C4 grasses in Australia. Elsewhere,

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research using stable isotopes to reconstruct past vegetation grew into a significant new field in the 1980s and ’90s.34 First, Johnson tackled modern emu eggshell, showing that the 13C signature of the calcium carbonate in the shell faithfully reflected the changing isotopic signature of the ‘ambient vegetation’ across the continent. 35 It could, she said, serve ‘as an indirect proxy for the effectiveness of the summer monsoon’: more summer rainfall meant more C4 plants and more 13

C in the eggshell. Johnson, Miller and colleagues then used a

sample of fossil emu eggshell from the Lake Eyre area, spanning the last 65 000 years, to reconstruct past changes in the boundary or strength of the summer monsoon. The results showed that monsoon rainfall weakened around 45 000±7900 years ago, somewhere within the long interval between 52 900 and 37 100 years ago.36 Before this, the monsoon had fluctuated in strength, but after this time it progressively declined in effectiveness until the last glacial maximum, when there was ‘a near total absence of C4 grasses’. The next step was to look at the isotopic signature of Genyornis eggshell. What was the bird eating? There were some surprises. In emu and ostrich, the organic and carbonate fractions of the eggshell have slightly different

13

C signatures.37

Partly this is a consequence of a bird’s physiology. But it is also because the carbonate in eggshell reflects a bird’s diet three to five days prior to laying, whereas the organic (protein) component reflects the composition of the bird’s tissues, averaging a dietary signature over a few months. In emu eggshell, the

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offset between the two values is 9‰.38 However, in Genyornis eggshell, the first surprise was that the offset between the organic and carbonate fractions was not a constant, nor was it random: it decreased when the organic component was closer to the C4 end of the range. Genyornis, said Miller, must have been a summer breeder.39 When it laid eggs towards the end of summer the divergence between its daily diet and its monthly average was greatest. When Genyornis nested at the start of summer, the isotopic differences between spring feed and its nesting diet were reduced. If true, this helps explain why the eggshell of Genyornis and emu are often found at the same nesting sites: there was a seasonal offset between the winterlaying emu and the summer-nesting Genyornis. The second surprise was that Genyornis was a more specialised, or at least much more selective feeder than emu: 13C values for Genyornis eggshell are more tightly distributed around a mean value (–19.60±2.39‰) than emu (–18.92±3.84‰), showing that this species had a narrower dietary range during the nesting season.40 Although the ‘ecosystem collapse’ paper of Miller et al. in 2005 stressed the importance of C4 plants in the diet of Genyornis, both taxa are clearly mixed feeders using a range of trees, shrubs and grasses. An earlier paper by Miller and others, in 1999, had read the data with a different eye: ‘Genyornis ate primarily C3 browse’ and was likely to be ‘dependent on extensive shrubland’, they said.41 Whatever the case, such a large-bodied bird would require a significant biomass of nutritionally high-value plant food (new grass shoots, flowers, fruits

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and seeds) during the summer breeding season and also during autumn when the chicks were growing fast. The productivity of its preferred habit during summer may therefore have been a strong limiting factor for Genyornis. How adaptable, then, were these birds in their diet? The isotopic data suggests they were selective rather than obligatory specialised feeders:

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show a shift towards more C3 plant foods in temperate areas.42 And these average values obscure some of the variability individual birds demonstrate. Some Genyornis in the Murray-Darling sample ‘ate exclusively C3 plants’ (–26±1‰, n = 9).43 These birds were not as opportunistic as the emu, but nor were they inflexible in their diet. The most dramatic signal in the eggshell, however, was a significant narrowing of the diet of emu about 45 000 years ago. Prior to this time, the diet of Dromaius had ‘a broad dominance of C4 dietary sources and a subsidiary peak dominated by C3 plants’.44 After 45 000 years ago it fed exclusively on C3 plants (–23.64±1.96) and Genyornis, a picky eater, had become extinct.

‘The remaking of Australia’45 Miller put forward a bold synthetic interpretation of his eggshell data, elaborated in his 2005 paper, ‘Ecosystem collapse in Pleistocene Australia’.46 The initial entry of people into Australia around 50 000 years ago, he said, had led to a massive increase in fires that transformed the continent’s vegetation. The decline of C4 plants in the diet of emu, and the extinction of Genyornis, signalled not climate change but rather the

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destruction of a distinctive mosaic of palatable (C3) trees and shrubs in association with these (C4) tropical grasses. Genyornis, an obligate specialised feeder, could not cope with the destruction of this vegetation formation; the emu, an opportunist, simply ate whatever was available. This neatly stitched together ideas about fire and megafaunal extinctions from The Future Eaters by Tim Flannery, with the catastrophic fire histories of Peter Latz, and new research work by Murray and Vickers Rich that positioned the dromornithids as specialised feeders, dependent on a now vanished layer of woodland.47 This was a transformative scenario. It suggested that the contemporary desert landscape was a human artefact, a landscape irrevocably altered by the initial impact of human settlement. It was also stretching the evidence too far. Miller and his colleagues had already published the eggshell data as a record of climate-driven vegetation change; a record of the summer monsoon not human fires.48 There is little direct evidence for a vanished woodland: at Lake Callabonna, 75 000 years ago, Diprotodon died with saltbush in its gut not Bauhinia.49 The Exmouth GC-17 core, the only long pollen record for the arid zone, does not show radical restructuring of shrub or woodland formations 50 000 years ago, nor higher fire frequencies at this time. On the contrary, the Exmouth core records open arid vegetation with little variation in the tree/herb ratio over the last 100 000 years, except for a shift from eucalypts to chenopod shrubland as aridity intensified 35 000 years ago. 50 Archaeologists also have problems with

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Miller’s thesis. Aboriginal population densities in the arid interior appear to have been low throughout much of the late Pleistocene. 51 This means human fires would have been localised and patchy, and major impacts on desert vegetation are unlikely, except where the effects of human burning were amplified by climatic variability. All of this suggests the eggshell isotopic data is more simply explained as a record of a climate shift towards drier conditions, and declining summer rainfall in the Lake Eyre Basin, registered in different ways by Dromaius and Genyornis, two birds nesting in different seasons, one an opportunistic feeder and the other more selective in its diet. To see this, however, we need to uncouple the eggshell record from the tangled debate about pan-continental extinctions. But if the eggshell record is an incidental record of climate and diet, what did happen to this bird?

Blasted into desert The Lake Eyre Basin is a region where signs of Quaternary climate change are deeply etched into the landscape. In 1906, geologist JW Gregory coined the term ‘the Dead Heart of Australia’ for this region, commenting that ‘the once fertile basin of Lake Eyre’ had been ‘blasted into desert’ at some time in the past. 52 A century later, the history of this ‘desiccation’ has now been pieced together in some detail.53 The last significant imprint of river and lake activity in the region seems to have been during Stage 5, 70 000–130 000 years

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ago. At this time, the region was a maze of active river channels and lakes, floodplains and ephemeral wetlands, and a mosaic of riparian vegetation cut into an arid or semi-arid landscape.54 At the centre of this was Lake Eyre, a deep permanent brackish lake until 90 000 years ago, when lake levels began to oscillate and the lake was often dry or a shallow saline water-body. The last major still-stand of Lake Eyre – the final flutter of the ‘Dead Heart’ - was around 65 000 years ago when flooding of the lake threw back a small beach of Coxiellada shells. The rivers flowing into the lake show much the same record. During Stage 5 the channels were active with large permanent waterholes supporting freshwater turtle (chelids), saratogasized fish, large Velesunio mussels and crocodiles. The floodplains surrounding these channels supported a diversified marsupial fauna, including large herbivores like Diprotodon.55 This distinctive semi-arid ecosystem began to wind down as the last interglacial came to an end – and a critical environmental threshold appears to have been breached around 60 000–65 000 years ago. After the last great filling of Lake Eyre, the bed of the dry lake deflated and wind erosion salted the surrounding country with halites and gypsum. Regional dune fields in the Tirari and Strzelecki Deserts were re-activated.56 The Warburton, Cooper and Kallakoopah now became classic arid rivers: mostly dry channels with saline groundwater outcropping in places, but periodically re-activated during ‘boom’ years when they funnel large flows of floodwaters into otherwise ephemeral lakes and back swamps. 57

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With the collapse of this semi-arid and riverine ecosystem most of the megafauna disappeared from the region. The large marsupials appear to have contracted further up the drainage system, towards the margins of the arid zone. 58 However, the dramatic collapse of this ecosystem 65 000 years ago was only one stage of a series of adjustments to declining summer rainfall. Both Johnson’s eggshell data and the Exmouth GC-17 core show that the summer monsoon became significantly weaker after 45 000 years ago. By 35 000–40 000 years ago, increasing aridity had also led to a restructuring of vegetation communities in the northern part of the arid zone. All of this suggests an ongoing process of desertification, with declining summer rainfall, greater inter-decadal variability in rainfall, and a range of impacts on local landscapes and fauna. Genyornis survived longer than most of the megafauna in this region, perhaps because it tolerated saline water, but when summer rainfall declined past a critical point, it too disappeared from the Lake Eyre/Cooper Creek area. These climatic shifts are most evident in the Lake Eyre Basin, but are not restricted to that region. There appears to have been widespread landscape instability in south-eastern Australia around 50 000 years ago (early in Stage 3), changes in the structure of vegetation (with open herbaceous vegetation, predominantly Asteraceae, expanding at the expense of woodlands), and mass-death assemblages of megafauna (such as the 10 000 large macropods that died around a small swamp at Lancefield, Victoria). 59 This suggests Genyornis would have

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faced environmental pressures across much of its range, varying in degree from desert to coast, and by 45 000–50 000 years ago the distribution of this bird may have been contracting to the fringes of the desert. What were the likely effects on Genyornis? The most obvious is fragmentation of the bird’s riparian habitat, with the decline of local shrub and tree formations along the floodplains and lake margins. Substantial blocks of chenopod shrubland and samphire remain today, so this cannot be the full story, but in the sandy country favoured by Genyornis near Lake Eyre, habitat fragmentation would have been sufficient to affect the demographic structure of Genyornis populations and their long-term viability. Another problem may have been disruption of the bird’s nesting ecology. Declining summer rainfall and river flows, and greater inter-decadal variability in rainfall, would have interfered with the summer breeding of Genyornis, reducing the production of fruits, seeds and new growth and limiting the food available for newly hatched chicks.60 For such a large-bodied bird requiring highly nutritious feed, this would have been a strong limiting factor. And then there is the issue of competitive exclusion by the emu: Genyornis may have increasingly found itself in direct competition with the more adaptable and mobile emu for the remaining patches of seasonally productive habitat. Miller et al. discounted the role of climate change in the extinction of Genyornis because the bird had clearly survived earlier arid periods and recovered, and because its final

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disappearance was broadly synchronous across a range of habitats, some of which (like the Murray-Darling Basin) should have been buffered against increasingly arid conditions.61 But evidence now suggests that each interglacial period had its own character.62 And in Australia, at least, these cycles of wet and dry climates are superimposed on a longterm trend towards greater aridity, beginning about 300 000 years ago. ‘It seems,’ says Gerald Nanson, ‘that with each glacial cycle, northern Australia is becoming more arid and the monsoon a less effective driver of fluvial activity.’63 In the Lake Eyre Basin, the long record from rivers such as Cooper Creek ‘provides strong evidence of a trend to decreasing monsoon strength through successive interglacials.’64 These long cycles of boom and bust inexorably shredded the ecology of the desert and its margins.65 But there is probably more to the Genyornis story than climate change alone. For this we need to look outside the arid zone. ‘Evidence of direct predation on Genyornis by humans’, wrote Miller, ‘is limited to a single site’ – Wood Point.66

The last Genyornis? The Wood Point site is on mud flats on the eastern shore of Spencer Gulf, South Australia, in the shadow of the lead smelting town of Port Pirie. It is not a place to be caught in heavy rain. The Genyornis site is on an old mound of red sand but the samphire salt flats surrounding it are impossibly greasy after rain and steering a car becomes a purely nominal

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exercise. Here in October 1992, I excavated what may be one of the last Genyornis nesting sites, working under dark skies and taking refuge in a Port Pirie café when imminent heavy rain forced a halt.67 The presence of stone artefacts showed it was once also an Aboriginal campsite, but the key question was whether the now scattered Genyornis eggshell had formed part of this. The site centres on a low sand ridge, a few metres above samphire mud flats in the supra-tidal zone. Erosion and disturbance by cattle have torn out the centre of the dune, leaving the original deposits standing as ‘residuals’ on either side. My first task was to make a survey plan using the litter of eggshell and stone artefacts on the surface as a guide to the internal structure of the deposit. It quickly became clear that the stone artefacts were widely scattered, whereas, the eggshell was more spatially constrained. The scatter of Genyornis eggshell formed a long patch, some 25 m long, aligned with the strike of the sand ridge on one side of the blowout, in what once would have been the sheltered eastern slope of the dune. Some metres away, the emu eggshell centred on a single dense mass of broken and burnt shell, only a metre in diameter. Excavations showed that the Genyornis eggshell formed a thin (20–40 mm) horizon stratified within the dune, beneath two carbonate palaeosols, following the dip of the dune sediments.68 Some of this shell was scorched. A few pieces were spot burnt. Some pieces of eggshell had holes from the teeth of small animals (possibly dasyurids) that had robbed the nest.

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None of the excavations turned up any in situ stone artefacts: by the end of the dig it was clear to us that the stone artefact and other occupation debris had been let down from younger sediments higher in the dune profile. However, the presence of so much burnt eggshell suggested some human intervention. Burnt eggshell is rare in the Australian landscape.69 However, at Wood Point, nearly 8% by weight of Genyornis eggshell was scorched or burnt, and 24% of the emu eggshell was burnt.70 The mud flats surrounding the sand ridge are unlikely to have carried a wildfire. In this context, the presence of so much burnt eggshell suggests that people cooked and ate the eggs of both Genyornis and emu, in what is known in the trade as a ‘dinnertime camp’. But this interpretation remains circumstantial: clinching evidence of artefacts in association with Genyornis eggshell eluded me. It was almost a decade before we had a firm handle on the chronology of Wood Point but we can now reconstruct its history in general outline.71 As sea level fell from its interglacial high-stand, the Wood Point area became part of a wide plain bordering a shallow sub-tidal estuary.72 With a landscape inheritance of extensive saline mudflats, this area is likely to have been dominated by halophytes such as chenopods and samphire. The local Genyornis would have been part of a population inhabiting a huge area of chenopod shrubland that stretched from Wallaroo across to Whyalla and Port Augusta. As sea level fell, this area increased probably reaching its maximum extent around 50 000 years ago. At Wood Point, a linear

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dune built up over the coastal muds as aridity increased. Genyornis favoured this as a nesting site and laid its eggs in a shallow deflation hollow on the sheltered side of the dune, around 43 000–46 000 years ago. Judging by the amount of eggshell, local Genyornis must have re-used this nesting site over several seasons. Some eggs were lost to predation by small animals. Others were lost to people. The prominent sand ridge attracted the attention of Aboriginal foragers, who found the nests and lightly cooked at least one clutch of eggs on the spot. There is no sign of Genyornis after this. Another clutch of eggs was laid on the dune and again lost to the hunters, but these were emu eggs dating between 41 000 and 43 000 years ago. Between these two points in time, the Genyornis population in this area collapsed. The emu appears to have taken over the dune as a favoured nesting site from this time on and its eggshell is found in the modern sands at the top of the dune sequence. Finally, sometime in the last few thousand years, Aboriginal people used the old dune as a convenient raised campsite giving access to the surrounding supratidal flats. Wood Point illustrates some of the complications in reconstructing the pattern of events leading to the extinction of Genyornis. Unlike the Lake Eyre region, this was an area where suitable habitat for Genyornis was expanding rather than fragmenting. Other pressures may have been in play - disruption of summer nesting, competitive exclusion by the emu, and predation on the eggs and chicks – but there is the additional impact of people. The archaeology suggests that Genyornis

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abandoned the old nesting site at Wood Point once people discovered it was a place where eggs could be had. And yet, the impact of people cannot have been instantaneous for the Wood Point Genyornis persisted until 43 000–46 000 years ago, some 10 millennia after people first entered the continent. In a recent review of megafaunal extinctions, Chris Johnson noted that a small increase in mortality resulting from human predation on animals with low fecundity could eventually push them to extinction over a few millennia. In his words, ‘long-lived, slow-breeding and slow maturing species, living in situations that guaranteed high exposure to people, have been the most likely to disappear.’73 Thus, space and time conspired against Genyornis. By 50 000 years ago, it was under a range of environmental pressures and shared most of its fallback areas with a new opportunistic predator – humans. In these circumstances the line between resilience and ‘bust’ may have been a fine one. The eggs at Wood Point may well have been laid by the last bird of this species. Whatever the case, with the death of the last Genyornis, Australia lost the last member of a distinctive Australian lineage of large flightless birds.

Endnotes 1

The species was described and named by Stirling and Zietz in 1900 from several skeletons recovered from excavations at Lake Callabonna, a salt lake in arid northern South Australia. The name commemorates Alfred Newton, then professor of zoology at the University of Cambridge, who was enlisted to announce the Lake Callabonna finds in Nature, in 1893 (Newton A (1893) Palaeontological discovery in Australia. Nature 47, 606). ‘Genyornis’ refers to the large size of the mandible (‘genys’ jaw; ‘ornis’

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bird). Although there were earlier finds of dromornithid fossils – by Thomas Mitchell at Wellington Caves in 1836 and Julian Tenison Woods near Penola in 1865–6 – most appear to belong to an older, late Miocene genus, Dromornis. Stirling and Zietz published a detailed monograph on their Callabonna finds, Fossil Remains of Lake Callabonna, in the shortlived series Memoirs of the Royal Society of South Australia 1 (parts 1–4), 1899– 1913, pp. 1–178. These birds then received little scientific attention until the late 1970s, when Pat Vickers Rich reviewed their systematics (Vickers Rich P (1979) The Dromornithidae. Bulletin of the Bureau of Mineral Resources, Geology and Geophysics 184, 1–196.) 2

Stirling EC and Zietz AHC (1899–1913).

3

The Quaternary was marked by a repeated cycle of rapid global warming followed by more gradual cooling. The oscillations form a distinctive saw tooth pattern with the warm peaks roughly 100 000 years apart. Cold stages are given even numbers. Warm stages have odd numbers. The original stage numbers were allocated by C Emiliani. As the palaeoclimate record became more detailed, the scheme had to expand to include lettered substages (e.g. 5e). These were later replaced by a decimal system (e.g. stage 5.5), which allowed for even more detail. See Aitken MJ (1990) Science-Based Dating in Archaeology. Longman: London.

4

Archaeological research in western Arnhem Land in the late 1980s suggested that Australia was initially colonised by people somewhere between 50 000 and 60 000 years ago (see Roberts RG, Jones R and Smith MA (1990) Thermoluminescence dating of a 50 000 year-old human occupation site in Northern Australia. Nature 345, 153–56, and Roberts et al. (1994) The human colonisation of Australia: optical dates of 53 000 and 60 000 years bracket human arrival at Deaf Adder Gorge, Northern Territory. Quaternary Science Reviews (Quaternary Geochronology) 13, 575–83). This later received support from luminescence and ABOX 14 C dating of the Devils Lair cave site (south-west WA) and Lake Mungo (western NSW). JF O’Connell and Jim Allen argued in 2004 that none of the field evidence necessarily indicated occupation prior to 45 000 BP (about 48 000 years ago) (see ‘Dating the colonisation of Sahul (Pleistocene Australia-New Guinea): a review of recent research’. Journal of Archaeological Science 31, 2004, pp. 835–53). Most recently, Peter Hiscock (2008) in Archaeology of Ancient Australia, Routledge: London, argues that people must have reached Australia between 50 000 and 60 000 years ago.

5

Rich PV (1979). Throughout the 1980s, doubts about identification of dromornithids as ratites grew. In 1984, Rich and Gerry van Tets com-

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mented (in: What fossil birds contribute towards an understanding of origin and development of the Australian avifauna. In Vertebrate Zoogeography and Evolution in Australasia; Animals in Space and Time. (Eds. M Archer and G Clayton) pp. 421–26. Hesperian Press: Carlisle, Western Australia.) that: ‘the dromornithids are quite distinct from the Casuariidae, especially in the structure of the quadrate … and may in fact not even be ratite birds.’ (p. 432). See also Rich PV and Balouet J, The waifs and strays of the bird world or the ratite problem revisited, one more time. In Archer M and Clayton G (1984) pp. 447–55; and Noble JC (1991) who in ‘On ratites and their interactions with plants’ (Revista Chilena de Historia Natural 64, 85–118) commented that ‘while palatal evidence is still lacking, these birds may not have been ratites…’(p. 95). 6

For the reconstruction by Henry Galiano see Rich PV (1979) (cover, frontispiece and Fig. 7); for that by Frank Knight see Rich PV and van Tets G (1985) (Eds.) Kadimakara: Extinct Vertebrates of Australia. Pioneer Design Studios: Lilydale, Victoria, p. 189.

7

Murray PF and Megirian D (1998) The skull of dromornithid birds: anatomical evidence for their relationship to Anseriformes (Dromornithidae, Anseriformes). Records of the South Australian Museum 31, 51–97. Murray PF and Rich PV (2004) Magnificent Mihirungs: The Colossal Flightless Birds of the Australian Dreamtime. Indiana University Press: Bloomington. Classification of the ratites has also changed. Genetic data shows that these birds diverged rapidly following the break up of Gondwanaland. Cassowary, emu, kiwi and ostrich form a cluster of related lineages, the various genera of moa form another, and the South American rhea forms an outlier. See Cooper A et al. (2001) Complete mitochondrial genome sequences of two extinct moas clarify ratite evolution. Nature 409, 704–7.

8

These are by Peter Murray. See Murray P and Rich PV (2004) p. 189 and 205, and Johnson C (2006) Australia’s Mammal Extinctions: A 50,000 Year History. Cambridge University Press: Melbourne, p. 64.

9

Information about the ecology of Genyornis is summarised from Murray P and Rich PV (2004) with additions from Williams DLG (1981) Genyornis eggshell (Dromornithidae; Aves) from the Late Pleistocene of South Australia. Alcheringa 5, 133–140; Miller GH et al. (1999) Pleistocene extinction of Genyornis newtoni: human impact on Australian megafauna. Science 283, 205–8; and from my own observations of fossil eggshell in the field.

10 Murray P and Rich PV (2004) Chapter 10, use several different methods to estimate body mass with estimates ranging from 250–515 kg.

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11 Stirling EC and Zietz AHC (1900) Fossil remains of Lake Callabonna. Part II, p. 55. Later work suggests it may have been more nimble than this allows: Murray P and Rich PV (2004) describe the large dromornithids as ‘efficient pacers that could sustain moderate speeds for long periods’. p. 231. 12 Murray P and Rich PV (2004) p. 263. This is another point of difference with emus and cassowary, which do not have nasal salt glands. 13 Williams DLG (1981) Table 1. 14 At Lake Callabonna, several individuals were found bogged in the lake muds together, suggesting these birds may have lived in small groups like emu, rather than as solitary birds, like cassowary. See Stirling EC and Zietz AHC (1899–1913) Fossil remains of Lake Callabonna; and Rich PV (1985) ‘Genyornis newtoni Stirling and Zietz 1896. A Mihirung’. In Kadimakara: Extinct Vertebrates of Australia. (Eds. PV Rich and GF van Tets) pp. 188–194. Pioneer Design Studios: Lilydale, Victoria, esp. p. 190. 15 For the distribution of skeletal fossils of Genyornis newtoni, see Horton DR (1984) Red kangaroos: last of the Australian megafauna. In Quaternary Extinctions: A Prehistoric Revolution. (Eds. PS Martin and RG Klein) pp. 639–80, Fig. 29.8b. University of Arizona Press: Tucson; and Rich PV (1985). For the distribution of Genyornis eggshell see Williams DLG (1981); Miller GH et al. (1999). Information on the Lake Lewis and Ningaloo eggshell is Miller pers. com., August 2007. Genyornis does not appear to be present in Pleistocene faunal assemblages from the Nullarbor Plain, Prideaux et al. (2007) An arid-adapted Middle Pleistocene vertebrate fauna from south-central Australia. Nature 445, 422–25. Genyornis eggshell has not been found in the Great Victoria Desert or around Lakes Gregory or Mackay, despite several field surveys. 16 For south-western Australia, the finds include a single dromornithid bone at Mammoth Cave – a sacral vertebra of the right size for Genyornis. 17 Atlas of Australian Resources, Volume 6. Vegetation. Third series. AUSLIG, Canberra, 1990, p. 42. An association with chenopod shrubland is my suggestion. 18 Field JH and Boles WE (1998) Genyornis newtoni and Dromaius novaehollandiae at 30 000 BP in central northern New South Wales. Alcheringa 22, 177–88; Field JH, Dodson JR and Prosser IP (2002) A Late Pleistocene vegetation history from the Australian semi-arid zone. Quaternary Science Reviews 21, 1023–37.

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19 Rich PV (1985). The age of the Lake Callabonna fossils is given by an OSL date (75±9 ka) on sediments inside a Diprotodon skull. See Roberts RG et al. (2001) New ages for the last Australian megafauna: continentwide extinction about 46 000 years ago. Science 292, 1888–92, Table 2. 20 Davies SJJF (1968) Aspects of a study of emus in semi-arid Western Australia. Proceedings of the Ecological Society of Australia 3, 160–66; Davies SJJF (1976) The natural history of the emu in comparison with that of other ratites. In Proceedings of the 16th International Ornithological Congress. Canberra. (Eds. HJ Frith and JH Calaby) pp. 109–20. Australian Academy of Science: Canberra. 21 See Calvery HO (1933) Some analyses of egg-shell keratin. The Journal of Biological Chemistry 100, 183–86; Mann K (2004) Identification of the major proteins of the organic matrix of emu (Dromaius novaehollandiae) and rhea (Rhea americana) eggshell calcified layer. British Poultry Science 45, 483–90; Panheleux M et al. (1999) Organic composition and ultrastructure of eggshell: a comparative study. British Poultry Science 40, 240–52. 22 Williams DLG (1981). 23 Williams also considered ostriches, which were introduced into this region in the 1880s, but ruled them out. The fossil eggshell is thinner than ostrich eggshell (1.15±0.12 mm compared to 2 mm), has a different pore structure, and occurs in late Pleistocene sediments. 24 This identification has not been challenged, but remains circumstantial as a skeleton of Genyornis has never been found with its eggs; eggshell is almost never found in the bone beds that contain the remains of these birds; and no-one has yet tested the attribution by comparing DNA in the bones and eggshell. 25 Miller’s work on eggshell was published in a series of papers, including three in Science and one in Nature, which collectively represent a remarkable contribution to understanding the Quaternary history of the Australian arid zone. See also Smith MA, Miller G and van Tets GF (1994) Burnt ratite eggshell from Pleistocene Aeolian sediments. Records of the South Australian Museum 27, 228; Kaufman DK and Miller GH (1995) Isoleucine epimerization and amino acid composition in molecularweight separations of Pleistocene Genyornis eggshell. Geochimica et Cosmochimica Acta 59, 2757–65; Miller GH, Magee JW and Jull AJT (1997) Low-latitude glacial cooling in the southern hemisphere from aminoacid racemization in emu eggshells. Nature 385, 241–44; Magee JW and Miller GH (1998) Lake Eyre palaeohydrology from 60 ka to the present:

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beach ridges and glacial maximum aridity. Palaeogeography, Palaeoclimatology, Palaeoecology 144, 307–29; Johnson BJ et al. (1999) 65,000 years of vegetation change in Central Australia and the Australian summer monsoon. Science 284, 1150–52; Miller GH et al. (1999); Miller GH et al. (2005) Ecosystem collapse in Pleistocene Australia and a human role in megafauna extinction. Science 309, 287–90; Miller G et al. (2005) Sensitivity of the Australian monsoon to insolation and vegetation: implications for human impact on continental moisture balance. Geology 33, 65–68. 26 SUA-1337 (40 680 BP) is from Williams DLG (1981) p. 138. No error is quoted. For other dates, see Miller GH et al. (1999). 27 Brooks AS et al. (1990) Dating Pleistocene archaeological sites by protein diagenesis in ostrich eggshell. Science 248, 60–64; see also Johnson BJ et al. (1999). This technique uses isoleucine, a common amino acid and a component of organic keratin, to estimate the age of the eggshell. It works because isoleucine has several isomers: L-isoleucine gradually degrades to D-alloisoleucine, at a rate dependent on temperature. If the thermal history of a piece of eggshell can be estimated, the D/L ratio will give its age. For a general account of AAR, see Aitken MJ (1990). 28 On its own, AAR is a ‘quick and dirty’ technique with a precision no better than 40–50%. However, if it can be calibrated against pieces of known age, the error reduces to 15% or about ±8000 years for samples of late Pleistocene age (see McCoy WD (1987) The precision of amino acid geochronology and paleothermetry. Quaternary Science Reviews 6, 43–54). Miller did this using 90 paired AAR/AMS 14C samples on emu eggshell within the range of radiocarbon dating, and using U-series dates (on Genyornis eggshell) or luminescence dates (on sediments containing eggshell) to extend the range of this to include the older Genyornis eggshell. 29 For these Genyornis remains, see Field JH and Boles WE (1998). Cuddie Springs is one of the most famous late Quaternary megafauna sites in Australia. The site has attracted much controversy regarding its age and interpretation (see, for example, Gillespie R and Brook BW (2006) Is there a Pleistocene archaeological site at Cuddie Springs? Archaeology in Oceania 41, 1–11), but has been forcefully defended by its excavator, Judith Field in Field JH and Dodson JR (1999) Late Pleistocene megafauna and archaeology from Cuddie Springs, south-eastern Australia. Proceedings of the Prehistoric Society 65, 275–301; Field J, Fullagar R and Lord G (2001) A large area archaeological excavation at Cuddie Springs. Antiquity 75, 696– 702; Field JH et al. (2002); Trueman CNG et al. (2002) Prolonged coexist-

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ence of humans and megafauna in Pleistocene Australia. Proceedings of the National Academy of Science 102, 8381–85. I was fortunate to be able to work at the site during the extensive open-area excavations in 1997, described in Field JH et al. (2002). My own reading of the site is that the crucial layers (stratigraphic units 6a and 6b) that contain the most recent megafaunal remains probably represent a palimpsest of archaeological occupation dating to 30 000–35 000 years ago and a slightly older bone bed, perhaps dating 40 000–50 000 years ago. This interpretation is in line with recent ESR dating of megafauna teeth, giving ages 10 000 years older than 14 C on charcoal from the archaeological layers, or OSL on the sediments; Field J (2007) Comment at The Selwyn Symposium (Geological Society of Australia) on Climate Change or Human Impact? Australia’s Megafaunal Extinction, 27 September 2007, University of Melbourne. 30 Plenary address by Miller GH (2007) Separating the impacts of climate change and human colonisation on the flora and fauna of Pleistocene Australia. 3 August 2007, XVII INQUA Congress, Cairns – see Quaternary International 2007, 167–168 (Supplement), pp. 282–3. 31 The phase is from Johnson BJ, Fogel ML and Miller GH (1998) Stable isotopes in modern ostrich eggshell: a calibration for paleoenvironmental applications in semi-arid regions of Southern Africa. Geochimica et Cosmochimica Acta 62, 2451–61. 32 See Johnson BJ et al. (2005) Carbon isotope evidence for an abrupt reduction in grasses coincident with European settlement of Lake Eyre, South Australia. The Holocene 15, 888–96; Miller GH et al. (2005). 33 For the distribution of C3 and C4 grasses see Hattersley PW (1983) The distribution of C3 and C4 grasses in Australia in relation to climate. Oecologia 57, 113–28. Miller GH et al. (2005), Table S6, list the carbon isotope composition of plants in the Lake Eyre region. Jacobs SWL (2001) Review of leaf anatomy and ultrastructure in the Chenopodiaceae (Caryophyllales). Journal of the Torrey Botanical Society 128, 236– 53, Table 1, gives a useful listing of C3/C4 pathways in the chenopodiaceae, with good coverage of Australian taxa. 34 See Ehleringer JR (1991) 13C/12C Fractionation and its utility in terrestrial plant studies. In Carbon Isotope Techniques. (Eds. DC Coleman and B Fry) pp. 187–200. Academic Press: New York; Grocke DR (1997) Distribution of C3 and C4 plants in the Late Pleistocene of South Australia recorded by isotope biogeochemistry of collagen in megafauna. Australian Journal of Botany 45, 607–17. For African research see Johnson BJ et al. (1998); Von Schirnding Y, Van Der Merwe NJ and Vogel JC (1982) Influence of diet and age on carbon isotope ratios in ostrich eggshell. Archaeometry 24, 3–20. 176

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35 Johnson BJ et al. (1999). 36 The eggshell series was dated by AAR. The error for samples dating between 45 000–65 000 years old could not be formally determined, but may be up to 7900 years, Johnson BJ et al. (1999), note 15. This is comparable to errors of 15% given for the AAR technique. See McCoy WD (1987). 37 The organic component of eggshell is enriched by 2–3‰ relative to a bird’s diet; whereas eggshell carbonate is enriched by 10‰ in emu and 15‰ in ostrich. 38 Most dating and isotopic analyses use eggshell carbonate because this makes up the bulk of the eggshell and is relatively easy to extract. The usual reason for then measuring the organic component is to check that fossil eggshell has been chemically stable: if the difference in 13C between the fractions is not within 2‰ of the average difference (9‰ in emu) then one of the fractions may have been diagenetically altered (making it unsuitable for 14C or AAR dating). 39 This argument is developed in the online supplementary data for Miller GH et al. (2005). 40 Miller GH et al. (2005). These are 13C carb for eggshell (>50 ka) from Lake Eyre (Table S1). Another difference between the taxa is that Genyornis eggshell has a clear unimodal distribution of 13C values. In contrast Dromaius 13C has a bimodal distribution, showing ‘a broad dominance of C4 dietary sources and a subsidiary peak dominated by C3 plants’, (p. 289). This must reflect interannual variability in rainfall, with pulses of C4 production extending into the laying season. 41 Miller GH et al. (1999) pp. 207–8. 42 From –19.6‰ at Lake Eyre to –20.42‰ and –22.49‰ in the MurrayDarling and Port Augusta areas respectively Miller GH et al. (2005) Table S1. 43 Miller GH et al. (1999) p. 208. 44 Miller GH et al. (2005) p. 289. 45 Johnson CN (2005) The remaking of Australia’s ecology. Science 309, 255–56. 46 Foreshadowed in Miller GH et al. (1999), and developed in Miller GH et al. (2005). 47 Flannery T (1994) The Future Eaters; An Ecological History of the Australasian Lands and People. Reed: Melbourne, suggested that overhunting led to rapid extinction of the megafauna. The removal of this guild of large 17 7

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herbivores from the Australian ecosystem had allowed massive build up of fuel loads, leading to destructive large fires, which transformed the landscape by permanently altering soils and nutrient cycling. By contrast, Miller placed more weight on destruction of habitat by human fires as the proximate cause of these extinctions. For ideas about fire by Peter Latz (1995), see Bushfires and Bushtucker: Aboriginal Plant Use in Central Australia. IAD Press: Alice Springs, and (2007) The Flaming Desert: Arid Australia – A Fire Shaped Landscape, The Author: Alice Springs. On the diet of early dromornithids, Murray P and Rich PV (2004) argue these birds were originally adapted to closed, low deciduous woodland and vine thicket, composed of fruit-producing species such as Brachychiton, Dichrostachya, Owenia, Swainsonia, Terminalia and species in the bullwaddie association (Macropteranthes keckwickii and M. leichardtii). They note, however, that Genyornis probably survived the long-term decline in this vegetation because it was better suited for ‘bulk feeding on less nutritional plants’ (p. 301). 48 Johnson BJ et al. (1999). 49 See Stirling EC and Zietz AHC (1900), described as the masticated remains of ‘stems and twigs of plants belonging chiefly to to the order Salsolaceae, or to the allied orders Amarantaceae or Nyctagineae.’ (pp. xii–xiii and p. 36). 50 Kershaw P et al. (2006) Environmental change and the arrival of people in the Australian region. Before Farming 2006/1, pp. 1–24. GC-17, an offshore ocean core, provides a record of vegetation change in the northern (summer rainfall) sector of the desert. Although it is the longest record of palaeo-vegetation for the arid zone, it is missing data for the key time period 41 000–64 000 years ago. Kershaw et al. rather coyly conclude: ‘The record has bearing on the interpretation of sequences of fossil egg shells of flightless birds derived from aeolian dune systems in similarly arid environments within central Australia.’ (p. 19) (my italics). In an earlier paper (Kershaw AP, van der Kaars S and Moss P (2003) Late Quaternary Milankovitch-scale climatic change and variability and its impact on monsoonal Australia. Marine Geology 201, 81–95), Kershaw more directly challenged Miller’s hypothesis, arguing that because ‘the environmental drying trend [evident in vegetation records like GC-17] was initiated well before evidence for the presence of people on the continent,’ and because there was also ‘strong MIS3 vegetation change … after the earliest dates for the presence of people,’ that ‘it is unlikely that anthropogenic burning was the trigger. Instead, it may be the case that summer monsoon activity has weakened generally along with increased ENSO activity over the late Quaternary period …’ (p. 93).

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51 The earliest human movements into the Australian desert may have taken place sometime between 35 000 and 45 000 years ago, see Smith MA (2005) Moving into the southern deserts: an archaeology of dispersal and colonisation. In 23°S: Archaeology and Environmental History of the Southern Deserts. (Eds. MA Smith and P Hesse) pp. 92–107. National Museum of Australia Press: Canberra. There is currently no direct evidence for human occupation of the Lake Eyre region before about 20 000 years ago. 52 Gregory JW (1906) The Dead Heart of Australia. John Murray: London, p. 151. 53 See Hesse PP, Luly JG and Magee JW (2005) The beating heart: environmental history of Australia’s deserts. In MA Smith and P Hesse (2005), pp. 56–72. The key papers are Johnson BJ et al. (1999); Magee JW et al. (1995) Stratigraphy, sedimentology, chronology and palaeohydrology of Quaternary Lacustrine deposits at Madigan Gulf, Lake Eyre, South Australia. Palaeogeography, Palaeoclimatology, Palaeoecology 113, 3–42; Magee JW and Miller GH (1998); Magee JW et al. (2004) Continuous 150 k.y. monsoon record from Lake Eyre, Australia: insolation-forcing implications and unexpected Holocene failure. Geology 32, 885–88; Nanson GC, Price DM and Short SA (1992) Wetting and drying of Australia over the past 300 ka. Geology 20, 791–94; Tedford RH and Wells RT (1990) Pleistocene deposits and fossil vertebrates from the ‘Dead Heart of Australia’. Memoirs of the Queensland Museum 28, 263–84. 54 See DeVogel SB et al. (2004) A GIS-based reconstruction of Late Quaternary paleohydrology: Lake Eyre, arid Central Australia. Palaeogeography, Palaeoclimatology, Palaeoecology 204, 1–13, for maps of the extent of lakes and floodplains at this time. 55 Tedford RH and Wells RT (1990). 56 Fitzsimmons KE et al. (2007) The timing of linear dune activity in the Strzelecki and Tirari deserts, Australia. Quaternary Science Reviews 26, 2598–616. 57 The early part of Stage 3 (45–60 000 years ago) saw water return to lakes in Willandra in western NSW (such as Lake Mungo), but there was only limited fluvial activity in central Australia during this period. There were episodic strong seasonal flows along Cooper Creek, but these were of short duration and did not rework the extensive floodplains. Shallow saline lacustrine conditions may also have returned to Lake Eyre sometime between 50 000 and 25 000 years ago – but stratigraphic evidence for this is ambiguous and the dating uncertain (Magee JW et al. (1995)

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p. 38). In general, Stage 3 seems to have been ineffective in reactivating rivers and lakes in the arid zone: The picture is more of an arid environment marked by high inter-decadal or sub-millennial variability and strong seasonality. 58 For an accessible summary of this fauna see Webb S (2006) The First Boat People. Cambridge University Press: Cambridge, UK, pp. 134–161. The Lake Callabonna diprotodons date to 75 000 years ago (Roberts RG et al. (2001) Table 2): the preservation of numerous skeletons of these animals on the surface of the modern playa is strong indication that this lake has not seen significant lacustral conditions since that time. Although most of the megafauna had disappeared from the Cooper Creek/Lake Eyre area by 65 000 years ago, the isolated find of an articulated Diprotodon skull on the shores of Lake Eyre, dated by OSL to 46.6±3 ka, indicates that these animals survived further upstream and were able to disperse widely during ‘boom’ years, perhaps following the river channels after episodic flooding (unpublished research by Steve Webb, in plenary INQUA address by GH Miller, pp. 282–83). 59 See Cupper ML and Duncan J (2006) Last glacial death assemblage and the early human occupation at Lake Menindee, south-eastern Australia. Quaternary Research 66, 332–41, for a summary and references (pp. 339– 341). The Cuddie Springs bone bed (especially unit 6b) may be a contemporary mass-death assemblage associated with an increasingly variable climate at this time. It is relevant that the remains of Genyornis are not found after 6b. 60 Even the relatively hardy chenopods and samphire communities decline in prolonged drought conditions: mature plants survive perhaps for a century or more but are not replaced when they become senescent. For instance, at Cuddie Springs, increasing aridity saw the replacement of chenopods by grass as aridity intensified about 35 000 years ago – Field JH et al. (2002). A decline in summer rainfall would also affect the production of fruit and new growth as most chenopods respond opportunistically to rain. 61 Miller GH et al. (1999) and (2005). 62 Paper by Wolff EW, Fischer H, Lüthi D and Masson-Delmotte V (2007) The occurrence and structure of interglacials in the Late Quaternary. XVII INQUA Congress, Cairns, 2007 – see Quaternary International 167– 168 (Supplement), 2007, pp. 451–52. 63 Gerald Nanson, keynote address (co-authored with J Maroulis and DM Price) ‘Climate and flow-regime changes during the last 300 ka in monsoonal and arid northern and central Australia’. Quaternary Deserts and Climate Change, IGCP 349 Conference, 30 June–3 July 1997, University 18 0

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of Wollongong. The idea of a long-term trend towards greater aridity, beginning with the ‘mid-Brunhes climatic event’ about 300 000 years ago, appears to have wide support amongst Australian Quaternarists from a range of disciplines. Nanson and his colleagues (for example, Nanson GC et al. (1992)) base their interpretations on the fluvial histories of rivers in the arid zone and elsewhere in south-eastern Australia. Palynologists (for example, Kershaw AP et al. (2003); Martin HA (2006) Cenozoic climatic change and the development of the arid vegetation in Australia. Journal of Arid Environments 66, 533–63; Longmore ME and Heijnis H (1999) Aridity in Australia: Pleistocene records of palaeohydrological and palaeoecological change from the Perched Lake sediments of Fraser Island, Queensland, Australia. Quaternary International 57/58, 35–47) note corresponding trends in palaeo-vegetation records, especially a shift towards drier conditions and more open vegetation after this time. Hesse PP et al. (2005) notes an increase in continental dust flux, recorded in Tasman Sea sediments and a decrease in the size of ‘megalakes’ in northern Australia (e.g. Lake Gregory and Lake Woods). 64 Hesse PP et al. (2005) p. 64. 65 Australian megafauna, say Wroe et al. (2004) (Megafaunal extinction in the Late Quaternary and the global overkill hypothesis. Alcheringa 28, 291–331) ‘were subjected to increasingly severe and frequent episodes of regional aridity over the last three glacial cycles, with the last being the driest.’ Other studies confirm that these climate cycles affected megafauna populations, and the range of individual species, but disagree as to whether this involved long-term changes (and loss of species) or whether populations recovered locally when climate ameliorated – see Prideaux et al. (2007) An arid-adapted Middle Pleistocene vertebrate fauna from south-central Australia. Nature 445, 422–25; and (2007a) Mammalian responses to Pleistocene climate change in southeastern Australia. Geology 35, 33–36. (Naracoorte and Nullarbor Plain); Hocknull SA et al. (2007) Responses of Quaternary rainforest vertebrates to climate change in Australia. Earth and Planetary Science Letters 264, 317–31 (Mt Etna region); Price GJ and Webb GE (2006) Late Pleistocene sedimentology, taphonomy and megafauna extinction on the Darling Downs, southeastern Queensland. Australian Journal of Earth Sciences 53, 947–70 (Darling Downs). 66 Miller GH et al. (1999) p. 207. 67 Smith MA et al. (1994). Wood Point was originally found by Dominic Williams, by then a post-doctoral Fellow at the Australian National University. He recorded it as a locality where fossilised marine shell, emu and Genyornis eggshell and stone artefacts were all exposed in a blowout in an old dune. Some years later Giff Miller relocated the site. As an archaeolo18 1

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gist, I was brought in to see if I could determine whether or not the eggshell and the artefacts were in primary association. Looking back, that small dig assumes an importance that it did not have at the time: it is the only stratigraphic excavation to have established the configuration of a deposit of Genyornis eggshell in any detail; it is one of the few sites were a multi-method dating strategy has been deployed; it is unusual in that burnt eggshell is present; and it is the youngest deposit of Genyornis eggshell found so far, at 43 000 to 46 000 years ago it dates 5–10 thousand years after people had dispersed across the Australian landmass. 68 These are soil horizons marked by an accumulation of calcium carbonate nodules. They are often the only internal stratigraphic markers within a sequence of aeolian sediments and so are invaluable in determining the relative position of features in different parts of the dune. The emu eggshell was stratified beneath the lower of the two palaeosols but was a little higher in the dune sequence than the Genyornis horizon and therefore probably a little younger. My field estimate of the relative ages of the Genyornis and emu eggshell was later borne out by the 14C dates. 69 In a decade of fieldwork, Miller found burnt eggshell at less than 70 of the 1600 sites he recorded. The actual figures for collections containing some burnt eggshell are 62 out of 973 (emu) and 5 out of 635 (Genyornis) – Miller GH (2007) Tracking late Quaternary environmental and climate histories using C, O and N isotopes preserved in avian eggshells, and contrasting megafaunal extinctions in Madagascar and Australia. 27 September 2007, Selwyn Symposium. 70 Dromaius – total weight of eggshell 464.6 g, 24.3% burnt. Genyornis – total weight of eggshell 235.6 g, 7.6% burnt. 71 For the radiocarbon chronology see Bird MI et al. (2003) Radiocarbon dating of organic- and carbonate-carbon in Genyornis and Dromaius eggshell using stepped combustion and stepped acidification. Quaternary Science Reviews 22, 1805–12. For the OSL age of the dune sediments, see Roberts RG et al. (2001). The OSL age of the dune sediments directly beneath the Genyornis horizon is 55±5 ka. TIMS U-Th dating on the eggshell was attempted but unsuccessful given the very low U content of the eggshell. AAR dates on excavated Genyornis eggshell give an age of 47+/-5 ka. ABOX 14C ages for Genyornis eggshell are 43.0–45.6 cal kyr BP (pooled age, combining oldest carbonate and organic fractions). For Dromaius these ages are 41.1–43.2 cal kyr BP. 72 In Spencer Gulf, local sea level was +1 m 125 000 years ago (see Hails JR, Belperio AP and Gostin VA (1984) Quaternary sea levels, northern Spencer Gulf, Australia. Marine Geology 61, 373–89). At –30m the north-

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ern sector of the Gulf becomes dry land. However, during Stage 3 (50 000 years ago) there appears to have still been shallow water (~2 m deep) in northern Spencer Gulf, see Cann JH, Belperio AP and MurrayWallace CV (2000) Late Quaternary palaeosealevels and palaeoenvironments inferred from Foraminifera, northern Spencer Gulf, South Australia. The Journal of Foraminiferal Research 30, 29–53. 73 Johnson C (2006) p. 114. 74 Estimates for Genyornis are from Murray P and Rich PV (2004) and Williams DLG (1981). Data for Dromaius is from Marchant S and Higgins PJ (1990) (Eds.), Handbook of Australian, New Zealand and Antarctic Birds. Volume 1 – Ratites to Ducks. Oxford University Press: Melbourne; and Noble JC (1991).

18 3

R AINBIRDS:

ORGANISING THE

COUNTRY Deborah Bird Rose

Connections ‘When the brolga sings out, the catfish start to move,’ said one of my Aboriginal teachers, Daly Pulkara. We were a small group, sitting together on the bank of the Wickham River in the north-west corner of the Northern Territory, chatting over our midday dinner of fish and damper. Daly had heard the call of the brolga, and he offered this little story of connectivity. He was speaking about one of the three important rainbirds in the Victoria River country: along with the brolga

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or native companion (Grus rubicunda), there is the eastern koel (Eudynamus orientalis), and the channel-billed cuckoo (Scythrops novaehollandiae). They arrive around the time of the rains, making noise, bringing the rain behind them, calling up rain, singing up rain, bringing life to a very dry country. Rainbirds are key actors, calling out for the rising and falling of rivers, influx and outflow of water, the changing and returning of relationships between sky and earth, water and land. ‘You didn’t know that?’ he asked me. ‘That’s really culture, that one.’ My long-term research has been with Aboriginal people who live in the monsoon savanna country, inland along the Victoria River around the junction of the Wickham and Victoria Rivers, and on upstream along both rivers. As an anthropologist I began by being particularly attuned to issues of identity, and my research with Aboriginal people in Australia had required me to become especially engrossed in kinship. Australian Aboriginal people have one of the most complex kinship systems on earth, and theirs are truly kin-based societies. All the major aspects of human life are organised through kinship – governance, justice, economy, significant life events, and philosophy. Their system of kinship confounds Western expectations along many parameters, one of which is that it does not set up a hierarchically separated dualism between human culture and the rest of the natural world. Rather, most living things are included within the kinship system, and because kinship is a domain of ethics, most living things are included within an ethical system. 18 6

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Kinship relations between humans and other living beings are organised into patterns of cross-species kin groups that were created by the great creative beings known as Dreamings. Thus, for example, the Dreaming brolga is ancestral to contemporary brolga people and brolgas. Daly’s wife Mirmir was a brolga person; her father’s Dreaming was brolga and so was hers.1 Brolgas are her kin, her countrymen. Daly was a catfish person through another type of kinship connection: his mother was a catfish person and so was he. There was a connection between catfish and brolgas in contemporary life, as his story told us, and there was a connection between catfish and brolga in their marriage as well. Patterns replicate across human and other-than-human systems. I came to learn that issues of who anyone is (identity) are also issues of time. ‘When’ one is, so to speak, is part of who one is. Take the catfish and brolga story. When the brolgas first sing out they are coming into this country along with the early rains. They are singing out rain, and their song calls up catfish. The catfish start to move because the rivers are warming up and the water is beginning to run again after having been confined to waterholes during the long dry season. The ‘when’ of their connected identity is late dry – early wet. As I will discuss below, variability and patchiness are integral to climate, and Aboriginal people’s knowledge is attuned to the system itself, rather than to an external frame such as a calendar, as the brolga-catfish story makes clear. Along with issues of time, I also came to learn that identity entails issues of where one is. We were sitting on the bank of 18 7

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the Wickham River when we had that conversation about the brolga and the catfish. The Wickham is a tributary of the Victoria River, and the Victoria is one of the great tropical rivers of north Australia. From its headwaters on the fringes of the Tanami desert to the salty mudflats of the Bonaparte Gulf, the Victoria catchment encompasses a range of bioregions and micro-habitats. From an Aboriginal perspective, the Victoria catchment encompasses four major zones or bioregions. Starting from the inland, there is the dry desert zone, and then there is the golden savanna zone with its jungly fresh-water rivers, known as ‘big river’ country. Further toward the sea there is greener country, saltwater (estuarine) country, where salty water comes upstream and saltwater creatures also live. And then there is the sandy, muddy, salty country abutting the ocean. Bioregions are a critical factor in people’s identity. A person would say, for example, I’m a freshwater person, or I’m a saltwater person. The identity says a lot about the person: the kind of knowledge they’re entitled to and, usually, are most competent in, the foods that grew them, the numerous skills that enable them to live their lives. Some plants and animals are restricted to a given bioregion and thus become indicator species, signalling home in a profound sense. Big Mick Kangkinang, a man I worked closely with, explained that everything comes from Dreaming, a statement that includes the idea that everything is emplaced by creation. He himself was a saltwater man who had been living

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for years in big river country. He was close to 90 and told me once that he had been away for years and years and had had enough. ‘I’m really old people now,’ he said. ‘I want to go home to kagawuli’ (Vigna lanceolata, yam).

Life in country All these issues – who, when, where – come together in the Aboriginal concept of country. Country is a spatial unit – large enough to support a group of people, small enough to be intimately known in every detail, and home to the living things whose lives come and go in that place. The origins of country are in creation. The Australian continent is criss-crossed with the tracks of the creator beings, called Dreamings in Aboriginal English. Walking, slithering, crawling, flying, swimming, chasing, hunting, weeping, dying, giving birth, Dreamings were performing rituals, distributing plants and marking the zones of animal and plant distributions, making the landforms and water, and making the relationships between one place and another, one species and another. They shifted their shape from animal to human and back to animal again, and as they acted they were becoming ancestral to life on Earth. Dreaming creation gives us a world in which most living beings are sentient agents, and sentience takes us into a world of animism. Graham Harvey has recently brought fresh insight to this term, rehabilitating it from the old anthropological evolutionary view that animism was a primitive and erroneous form of understanding the world. Harvey’s definition is succinct:

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animism involves the recognition ‘that the world is full of persons, only some of whom are human, and that life is always lived in relationship to others’.2 Indigenous Australian thought seeks out pattern, difference, relationship and connection among persons. A person in this context is both autonomous and connected, enmeshed in relations of interdependence, always bearing responsibilities for others, and always the beneficiary of the actions of others. Persons, whether human or not, act according to their own culture. They have their own ways of doing things, their own languages and their own ceremonies: ‘birds got ceremony of their own – brolga, turkey, crow, hawk, white and black cockatoo – all got ceremony, women’s side, men’s side, kids too, everything.’ Plants as well as animals are sentient, and, as Daly and others told me, the Earth itself has culture within it. My research took a turn toward ecological knowledge for two reasons. My early and ongoing interest in philosophy and questions of the meaning of life took me directly into efforts to understand Dreaming, and Dreaming is all about the living world. It does not refer to some other world, but is fully engaged with the patterns and connections of this world. Daly’s brolga/ catfish story is a good example. The link between brolga and catfish is given in creation, exists in everyday life, and indicates both a temporal pattern and a spatial connection. As I began to realise how little I understood of the living world in north Australia, I began learning the names of plants and animals, learning about habitats and communities. I soon

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realised that old people’s knowledge included a wealth of detail about plants, animals and habitats that were no longer readily accessible, and that might even be locally (or entirely) extinct. In 1986, at the height of the BTEC program (Brucellosis-Tuberculosis Eradication Campaign) when the country was nearly cleared of cattle, I saw flowering plants that I had never seen before. I saw billabongs with lilies that hadn’t been there before. I saw country that was much closer to people’s descriptions than it had been previously in my experience. I got grants that enabled me to employ a botanist, and I started documenting the plant world as it is known and identified by my teachers. The work kept expanding, both because Aboriginal people don’t think in terms of academic disciplines and, more importantly, because for them the process of naming the world is far less interesting than talking about how the world works, how things fit together. So the name of the plant, and the standard kind of information – habitat, flowering, for example, was kid’s stuff to them; they were happy enough to share it, but what really interested them was a whole set of other questions. They talked about what uses they put the plant to, and how it figures in kinship, and song and ceremony. They talked about the plant’s own kinship with other plants, and also about all the other creatures who also have an interest in the plant and who benefit from that plant’s existence. And they talked about the plant’s communicative agency: what, if anything, does this plant tell? In short, they were interested in systems, not labels.

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From this extended work I am developing an account of a philosophical ecology that works with several major principles: pattern, connectivity, patchiness and flux. Patterns are broad characteristics that are relatively unchanging. Bioregions are a good example. Connectivities are relationships within and across broad patterns. Patchiness identifies the fact that while broad patterns are identifiable they do not define internal homogeneity. Thus, for example, the wet season is broadly characterised by rain, but there are many sunny days with no rain, and some times rain comes in the dry season.3 Flux indicates processes situated in time and place. Whatever the broad characteristics, the actual flow of life is full of variability, uncertainty, patchiness. In real life what holds patterns and connections stable enough to be identifiable are the life processes that are in flux: the coming and going of rain, sun, wind, cold, dry; the arrival of rainbirds, the stirring of catfish, the flowering of trees and shrubs. Subtending it all is communication.

‘Tellers’ This monsoon region experiences an annual boom and bust, from hot, desiccated country in the late dry season, to lush green country during the wet season. The monsoon terminology of wet and dry suggests a certain predictability, and that suggestion is accurate up to a point, but meteorological records from Victoria River Downs Station over many decades show a lot of variability. Station record books provide day by day information on some years when rain basically never came in the wet season, years when the dry season was miserably cold and 19 2

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wet, times of massive flood and times of incredible absence of water. In short, even with the predictability of the monsoon, the classic Australian uncertainty prevails in this country. One of the big questions for any being who learns to live with these extremes is the question of time: where in the seasonal round are we at any given moment? How do we know what is going on? Some of my teachers used the term ‘tellers’ to identify creatures whose actions tell of what is going on in the world. Rainbirds tell about rain, and they do so on their own and in connection with others, as in Daly’s story of brolga and catfish. Let us pause for a moment to consider the catfish. They slow down in the cold dry season when their food becomes scarce. Catfish also eat mud. When you cut open a catfish and see mud in its guts, you know cold weather is coming because they have started eating dirt along the side of the river. Another teacher explained: Even the catfish, when we get it from the river, when we open im up, we can tell from the guts there. Dirt there, when they eat dirt, that’s the cold weather coming up. They tell you earlier [in advance] … Because winter time they got no choice, they have to show up and come in to the side. Because it’s hard to find their feed in the winter time. The only feed they can have is that mud.

Catfish tell about cold weather coming. Brolgas tell that the rains are coming, and by that time the cold is finished and the heat has kicked in. The early rains bring water that runs 193

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hot off the desiccated land, and the rivers start to flow. The first flush will be so hot that many fish will flounder and some will die. Provided the rains keep coming regularly enough, the temperature will stabilise, the rivers will come up and keep on flowing, and later the big rains will come and the floodwaters will rise, bringing the river out across the land. Arriving from far away, rainbirds bring news of what is happening elsewhere, and what is going to happen here. Tellers extend human knowledge into the knowledge systems of other creatures, and they offer a degree of local certainty in what otherwise appears to be a very uncertain climatic regime. Some of them, like some of the rainbirds, are linked into systems that extend beyond Australia. Most of the indicators, though, are extremely local, and tell at a very fine scale what is going on. One type depends on concurrence. When the march flies start biting, the crocodiles are laying their eggs. When the seed pods of the bauhinia tree (Lysiphyllum cunninghamii) turn dark red the really hot weather is here, thus signalling the final stages of the dry season. Other tellers, like the rainbird, are sequential; they tell what is going to happen. People are quite explicit in saying that other creatures know things that we don’t know because they inhabit regions that we do not inhabit. Another teacher explained with reference to a bird that is probably a swift (species unidentified): And when the wet season begins he flies low, but when the wet season is over he flies really high now. That’s the 19 4

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time for him, he’s feeling that air on top for that winter, you know. When that winter’s coming, he goes back low again and starts to make that camp in the side [of the cliff] there.

One of the questions I investigated was the extent to which this knowledge remains constant. I did interviews that took me across bioregion boundaries, and across a number of different languages within one bioregion. Did the same system of tellers exist, and was the information the same? In summary form, the system as a system is extensive, and there is a lot of patchiness; many of the same signs exist but often they index different events. So, for example, at Yarralin in the big river country the march flies tell that that the crocodiles are laying their eggs. Slightly further south but still in big river country, a tree called Jangarla white dragon tree (Sesbania formosa) tells that the crocodiles are laying their eggs. Back at Yarralin, this same tree, when it flowers, tells that the barramundi are biting. In Yarralin the bauhinia tree tells that the really hot weather is here, while to the south-east in desert country the same tree tells about hot weather, but it tells in a different way. Being human persons, it is easy to see how tellers communicate and humans take notice. But it is not only humans who pay attention. In the Victoria River country, the emu is associated with a little finch.4 Finches stay close to water and their alert and nervy action tells of other presences. When emus comes in for water, they stop to listen, and if the little birds are not calling out, they know they should stay away because there 195

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are people there. When the emu drinks, the little finches warn if somebody is coming. As I have shown, many tellers are animals. In general, plants offer highly localised information, whilst birds’ information is more consistent across much larger areas. Again, rainbirds are a good example. The common koel, for example, is known as a rainbird or storm bird in north Australia. Most of them come flying in from Papua New Guinea or Indonesia, calling out the news of rain. They travel across a lot of Australia and in southeast Australia, too, they are rainbirds and stormbirds. 5 In sum, the fact that this system is widespread ensures that people know that there is a system here, and when they go beyond the bounds of their knowledge, they still know that they are in the presence of a system, they just don’t understand it. Both widespread patterns and local patchiness are important within the communication system, and from a human point of view both are important to a knowledge system: the widespread consistencies ensure the recognition of systems; the highly localised specifics ensure that knowledge is country-based, and finely attuned to local events.

Organisation Bioregions, like seasons and like communication, are both broadly consistent and internally patchy. For example, one day we were up on top of a mesa, and I asked if there was a term for this country. I was told: ‘Kaja. It’s kaja country.’ This surprised me because earlier I had learned the term kaja as the

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term for desert, and we were in big river country. It seemed that the organisation of landforms included both broad regions and internal patchiness. I became fascinated, then, by two interrelated issues concerning regions. The first was how you would make such distinctions: what is it that characterises both the mesa-top and the desert? The answer in this region is surface water – an area that is significantly without surface water is kaja. The second was: how would you know, if you were walking along, when you had left the big river country and entered kaja country? What, if anything, would tell you that you were entering a region, not just a patch? Are there any sign posts to say ‘you are now entering kaja country, the bioregion’? The answers here combine plant and animal indicators. So for example, if you were walking east-south-east into kaja country, along the Muranji track, for example, the most visible indicator would be a species of brachychiton (B. paradoxa). It became clear that bioregions line up relatively well with the isohyets. In fact, however, local knowledge is more detailed and more interesting than diagrams of isohyets precisely because it works with relevant local indicators rather than with highly generalised information. On a mapped diagram, a line is drawn that marks the point between 600 and 800 mm. The locally relevant boundary is a fair bit north of this line; a small coastal range intercepts the rain and makes a more interesting pattern than the averages as extrapolated between measuring points and the differences measured by the decimal system.

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The main indicator species is kagawuli (Vigna lanceolata), the yam Big Mick Kangkinang wanted to return to. It grows in the saltwater country, but not in the big river country. Its distribution is almost certainly limited by rainfall, and it is very precise. A diagram shows averages, and uses a system that is abstracted and disembedded. Living things survive by adapting to the local rather than the abstract, and by adapting to the worst conditions, not the average. Old Jimmy Manngaiyarri was one of my most fascinating and knowledgeable teachers. He was notorious for having tirelessly walked everywhere, long and far, north and south. His knowledge was extremely broad, and at the age of 90 or so when I interviewed him in depth on bioregions he still possessed an excellent memory. He was a big river man, and in spite of his wide travels, he expressed distrust: When you go some place, well some place got a different tucker. You know, man might eat some poison one. Well, we can’t do nothing. We got to eat the right feed that we bin raised up on, that grew us up. Well, we got to go on that same tucker. Another end is using different tucker. They got different different tucker … That’s why we got to go with the right feed for us. We might be go somewhere, I might see some tucker there, I don’t know. Well I’m frightened to eat there. Very bad, you might eat some sort of tucker, poison one tucker. You see, that’s why that tucker what you were raised up on, that tucker, you gotta stick with that tucker. You get the picture? 19 8

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An important part of the organisation of bioregions is that things don’t go and get mixed up. The indicators hold good. The organisation of bioregions is the product of Dreaming action. Old Jimmy spoke of a little lizard called latatj which, I think, is Storr’s monitor (Varanus storri ocreatus).6 According to Old Jimmy, Latatj was responsible for organising the plant and other communities that go to make up this region. I asked him once about a plant I was hoping to be able to identify called tipil, wondering if it might also have grown in some remote corner of his country. He rejected the idea that I would find tipil in his country: No, nothing. That latatj didn’t want that kind of tucker. All kinds of tucker, he pushed them back, right back that way, on the sea side. He must have a good fish, good crocodile, good sugarbag [native honey], everything, good way … He’s boss for every tucker there. Whatever he don’t want, he push it away: ‘You take it back’, he said. ‘I don’t want that one.’ He don’t want it. Latatj don’t want it: ‘Keep it out on that side on the sea side.’

When latatj was saying ‘you take it back,’ he was talking to one of the rainbirds – the channel-billed cuckoo.7 The rainbird assisted latatj; it took back into the saltwater country those species that latatj did not want in the big river country: Another end (saltwater side), is using different tucker from us. They got different different tucker. That’s what 19 9

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that rainbird pushed out. It took it back. Rainbird took it back. Latatj don’t want it. Latatj don’t want it, Rainbird took it back, right back. That’s the way. Make it good, you know.

The lizard’s range is a region that roughly corresponds to the family of languages in the big river bioregion of the Victoria River catchment, and the rainbird, of course comes with the rain, announces the rain, and in some sense can be thought to bring rain. So in this organisation of bioregions we see a kind of tussle in which plants and animals from the saltwater side come creeping into the big river country, and get pushed back. The key figures in sustaining the organisation embody those familiar boom and bust forces: dry versus wet, earth versus sky. Latatj lives in dry stony areas, areas with dead trees and spinifex. The little lizards embody earth and dry. The channel-billed cuckoo comes soaring in from Indonesia and Papua New Guinea, flying along, calling out, announcing rain. The combined actions of latatj and cuckoo work with the monsoon booms and busts: on the one hand, the dry spinifex country, the hot stony ground, the dead trees, and on the other hand, the travels, the sky, the rain. The rainbird brings the necessary rain, but it is the dry earth that sets the limits.

Climate change There are big questions about how this system is going to fare with climate change, particularly with the apparently increasing

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amounts of rain in this region. Thus far the increased rain does not seem directly to be generating disorganisation in the information system, but it is hard to tell because other factors are already generating huge amounts of disorganisation. There is a process of intensification of vegetation, referred to in this region as ‘thickening’. In other regions it is called woody weed invasion; radically disturbed land is taken over by opportunistic native and introduced scrub (mainly Acacia spp.). In the Vic River District, the woody weed invasion is accompanied by an increase in the number of trees. The savanna is rapidly losing its ‘clean, clear’ character.8 Current scientific studies are documenting the changes, but the causes are not yet fully understood by anyone. It is probable that numerous factors are involved, including global warming, suppression of indigenous fire regimes, intensification of land use, and new breeds of cattle, among others. The increasing rainfall may well be implicated, and if it is, how ironic that both the rainbird and the polluted air that is causing the increased rainfall both come from overseas. To top off the list of changes, along with thickening, there is also loss – of topsoil, riverbanks, and, ultimately, of many of the species and habitats that supported Indigenous life. Daly’s son-in-law, Riley Young Winpilin, had a set of explanations, and while a lot of the people listening to him laughed at his choice of words and may (I think) have been sceptical of his effort to find an explanation for things that were looking pretty inexplicable, they did not fault the underlying logic:

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Used to be this country bin all clean, and today, round about two years or a year, you can see everything, different plants growing. And why? We got too different cattle la [here in] this country. Brahman, Santa Gertrude, all kinds of breeds. All kinds of horse breeds. And country was change … You can see everything all changed. And you can see too many white man. Too different white man. Too different … some of them good eye, some of them small eye, some of them big eye [laughter]. That kind make you wrong.

Riley contrasted the state of the country before the 1970s with its current state: And when I bin go long this land, land was really good. He was really good … But now … when we bin start again, country was little bit funny that day, I bin looking at country was little bit funny that day. I bin looking at, ‘what’s wrong this one? Something wrong.’ And after that I bin look now, one year’s time I bin see em plants bin get up. You go longa bush now looking for fruit, you can’t see em fruit. You see em all these trees now … And even if you go round la bush here, you can’t see that karil, gooseberry, kilipi, tipil, purlkal, ngaringari, that kind bin too much longa this – yarkalayin, mintarayij, that bin already bin clear. But you can’t see em this time now. That’s from what I bin say: ‘country bin change. Ground bin change.’

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Because no fruit now. Where him gottem good fruit longa this ground, country was look good. But fruit going away, country gone, finish.9

Riley spoke of a process that I would gloss as intensification. The social aspect of intensification concerns land use and social services. Tourism is increasing rapidly since the conversion of a few stations into national parks (Gregory National Park, Keep River National Park). In addition, there is a growing scientific presence, and investigations and activities by people from numerous branches of government. As we talked about ecological change in 2001, we were surrounded by Army personnel who were building houses and roads as part of a government effort to standardise living conditions in the outback. As I write in 2007–2008, there is another wave of Army interventions. What is being lost? Communication, patterns, connectivities, patches and flux are all under threat. The implications are huge. For humans to live anywhere they have to be able to plan. To plan well, they have to understand how ecosystems work. To understand systems in action there has to be communication, and living beings need to pay close attention. In order for there to be communication and attention, biodiversity is crucial. The indicators I’ve discussed are not just rules of thumb; they are a body of knowledge about patterns and connections. They indicate a further point: it takes the interactions of a multitude of living beings in local ecosystems and

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across wider global systems to know what is going on in the world. Without communicative complexity, as Riley said, country is gone, it’s finished.

Endnotes 1

Mirmir adopted me as her sister. Her other main Dreaming is flying fox – another creature that is associated with rain. Her home country is in the desert – a place where rain really matters. She is a rain woman, a desert woman, a rainbird women, a flying fox woman. I dedicate this paper to her and to her oldest daughter Aileen.

2

Harvey G (2006) Animism: Respecting the Living World. Columbia University Press: New York, p. xi.

3

Rose D (2005) Rhythms, patterns, connectivities: indigenous concepts of seasons and change, Victoria River district, NT. In A Change in the Weather: Climate and Culture in Australia. (Eds. T Sherratt, T Griffiths, and L Robin) pp. 32–41. National Museum of Australia: Canberra.

4

One of the most common finches is the zebra finch (Taeniopygia guttata), but all the finches in the region are referred to by the same term nini.

5

http://www.austmus.gov.au/factsheets/common_koel.htm.

6

I was unable to make a positive identification of this species; it could perhaps be V. acanthurus.

7

This identification was made in the absence of a sighting or specimen, but I feel quite confident. The description of the channel billed cuckoo, and its identification as a rainbird, make it unlikely to be confused with any other bird.

8

Lewis D (2002) Slower Than the Eye Can See: Environmental Change in North Australia’s Cattle Lands. A Case Study from the Victoria River District of the Northern Territory. Co-operative Research Centre for the Sustainable Development of Tropical Savannas: Darwin.

9

The plants Riley discusses are: Karil: Cucumis spp., probably trigonus; Gooseberry: Physalis minima; Kilipi: Leichardtia australis; Tipil: unidentified (unable to locate a specimen); Purlkal: Vitex acuminata; Ngaringarin: Pterocaulon serrulatum; Yarkalayin: unidentified water plant (unable to locate a specimen); Mintariyij: Nymphaea violacea.

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WOODSWALLOWS: A LONGER TERM, EVOLUTIONARY VIEW OF BOOM AND BUST Leo Joseph

Introduction A swarm of budgerigars (Melopsittacus undulatus) alights on branches overhanging a temporary desert waterhole. There are so many that some birds fall in and drown. Black-tailed native-hens (Tribonyx ventralis) appear in southern Australia in summer, and for a few months people see them scurrying to shelter at the edge of a lagoon. Letter-winged kites (Elanus

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scriptus) appear in coastal areas looking thin and hungry after a population explosion in the interior in response to a sudden increase of their favoured food item, a native rodent, the longhaired rat (Rattus villosissimus). These are manifestations of the boom and bust theme of this book. They are very much examples of how we in the hereand-now see the phenomenon. The aim of this chapter is to remind the reader that long-term evolutionary and ecological processes underpin the boom and bust phenomenon. They explain why it is that we see irruptive flocks of budgerigars but never of secretive understorey species such as striated grasswrens (Amytornis striatus), or why it is that coastal, urban birdwatchers encounter the pathetic and perhaps confronting sight of a starving bird of prey away from its normal interior, arid zone habitats. The basic idea of this chapter is that the biology of the species, populations and individuals that we see today is the result of a moulding by evolution in response to long-term environmental changes. That process of evolution leaves a footprint in the genetic make-up of a species, its DNA. One task of some of evolutionary biology’s many branches, specifically phylogeny and phylogeography, is to learn how to read that footprint and interpret the longer term history of a given species. Then we will be in a better position to build a full evolutionary picture of a species by integrating that knowledge with what is learned from study of the ecology and natural history of a species in its natural environment. In this chapter, I explore this by looking at one group of birds, the

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woodswallows. I focus on the white-browed woodswallow (Artamus superciliosus) and masked woodswallow (A. personatus). As we will see, they have a boom and bust story to tell that is both unique in some aspects but also shared with some other widespread Australian birds of the arid and semi-arid zones.

What is a woodswallow? If I were to say that woodswallows are corvoid oscines mostly endemic to the Australo-Papuan region and that within that group they are among the core Corvoidea … and that they have powder downs, I would get a well-deserved look of puzzlement accompanied by the typical Australian expression of incredulity: ‘Come off it, sport!’ So let’s approach the question of what a woodswallow is in another way. Most will agree that gum trees and kangaroos are among the more unique evolutionary products of this continent. What many are still surprised to learn is that until 1985, many of Australia’s birds were thought to be closely related to ecologically similar counterparts in the northern hemisphere. Thus the treecreepers of the genera Climacteris and Cormobates that one sees foraging on tree trunks in Australia and New Guinea were for many years thought to be closely related to the northern hemisphere’s creepers, such as the brown creeper (Certhia americana) of North America. The names of other Australian birds, such as robins, cuckooshrike and the familiar term ‘wren’ also indicated this. In 1985, however, North American ornithologist Charles Sibley

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and his long-time collaborator Jon Ahlquist published a paper that overturned this notion for the Australian representatives of the world’s largest taxonomic Order of birds, the Passeriformes, or the passerines.1 Passerines are often described in Australia as bush birds. They include the lyrebirds, bowerbirds and birds-of-paradise, as well as many other usually small to medium-sized birds one sees in trees or undergrowth such as (and notice their often northern hemisphere-centric names) magpies, fairy-wrens, magpie-larks, cuckoo-shrikes, robins and honeyeaters, which include familiar species like the noisy miner (Manorina melanocephala). To be sure, though, some Australian passerines are very Australian in name and attitude, and currawongs come to mind here. There are so many passerines that it is convenient to refer to a bird as being passerine or non-passerine. Familiar nonpasserines are parrots, kingfishers, pigeons, waterbirds and marine groups such as penguins, albatrosses and petrels. Woodswallows are passerines. From here on, my description of woodswallows is based on work of a number of colleagues who have built on the pioneering research of Sibley and Ahlquist.2 Passerines are divided into three groups. At the base of their evolutionary tree, or phylogeny as it is more strictly known, are the first of these, the peculiar New Zealand wrens. The most familiar of these to most ornithologists is probably the rifleman (Acanthisitta chloris). The New Zealand wrens are understood at present to be the closest living

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relative of all other passerines. That is, they are the sister group of all other passerines. The date of the divergence of the lineage that led to the New Zealand wrens on one hand and the sister lineage that led to all other passerines on the other is not certain but has been estimated at 80 million years ago (mya), well before the K-T (Cretaceous-Tertiary) boundary at 65 mya. For the purpose of this chapter, the exact date or even range of dates is not as important as the point that what we see today as woodswallows have been evolving with environmental changes on short- and long-term time scales for a very long time. The remaining two groups of passerines are the oscines and suboscines. The differences between the two are primarily anatomical and the details again need not concern us here. More relevant is that the vast majority of suboscines occur in the Neotropics, with smaller numbers in the IndoPacific and South East Asia. In Australia, the only suboscines are the pittas. Woodswallows are oscine passerines. The divergence between oscines and suboscines has been estimated to have occurred around 77 mya. But again uncertainty in that date need not obscure the point that these lineages have evolved with many environmental changes over a long time. These include shifting positions of continents and increasing aridity of the Australian continent. Oscine passerines are divided into two groups, the corvoid (crow-like) oscines, including woodswallows, and the

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passeridans. Indeed, the corvoid oscines are where passerines of the Australo-Papuan region start to come into their own. Almost all passerines in the region today are members of the corvoid oscines. To appreciate their uniqueness to this region, and also that of woodswallows, a brief discussion of the passeridans will bring some perspective. Passeridans appear to have evolved from a divergence within the corvoid oscines around 45 mya. This divergence gave rise on one hand to a small branch of the corvoid oscines thought currently to have led to the Australo-Papuan robins (though some doubt exists).3 The other branch evidently dispersed out of the southern hemisphere. Details of how that dispersal actually happened in relation to how landmasses were distributed at the time are still being investigated. However it happened, it was a branch of that corvoid oscine ancestral stock that diversified into the enormous radiation of many hundreds of species now known as the passeridans, or Passerida. They comprise almost all the oscine passerine birds of the northern hemisphere. A few appear to have secondarily reentered the Australo-Papuan region. Among these are true swallows and martins, reed-warblers, and white-eyes (the familiar silvereye Zosterops lateralis, for example). All remaining oscines are corvoid oscines and they are almost unique to the southern hemisphere and, in particular, Australo-Papua. Just a few groups, such as the crows and jays of the northern hemisphere, occur outside it. Woodswallows are corvoid oscines.

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Within corvoid oscines there are two groups. One is the old Australo-Papuan oscines, which includes lyrebirds, bowerbirds, logrunners, fairy-wrens, honeyeaters and, impressively enough, the treecreepers mentioned earlier. The other group, which is where woodswallows fall, is known as the core Corvoidea. Among them are monarch flycatchers and the crow-like passerines such as the crows and ravens themselves, birds-ofparadise, and, at last woodswallows. The core Corvoidea is thought to have radiated some 50 mya. Woodswallows belong to a taxonomic Family of core Corvoidean oscine passerines, the Artamidae, which includes magpies and currawongs, butcherbirds, two New Guinean birds of the genus Peltops, woodswallows and for now, at least as we currently understand things, an enigmatic red and black bird of Borneo, the Bornean bristlehead (Pityriasis gymnocephala). Within the artamids, woodswallows are a numerically small genus, Artamus, of about 10 species (allowing for some taxonomic uncertainty). They are almost unique among passerines in having feathers known as powder downs from which they can extract a ‘powder’ that results from decay of the feather. With this they dust their feathers, giving them a powder-like bloom. Six species occur in Australia and a few range from India in the west to the islands of the south-west Pacific in the east.

White-browed and masked woodswallows With a deep evolutionary perspective on woodswallows established, I now focus on our two species of interest and the

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special evolutionary perspective they bring to the boom and bust phenomenon. Of the six Australian woodswallows, the masked and white-browed woodswallows are remarkable in a number of ways.4 The adult males of the two species are very different. The masked male is a predominantly grey bird with a black facial mask and throat, and a white spot above and behind the facial mask. Male white-browed woodswallows in contrast are blue-grey above, mostly chestnut below but with a broader black mask than that seen in the masked woodswallow. They also have a prominent long white eyebrow, hence their name. Females of the two species are far more similar to each other, though certainly different, being predominantly grey birds with some rufous toning in their underparts, especially in the white-browed. The two species range over most of the continent but the masked woodswallow is the most common of the two in western parts. In eastern Australia, their occurrence in mixed flocks, often of several hundred individuals, is well-known to bird observers. These flocks are often heard high above before they are seen against the backdrop of a glaring sky where they hunt flying insects. Furthermore, they often nest in the same groups of trees if not occasionally in the same tree. Hybridisation, however, has been rarely reported either by direct observation of mixed pairs at nests or from possible hybrid specimens in museum collections. On balance the evidence that they hybridise is tenuous although some low frequency of hybridisation cannot be ruled out. Even more remarkably, in a survey of ecological

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differences among closely related pairs of bird species in south-eastern Australia, these were the only two that could not be separated at all.5 And finally, their vocalisations are also essentially indistinguishable.6 This almost certainly is a unique feature of this pair of birds. I know of no reference to any other pair of bird species as different in appearance as these two are for which it is so hard if not impossible to distinguish vocalisations. A third species, the dusky woodswallow (A. cyanopterus), is known for some regular seasonal movement but the masked and white-browed species show more of a boom and bust trend in their movements, being in vast numbers in some years in southern Australia and not so in other years. The ecological and environmental controls on their movements are still poorly understood. Through an interest in the evolution of migration, I was drawn to study of these birds using DNA sequence-based methods to understand their evolutionary history.7

DNA and woodswallow evolution: the species level My research used DNA sequences from the mitochondria of cells, mitochondrial DNA (mtDNA). MtDNA is thus physically separate from the main part of the genome in the nucleus of the cell. It is further distinguished from the DNA of the nucleus by a number of properties, the details of which can be examined elsewhere.8 These properties make it a highly useful genetic marker for studying evolutionary problems that concern closely related species or groups of species.

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MtDNA sequences were obtained from all six species of Australian woodswallows to test whether masked and whitebrowed were each other’s closest relatives. It was not feasible to include representatives from outside Australia but this was not a major concern. On grounds of plumage and morphology, those woodswallows that occur outside Australia can reasonably be assumed to be more closely related to species other than the two of interest. What emerged first was that masked and whitebrowed woodswallows are indeed each other’s closest relatives, more or less as expected. The more interesting implication of their close relationship was that their capacity for long-distance movements across the continent need only be postulated to have evolved once in their common ancestor, not twice independently in the two species (though that cannot be totally discounted). Again, this is not in itself a very momentous conclusion. What was far more unexpected was that the mtDNA sequences from the two species did not fall into two discrete groups that showed any kind of neat, one-to-one correspondence with the identity of the birds from which they had come. In fact, the mtDNA of an individual of one species was often more closely related to the mtDNA from an individual of the other species than to one from its own species. This phenomenon is known as paraphyly, or in an extreme case such as this, polyphyly. Paraphyletic relationships of the mtDNA sequences in two distinct species can arise in a number of ways, including hybridisation between the species. The simplest explanation to accommodate the details of this particular

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case of paraphyly, however, is that masked and white-browed woodswallows diverged from their most recent common ancestor in very recent evolutionary terms. By this I mean well within the last 1.8 million years, the period known as the Pleistocene. A corollary of this is that their mtDNA gene pool would still have a lot in common with that of their most recent common ancestor. Their plumage is most likely to have been changed – evolved – much more rapidly through the evolutionary processes of natural and/or sexual selection. That they still share the same ecology and vocalisations as well as mtDNA gene pool suggests the merit of testing the role of sexual selection. This description of the evolution of white-browed and masked woodswallows sets the scene for the unique perspective they bring to the boom and bust phenomenon. Woodswallows, as we have seen, are members of a lineage of passerine birds, the Artamidae. That lineage also includes the magpies, butcherbirds and currawongs. It diverged from its nearest relatives sometime in the Tertiary. In the more recent evolutionary history of that lineage, the woodswallows themselves have diverged from their nearest relatives. But far more recently in the history of that lineage, masked and white-browed woodswallows have diverged from other woodswallows and evolved their way of living in the present-day Australian environment. During the course of their history, therefore, they have been subjected to evolutionary challenges such as aridification of the Australian continent and climatic changes that dwarf the inter-annual changes with which we are familiar

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and which we associate with the boom and bust phenomenon. The drying out of the Australian continent that began in the Miocene (from 24 to 5 mya). The glacial cycles that led to repeated cooling and warming during the Pleistocene, which lasted from 1.8 mya to 10 000 years ago, were also a phenomenon to which they were subjected. Most recently, in the Late Pleistocene, it may well have been these very same two species that we see today, not their more immediate ancestors, that survived the Last Glacial Maximum (LGM) of 18 000 years ago.9 Of course, all of the endemic fauna and flora of Australia are descended from lineages that have had to deal with these changes. The whole Australian biota, therefore, brings a deep evolutionary perspective to boom and bust and the woodswallows have so far been just one example. It is how species dealt with and were shaped by the cyclical swings of cooling and warming in the Pleistocene, especially that of the LGM (which was most severe), that brings us to a more unique perspective offered by these woodswallows on the boom and bust phenomenon. The LGM was a time of aridity that was windy and cold. Ornithologists have long felt that the geographic ranges of Australian birds would have been severely bottlenecked by the LGM.10 The mtDNA data from the masked and white-browed woodswallows show a genetic footprint of what appears to have been a response to the aridity of the LGM as well as to the warming that followed it; other similarly distributed wide-

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spread Australian birds of the arid and semi-arid zones have been studied with the same methods and show very similar results.11 We now examine what the mtDNA data had to say about that in the woodswallows.

DNA and woodswallow evolution: the population level When a population expands in size from being very small to very large, particular patterns of genetic diversity are expected to arise. The basis for this expectation lies in two principles. One is that mutations in DNA, which alter the sequence of base pairs or ‘letters’ in DNA molecules, are rare events that take place at random locations within a given piece of DNA. The other is the relationship between this rare process and the size of the population in which it occurs. Consider a small population and a particular gene, or piece of DNA, that we can sequence in each individual of that population. Consider also that most individuals in this small population share the same version of that gene, that is the exact same DNA sequence. Perhaps a few individuals have one or two rarer forms of the gene that differ from the main one by just a base pair or two. If the population undergoes a sudden increase in size, as happens in the boom phase of the boom and bust phenomenon, the frequency of mutation itself is not expected to increase. It will remain a rare event. But it now has a bigger pool of genes in which to occur because there are now many more individuals in the population due to the boom it has experienced. Recall also that mutation is random as well as rare, which means that

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it will not always occur at the same spot in the piece of DNA that we are considering. This sets up the expectations for how genetic diversity should appear in a population that has undergone a rapid expansion. First, we will expect most individuals in the newly increased population to have the same version of the gene that was most frequent when the population was of a smaller size. That is because mutation is and has remained a rare event. Second, we now expect that a scattering of individuals will differ from that main, most common form of the gene, and from each other, by one or two base pairs. That is because rare mutations strike randomly in different places within the piece of DNA under study. The more complete analysis of the woodswallow data suggests a twist to the boom and bust story. It is that evolutionary divergence from their common ancestor into what we now see as white-browed and masked woodswallows may have been so recent as to have been after the population expansion the molecular data suggest they have undergone. Therein lies our next challenge in this story: to piece together how they diverged when so much of their biology remains so similar.

A note on natural selection and boom and bust This chapter has focused on what we can learn from the ‘footprint’ provided by DNA sequences of the booms and busts a species has undergone in the course of its history as a species, and even earlier. In every minute of every day of its existence as a species, however, the individuals that make up its popula-

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tions are dealing with the environment, and are exposed to natural selection. So I wish to bring here a further long-term evolutionary perspective to the boom and bust phenomenon, this time from thinking about the relevance of a population explosion to the process of natural selection. Natural selection is often paraphrased as survival of the fittest, or as John Endler has pointed out, ‘nothing succeeds like excess’.12 These are appropriate ways of thinking about how evolution is at work in species that undergo periodic booms and busts such as the more obvious one with which I opened this chapter and, of course, the woodswallows on which I have focused. As is a basic prerequisite for natural selection, populations often exceed the number of individuals that the environment can support and this is very much the case with boom and bust species. The boom and bust strategy appears to work by producing as many individuals as possible in as short a space of time as possible during the boom times. Then natural selection has a bigger pool of individuals from which to select for the next generation. Each round of boom and bust must result in evolution of the population because many individuals and their genes will be eliminated when the bust sets in. If anything, the immediate excess of the boom sets up longer term survival of the fittest in the bust.

Conclusion White-browed and masked woodswallows and several other similarly distributed widespread species have genetic footprints

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that show they have undergone booms and busts in the last tens of thousands of years. The lineages from which they have evolved may have done so over the last tens of millions of years. That the evolutionary lineages to which they belong, if not the actual species we see today, have survived these upheavals might on the one hand be an encouraging signal of the native fauna’s resilience. On the other hand, it makes us wonder about the species that have not survived, as we surely know from fossil and sub-fossil data that many did not. That is a whole other story that centres on the debate as to whether human predation or climate change caused the demise of the extinct Pleistocene Australian megafauna.13 Regardless of how that debate plays out, it is sobering that humans now appear to be causing the next round of climate change. In terms of the debate over whether humans or climate change caused extinction of the Pleistocene megafauna, this is nothing if not an ironic double punch. It is also widely predicted to lead to another cycle of boom and bust in populations of the species with which we share the planet.

Endnotes 1

Sibley CG and Ahlquist JE (1985) The phylogeny and classification of the Australo-Papuan passerine birds. Emu 85, 1–14.

2

Barker FK, Barrowclough G and Groth J (2002) A phylogenetic hypothesis for passerine birds: taxonomic and biogeographic implications of an analysis of nuclear DNA sequence data. Proceedings of the Royal Society of London B 269, 295–308. Barker FK et al. (2004) Phylogeny and diversification of the largest avian radiation. Proceedings of the National Academy of Sciences of the USA 101, 11040–45; Ericson P et al. (2002) A Gondwanan

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origin of passerine birds supported by DNA sequences of the endemic New Zealand wrens. Proceedings of the Royal Society of London B 269, 235– 41; Christidis L and Schodde R (1991) Relationships of Australo-Papuan songbirds (Aves: Passeriformes). Protein evidence. Ibis 133, 277–85; Edwards SV and Boles WE (2002) Out of Gondwana: the origin of passerine birds. Trends in Ecology and Evolution 17, 347–49. 3

Irestedt M and Ohlsson J (2008 in press) The division of the major songbird radiation into Passerida and ‘Core Corvoidea’ (Aves: Passeriformes) – the species tree vs. gene trees. Zoologica Scripta.

4

Higgins PJ and Cowling S (2006) (Eds.) Handbook of Australian, New Zealand and Antarctic Birds. Volume 7: Boatbill to Starlings, Oxford University Press: Melbourne.

5

Loyn R (2002) Patterns of ecological segregation among forest and woodland birds in south-eastern Australia. Ornithological Science 1, 7–27.

6

Pizzey G and Knight F (1997) The Graham Pizzey and Frank Knight Field Guide to the Birds of Australia. Harper Collins: Sydney.

7

Joseph L et al. (2006) Implications of mitochondrial DNA polyphyly in two ecologically undifferentiated but morphologically distinct migratory birds, the masked and white-browed woodswallows Artamus spp. of inland Australia. Journal of Avian Biology 37, 625–36.

8

Avise JC (2000) Phylogeography: The History and Formation of Species. Harvard University Press: Cambridge, Massachusetts.

9

Williams MAJ (2002) Quaternary Australia: extremes in the last glacialinterglacial cycle. In Billion-year Earth History of Australia and Neighbours in Gondwanaland. (Ed. JJ Veevers) pp. 55–59. GEMOC Press: Sydney.

10 Keast JA (1961) Bird speciation on the Australian continent. Bulletin of the Museum of Comparative Zoology, Harvard 123, 303–495; Schodde R (1982) Origin, adaptation and evolution of birds in arid Australia. In Evolution of the Flora and Fauna of Arid Australia. (Eds. WR Barker and PJM Greenslade) pp. 191–224. Peacock Publications: Adelaide; Ford JR (1974) Speciation in Australian birds adapted to arid habitats. Emu 74, 161–68. 11 Joseph L and Wilke T (2006) Molecular resolution of population history, systematics and historical biogeography of the Australian ringneck parrots Barnardius: are we there yet? Emu 106, 49–62; Joseph L and Wilke T (2007) Lack of phylogeographic structure in three widespread Australian birds reinforces emerging challenges in Australian historical biogeography. Journal of Biogeography 34, 612–24; Joseph L, Wilke T and

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Alpers D (2002) Reconciling genetic expectations from host specificity with historical population dynamics in an avian brood parasite, Horsfield’s bronze-cuckoo Chalcites basalis of Australia. Molecular Ecology 11, 829–37. 12 Endler JA (1986) Natural Selection in the Wild: Monographs in Population Biology, 21. Princeton University Press: Princeton, NJ. 13 Prideaux GJ et al. (2007) An arid-adapted Middle Pleistocene vertebrate fauna from south-central Australia. Nature 445, 422–25.

WHITE-WINGED

CHOUGHS :

THE SOCIAL CONSEQUENCES OF BOOM AND BUST Robert Heinsohn

Australia is a dry continent. It is particularly dry in the arid interior, and therefore it is not surprising that most Europeans settled in the wetter belts of the eastern and western seaboards. However even the wetter regions go through extreme droughts, which have devastating effects on both agriculture and wildlife. In recent decades, we have started to understand that wet years and dry years are driven by the El Niño southern oscillation (ENSO). The dry El Niño years in Australia occur when

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weakened trade winds in the western Pacific allow warm water to flow toward South America. This draws the rainfall that would normally fall on Australia away to the east. There is a growing recognition that drought in Australia is both normal and common, and indeed we should expect one or two bad years every decade or so.1 Alongside major losses to agriculture, drought imposes severe limitations on our wildlife. In less severe droughts animals may simply fail to breed, but in more severe droughts adult and juvenile mortality can rise alarmingly. Whereas science has learned a great deal about how animals have adapted to climatic uncertainty in the arid zone, there is less appreciation that many Australian animal populations living in the wetter regions have also evolved to deal with boom and bust conditions.2 In this chapter I outline how one of Australia’s very social and charismatic birds offers something of a metaphor for humans living in such highly variable climatic conditions. White-winged choughs (Corcorax melanorhamphos) are large (350–380 g), black and white, comical and endearing birds found in the eucalypt woodlands of eastern Australia. They have been the subjects of our research in the nature reserves near Canberra for over 20 years. Choughs are unusual because they simply must live in groups from four to 20 birds in order to survive and breed.3 All of the group members contribute to building their remarkable mud nest and to caring for the young, and their intense sociality is illustrated by the fact that group members are rarely more than a few metres apart.4 The

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apparent metaphor they provide for humans is that their intense sociality can lead to two entirely different outcomes in the harsh Australian landscape. Group living allows these birds to eke out a stable and mutually beneficial living in the good years, but also leads to gang warfare and group-based power struggles for limited resources when ecological conditions deteriorate.

Cooperative breeding – an Australian phenomenon? Choughs exhibit an intriguing phenomenon called cooperative breeding, a form of social organisation that is relatively rare on the world stage but surprisingly common in Australia. Whereas less than 3% of birds do it worldwide, cooperative breeding occurs in over 20% of Australia’s old endemic birds, those birds that evolved in ancient Gondwana rather than those that found their way to Australia later.5 Cooperative breeding is now considered an especially Australian phenomenon, and researchers who want to understand it often choose to study Australian species, irrespective of their own background. Cooperative breeding is unusual because some individuals forgo breeding themselves. Instead they act as helpers, and assist others to raise their young.6 This usually means bringing food to the nest but it can also entail incubating the eggs and chasing away predators. To evolutionary biologists, the idea of forgoing one’s own reproduction is anathema. Our understanding of natural selection acting on individuals to produce as many offspring as possible means that the last

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thing an individual should do is give up and help someone else to raise their young. It was after long-term studies of individually marked birds, many of them Australian species, that we worked out that birds only behave this way when they are forced to ‘make the best of a bad job’. Often environmental circumstances mean that individuals cannot get a breeding position themselves. There might be no room to set up a new territory in an overcrowded habitat, or environmental conditions might make it too hard to find enough food to feed offspring.7 When this happens, young birds that might otherwise disperse to breed themselves have the option of staying home and helping their parents. If they can boost the output of related offspring, they get a reward known as ‘inclusive fitness’.8 Because the offspring share many of the same genes, this can be thought of as producing ‘offspring equivalents’. It is interesting that Charles Darwin himself worried that the equivalent altruism shown by non-reproductive workers in insects (helping in birds had not yet been fully appreciated) was ‘possibly fatal’ to his theory of natural selection.9 Hamilton’s theory of inclusive fitness, published 105 years after Darwin’s, smoothed over this potential pitfall in one of history’s greatest ideas.10 Australian ornithologists have featured prominently in the initial identification of, and subsequent explanations for, cooperative breeding. This probably relates to the fact that many of Australia’s cooperatively breeding birds are also our most common and easily observed species. Unfortunately much of

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this pioneering work has not been appropriately noted in the literature, and many authors have mistakenly attributed the discovery of cooperative breeding to Alexander Skutch, an expert on Central American ornithology, who published an important paper about this in the North American journal Auk in 1935.11 For example, the fact that fairy-wrens of the genus Malurus occur in small parties was first noted as long ago as 1808, with the suggestion of polygamy within these groups in 1827.12 The regular attendance at the nest by at least two males was unambiguously described by Graham and noted by Australian ornithologist Robert Hall in a popular book in 1907.13 The extreme gregariousness of white-winged choughs was also noted as early as 1827, and the fact that many birds operated from one nest was clearly described by John Gould in 1846 as part of his study The Birds of Australia.14 The most likely reasons for the slow recognition of the phenomenon of cooperative breeding, and the early Australian descriptions of it, include the ‘theoretical blinkering’ of early European naturalists (in the same vein as the original denial of the existence of the platypus), and Australia being a ‘victim of geography’ with a low density of naturalists to study its vast and distinctive biota.15 Australian researchers were also in the forefront of post-war studies on the behaviour and ecology of birds conducted against the backdrop of the modern evolutionary synthesis. Ian Rowley, in particular, was amongst the first to mark birds using coloured leg bands and to follow the behaviour and strategies of individuals in the broader population. His pioneering studies of superb

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fairy-wrens (Malurus cyaneus) and white-winged choughs showed that helping behaviour is a common and advantageous pathway for individuals to take as they wait for breeding positions to become available.16 Rowley was also one of the first researchers to shatter the myth of widespread monogamy in birds. His later study of splendid fairy-wrens (Malurus splendens) in Western Australia showed that most members of the population indulged in high rates of extra-pair copulations.17 More recent techniques using DNA-fingerprinting have shown this to be widespread amongst bird species even if they are socially monogamous.

Obligate cooperative breeding in choughs Cooperative breeding is usually facultative, that is it occurs when environmental conditions render independent breeding difficult for some, usually the youngest, individuals.18 This compares with favourable environmental conditions when individuals virtually always prefer to breed independently. However an interesting recent revelation based on new DNAbased phylogenies is that cooperative breeding may be an ancestral condition in many bird taxa. That is, cooperative breeding may have been the norm before many species later became pair-dwelling.19 White-winged choughs are ‘obligately cooperative’, that is they always have to be in groups to reproduce successfully.20 Obligate sociality is rarer in nature but occurs in some birds and primates (including humans) and a few other mammals such as wild dogs (Lycaon pictus), meerkats (Suricata suricatta) and lions (Panthera leo).21 The obligate

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group living found in primates is believed to be the ultimate driving force behind the evolution of higher intelligence. The conflicting demands of both needing, and being in competition with, your group members sets the scene for complex ‘Machiavellian’ social interactions.22 The ultimate cause of obligate sociality in choughs appears to relate to their foraging niche. They have a very difficult method of foraging in which they sift leaf litter and dig into the ground to get beetle larvae and other invertebrates such as earthworms. Knowing where to dig is a skill that takes young choughs years to master, to the extent that they have a mandatory four year apprenticeship before they reach adulthood. That is, they fail to develop sexually until they have had enough years to practise foraging. They undertake their apprenticeship in their parents’ territory and contribute to helping to raise new clutches of their siblings as far as their limited abilities let them.23 Even when they reach adulthood after this long apprenticeship, choughs still cannot raise young without help. They need at least two ‘helpers’ to get one or more nestlings through to fledging. Thus young birds need to remain in their parents’ social group while they slowly mature, but equally their parents need them to stay in order to increase their subsequent reproductive success. So living in groups is essential for both parents and grown offspring. When parents die, the older offspring vie for the breeding positions, and the workforce of younger individuals becomes a hotly contested resource.24

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Societal collapse The choughs’ chronic need for help leads to fascinating politics, especially when the weather turns bad. In good years, when the ground is soft and invertebrates are in good supply, chough groups continue to grow in size and young birds tend to stay at home to help their parents. Even older offspring who have completed their apprenticeships, and who are now sexually mature, tend to stay home and help their parents. Thus, in good weather all choughs are either dominant breeders or subservient helpers. Life in the chough group, at least to the outside observer, seems peaceful and harmonious.25 In drought years, the ground becomes hard and dry, the beetle larvae and earthworms die off, and food is harder to find. Breeding is more difficult and the youngest nestlings die so that brood size is closely matched to group size.26 Mortality amongst adult choughs also increases. At least some of the old breeding birds controlling the groups are likely to die, and breeding aged but subordinate birds at last get their chance to try and reproduce for themselves.27 They squabble with other contenders over the breeding positions, and power struggles ensue. Large groups often break up along factional lines into smaller units. An older female might take a younger sibling with her. Three brothers might leave together to try their luck in the outside world. These changes ripple through the population in a domino effect. Even in groups where the breeders have not died, the older helpers see these floating factions as new potential collaborators for breeding. They often disperse

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from their natal group, sometimes taking younger birds with them as helpers.28 Thus El Niño-induced drought tears the fabric of chough society. Large stable groups are replaced by small floating factions that are rarely big enough to breed on their own. The oldest bird in each of these factions looks for other factions with another bird of appropriate sex and age as a mate. Prospective mates are more attractive if they come with an entourage of helpers. The two factions together must comprise at least four birds to be successful, but of course the larger the new group the better.29 The previous harmony of chough society disappears as choughs become pitted against each other in these new harsh circumstances, and start indulging in dastardly deeds against their own kind. We see a version of gang warfare more reminiscent of the streets of Mogadishu or Melbourne Docklands than anything we are accustomed to in the Australian bush. In general, larger groups bully and harass smaller groups. My research showed that groups needed to comprise six or more individuals in order to be relatively safe from this activity.30 Choughs engage in three forms of aggression when pitted against each other. First, in normal contests entire groups will line up along branches and display vigorously, and fly at, and peck each other. Their wing-wave-tail-wag display is spectacular and bizarre to behold. The birds wave their wings and tails and engorge the conjunctiva of their eyes with blood to the extent that their eyes appear to bulge out of their skulls. 31

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Second, if one group is trying to destabilise another, its members will fly to the newly made mud nest and dislodge it from the tree.32 If the nest is empty it will cost the resident group the week or so it took to make it. If the group has eggs or chicks, it may lose weeks of precious breeding time. Sometimes they start again, but often in these turbulent times the group will break up into its original factions which then fly off in separate directions in search of better breeding opportunities. One temporary amalgamation of 13 males in our study population near Canberra, whom I referred to as the ‘Mafia’, ruined the nesting attempts of over 10 other groups. They appeared most concerned with finding females to breed with, and some of them eventually found mates in the fragments of the groups they harassed. Third, warring chough groups kidnap each others’ fledglings. The quest to find helpers is so intense that choughs often resort to forcibly taking the newly fledged young from other groups. They fly in, attack any of the babysitters of these young helpless birds, and then entice the fledglings away using the wing-wave-tail-wag display. They then proceed to feed the youngsters and care for them as if they were the group’s own. This investment in offspring to whom they are not related makes sense in the long-term, as eventually after a year or so, the young birds begin to pay their own way as helpers.33 The explanation for this ‘crime wave’ that sweeps through chough society every drought year may be particularly pertinent to human societies living under ecological stress.

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Diamond’s chapter on Rwanda, in his book Collapse, paints a remarkably similar picture of social mayhem in an ecologically stressed human society. In Rwanda, population densities over a long run of relatively good years had crept higher and higher. Farm sizes were getting smaller and smaller through inheritance, to the point that many children were not inheriting any land. Many people were forced to beg for food, or at least to rely very heavily on relatives. There was also a mix of different ethnic groups adding further complexity and fragility to the political situation. In the case of Rwanda, the trigger for large scale violence, against the backdrop of an ecologically strained population, was indeed political, but once it started mob violence spread across the country in domino-like fashion. 34 The meltdown of chough society that I observed over two droughts, one in 1994 and the other in 2002, was in some ways analogous to that among humans in Rwanda. In good times, the dominant birds ruled and the subordinates stayed in their place through a combination of social dominance hierarchies and direct rewards such as high inclusive fitness and eventual inheritance. Once drought set in, especially after many good years, the choughs found themselves living at high density in an environment that could no longer support so many birds. When dominant birds died, opportunities were created, and restless subordinates seized their chance for greater gains. Violence erupted amongst the ensuing competition, and some individuals emerged victorious, albeit in circumstances where it was almost impossible to live and breed without severe

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harassment. Like Rwanda during its turbulent times, a handful of aggressive individuals did very well, but the society as a whole seemed to suffer. Thus a system which entailed positive rewards for most players during good times got replaced by a harsher system with high rewards for a few, and little gain and much hardship for most of the population.35

The metaphor How much of a metaphor do white-winged choughs offer to humans living in variable and sometimes stressed ecological conditions? Constant group-living brings both constant benefits and a buffer against a capricious environment.36 In this sense the obligate communal lifestyle of choughs appears to have similar benefits as that of human hunter-gatherer societies.37 The benefits span those from simply living in a group per se, such as having many eyes to watch out for predators, to further forms of active cooperation that evolve when individuals are always together and have the opportunity to learn whom they can trust.38 Cooperative hunting and sharing of food are common benefits for human and animal societies. The joint care of young is another benefit of cooperation when foraging or predation make rearing offspring difficult. Modern humans took the fundamental advantages of group-living and cooperation, including the specialisation of roles and trading of goods and services, to ever greater heights after the advent of intensive agriculture. Of equal interest is how individuals behave when complex societies collapse, and in this context the chough metaphor

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may illustrate some basic principles behind the less attractive behaviour of humans under social and ecological duress. Even at the best of times, life in permanent groups is a finely tuned mix of cooperation and competition in which it pays all members to be there, but some individuals clearly do better than others. Usually it is the higher ranking individuals that gain most, but those of lower rank benefit nonetheless. However, when the potential pool of rewards is reduced (for example, through over-crowding and/or deteriorating environmental conditions) both within and between-group competition must increase. Subordinate individuals might also find the ground rules for their own behaviour have changed when other dominant and controlling individuals are removed through death or social unrest. The formation of small coalitions or gangs to enhance the competitive abilities of individuals is a feature observed in both choughs and humans when their societies collapse. Warfare between larger groups, tribes, and nations also increases when social and ecological conditions deteriorate.39 Perhaps the simple lesson to be learned by watching the remarkable differences in white-winged chough society in boom and bust years is that the bust years place great strain even on societies that are otherwise functioning well. In Australia, there are natural fluctuations beyond the annual seasons, and perhaps understanding and working with these extremes would be to the benefit of all. We have transplanted cultures and technologies developed elsewhere into a landscape with sometimes fundamentally different ecological

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conditions, and need to accept that much of the growth and prosperity that we experience in boom years (for example, through agriculture) will not be available in the bust years.40 We should also expect extreme ecological strains on both our society and our wildlife in the bust years, and be prepared for varying levels of societal meltdown and bullying of the smaller players. As illustrated by choughs there is an inevitable shakedown of all players, both human and wildlife, whenever an El Niño-induced drought occurs. Just as the chough groups cannot grow forever, the carrying capacity of this continent for all creatures is kept in check by our unpredictable weather. With global warming and shrinking habitats the metaphor suggested by choughs coping with booms and busts may only become more relevant and to a wider range of places.

Endnotes 1

Bureau of Meteorology, Living with Drought. Commonwealth of Australia: Melbourne, 2006. http://www.bom.gov.au/climate/drought.

2

Manning A, Lindenmayer D, Barry S and Nix H (2007) Large scale spatial and temporal dynamics of the vulnerable and highly mobile superb parrot. Journal of Biogeography 34, 289–304; Saunders D and Heinsohn R (2008) Winter habitat use by the endangered, migratory swift parrot (Lathamus discolor) in New South Wales. Emu 108, 81–89; Gardner J and Heinsohn R (2007) Probable consequences of high female mortality for speckled warblers living in habitat remnants. Biological Conservation 135, 489–99. On desert adaptations see Keast JA and Marshall AJ (1954) The influence of drought and rainfall on reproduction in Australian desert birds. Proceedings of the Zoological Society of London 124, 493–99; Serventy DL (1971) Biology of desert birds. In Avian Biology. Vol. 1. (Eds. DS Farner, JR King and KC Parkes) pp. 287–339. Academic Press: New York; Schodde R (1982) Origin, adaptation and evolution of birds in arid Australia. In Evolution of the Flora and Fauna of Arid Australia. (Eds. WR Barker and PJM Greenslade) pp. 191–224. Peacock Publications: Adelaide;

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Astheimer L and Buttemer W (2002) Changes in latitude, changes in attitude: a perspective on ecophysiological studies of Australian birds. Emu 102, 19–27; Perfita N, Zann R, Bentley G and Hau M (2007) Opportunism at work: habitat predictability affects reproductive readiness in free-living zebra finches. Functional Ecology 21, 291–301; Steve Morton, this volume, Chapter 4. 3

Heinsohn R (1992) Cooperative enhancement of reproductive success in white-winged choughs. Evolutionary Ecology 6, 97–114.

4

Rowley I (1978) Communal activities among white-winged choughs. Ibis 120, 178–97; Heinsohn R and Cockburn A (1994) Helping is costly to young birds in cooperatively breeding white-winged choughs. Proceedings of the Royal Society (London) Biological Sciences Series B 256, 293–98.

5

Russell E (1989) Cooperative breeding – a Gondwanan perspective. Emu 89, 61–62; Cockburn A (1996) Why do so many Australian birds cooperate: social evolution in the Corvida? In Frontiers of Population Ecology. (Eds. RB Floyd, AW Shepherd and PJ De Barro) pp. 451–72. CSIRO Publishing: Melbourne.

6

Cockburn A (1998) Evolution of helping behaviour in cooperatively breeding birds. Annual Review of Ecology and Systematics 29, 141–77.

7

Brown J (1987) Helping and Communal Breeding in Birds. Princeton University Press: Princeton; Koenig WD, Pitelka FA, Carmen WJ, Mumme RL and Stanback ML (1992) The evolution of delayed dispersal in cooperative breeders. Quarterly Review of Biology 67, 111–50.

8

Hamilton W (1964) The genetical evolution of social behaviour I & II. Journal of Theoretical Biology 7, 1–52.

9

Darwin C (1859) On the Origin of Species. Murray: London.

10 Hamilton W (1964). 11 Skutch A (1935) Helpers at the nest. The Auk 52, 257–73. 12 Lewin JW (1808) Birds of New Holland with their Natural History Collected, Engraved and Faithfully Painted After Nature. White and Bagster: London; Vigors NA and Horsefield T (1827) A description of the Australian birds in the collection of the Linnaean Society, with an attempt at arranging them according to their natural affinities. Transactions of the Linnaean Society, London 15, 170–331. 13 Graham quoted in Hall R (1907) The Useful Birds of Southern Australia with Notes on Other Birds. Lothian: Melbourne.

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14 Vigors and Horsefield (1827); Gould J (1846) The Birds of Australia (in seven volumes), Vol. IV. Taylor: London. 15 Boland C and Cockburn A (2002) Short sketches from the long history of cooperative breeding in Australian birds. Emu 102, 9–17. 16 Rowley I (1964) The life history of the superb blue wren Malurus cyaneus. Emu 64, 251–97; Rowley I (1978). 17 Rowley I and Russell E (1990) Philandering – a mixed strategy in the splendid fairy-wren Malurus splendens. Behavioral Ecology and Sociobiology 27, 431–37. 18 Brown J (1987). 19 Cockburn A (1996). 20 Heinsohn R (1992). 21 Clutton-Brock T (2002) Breeding together: kin selection and mutualism in cooperative societies. Science 296, 69–72; Heinsohn R and Packer C (1995) Complex cooperative strategies in group-territorial African lions. Science 269, 126–62; Vucetich JA and Creel S (1999) Ecological interactions, social organization, and extinction risk in African wild dogs. Conservation Biology 13, 1172–82. 22 Whiten A and Byrne B (Eds.) (1997) Machiavellian Intelligence II. Cambridge University Press: Cambridge. 23 Heinsohn R, Cockburn A and Cunningham R (1988) Foraging, delayed maturation and advantages of cooperative breeding in white-winged choughs, Corcorax melanorhamphos. Ethology 77, 177–86; Rowley I (1978); Heinsohn R and Cockburn A (1994). 24 Heinsohn R (1992). 25 Heinsohn R (1992). 26 Heinsohn R (1995) Hatching asynchrony and brood reduction in cooperatively breeding white-winged choughs. Emu 95, 252–58. 27 Heinsohn R, Dunn P, Legge S and Double M (2000) Coalitions of relatives and reproductive skew in cooperatively breeding white-winged choughs. Proceedings of the Royal Society (London) Biological Sciences Series B 267, 243–49. 28 Heinsohn et al. (2000). 29 Heinsohn R (1992); Heinsohn R and Cockburn A (1994).

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30 Heinsohn et al. (2000). 31 Rowley I (1978). 32 Heinsohn R (1988) Inter-group ovicide and nest destruction in cooperatively breeding white-winged choughs. Animal Behaviour 36, 1856–58. 33 Heinsohn R (1991) Kidnapping and reciprocity in cooperatively breeding white-winged choughs. Animal Behaviour 41, 1097–100. 34 Diamond J (2005) Collapse: How Societies Choose to Fail or Succeed. Viking Press: New York, esp. pp. 311–28. 35 Heinsohn et al. (2000). 36 Rowley I (1965). 37 Anthropologist Deborah Rose suggested that this might have applied to Australian Aborigines before European contact (personal communication). 38 Clutton-Brock (2002); Whiten and Byrne (1997). 39 Diamond J (2005). 40 Robin L (2007) How a Continent Created a Nation. UNSW Press: Sydney, esp. pp. 1–6, 56–74.

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EMU :

NATIONAL SYMBOLS AND ECOLOGICAL LIMITS Libby Robin

Taking up arms One of the most prominent symbols of a nation is its coat of arms. Australia followed a common British tradition when it chose two animals to embrace a shield, a kangaroo and an emu. But the two animals are not as equal as one might expect. The emu is always second to the kangaroo, just as the British unicorn is second to the dominant British lion.1 The lion is the major symbol of Britain and in Australia, the kangaroo takes this role. Australia’s Department of Foreign Affairs lists a

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special web page on kangaroos amongst its ‘top eight’ recommended information sites, but does not have a similar one for emus.2 Even the famously unappreciative early colonial poet and judge, Barron Field regarded the kangaroo as: ‘the spirit of Australia/ that redeems from utter failure/ from perfect desolation/ and warrants the creation/ Of this fifth part of the Earth’.3 Australian nature poses a serious problem for writers seeking poetic resonance with European sensibility. As Judith Wright wrote: ‘there were aspects of nature here – like … the emu … – at which even a dedicated bard might falter in attempting to translate … into terms which might be admired in London’.4 Yet if Australian writers of the 19th century were to be recognised as serious, they needed to engage with the British market, because that was where the readers and the publishers were; London was the cultural reference point. The kangaroo is so much the ‘signature’ animal for Australia that Austrians, anxious to clear up tourist confusion, manufacture T-shirts and caps with a kangaroo enclosed in an international red circle with a bar through it and a slogan: ‘No kangaroos in Austria’. If Tim Flannery’s book had been about emus, he could hardly have called it Country, yet that was the sole title of the first edition of his popular kangaroo book.5 The successful challenge for the America’s Cup by Australia II in 1983 became inexorably entwined with the symbolism of the boxing kangaroo. And Australia’s airline Qantas eschewed the flightless bird in favour of the flying kangaroo in its logo, and ‘The Spirit of Australia’ motto that harks back to Barron Field.6 242

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In Britain, the lion became the essential symbol of the nation, the empire and patriotic pride. It was large and strong, like the Danish elephant and the American bald eagle. Less evident in British imperial rhetoric was the lion’s mythical counterpart, the unicorn. In one sense, both were mythical. Lions have not roamed the British Isles since before the last Ice Age, and certainly not in the era of imperial pomp and ceremony, although there is still enthusiastic searching for large felines in remote places like Dartmoor. Lions (Panthera leo) are, however, real animals and live today in countries like South Africa that used to be part of the former British empire. The unicorn is, by contrast, quintessentially fictional. It is probably based on the narwhal (Monodon monoceros) of the Arctic, but is more a horse (land animal) than a seal (sea animal). It came via Norse sagas to represent the Scots in the British union. The unicorn is an enigmatic national symbol. It quickly became the ‘optional’ part of the arms. The coat of arms for the British Antarctic Territories, for example, has the lion in its usual prominent place on the left, but the unicorn on the right is replaced with an emperor penguin (Aptenodytes forsteri).7 When the Australian colonies bid to come together to make a nation, they sought something for the new Coat of Arms that was authentically southern and separate, but which also embraced Britain in a traditional way. The tension between the old loyalties and the new was apparent in the words of a pro-Federation poster of 1900, where Australians were urged to ‘prove to men of every race that the decendants 243

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[sic] of Britons in AUSTRALIA HAVE NOT LOST THEIR CAPACITY FOR SELF-GOVERNMENT’.8 Having voted to federate, the ‘decendants of Britons’ chose a kangaroo and an emu as heraldic emblems for the new nation, to replace the lion and the unicorn of the old country. This combination was not new. In 1804 Robert Knopwood chose it for the arms for Hobart Town. On the left he drew a forester kangaroo (Macropus giganteus, known on the mainland as an eastern grey kangaroo), the largest mammal in Tasmania, and on the right the now extinct Tasmanian emu, currently regarded as an extinct population of the mainland emu (Dromaius novaehollandiae) and then the largest bird in Tasmania. Knopwood added a Latin motto Sic Fortis Hobartia Crevit (‘Thus by Industry Hobart Town Increased’), also typical of the time. James Boyce has argued that the kangaroo and the emu were not merely symbols; they were chosen because they formed a significant part of the diet of Van Diemonians and were therefore regarded as part of the ‘natural bounty’ of the place.9 The Tasmanian emu was driven to extinction before the colonies federated, and the forester kangaroo very nearly followed suit, being now confined to districts far from Hobart, but it is not clear whether this was because of over-hunting or changed land use patterns, or both. A hundred years after Knopwood, the Australian national coat of arms carried a motto in English rather than Latin: ‘Advance Australia’. In 1912, when the States were added to the shield, golden wattle (Acacia pycnantha), a national flower, was added as an additional

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feature. In Britain, the rose, thistle and shamrock (for England, Scotland and Ireland) were featured below the shield in the same way.10 In this chapter, I explore the emu’s national symbolism and its stories. I argue that the emu seems to have followed the unicorn in being a creature of myths and abstract ideas rather than understood as a living bird. Status as a national symbol has not helped its treatment under settler Australian rule. At times, national myths have been disadvantageous to its survival. The ‘yeoman ideal’, where land is allocated in small parcels to enable each farming family to own its own land, was transferred from the land of the unicorn to the land of the emu, and underpinned much national pride in the latter. Indeed, the politics of small farms dates back to well before Federation. In New South Wales, the Robertson Land Acts of 1861 were designed to give the land titles to ‘selectors’, people who undertook to live on their properties, rather than the earlier situation where wealthy squatters controlled vast open tracts of country from a distance.11 The coat of arms chosen in 1901 represented a national economy and a nation with pastoralism at its core. But the emu’s nomadic habits were not well suited to the fences and small farms of the yeoman settlement ideal. The emu was unpopular and even deemed to be ‘unpatriotic’ when it threatened nation-building pastoral activities with its nomadic wanderings. Like the kangaroo, the emu enjoys the favoured pastoral country of ‘Australia Felix’: open grasslands and scattered

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tree-cover, in the temperate south. But both our national faunal emblems are, perhaps appropriately, roaming opportunists. Their peregrinations are not limited to the good pastoral country. Each is quick to respond to booming resources wherever they may be and each is prepared to move great distances to avoid bust situations. Emu populations can appear apparently from nowhere and expand rapidly as birds chase down the new resources, especially the ones provided by the developing pastoral and agricultural economy. Taking an emu’s perspective and watching how it adapts to the rhythms of Australia’s boom and bust ecology can provide settler Australians with alternative models for building an Australian economy and way of life that works ecologically in this ancient land.

Distribution, food and movements The emu occurs naturally throughout the Australian mainland, except in tropical rainforests, big cities and peri-urban areas. It is generally absent from cleared farmland, as it needs good cover to breed.12 It is relatively uncommon in the uninhabited inland, but after heavy rains, concentrated populations may appear moving in quickly from distant places. CSIRO scientist Stephen Davies, who studied emus extensively in the 1960s and ’70s, reported recapturing emus up to 500 km from where they were banded. In successive aerial surveys of a 2500 km2 area conducted over an eight year period, he discovered that their numbers fluctuated between 20 and 576 birds.13 Although they spread over large ranges, they also have a sense

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of ‘territory’ when they are breeding. They do not like other emus nearby. Over his huge study area, Davies found that each breeding pair of emus had approximately a 30 m2 territory, and that when they met other emus they tended to disperse. So once they are moving, they tend to keep moving because they all move in the same direction, usually following watercourses, so they keep encountering each other. The watercourses support higher numbers when required, but the birds prefer more territory and less movement if they can find it.14 The idea that emus are a monogamously pairing species has been challenged in more recent work, which shows that like all other ratites except kiwis, emus exhibit monogamy, polyandry (it is the male that takes primary care of the chicks) and promiscuity depending on circumstances and resources.15 Davies’ finding that emus prefer a larger, more open territory is consistent with this, as a larger area better supports opportunities for mating beyond the primary pair. The emu is generally regarded as a winter breeding species, while most specialist arid-zone birds (slightly) favour spring. But emus also have the capacity to breed at other times and in both semi-arid and temperate places. Studies of the arid zone, often conducted after heavy rain, have lent support to the idea that arid zone birds breed ‘erratically’ – like the zebra finches discussed in Chapter 4. Generally this is not true, but there appears to be an option for birds like emus that move in and out of the unpredictable and variable outback to take advantage of unexpected good resources at times other

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than their preferred breeding times.16 As early as 1908, one of the closest observers of emus in Mallee country in Victoria, Charles M‘Lennan, commented: they seemed to have some strange foreknowledge of the weather … because I have noticed that in seasons which have turned out very wet the bird frequently builds its nest on high ground … before the winter and spring rains have fairly set in … but in seasons which afterwards turned out to be exceptionally dry, I have found the nest on low ground, even in depressions.17

Emus feed on different things at different times of year, depending on seasonal abundance. In Western Australia, Stephen Davies observed that: ‘they ate fruits, flowers, seeds, succulent green shoots of grass, herbs and shrubs and also took insects’.18 He also commented that other studies showed that they ate different foods in different parts of Australia. Although the range of food is wide, ‘all the types represent concentrated sources of nutrient materials when contrasted with the mature or dry grass and browse taken by many herbivores’.19 This was consistent with other ratite birds, such as rheas and ostriches. It is also a pattern evident from studies based on fossil egg shells from 45 000–55 000 years ago, a transitional time when the landscape was changing fairly rapidly. At a time when most of Australia’s largest mammals became extinct, including over a hundred species of kanga-

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roo, the emu survived.20 The landscape was changing quite rapidly at this time, possibly because of similarly changing natural rainfall patterns or possibly because of regular fires lit by the first wave of humans as they colonised the continent. The mosaic of drought-adapted trees, shrubs and rich grasslands shaped by the seasonal rainfall that was typical of 60 000 years ago converted to fire-adapted grasslands and chenopod desert scrub over a period of about 10 000 years. Recent studies have shown that in the river systems of the south-eastern arid zone where emus and the extinct Genyornis co-occurred, there was a major reduction in the plants that used the C4 photosynthesis path before this change and that the C4 plants that remained after this time (for example, spinifex) were much less nutritious. Palaeobotanists can tell what the class of plants various animals ate by looking at the C3 and C4 traces in eggshells (for birds) and teeth (for mammals). While Genyornis specialised on the nutritious C4 synthesising plants of the earlier era died out (see Chapter 8), generalists like emus adapted from a mixture of C3 and C4 plants to a diet dominated by C3 (woody shrubs) and survived.21 Such adaptability has been documented more recently in different ways, in the emu’s ability to make ‘specialties’ of newly available food sources such as the pest plant, prickly pear (Opuntia spp.) as well as farmers’ crops, such as wheat. Their ability to move around quickly and eat and breed opportunistically makes emus a classic boom and bust species. They are a symbol of Australia in more than one sense.

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Emu myths at the time of federation From the start emus were a significant feature of Australia. An emu was shot within a few weeks of the first British settlement, less than two miles from Sydney Cove. It was dissected carefully and finally eaten by Watkin Tench, who described the flesh as tasting ‘like beef’. The skin was sent to Lord Sydney by Governor Arthur Phillip who thought it a significant bird, meriting attention.22 The Tasmanian emu and the much smaller King Island emu and Kangaroo Island emu (D. baudinianus) lived in similar country to that preferred by the mainland emu. They were shy and much more difficult to hunt than kangaroos, according to Baudin’s 1803 account of Kangaroo Island emus but despite this – or perhaps because of it – they were prized as meat.23 After the arrival of Europeans, extensive clearing and hunting caused their quick demise, particularly in smaller islands where there was little alternative country where they could re-establish populations.24 Settler Australians were drawn to emu stories round the time of Federation when the emu became formally part of the arms, perhaps because the unicorn had such a strong status as a mythical animal in Britain. Katie Langloh Parker (later Catherine Stow) actively promoted stories about emus as part of her work on Aboriginal ideas about the Australian environment. She was one of Australia’s earliest female ethnologists, an author of children’s books and of a magazine of Aboriginal myths and culture based on the tales she heard from local Noongahburrah people in and around Walgett in western New

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South Wales.25 She grew up a fluent speaker of Noongahburrah. When she was six, an Aboriginal girl saved her and two of her sisters from drowning in the Darling River. As an adult, an isolated wife on Bangate station for over 20 years, Catherine Stow had several Aboriginal women working in her home and found opportunities to talk about all sorts of stories. In Woggheeguy she relates the story of Dinewan, the emu who was tricked by Goomblegubbon (wild turkey) into having her wings cut off, and how this taught the Laughing Jackass (kookaburra) to laugh.26 These stories were more about the social relations between the birds than their ecological niches, but as Deborah Rose showed in Chapter 9, there were also many Aboriginal stories that explicitly considered the ecological relations between birds and their environments.

The emu’s symbolism Why was the emu chosen to represent ‘nationalist’ ventures? It was not just part of the coat of arms, but a national symbol in a range of places at the time of Federation. For example, the Australasian Ornithologists’ Union, founded in 1901, adopted Emu as the name of their learned journal.27 Its almost dinosaur-like head is strongly suggestive of the ‘primitive’, a concept of Australia popular at the turn of the 20th century. Indeed, evolutionary biologists still regard the ratites, including the emu, as having particular interest. Recent studies have shown that the ratites are among evolutionary lineages that date back to Gondwanan times. In the early 1900s classifications were not based

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on cladistics and today’s genetic technologies of course, but on Eurocentric ideas based on form (morphology). Nonetheless biologists were very concerned with the idea of what was ‘primitive’ in the system. The emu was part of the discourse where so-called primitive life-forms were regarded as inferior, or only ‘halfway developed’ towards the better-developed forms of other places. Australia was seen as something of a ‘continental museum’ and the emu was one of the animals regarded as more a ‘curiosity’ than a functional animal. Taxonomy, founded in Europe, was written in a Eurocentric language, as Stephen Jay Gould has observed: Prototheria (monotremes) were ‘premammals’; Metatheria (marsupials) were ‘middle mammals – not quite there’; and ‘Eutheria’ (the warm-blooded animals of the North) were the ‘true mammals’.28

Australia, the continental museum, had curious ‘missing links’ that supported some theories of progressive evolution. It was also ‘a place of refuge for mediaeval types’. This particular idea, put forward in 1910 by passing English journalist Joseph McCabe, became part of the mythology that justified an active campaign of ‘improvement’ as part of nation-making.29 This was a land where the development of the natural creatures had stalled. It was a land of ‘missing parts’ in need of new and useful biota from elsewhere as well as new technologies that could to guide it into modernity and participation in international markets. 252

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Misunderstanding the emu The emu was not famously noble like the lion or the unicorn, but it did gather a reputation for being fast and difficult to catch. Emu feathers were sought-after symbols of riding competence, especially among mounted infantry units during the years of the First World War. 30 The Australian soldiers in the Australian Fourth Light Horse Brigade wore emu feathers in their hats with pride, but only one year after the end of the war, some Western Australians began to find the bird a nuisance in the new wheat and sheep country being opened up by returned soldiers. In 1919, the Upper Chapman Road Board at Nanson requested that the emu be removed from the protected list and by 1923, emus north of latitude 30ºS were classified officially as ‘pests’. As the farmers in Western Australia battled to make a new life for themselves in the post-war years on new blocks in the ‘wheat belt’, emu numbers expanded rapidly, especially around the Number 3 Rabbit-proof Fence running east-west from just south of Kalbarri to join up with the earlier Number 2 north-south Fence (see Figure 12.1).31 The Number 3 Fence increasingly became known as the Emu Fence, as bird numbers increased exponentially. From 1923, district vermin boards started to levy a bounty on emu heads from about 300 km north of Perth – between latitude 30ºS and the fence at 28ºS. The birds were no longer protected by the Game Act, and this area was being rapidly developed for sheep farming. Dominic Serventy estimated that in 1928, 3000 to 4000 emus were destroyed in just one district. He had a report from a 253

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Figure 12.1 Western Australian wheatbelt, showing the vermin fences constructed in the early years of the 20th century. (Illustration by Clive Hilliker)

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farmer stating that ‘he had seen a heap of dead emus 16 ft (3 m) high and 4 ft (1.3 m) square’. The next year, in response to an outbreak in the Northampton district, the Government subsidised the bounty, and 2/6 (25 cents) per head was paid on 2148 birds.32 The Depression years made margins tight and bounties valuable, but it appears that many birds perished on the Fence before they made it into populated areas where bounty hunters awaited them. According to a local vermin inspector in November 1930, ‘they are all lying dead along the fence between Ajana and the junction of the No. 2 and No. 3 fences … the farmers on the south side of the fence are jubilant’.33 The bounty system had also been adopted in the late 1920s in Queensland. The issue here was not emus but the noxious prickly pear, and because emus consumed the fruit, they were rightly regarded as a major vector, spreading the plant as they roamed. The consequences for emus were just as bad as in the West; bounties were paid on 330 000 birds and eggs of the emu, crow and even the cassowary in Queensland.34 The Emu Fence did not hold the birds for long in Western Australia. In 1932 an outbreak was reported further south and further inland, in the north-eastern wheatbelt country not far from the town of Southern Cross. This angered farmers who resented the ‘invasion’ and its effects on the important new wheat economy. The Commonwealth Minister for Defence, Sir George Pearce, was persuaded by the complaints of Western Australian

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farmers to send in the Army. He thought the emu was a scourge to be routed, and that this would make excellent military target practice. Pearce declared an ‘emu war’. The Seventh Heavy Battery of the Royal Australian Artillery brought two Lewis guns and 10 000 rounds of ammunition to the ‘war’, which was conducted without discussion with the local Department of Agriculture, which was the main promoter of the wheat belt development. Fifty local settlers organised a 35 km drive of emus to an ambush point on the No. 1 Rabbit Proof Fence. 35 But they had not reckoned on the response of the emus. Alas for the settlers and the honour of the Army, the birds knew how to split into small parties and ran zigzagging across the country completely foiling the stationary and inflexible ambush. A ploy to trap about 1000 emus at a watering point just before daylight on 4 November 1932 resulted in just 12 birds being shot before the gun jammed. Two days later the Army managed 50 birds, but at a total cost of more than 2500 rounds of ammunition. The miss rate was spectacular, and when this was mentioned in speeches in the House of Representatives in Canberra, the Minister ignominiously withdrew the Lewis guns on 8 November, just four days after the ‘war’ had begun. As the emu war became an evident failure, the Western Australian Department of Agriculture quickly distanced itself from the venture, leaving the Minister with the ignominy. The Western Australian government continued to prefer a bounty system, which effectively outsourced the problem and was popular with farmers and itinerant farm workers. Even at

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just a shilling a head for emus, skilled emu-catching was a bonus in tough times and the slaughter of emus provided extra income to workers on the edge. In 1935, bounties were paid on 57 034 birds in six months. Small mobile parties and emu drives were much more ruthlessly effective than the Army’s heavy-duty attack, which failed to engage with the way emus run across country. Both Aboriginal people and rural settlers had developed better methods for catching emus and had employed them with some success for many years. But even with the bounty-hunters, emu populations continued to be considered a problem for the farming economy. In 1944, at a time when the bounty fund had become permanent and centrally funded (rather than the separate initiatives of local boards), the Western Australian Minister for Agriculture FJS Wise declared the emu ‘vermin throughout the state’. It took naturalists until 1947 to get the message across that that it was a localised pest and quite rare in the southwest corner, where it was ultimately restored to the protected list.36

New science for an old land The emu’s status as a pest eventually became a blessing of sorts. Its biology and behaviour would have remained poorly understood if it had simply been regarded as a curiosity. There has traditionally been little funding for non-rare species that are not of economic value – either positively or negatively. The Agriculture Protection Board of Western Australia in 1958 approached Francis Ratcliffe, head of the CSIRO Wildlife

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Section, about studying the control and ‘bionomics’ of pest species including emus and black-cockatoos (now Baudin’s black-cockatoo Calyptorhynchus baudinii and Carnaby’s blackcockatoo C. latirostris).37 The thinking was that if its biology were better understood, methods to control numbers and behaviour of problem birds would be more effective. It is hard enough to catch an emu, but to catch it so it can then be released and followed, especially in the years before the advent of satellite tracking, posed a particular problem. In 1969, 154 fast-running emus were ‘noosed’ by the enterprising Stephen Davies and his CSIRO colleagues, one of whom was sitting on the bonnet of a moving land rover. These birds were then banded and released. Such ‘mark-recapture’ studies allow scientists to monitor individual characteristics as well as their movement and behaviour. Thirty bands were recovered over the next year, two on emus more than 400 km from where they were banded, showing the extent of movement of individual birds.38 Davies reported accounts from farmers and other observers of particular emus with distinguishing marks moving up to 800 km in a matter of a few days.

The idea of nomadism Nomadism has been closely studied by biologists internationally, particularly since the technique of radio tracking has allowed more detailed mapping of movements. It is common in places where rainfall and resources are highly variable. Some migrations are also ‘nomadic’ where birds travel to

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breed or survive on wintering grounds, and where their breeding is constrained by a resource pulse.39 Craig Allen and Denis Saunders define nomadism as ‘unpredictable movements in space and time that track unpredictable changes in resource distribution and abundance’, and they noted that ‘nomadic species tend to occur disproportionately in arid or semi-arid ecosystems’.40 Americans Robert Bennetts and Wiley Kitchens, reviewing the literature of nomadism, commented that it can be a response to a local depletion of food or other resources, but this is not always the case. Some animals ‘exhibit exploratory movements during time of high resource abundance’.41 That is, nomadism can be a response to bust conditions or it can equally be facilitated by boom. While not moving when there is no food is clearly a ‘cost’ for an animal, it is also an effort to move and extra resources can make this possible. Emus move in times of scarcity, but they also respond to unexpected resources. They responded quickly and abundantly to the unexpected opening up of the Western Australian wheat belt: the planted crop in 1932 supported 20 000 emus in an area of about 40 square miles near Southern Cross, something we know because the birds frustrated struggling farmers. So while nomadism is a boom and bust phenomenon, it is not always clear which motivation drives which movements, particularly for opportunistic feeders like emus. The international literature is however, agreed that uncertainty of resources is a major factor in nomadism. The Australian interior’s highly unpredictable ecosystems with boom and bust cycles set off by

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irregular rain and fire make nomadism, not sedentary behaviour, an evolutionarily advantageous strategy for many creatures, including emus.

A new symbolism based on the real emu The demise of the smaller emus in Tasmania and Kangaroo Island in the 19th century showed that populations can collapse suddenly, especially under pressure from hunting and habitat loss. Even on the mainland, habitat loss and restricted movement can threaten emus. Emus can also be adaptable survivors, sometimes the first animals to recolonise a landscape after disaster, moving quickly to take advantage of places where competition has been eliminated. Nomadism has a cost however, and emus may be particularly vulnerable to other pressures when they are moving. For example, many emus still perish at the Emu Fence in Western Australia on their spring migration coastward after winter breeding in the inland. This Fence, originally built in 1908, was still being extended by the Western Australian Department of Agriculture in the late 1950s as farming expanded into new country.42 In the 21st century mainland emu populations appear to be on the decrease again. Only half the population of the 1984 Atlas of Australian Birds was reported in the New Atlas of Australian Birds published in 2003, but this figure may be complicated by the different sampling techniques used in the two surveys, and the different environmental conditions that prevailed at the time each was undertaken.43 There is still only limited

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understanding of the problems posed by fences, roads and other obstacles to cross-country movement for emu populations.44 Nomadic strategies dictate that a ‘home country’ should have fluid edges. Rigid barriers like fences and highways will always pose a challenge to a nomadic animal like an emu. A typical emu population will spread its net as widely as possible across country to gain advantage in movement, rather than in investing in defending a single territory. Nomadism is a diametrically opposite strategy of settled yeomanry, the European ideal for settler Australia. Yeoman farmers develop small areas intensively, settle close to each other and create their society quickly and definitively. They need fences to set legal and territorial limits. By contrast, emu groups prefer not to live too close to each other or in fixed places. They need to be free to move quickly to take advantage of resource opportunities as they arise. From an emu’s perspective, a fence is major obstacle to a survival strategies developed over many generations of living within the limits imposed by arid country with a highly variable climate.

Postscript The emu is a diurnal creature and Western science has concentrated on its daytime activities. For Aboriginal people, the emu is very important; indeed it is kin for some people. And it is not just a bird of the day. It is also a story of the night. The emu is closely associated with Aboriginal astronomical ideas. Australia has glorious night skies, with an extraordinary

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number of stars brilliantly visible in the Milky Way. You need a Gestalt switch, however, to find the emu. Instead of looking at the stars, you have to look at the black between them and there you will find, silhouetted by stars, a very large blue-black emu. The emu in the sky complements the emu-hunting of the day. The idea of putting the emu and kangaroo together on the Australian coat of arms also perhaps has origins in Aboriginal ideas. As Bluey Roberts, a Ngarrindjeri Kukutha boomerang artist, from the region between the Murray and the Coorong in South Australia, explained it: Emus and kangaroos always eat together – they look out for each other and they protect each other and give warning to each other – they are always on watch always looking around.45

Where booms and busts can change country dramatically, a nation needs watchful guardians, day and night.

Endnotes 1

Souter G (2000) Lion and the Kangaroo: The Initiation of Australia. Text: Melbourne.

2

http://www.dfat.gov.au/geo/australia/. The others were (1) Australia’s system of government (2) Australia’s national symbols (3) Australia’s Coat of Arms (4) The Australian National Anthem (5) The Australian currency (6) Muslims in Australia – A long & vibrant history (7) Australia’s kangaroos (8) Australia’s World Heritage properties.

3

Field B (1819) First Fruits of Australian Poetry. George Howe: Sydney. Online: http://image.sl.nsw.gov.au/cgi-bin/ebindshow.pl?doc=dixson/ a319;seq=75.

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4

Wright J (1975) Romanticism and the last frontier. In Because I Was Invited. pp. 59–80. Oxford University Press: Melbourne.

5

Flannery T (2004) Country. Text: Melbourne. Later editions were called Country: A Continent, A Scientist and a Kangaroo (Text, 2005) and Chasing Kangaroos: A Continent, a Scientist, and a Search for the World’s Most Extraordinary Creature (Grove Publications: New York, 2007).

6

There was an Emu Airlines, however. One of its principal routes was to Kangaroo Island.

7

http://www.mapsofworld.com/f lags/british-antarctic-territory-f lag. html.

8

Poster ‘To the Australian Born’ (for vote for Federation), William Brooks and Co. Printers, Sydney, 20 June 1900.

9

Boyce J (2008) Return to Eden: Van Diemen’s Land and the early British settlement of Australia. Environment and History 14(2), 289–307.

10 The first arms (drawn in 1908) were just the St George Cross on the shield with the kangaroo and emu, but the 1912 version with a division for each State was much more elaborate, and has persisted. The wattle under the shield is optional, as is design of rose, thistle and shamrock in Britain. See Robin L (forthcoming 2009) Wattle. (Eds. M Harper and R White) Australian Symbols. 11 Strictly these were The Crown Land Acts 1861 (NSW) – and the Premier who introduced them was John Robertson – but they are generally referred to as the Robertson Land Acts. 12 Davies SJJF (1976) The natural history of the emu in comparison with that of other ratites. Proceedings of the 16th International Ornithological Congress. Canberra. (Eds. HJ Frith and JH Calaby) pp. 109–20. Australian Academy of Science: Canberra, ACT. 13 Davies SJJF (1976) p. 112; Davies SJJF, Beck MWR and JP Kruiskamp (1971) The results of banding 154 emus in Western Australia. CSIRO Wildlife Research 16, 77–79. 14 S Davies, interview with the author, 3 October 1998, Tape 1, side 1; Transcript, p. 14. 15 Coddington CL and Cockburn A (1995) The mating system of freeliving emus. Australian Journal of Zoology 43, 365–72. 16 Davies SJJF (1976) Environmental variables and the biology of Australian birds. In Proceedings of the 16th International Ornithological Congress.

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pp. 481–99; Davies SJJF (1976) p. 113; Allen CR and Saunders DA (2002) Variability between scales: predictors of nomadism in birds of an Australian Mediterranean-climate ecosystem. Ecosystems 5, 348–59. 17 [Report from the Australasian 21 March 1908] ‘The ways of the emu’. Emu 8, 42. 18 Davies SJJF (1976) p. 111. 19 Davies SJJF (1976) p. 111. On earlier studies, see Lea AM and Gray JT (1934) The food of Australian birds: an analysis of the stomach contents – part I. Emu 34, 275–92 (mostly South Australia, with one Queensland observation); Long JL (1964) Weights, measurements and food of the emu in the northern wheatbelt of Western Australia. Emu 64, 214–19; Noble JC (1975) The effects of emus (Dromaius novaehollandiae Latham) on the distribution of the nitre bush (Nitraria billardieri DC). Journal of Ecology 63, 979–84 (data from South Australia and New South Wales). 20 Flannery T (2004) p. 5. See also Mike Smith, ‘Genyornis’, this volume, Chapter 8. 21 Johnson BJ et al. (1999) 65 000 years of vegetation change in central Australia and the Australian summer monsoon. Science 284, 1150–52; Johnson BJ et al. (2005) Carbon isotope evidence for an abrupt reduction in grasses coincident with European settlement of Lake Eyre, South Australia. The Holocene 15(6), 888–96; Miller GH et al. (2005) Ecosystem collapse in Pleistocene Australia and a human role in megafaunal extinction. Science 309, 287–90. 22 Whittell HM (1954) The Literature of Australian Birds: A History and a Bibliography of Australian Ornithology (Part 1). Paterson Brokensha: Perth, p. 24. 23 Whittell HM (1954) p. 68. 24 One of the earliest reported sightings of the King Island emu was by Captain Péron of the Géographe. It was one of ‘five or six’ on a ‘type of butcher’s hook’ in a sealers’ camp. See Whittell HM (1954) p. 66. 25 Muir M (1990) Catherine Eliza Somerville Stow (1856–1940). In Australian Dictionary of Biography. Vol. 12. (Ed. J Ritchie) pp. 113–14. Melbourne University Press: Carlton. Her journalism was published by the Bulletin, Lone Hand and the Pastoralists’ Review. Her most serious anthropological work (on the Narran River Aboriginals) was The Euahlayi Tribe: A Study of Aboriginal Life in Australia, Archibald Constable, London, 1905. The Noongahburrah people was a branch of the Euahlayi group. See also Grimshaw P and Evans J (1996) Colonial women on intercultural fron-

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tiers: R Campbell Praed, M Bundock and K Langloh Parker. Australian Historical Studies 106, 79–95. 26 Stow C (1930) Woggheeguy: Australian Aboriginal Legends. F.W. Preece and Sons, Adelaide, pp. 1–6. (Illustrations by Nora Heysen.) The story goes on to relate the revenge of Dinewan on Goomblegubbon and her friend Bralgah (brolga), and that this is why these birds are still enemies. 27 Libby Robin (2001) The Flight of the Emu: A Hundred Years of Australian Ornithology. Melbourne University Press, Carlton. The journal Emu continues today – now with a subtitle, Austral Ornithology. 28 Gould SJ (1980) Sticking up for marsupials In The Panda’s Thumb. Penguin: Ringwood, pp. 241–2. 29 McCabe J (1910) Australia – A Museum of Living Antiquities. Lone Hand. November, p. 44. 30 Dart J (2007) ‘Emu feathers fired spirits at Beersheba’. Canberra Times, 27 October, ‘Panorama’ p. 21. 31 Fence No. 1, built in 1901–1907 went from just west of Esperance due north to the coast of the Pilbara, but before it was complete the rabbits had broken through, and fence No. 2 was started in 1905, beginning from Bremer Bay, just west of Albany northwards, marking the edge of the wheat belt. 32 The Ajana district, near Kalbarri was where the 1928 estimate and the farmer’s report came from. Ajana is right on the fence-line. Serventy DL and Whittell HM (1962) Birds of Western Australia. Paterson Brokensha: Perth, p. 66. 33 Serventy DL and Whittell HM (1962) p. 66. 34 Information on the prickly pear wars kindly supplied by Henry Nix, pers. com. November 2000. 35 Serventy DL and Whittell HM (1962) pp. 68–69. See also Johnson M (2006) ‘Feathered foes’: soldier settlers and Western Australia’s ‘Emu War’ of 1932. Journal of Australian Studies 88, 147–57, 185–87. 36 Serventy DL and Whittell HM (1962) p. 68. 37 Ratcliffe to GK Baron Hay (Chairman of the Agriculture Protection Board), ‘Emus and black cockatoos’, 27 March 1958. Ratcliffe letters (private collection of the estate of JM Calaby). The later division into short and long billed black cockatoos was the result of the work of Denis Saunders. See Saunders DA and Ingram JA (1998) Twenty-eight years of

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monitoring a breeding population of Carnaby’s cockatoo. Pacific Conservation Biology 4, 261–70. 38 This story is told by Rowley, ‘Nomads of the inland’, in Bird Life. The technical report on the study is Davies, Beck and Kruiskamp, ‘The Results of Banding 154 Emus…’. 39 See David Roshier, this volume, Chapter 5. 40 Allen CR and Saunders DA (2002) p. 348. 41 Bennetts RE and Kitchens WM (2000) Factors influencing movement probabilities of a nomadic food specialist: proximate foraging benefits or ultimate gains from exploration? Oikos 91, 459–67. 42 The Lake Moore Emu Fence was constructed between 1957 and 1959. Serventy DL and Whittell HM (1962) pp. 39, 367. 43 Blakers M, Davies SJJF and Reilly PN (1984) (Eds.) The Atlas of Australian Birds. Melbourne University Press: Carlton; Barrett G et al. (2003) (Eds.) The New Atlas of Australian Birds. Royal Australasian Ornithologists Union: Hawthorn East. On the caveats, see Dunn AM and Weston MA (2008) A review of terrestrial bird atlases of the world and their application. Emu 108, 42–67, esp. p. 52. 44 Davies SJJF (1972) Results of 40 hours’ continuous watch at five waterpoints in an Australian desert. Emu 72(1), 8–12. See also Davies SJJF (1963) Emus. Australian Natural History 14, 225–29; Davies SJJF (1968) Aspects of a study of emus in semi-arid Western Australia. Proceedings of the Ecological Society of Australia 3, 160–6. 45 http://www.didgeridoos.net.au/Bluey’s-painting-story.html. D’Arcy P (1994) (Ed. M Sutton) The Emu in the Sky. National Science and Technology Centre: Canberra; Bhathal R and White G (1991) Under the Southern Cross: A Brief History of Astronomy in Australia. Kangaroo Press: Kenthurst; Rose D, James D and Watson C (2003) Indigenous Kinship with the Natural World. National Parks and Wildlife Service (NSW), Sydney.

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INDEX Aboriginal astronomy 261–2 campsite 167, 169 country 185–204, 262 Dreaming 187–90,199, see also Tjukurpa fire 141, 160–2, 201, 249, 260 forager 169 hunting 103, 169, 189, 234, 262 knowledge see Indigenous knowledge languages 8 population density 162 stone artefact 9, 167 Acacia spp. 47, 201 Adelaide SA 10, 12, 36–7, 104–5, 127–8, 132–3, 141 Adelaide Zoo 105 Africa 15, 51, 53, 90, 155, 243 Africa, northern 15 Africa, South see South Africa agriculture 9, 15, 19–20, 23, 25, 78, 86, 223–4, 234, 236, 246, 256–7, 260 Agriculture Protection Board of Western Australia 257 Ahlquist, JE 208, 220 Aitken, MJ 171, 175 Ajana WA 255 Åkesson, S 94 Albatross 208 Albury NSW 36 Alerstam, T 93–4 Alexander, F 73 Alice Springs NT 16, 26, 28, 55, 59, 61, 135 Allan, R 92 Allen, Craig 259, 264, 266 Allen, J 171 Alpers, D 222 amino acid racemisation (AAR) 155, 175 Anas gracilis see grey teal Andracunie Swamp SA 79

Andrews, Frederick William 123, 129–35, 137, 139, 142–3, 145–6 animism 189–90, 204 anseriform birds (ducks, geese and swans) 149, 162 anthropology 8, 186, 189 Arafura Sea 133 archaeology 8, 161, 169 Archer, M 71, 172 Arctic 12, 25, 90, 155, 243 arid zone 9–11, 13–14, 17, 21, 23, 25–8, 63–4, 130, 147, 150, 157, 161, 164, 166, 206–7, 217, 224, 247, 249 definition 25–7 aridification (continental) see drying out Aristida 58 Armidale NSW 55–6 Artamidae 211, 215 Artamus cyanopterus see dusky woodswallow Artamus personatus see masked woodswallow Artamus superciliosus see white-browed woodswallow Aru Islands 133 Asmus, Martin 78, 92–4 Astheimer, LB 31, 70, 72, 237 Atlas of Australian Birds 260 Atriplex see saltbush, chenopod Auk Journal of the American Ornithologists’ Union 227 Austin, Claude 41, 43 Austin, Robert 125–6, 130, 144 Australasian Ornithologists’ Union see Royal Australasian Ornithologists Union (RAOU) (name changed in 1910) Australia Felix (term used by Thomas Mitchell, 1836) 245 Australia II (America’s Cup) 1983 242 Australian Fourth Light Horse Brigade 253 Australian magpie 208, 211, 215

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Australian Museum (Sydney) 18, 135 Australian National University ACT 16, 157 Australian pelican Pelecanus conspicillatus 11, 75, 77, 95–120 Australo-Papuan region 207, 210 Austria 242 Avise, JC 221 avocet see red-necked avocet

biogeography 17–18, 24 bioregion 188, 192, 195–200 Bird Rose, D see Rose DB Bird, MI 182 bird-of-paradise 208, 211 Birdsville track 3, 80 Birt-Friesen, VL 118 black box Eucalyptus largiflorens 39, 41 black-cockatoo 258 black-tailed native-hen Tribonyx ventralis 23–4, 35–43, 122, 143, 205 black-winged stilt Himnatopus himantopus leucocephalus 12 Blakers, M 266 Bobuna NSW 40 Boland, C 238 Boles, WE 146, 173, 175, 221 Bolger’s Soak 139 Bonaparte Gulf 188 bony herring Nematalosa erebi 102,107 boom and bust (definition) 1–5, 7, 27–9, 223–49 Bornean bristlehead Pityriasis gymnocephala 211 Borneo 211 Both, C 94 bounty system 253, 255–7 Bourgoin, MA 138–9, 142 Bourke’s parrot Neopsephotus bourkii 8 bowerbird 208, 211 Bowles, SW 23–4 Boyce, James 244, 263 Brachychiton paradoxa 197 breeding biology 8, 12, 17–18, 22–23, 28, 50–1, 53–64, 69, 77, 81, 84, 89–91, 98–103, 107–9, 113–14, 122, 130, 142, 160, 165, 170, 225–30, 232, 247, 259–60 breeding site 11–14, 17–18, 30, 56–7, 84, 89–91, 97–8, 105–6, 247, 259 Breen, G 115, 117 Brewarrina NSW 156 Briggs, S 94 brine shrimp 14 Brisbane Qld 10, 134 Britain 22, 50, 241, 243, 245, 250 British Antarctic Territories 243

Badman, FJ 119 Baker, John 50 Baker, L 70 bald eagle 243 Ballarat Vic 41 Balouet, JC 172 banded stilt Cladorhynchus leucocephalus 11–14, 18, 30 Bangate NSW 251 Barcoo Bantam see black-tailed native-hen Barker, FK 220 Barker, WR 72, 221, 236 Barkly Lakes NT 102 Barkly Tableland NT 82 Barnard islands Qld 133 Barnard, P 93 barramundi 195 Barrenbox Swamp NSW 86 Barrett, G 266 Barrier Range NSW 80 Barrowclough, G 220 Barry, S 236 Bartlett, Mr 125 Baudin, Nicolas 250 Baudin’s black-cockatoo Calyptorhynchus baudinii 250, 258 bauhinia Lysiphyllum cunninghamii 161, 194–5 Bedourie, Qld 58 beetle larvae 229–30 Belperio, AP 182–3 Bennetts, RE 93, 259, 266 Bentley, G 237 Berthold, P 93 Bhathal,R 266 Big Desert Vic 41 big river country 188–9, 195, 197–200

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British Museum, London UK 129, 134 British National Archives 141 Broken Hill NSW 53 brolga Grus rubicunda 185, 187–8, 190, 193 bronzewing, flock see flock bronzewing Brook, BW 175 Brooks, AS 175 brown creeper Certhia Americana 207 Brown, J 237–8 Brucellosis-Tuberculosis Eradication Campaign (BTEC) 1986 191 brush turkey 18 budgerigar Melopsittacus undulates 49, 51, 122, 205–6 Bullocky (Aboriginal man) 138 Bulloo, NT 102 Burbidge, AA 73 Bureau of Meteorology 236 Burgin, S 34, 270 Burgman, M 5 Burley, NT 60, 72 butcherbird 211, 215 Buttemer, WA 31, 70, 72, 237 Byrne, R 238–9

Central Australia 10–11, 42, 49, 63, 100, 107, 135–6, 140, 149, 153 Centre for Arid Zone Research (CAZR) Alice Springs NT 16, 26, 28 Channel Country Qld 39, 114 channel-billed cuckoo Scythrops novaehollandiae 186, 199–200 Chapman, FRH 115–16, 118 chenopod shrubland 153, 161, 165, 168, 249 China 16 chough, white-winged see white-winged chough Christidis, L 221 Cladorhynchus leucocephalus see banded stilt Clare valley SA 80 Clarke, MF 72, 93 Clayton, G 71, 172 Cleland, JB 144–5 climate change 3, 5, 91, 147–8, 160–2, 165–6, 200–4, 220 Cloudsley-Thompson, JL 25, 33 Clout, MN 114 clutch size 99, 150 Clutterbuck Hills WA 64 Clutton-Brock, T 238–9 Cobourg Peninsula 10 Cockburn, A 237–8, 263, 265 Cockerell, James 134–5 Cockerell, John Thomas 134 Coddington, CL 263 Cold War 16 Coleman, DC 176 Coleraine Vic 41 Collier, R 31 Collis, B 32 Commonwealth Scientific and Industrial Research Organisation see CSIRO connectivity 185, 192 coolibah Eucalyptus coolabah 65 Coonawarra SA 152 Coongie Lakes 78, 82, 95–7, 101–2, 111 Cooper Creek 77, 79, 81, 101–2, 105–6, 113, 128, 130, 163, 164 Cooper, A 172 cooperative breeding 225–30, 232

Calaby, JH 20, 52, 71, 174, 263, 265 callop Macquaria 107 Calvert Expedition 1896 12, 135 Calvery, HO 174 camel 135, 141 Campbell, Archibald James 138, 146 Canberra ACT 24, 31, 70, 224, 232, 256 Cane, S 70 Cann, JH 183 Canning Stock Route 139 Cannon, BE 12–13 Carmen, WJ 237 Carnaby, IC 13, 23, 31, 50, 70 Carnaby’s black-cockatoo C. latirostris 258 Carnarvon WA 50 cassowary 148, 255 cat 135–6 catfish 185, 187–8, 190, 192–3 cattle (livestock) 2, 8, 37, 46, 136–7, 140, 142, 167, 191, 201–2 Central America 227

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Coorong SA 80, 103–4, 115, 262 Coppack, T 94 Corcorax melanorhamphos see white-winged chough Corney (vermin control officer) 20 Costelloe, JF 116–17, 119–20 Council for Scientific and Industrial Research see CSIR courtship 48, 59, 99 Cowling, S 221 creching 99, 103 crested bellbird Oreoica gutturalis 48 crocodile 163, 194–5, 199 crow 190, 209, 210–11, 226, 255 CSIR 16–17 CSIRO 16–17, 19–21, 24, 28, 48, 55, 59, 78, 246, 257–8 cuckoo, channel-billed see channel-billed cuckoo cuckoo-shrike 207–8 Cuddie Springs NSW 153, 156 Cunnamulla Qld 86 Cupper, ML 180 currawong Strepera spp. 208, 211, 215 Curtin AL 92, 116 cyclone 29

DeVogel, SB 179 Diamantina River 77, 80 Diamond, Jared 233, 239 Dickman, CR 34 dinewan see emu Dingle, H 92–4 dingo 135–7 diprotodon 161, 163 DNA 8, 30, 206, 213–18, 228 Dodson, JR 173, 175 Doerr, E 94 Doerr, V 94 Donaldson, T 73 Double, M 238 Dowerin WA 23 Drake, VA 94 Dromaius novaehollandiae see emu dromornithid 15, 147–183 drought 1–3, 37, 68, 78, 86, 92, 98, 104–5, 108, 110, 130, 136–7, 223–4, 230–3, 236, 249 drying out 216 Drysdale, Russell 23, 33 dualism (separation of nature and culture) 186 Dublin, Ireland 132 duck 29, 77, 79, 82, 90, 91, 108, 149, 155 Duncan, J 180 Dunlop, N 118 Dunn, AM 266 Dunn, P 238 dusky moorhen Gallinula tenebrosa 38 dusky woodswallow Artamus cyanopterus 213 dust bowl (soil erosion) 16

D’Arcy, P 266 Darling River 10, 39, 87, 102, 106, 152, 154, 156–7, 160, 166, 251 Dart, J 265 Dartmoor UK 243 Darwin, Charles 121, 143, 226, 237 dasyurid 167 Davies, PH 72 Davies, SJJF 55–7, 71–2, 174, 246–8, 258, 263–4, 266 Davis, J 33 Dempsey’s Lake SA 155 Denmark 243 Department of Agriculture, Western Australian 256, 260 Depot Glen 127 Derby, Lord 128–9 desert country 4, 7–34, 195 desert river 28 desertification 15, 164

earthworm 229–30 eastern grey kangaroo Macropus giganteus 229 eastern koel Eudynamus orientalis 186, 196 Eckert, J 115 ecology 8, 16, 22, 29, 54, 57, 88, 91, 95, 104, 121, 148–9, 151, 165–6, 190, 192, 203, 206–7, 212–13, 215, 225, 227, 232–6, 241–62 Edenhope Vic 41 Edwards, SV 221

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I n de x

eggshell, fossil 147, 151–63, 165, 167–9, 249 Egypt 16 Ehleringer, JR 176 El Niño Southern Oscillation (ENSO) 4, 223, 231, 236 Australia as ‘El Niño continent’ 4 Elanus scriptus see letter-winged kite elephant 243 Emiliani, C 171 emperor penguin Aptenodytes forsteri 243 emu Dromaius novaehollandiae 11, 17, 123, 148–51, 153–61, 165, 167–9, 195–6, 241–62 Emu Fence WA 20, 253, 255, 260 Emu War 256 emu, Kangaroo Island Dromaius baudinianus 250, 260 emu, King Island Dromaius ater 250 emu, Tasmanian Dromaius novaehollandiae diemenensis 244, 250 Emu: Austral Ornithology (formerly Emu) 251 Endler, JA 219, 222 Engel, S 93 Enneapogon 58 Eragrostis 58 Eriachne 58 Ericson, P 220 erosion, soil/wind 16, 163, 167 Estrildidae (grass finches) 57 estuarine country 188 ethics 186 ethology 23, 45 eucalypt woodland 153, 224 Eudynamus orientalis see eastern koel Eurocentric thinking 25, 252 Europe 2–4, 10, 75, 83, 96, 104, 123, 223, 227, 242, 250, 252, 261 Eutheria see mammal, placental Evans,J 264 evolutionary biology 4, 8, 28, 30, 38, 113, 205–22, 225, 227, 229, 251–2, 260 Exmouth GC-17 core 161, 164 extinction 3, 9, 15, 27, 30, 140, 147–70, 191, 220, 244, 248–9 Eyre Creek 82

Eyrean fauna 136 fairy-wren 208, 211, 227–8 fairy-wren, splendid see splendid fairy-wren Falco hypoleucos see grey falcon falcon, grey see grey falcon Farmer, Charles 126 Farner, DS 22–5, 33, 71, 236 Faulkner, A 118 Federation (of Australian colonies) 1901 243, 245, 250–1 Field, Barron 242, 262 Field, JH 173, 175–6, 180 finch, zebra see zebra finch Finke River NT 153 fire 121, 141, 160–2, 168, 201, 249, 260 Fischer, H 180 fish 77, 96–8, 100–3, 105–8, 113–14, 163, 185, 187–8, 190, 192–4, 199 Fisher, CT 31 fishery 17, 104 Fitzsimmons, KE 179 Fivebough Swamp NSW 86–7 Flannery, TF 161, 177, 242, 263–4 Flinders Ranges SA 80 Flinders University SA 105, 155 flock bronzewing Phaps histrionica 11 flood 1–3, 14, 39, 65, 77, 80–2, 84–7, 92, 97, 100–2, 107–15, 121, 136, 193–4 flux 192, 203 Fogel, ML 176 follow the leader behaviour 111 forage 11, 39, 59, 67, 105, 151, 207, 229, 234 Foran, BD 5 Ford, HA 63, 72, 93 Ford, JR 221 forester kangaroo Macropus giganteus 244 Forshaw, JM 144–5 Fowlers Gap Research Station NSW 53–4, 58, 80 Fremantle WA 10 Friedel, MH 5, 73 Frith, HJ 19, 21–2, 24, 33, 45, 55, 71, 78, 84, 92, 174, 263

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frog 133, 139 Fry, B 176 Fulbright Scholarship 14 Fullagar, PJ 144 Fullagar, R 175 Fuller, PJ 73

grasswrens, striated see striated grasswrens grebe 108 Greenberg, R 93 Greenslade, PJM 72, 221, 236 Gregory National Park NT 203 Gregory, JW 162, 179 grevillea 47 grey falcon Falco hypoleucos 40–1 Grey Range 127 grey teal Anas gracilis 11, 39, 51, 75–94 Griffioen, PA 93 Griffith NSW 21, 55, 86 Griffiths, TR 31–2, 73, 204 Grimshaw, P 264 Grocke, DR 176 Groth, J 220 ground parrot 124–6, 128, 138, 141 Grus rubicunda see brolga gum tree Eucalyptus spp. 39, 207 Gunbower NSW 87 Gwinner, E 93

galah 20 Galiano, Henry 148, 150, 172 gallinule 38, 40 Gardner, J 236 Gascoyne River WA 50, 125 Gawler Ranges SA 129–30, 133, 137–8, 142 Gaynor, A 73 Geelong Vic 36, 41 Gentilli, J 17–18, 32 Genyornis newtoni 15, 147–83, 249, 264 geography 17–18, 24, 68, 147, 153, 206, 216, 227 Geopsittacus 125 Georgina River 97 German Federal Research Organisation 23 German language 49 Germany 23, 45, 49, 53 Gibney, HJ 144 Gibson Alfred 46, 68 Gibson Desert WA 46–8, 68 Gilbert, John 10, 31, 36, 43 Giles, E 70 Gillen, J 120 Gillespie, R 175 Gilmore, D 71 Glauert, Ludwig 12–13, 18, 31 gnow see malleefowl golden perch 107 golden wattle Acacia pycnantha 244 Gondwana 18, 148, 225, 251 goomblegubbon see bush turkey gooseberry (NT) Physalis minima 202 Gostin, VA 182 Gould, John 10–11, 31, 36, 43, 57, 124–6, 128–9, 143–4, 227, 238 Gould, Stephen Jay 252, 265 Graham (ornithologist) 227, 237 granivory 54–5, 57

Haddon, South Australia 3 Haebich, A 73 Hagstrum, JT 93 Hails, JR 182 Hall, Robert 227, 237 Halosarcia see samphire, chenopod Hamilton Vic 41 Hamilton, W 226, 237 hardyhead Craterocephalus eyresii 107 Harper, M 263 Harris, JI 144 Harrison, P 117 Harvey, Graham 189, 204 Hattersley, PW 157, 176 Hau, M 237 hawk 65, 190 Hedenström, A 94 Heijnis, H 181 Heinsohn, RG 223, 236–9 Hercus, L 73, 115 Hermannsburg Mission NT 141 Hesse, PP 179, 181 Higgins, PJ 31, 93, 116–17, 119, 143, 183, 221

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I n de x

high-performance liquid chromatograph (HPLC) 155 Himnatopus himantopus leucocephalus see blackwinged stilt Hiscock, P 171 history 4, 8–10, 15, 29, 52, 130, 137, 148, 154, 162, 168, 215, 226 Hobart Tas 10, 244, 256 Hobbs, John 40–1, 43 Hocknull, SA 181 Hodges, F 73 Holling, CS 27, 34 honeyeater 49, 208, 211 Hopkins, AJM 5 Horn Expedition 1894 135–6 horse 3, 38, 126, 202, 243 Horsefield, T 237–8 Horsham Vic 41 Horton, DR 152, 173 How, RA 5 Hudson, PJ 116 human predation 148, 153, 166, 170, 220 human settlement (pre-European) 154, 161 hybridisation 212, 214

Jacobs, SWL 176 Jaensch, RP 114, 116–7, 120 James, D 266 Jamieson, IG 93 Jangarla see white dragon jay 210 Jenkins, CFH 13, 31 Jennacubbine WA 24 Johnson, BJ 157–8, 164, 175–9, 264 Johnson, CN 170, 172, 183 Johnson, M 265 Johnston, G 105, 117–18, 120 Jones, J 31 Jones, KR 60, 72–3 Jones, R 171 Joseph, L 7, 33, 120, 145, 205, 221 Journal of Arid Environments 25 Jull, AJT 174 kagawuli see yam kaja (NT desert country) 196–7 Kalbarri WA 253 Kalkannie WA 20 Kallakoopah Creek 163 kangaroo 17, 52, 207, 241–2, 244–5, 250, 262 Kangaroo Island 36, 250, 260 Kangkinang, Big Mick 188, 198 karil Cucumis spp. (probably trigonus) 202 Kaufman, DK 174 Keartland, George 12, 135–6 Keast, JA 50, 70, 221, 236 Keep River National Park NT 203 Kerle, JA 70 Kershaw, AP 178, 181 Kerwin, B 115, 117 Kew UK 141 Kiata Vic 19 Kikkawa, Jiro 55–6, 71 kilipi Leichardtia australis 202 Kimberley region WA 12, 48 King, JR 33, 71 kingfisher 208 Kingsford, RT 28, 34, 79, 92–4, 116–19 Kinsella, John 139, 146 kinship system 186–7, 191 Kitchens, WM 93, 259, 266

Ice Age see last glacial maximum Immelmann, Klaus 23, 25, 45, 48–50, 52, 55, 59, 61–2, 70 inclusive fitness 226, 233 India 15–16, 211 indicator species 24, 188, 198 Indigenous knowledge 19, 67, 190, 201 Indonesia 196, 200 infrasound 30, 85 Ingram, JA 265 inland sea 75, 97, 127 Innamincka SA 81, 101–2, 138 interglacial 148, 163, 166, 168 International Ornithological Congress 21, 24 Irestedt, M 221 irruption 14, 22–3, 37, 41, 68, 206 isotopic signature 30, 154, 158–60, 162 Israel 15, 27 Ivanhoe NSW 40, 42

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kiwi 247 Klaassen, M 93–4 Klein, RG 173 Klomp, M 92 Knight, Frank 43, 148, 172, 221 Knopwood, Robert 244 koel, eastern see eastern koel Koenig, WD 237 kookaburra 251 K-T (Cretaceous-Tertiary) boundary 209 Kukutha people 262 Kununurra WA 48

letter-winged kite Elanus scriptus 122, 205 Lewin, JW 237 Lewis, Darrell 204 Lewis, JW 130, 145, 153 Lindenmayer, DB 5, 236 lion Panthera leo 228, 241, 243–4, 253 Lipp, HP 93 Little Desert Vic 19 logrunner 211 London UK 22, 123, 128–9, 134, 242 London Zoological Gardens 123 long-haired rat (native rodent) 206 Longmore, ME 181 Longmore, NW 146 Lord, G 175 lowan see malleefowl Loyn, R 221 lucerne (crop) 23 Luly, JG 179 Lunney, D 34 Lüthi, D 180 lyrebird 208, 211 Lysiphyllum cunninghamii see bauhinia

La Niña (high rainfall) episodes 101, 113 La Trobe University Vic 56 Lachlan River 39 Lake Blanche SA 80 Lake Callabonna SA 13, 147, 153, 161 Lake Cawndilla 97 Lake Eyre basin 39, 77, 102, 106–7, 112, 114, 152, 156, 162, 164, 166 Lake Eyre 97, 106–7, 131, 156–8, 162–5, 169 Lake Frome SA 13, 156–7 Lake Goolangirie 197 Lake Grace WA 13 Lake Gregory WA 102 Lake Hope SA 79–80, 106–7 Lake Lewis NT 153 Lake Machattie 97, 114 Lake Toontoowaranie 82 Lake Torrens SA 80, 152 lakes, ephemeral 14, 28, 109, 140, 151, 163, see also wetland habitat Lambourne, M 31 Lancefield Vic 164 last glacial maximum (LGM) 158, 216, 243 latatj see Storr’s monitor Latz, Peter 161, 178 Laughing Jackass see kookaburra Lawson Whitlock see Whitlock, FB Lawson Le Souëf, Dudley 142 Leckey, T 73 Leeton NSW 86–7 Legge, S 238 Leipoa ocellata see malleefowl Lena River, Siberia 12 Letnic, M 121, 143

M‘Lennan, Charles 248 MacGillivray, DWK 73 MacGillivray, W 116 MacKellar, Dorothea 1–2, 5 Maclean, GL 53–4, 71 MacMillen, RE 54, 71–2 macropod 164 Magee, JW 174, 179 magpie 208, 211, 215 magpie-lark 208 mallee country 11, 17–19, 41, 248 malleefowl (Leipoa ocellata) 11, 18–22 Malurus 227 Malurus splendens see splendid fairy-wrens mammal 3, 52, 55, 82, 228, 244, 248–9, 252 Mann, K 174 Manngaiyarri, Old Jimmy 198 Manning, A 236 Manorina melanocephala see noisy miner march fly 194–5 Marchant, S 31, 93, 116–17, 119, 183 mark-recapture studies 258 Maroulis, J 180 Marra, P 93

292

I n de x

Marshall, AJ 22–3, 28, 33, 45, 50, 62, 70–1, 236 marsupial 3, 23, 52–3, 163–4, 252 martin 210 Martin, HA 181 Martin, PS 173 masked woodswallow Artamus personatus 211–16, 218–19 Masson-Delmotte, V 180 McAllan, IAW 116, 119 McCabe, Joseph 252, 265 McCoy, WD 175, 177 McDonnell, R 94 McGilp, JN 13–14, 31–2 Meekatharra WA 55 meerkat Suricata suricatta 228 megafauna 161, 164, 170, 220 megapode 18–19 Megirian, D 172 Melbourne Vic 10, 36, 41, 60, 231 Menindee Lakes 97 mesembrianthemum 128 Metatheria see marsupial migration, seasonal 12, 17, 67, 76, 78, 81, 84, 88–9, 91–2, 108, 111, 115, 213, 258, 260 Mileura WA 55–6 Milky Way 262 Miller, Alden 15 Miller, GH 152, 155–62, 165–6, 172, 175–82, 264 mineritchie Acacia cyperophylla 7–8, 29 Minister for Defence, Commonwealth 255–6 Minna Minna WA 65 mintariyij Nymphaea violacea 202 Miocene (era) 149, 216 Mirmir (Aboriginal woman) 187 mitochondrial DNA (mtDNA) 213–17 Mogadishu 231 mollusc shell Coxiellada 163 monarch flycatchers 211 Monash University Vic 23 monitor, Storr’s see Storr’s monitor monotreme 3, 252 monsoon, summer 29, 76, 84, 158, 161, 164, 166, 186, 192–3, 200 Moolawatana SA 13 Moonaree SA 131

Moree NSW 86 Moreton Bay 10 Morgan, AM 13–14, 31–2 Morton, SR 28, 34, 45, 70, 72, 117, 120, 237 Moss, P 178 mound-building 17–20, 166 Mt Farmer district WA 125 Mt Jagged SA 132 Muir, M 264 mulga Acacia aneura 46, 65, 135 Mulka SA 80, 119 Mumme, RL 237 Muranji track 197 Murie, James 143–4 Murnia rockholes SA 131 Murray river 39, 87, 102–4, 106, 152, 154, 156–7, 160, 166, 262 Murray, Peter 149–50, 161, 172–3, 178, 183 Murray-Darling basin 87, 102, 106, 152, 154, 156–7, 160, 166 Murray-Wallace, CV 183 Murrumbidgee river 39, 71 Museum Victoria (Melbourne) 130 mussel Velesunio 163 Mutitjulu 70 myrrlumbing see night parrot Nagambie Vic 36 Namib Desert 51 Nanson WA 253 Nanson, GC 166, 179–81 Naracoorte SA 152 narwhal Monodon monoceros 243 National Museums, Liverpool UK 128 national symbol 241–66 native companion see brolga native-hen, black-tailed see black-tailed native-hen Tribonyx ventralis Natural Selection, Theory of 30, 122, 218–19, 225–6 Ned’s Creek 139 Nelson, B 117, 119 Neopsephotus bourkii see Bourke’s parrot neuroanatomy 30 New Guinea 78, 196, 200, 207, 211, see also Papua New Guinea New Hebrides (later Vanuatu) 22

2 93

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oscine (songbird) 207, 209–11 ostrich Struthio camelus 148, 151, 155, 158, 248 Outer Harbor SA 105 Overton, Gary 81 Oxford University UK 15

New South Wales 19, 21, 40, 42, 54–5, 78, 80, 86, 115, 128, 151, 245 New South Wales, western 40, 42, 54, 151 New Zealand wren 208–9 Newman, RJ 33 Newton, Alfred 170 Newton, I 94 Ngaanyatjarra people 46–7 ngaringarin Pterocaulon serrulatum 202 Ngarrindjeri people 262 Nichol Spring WA 138–9 Nicholls, FG 21, 33 night parrot Pezoporus occidentalis 11, 121–146 Ningaloo WA 153 Nix, HA 236, 265 Noble, JC 172, 183, 264 noisy miner Manorina melanocephala 208 nomadism 64, 76, 85, 88–91, 108, 245, 258, 260–1 Nonning SA 129–30, 137 non-passerine 15, 208 Noongahburrah people 250–1 Norman, FI 93, 116, 118–19 Norse saga 243 North America 4, 14, 207, 227 North, AJ 145–6 Northam WA 126 Northampton WA 255 Northern Territory 60, 82, 102, 185 Noy-Meir, I 27, 34 Nullarbor Plain 153 nyii nyii (WA), nini (NT) see zebra finch

Pacific region 209, 211, 224 palaeobotany 249 palaeoecology 149–54 palaeontology 9, 12, 15, 29, 155 palaeo-ornithology 15 Palmer River NT 140 Panheleux, M 174 panicum 59 Papua New Guinea 196, 200, 207, 210–11, 220 paraphyly 214–15 Parker, Katie Langloh 250, 265 Parker, P 71 Parker, SA 117, 119 Parkes, KC 71, 236 parnparnpalala see crested bellbird Paroo River 86–7, 102 parrot 8, 11, 121–46, 208 passerine 209–11, 215 pastoralism 9, 15–17, 19, 25, 41, 46, 80, 136–7, 142, 245–6 patchiness 187, 192, 195–7 Patjarr Creek WA 64–5 Patjarr WA 46, 64 Paton, DC 104, 118, 120 Paton, JB 144 Pearce, Sir George 255–6 Pelecanus conspicillatus see Australian pelican Pelican Point SA 105 pelican, Australian see Australian pelican Peltops 211 penguin 208, 243 Penguin Island WA 105 Perfita, N 237 Perrin, MR 72 Péron, Capt. 264 Perth WA 10, 12, 17, 105, 124, 141, 253 petrel 208 Petrie, S 94

O’Brien, M 118, 120 O’Connell, JF 171 Ohlsson, J 221 Oodnadatta SA 141 oology 18, 140 opportunistic breeding 14–18, 48–57, 61–4, 114, 249 opportunistic feeding 105, 114, 151, 160–2, 246, 259 Orientos Qld 82 ornithology 7–34, 41, 45, 50, 55, 63, 89, 125, 133–4, 137–8, 141–2 ,207–8, 216, 226–7, 251

294

I n de x

Pezoporus (formosus) 126 Pezoporus occidentalis see night parrot Phaps histrionica see flock bronzewing Phillip, Governor Arthur 250 philopatry 89 photosynthetic pathways 157 phylogeny 149, 206, 208 phylogeography 206 physiology 22–3, 25, 28, 50, 52, 54, 83, 88–90, 158 Piersma, T 94 pigeon 138, 208 pink-eared duck Malacorhynchus membranaceus 79 Pinyerinya Pool 139 Pitelka, FA 237 Pitjantjara people 8, 47–8 pitta 209 Pizzey, GM 35, 43, 221 platypus 227 Pleistocene (era) 160, 162, 215–16, 220 Policemans Point SA 104 pollen record 161 polygamy 227 polyphyly 214 population biology 11, 20, 27–8, 30, 57, 59, 77, 91, 97, 99–100, 102, 105–6, 108, 110–11, 142, 151–2, 165, 168–9, 206, 217–20, 224, 227–8, 230, 232, 244, 246, 250, 257, 260–1 porcupine grass Triodia 123, 131, 136 Port Adelaide 130, 133 Port Augusta SA 80, 130, 152, 155, 157, 168 Port Essington 10 Port Jackson 10 Port Pirie SA 157, 166–7 Porter JL 92, 116–17 Porter, Dorothy 140, 146 post-war period 14–16, 227, 253 powder down (feathers) 207, 211 Poynter, C 94 Preservation Creek 127 Price, DM 179–80 Price, GJ 181 prickly pear Opuntia spp. 249, 255 Prideaux, GJ 173, 181, 222

Pritchard, JC 116 Prosser, IP 173 Prototheria see monotreme Puckridge, JT 101, 116–20 Pulido, F 94, 119 Pulkara, Daly 185 Pullman USA 24 pulse and reserve model 27 purlkal Vitex acuminata 202 purple swamphen Porphyrio porphyrio 38 Qantas (airline) 242 Quaternary (era) 148, 154, 162 Queensland 7, 37, 39, 58, 77, 81–2, 86, 101, 130, 133, 152, 255 Queensland Museum 134 Queensland, south-west 58, 77, 81–2, 86, 101, 152 Quilpie Qld 82 rabbit 17, 125, 137–8, 253, 256 rabbit-proof fence 253, 256 radiocarbon dating 155 radio tracking (satellite) 30, 69, 78, 84–5, 90, 258 rail (bird) 35, 37–8, 108 rainbird 185–204 rangelands science 17, 25–8 Ratcliffe, Francis 16, 21, 24, 32, 50, 55, 257 ratite 148, 150, 247–8, 251, 269 raven 211 Read JL 109–10, 116–19 red-necked avocet Recurvirostra novaehollandiae 12 reed-warbler 210 Reid, JRW 28, 70, 73, 94, 95, 116–17, 119 Reilly, PN 266 resilience 25–8, 133, 148, 170, 220 rhea 248 Rich, Patricia Vickers 15, 149–50, 152, 161, 171–4, 178, 183 Ride, WDL 15 rifleman (New Zealand wren) 208 riparian zone 151, 163, 165 Rippey, E 118 Rippey, JJ 118

2 95

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Ritchie, J 264 river channel 151, 163 Riverina NSW 21, 55, 78, 86–7 Roberts, Bluey 262 Roberts, George 3 Roberts, RG 171, 174, 180, 182 Robertson Land Acts 1861 (John Robertson) 245, 263 Robertson, A 92 robin 207–8, 210 Robin, L 1, 7, 31–3, 70–1, 73, 204, 239, 241, 263, 265 Roby, DD 118 rodent 54, 57, 206 Rogers, K 94 Rose, DB 115, 185, 204, 239, 251, 266 rose, English 245 Roshier, DA 28, 73, 75, 92–4, 117, 119–20 Rottnest Island WA 12 Rowley, ICR 227–8, 237–9, 266 Royal Australian Artillery 256 Royal Australasian Ornithologists Union (RAOU) 251 Royal Commission into pastoral industry 1897 137 Royal Geographical Society (London) 128 Royal Society of South Australia 132 Ruello, NV 119 Russell, E 237–8 Rwanda 233–4 Ryan, Charles 129

Scythrops novaehollandiae see channel-billed cuckoo Sears, Paul 15–16, 32 seasonality 2, 4, 26, 28, 54–7, 61–4, 84, 88–91, 108, 151, 157, 159, 165, 193, 213, 248–9 Seattle USA 24 seed-eating 54–9, 61, 66–9, 121, 123, 131, 137, 151, 160, 248 Serventy, DL 17–20, 22–5, 28, 34, 45, 50–1, 53, 56, 62, 71–2, 236, 253, 265–6 Sesbania formosa 17 see white dragon sexual selection 215, 229–30 shamrock, Irish 245 sheep 127, 129, 137, 253 Sheldon, F 118 Shepparton Vic 57 Sherratt, T 31, 204 shorebird 11–12, 77, 108 Short, SA 179 Siberia 12–13 Sibley, CG 33, 207–8, 220 silvereye (Zosterops lateralis) 210 Simpson desert 82, 127 Simpson, J 73 Skutch, Alexander 227, 237 Smith, MA 1, 147, 171, 174, 179, 181, 264 soldier settlement scheme 11, 19, 253 Sonnenschein, E 93 Soobramoney, S 72 Souter, G 262 South Africa 53, 243 South America 224 South Australia 7, 11–13, 19, 36, 42, 77–8, 80, 95, 101, 103, 123, 128–32, 136, 152, 166, 262 South Australian Museum 123, 129–31 South Australian Ornithological Association 13 Southern Cross WA 255, 259 sparrow 125 Spencer Gulf SA 129, 166 Spencer, WB 136, 145–6 spinifex Atriplex spp. 46–7, 122, 130, 135, 138, 141, 200, 249 splendid fairy-wren (Malurus splendens) 228 Stafford Smith, DM 5, 28, 34, 72, 117

salsolae 128 saltwater country 188, 198–200 saltbush Atriplex 53, 157, 161 samphire Halosarcia 14, 128, 131, 151, 157, 165–8 Sandringham Qld 58 Sanford, WA 126 Sauer, Gordon 144 Saunders, Debra 236 Saunders, Denis A 5, 259, 264–6 savanna woodland 186, 201 Schodde, R 63, 72, 221, 236 Schrader NW 42–3 Schultz, M 94 Sclater, Philip Lutley 143–4 Scottish nationalism 243

296

I n de x

Stafford, A 146 Stanback, MT 237 stilt, banded see banded stilt Cladorhynchus leucocephalus stilt, blackwinged see blackwinged stilt Stirling, EC 144, 147, 150, 170–1, 173, 178 Stirton, RA 15 Stokes, Edward 92 Stonehouse, B 71 Storm Boy 103 Storr’s monitor Varanus storri ocreatus 199 Stow, Catherine 250–1, 264–5 Straw, B 71–2 Strepera spp. see currawong striated grasswren Amytornis striatus 206 Struthio camelus see ostrich Strzelecki desert 76, 78–80, 163 Stuart, John McDouall 10, 128, 138 Sturt Creek WA 102 Sturt, Charles 10, 36, 43, 75, 126–9, 138, 144 suboscine 209 sugarbag native honey 199 Sutton, M 266 Sutton, P 115 swallow 210, see also woodswallow swan 149 Swan Hill Vic 86 Swan River colony 10, 36 Switzerland 21 Sydney NSW 10, 17, 133–5, 250 Sydney, Lord 250 systems, natural 3, 27–9, 77, 84, 101–2, 108, 113–14, 147–8, 151, 191, 194, 196, 203–4, 249, 259

Tilt, RA 71 Timor Sea 10 tipil (plant) 199, 202 Tirari desert 163 Tjukurpa 47, see also Aboriginal Dreaming Tolcher, HM 5 treecreeper 207, 211 Tribonyx ventralis see black-tailed native-hen Trinca, M 73 Triodia 131, 141 Trueman, CNG 175 Truran, J 31 turkey 18, 190, 251 Turner, DB 119 turtle, freshwater Chelidae 163 Tyler, MJ 119 Tyndale-Biscoe, CH 71 Uluru-Kata Tjuta National Park 48 UNESCO 15 unicorn 241, 243–5, 250, 253 United States of America 16, 23 University of Adelaide SA 104 University of Bielefeld, Germany 23 University of California, Irvine USA 54–5 University of Cambridge UK 17 University of Colorado USA 155 University of Washington USA 22, 24, 157 University of Western Australia 17 Upper Chapman WA 253 van der Kaars, S 178 Van Der Merwe, NJ 176 Van Diemens Land (later Tasmania) 244 Van Tets, GF 171–4 Varanus storri ocreatus see Storr’s monitor variability 3–4, 26, 29, 46, 52, 84, 91, 98, 100–1, 113, 148, 153, 160, 162, 164–5, 187, 192, 224, 234, 247, 258, 261 Veitch, CR 118 vermin board 20, 253, 255, 257 vermin fence 20, 254 Vestjens, WJM 117–18 Vickers Rich, P see Rich, Patricia Vickers Victoria 11, 19, 26, 36, 41–2, 57, 105, 130, 164, 248 Victoria River 185–6, 188, 195, 200 Victoria River Downs NT 192

Taeniopygia guttata see zebra finch Tanami desert 188 Tasmania 244, 250, 260 teal, grey see grey teal Tedford, RH 15, 179 Tench, Watkin 250 Tenison Woods, Julian 171 Tertiary era 15, 209, 215 thickening see woody weeds Thiele, C 118 thistle, Scottish 245 Thompson, MC 146 Thursday Island 134

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Vigna lanceolata see yam Vigors, NA 237–8 vocalisation 213, 215 Vogel, JC 176 Von Schirnding, Y 176

White, R 263 White, Samuel 133–5, 137–8, 140 White, Samuel Albert 138, 140, 145–6 white-browed woodswallow Artamus superciliosus 207, 211–16, 218–19 white-eye 210 white-faced whistling-duck 90 Whiten, A 239 white-winged chough Corcorax melanorhamphos 223–39 Whitlock, FB Lawson 140–2, 145–6 Whittell, HM 145–6, 264–6 Whyalla SA 168 Wickham River NT 185–6, 188 Widmer, M 94 wild dog Lycaon pictus 228 Wilke, T 221 Williams, DLG 152, 155, 172–5, 181, 183 Williams, L 33 Williams, MAJ 221 Willunga SA 132 Wilson, H 143–4, 146 Wilson, M 94 Wilyurukuruku see Bourke’s parrots Wimmera-Mallee (north-western) Vic 19, 26 Windich Spring 139 Windorah Qld 81 Wingfield SA 105 Winpilin, Riley Young 201 Wise, FJS 257 Wodonga Vic 36 Woenne-Green, S 70 Wolff, EW 180 Wood Point SA 157, 166, 168–70 woodswallow 205–22 woodswallow, dusky see dusky woodswallow woodswallow, masked see masked woodswallow woodswallow, white-browed see white-browed woodswallow woody weeds 201 World War I 9, 18, 253 World War II 9, 14–18, 227 wren 206–9, 211, 227–8

Walgett NSW 250 Walker, Alfred 138 Walker, BH 34 Walker, KF 117–18 Wallaroo WA 168 Wallraff, HG 93 Walsh, F 73 Warburton River 163 Warren, P 20 waterbird 11, 29–30, 35, 53, 75–9, 84, 88, 98, 106, 108, 111, 113, 208 Waterman, MH 116–18 Watson, C 266 weather, extreme 5, 236 Webb, GE 181 Webb, S 180 Webling, DD’A 145 Wells, Mr 135 Wells, RT 179 Werribee Vic 41 Western Australia 11–12, 16–19, 23, 26, 46, 48, 50, 55, 78, 102, 105, 124–5, 130, 134, 138, 141, 153, 228, 248, 253–7, 259–60 Western Australia, south-west 13, 153 Western Australian Museum 12, 15 Western Desert NT 48, 100 Western District Vic 41 Westoby, M 27, 34 Weston, M 266 wetland habitat 77–8, 80, 82, 84–5, 87–8, 90, 98, 100–1, 103, 106, 110, 112, 114–15, 163, see also lakes, ephemeral wheat 11, 17, 26, 37, 249, 253–6, 259 wheatbelt WA 11, 13, 17, 26, 253–6, 259 Wheeler, Roy 41, 43 Whetton, PH 92 white cockatoo 190 white dragon (tree) Sesbania Formosa 195 White,G 266 White, HL 140

298

I n de x

Wright, Judith 242, 263 Wroe, S 181

yeoman ideal 245, 261 Yu, S 73

Yalgoo WA 20 yam Vigna lanceolata 189, 198 Yandruwandha people 96, 102, 115 Yardea SA 130 yarkalayin unidentified water plant 202 Yarralin NT 195

Zann, RA 49, 56–7, 59–60, 67, 70–3, 237, 277 zebra finch Taeniopygia guttata 23–4, 45–73, 122, 247 Zietz, AHC 147, 150, 170–1, 173, 178 zoology 12, 23, 28, 50, 52, 123, 129

299