- ——— . ae ee

Editors P. VICKERS-RICH, J.M. MONAGHAN R.E BAIRD & T.H. RICH

With the assistance of E.M. Thompson éz C. Williams

VERTEBRATE PALAEONTOLOGY OF AUSTRALASIA

Editors P. VICKERS-RICH, J.©M.-MONAGHAN, R.E BAIRD & T.H.RICH

With the assistance of E.M.Thompson & C.Williams

Graphics by D. Gelt Photography by S. Morton & F. Coffa

Pioneer Design Studio peration with th

in coope ith the Monash University Publications Committee, Melbourne

First published in 1991 by Pioneer Design Studio Pty Ltd 486 Maroondah Highway, Lilydale, Victoria, 3140 for and in co-operation with the Monash University Publications Committee, Melbourne © P. Vicers-Rich 1991

Typeset in Australia Printed and bound in Singapore ISBN 0 909674 36 1

Reprinted 1991

All rights reserved. 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 or otherwise without the prior written permission of the publisher.

Geologic Time Scale reproduced on page xv from: Harland, W. B., Armstrong, R. L., Cox, A.V., Craig, L. E., Smith, A. G., & Smith, D. G. 1989. A Geologic Time Scale. Cambridge Univ. Press, Cambridge.

PREFACE

In 1982 publication of this volume's predecessor, The Fossil Vertebrate Record of Australasia, was a milestone. That book provided a remarkably valuable reference for palaeontologists and biologists who have an interest in the evolution of vertebrates in Australasia. The dog-eared covers and almost fatally broken spine of my copy provide clear evidence of the many times it has been consulted by me and my colleagues.

The central theme of this new volume, Australasian vertebrate palaeontology, is the same as that of its predecessor, but the interpretations of the theme are richer and more diverse and the cast of authors has been enhanced. As the historical accounts opening this book vividly relate, the origins of the current, expanding activity in Australasian vertebrate palaeontology are complex, but two figures loom large. The scientific contributions and contagious enthusiasm of Profs. W.D.L. Ride and R.A. Stirton played a major role. Many of the chapters in The Fossil Vertebrate Record of Australasia were authored by David's and Stirt's students. In this new volume we find abundant evidence that not only does this "mob" continue to be active, but now they are being joined by the next generation including their students as well as palaeontologists with other academic backgrounds.

The fossil record of Australasia cannot be faulted for limitations in temporal range. Its oldest records include the Ediacaran fauna that documents the diversity of invertebrate life in the seas of the later Precambrian. The first traces of vertebrates are specimens of agnathan fishes of Middle Ordovician age. Footprints on an Early Devonian sandy river bank in eastern Australia illustrate the evolutionary emergence of tetrapods long before that group is known from skeletal remains. Other occurrences of fossil vertebrates, analyzed with equal insight in this volume, provide us with glimpses of the subsequent evolution of vertebrates in Australasia. However, it resembles a cheap grade of Swiss Cheese -- one with greater voids than substance; the fossil record available for study is disfigured by "ghastly blanks".

These blanks in the fossil record remain vexatious, but each year they shrink in number and duration. Long days spent by the authors and their colleagues in the quiet of the Outback to the west of the Birdsville Track, following the ebb and flow of the tides to recover fossils from rocks in sea cliffs, as well as in other areas of the continent are paying off in generous dividends. Their updated versions of compilations of basic data on fossil localities are starting points for future research. Many acknowledgements of personal communications from other palaeontologists, or to works in press, are a promising measure of the information that soon will appear in scientific publications.

Although any assessment of the status of Australasian vertebrate palaeontology must account for changes in the research data base and the cast of researchers, of greater significance is the development of the research questions that are being addressed. Palaeontologists are far from bashful in posing questions concerning the nature of the mechanism and causal factors that have directed the course of vertebrate evolution. It's easily as healthy a cottage industry as the generation of speculations about what killed off the dinosaurs. The chapters in this volume show that an impressive array of palaeobiological questions are being successfully addressed in research on Australasian vertebrates.

Early workers, Lamarck, Owen, and other 19th Century biologists, recognized that the unique character of the Australasian fauna reflected the much later survival of many groups in this area than on other continents. In his influential book, Climate and Evolution, William Diller Matthew argued that the terrestrial vertebrate faunas of Australia and other southern

continents had their origins in stocks that evolved in Holarctica and then were displaced southward by their descendants. Many years later Philip Hershkovitz dubbed this pattern the "Sherwin-Williams effect," a reference to that paint company's advertising symbol depicting a can of paint being poured over a globe. Further, in the context of a stablist view of continental positions, Matthew considered and rejected the possibility that the occurrence of closely related mammals or other members of the terrestrial faunas of Australia and South America reflected interchange across Antarctica.

Understanding of the changes in positions of continents, patterns of circulation of the oceans, currents, and continental climates through the course of earth history continues to expand. The following studies show that during the late Palaeozoic and early Mesozoic Australasia formed the tip of the southern peninsula of Pangaea; dispersal of vertebrates across this globally continuous supercontinent appears to have been little impeded. Later in the Mesozoic changes in climate regimes left Australasia partially cloistered biogeographically by the high latitude environments of South Polar Antarctica. During the late Mesozoic and early Cainozoic shifts in continental position and climatic change increased Australasia's biogeographic isolation and magnified its role as a haven for vicariantly isolated populations of lineages that had or would become extinct in the Northern Hemisphere. Australasia's continued northward movement first maintained its isolation but then brought it into closer proximity of southeastern Asia, so increasing the probability of chance dispersal of birds, bats and, later, rats and other terrestrial vertebrates.

Analyses of the evolution of Australasian biogeographic patterns have advanced beyond the level of debates over the primacy of dispersal or vicariance. Informative studies presented in this book reveal the complex interplay of these factors at continental and smaller scales as well as the environmental consequences of changing global climates and the latitudinal position of Australasia. Data on the avian and terrestrial vertebrate faunas of smaller islands of Australasia, for example, have increased to a point where they are pertinent to testing and qualifying the MacArthur-Wilson hypothesis of insular biogeography.

On a larger scale, additions to the available fossil record add substantially to an interpretation of the biogeographic history of marsupials, my particular pets, which until recently had fallen into disfavor. Recent discoveries strongly indicate that the "Sherwin- Williams effect" probably accurately describes the origin of the group in the Northern Hemisphere and its dispersal into South America, but not Australia. Dispersal of marsupials, but not their eutherian contemporaries, across Antarctica to Australia, a possibility rejected by W.D. Matthew, probably occurred very late in the Mesozoic or early in the Cainozoic. Then vicariant isolation set the physical stage for an extensive evolutionary radiation of Australasian marsupials. Further movement of the continent toward the Equator and southeastern Asia increasingly opened the door to dispersal of plants and animals into Australasia that, in turn, affected the course of evolution of its marsupial fauna.

Biogeographic analyses are only as strong as the understanding of the evolutionary interrelationships of the organisms being studied. The authors show that taxonomic research on Australasian vertebrates is being rapidly advanced on many fronts. Modern methods of analysis of phylogenetic relationships are being applied to a widening spectrum of data. Discoveries of members of new living and prehistoric species are yielding hitherto unknown data. Biomolecular studies bringing new data from living and recently extinct lineages are providing additional kinds of information for the taxonomic analyses. Inventive studies of form and function of marsupial dentitions and avian egg shells add not only data for studies of evolutionary interrelationships, but also provide a better appreciation of their ecological roles. Additionally, taphonomic studies of the fossil assemblages refine interpretations of the composition of the biotas from which they are drawn.

The modern terrestrial biota of Australasia, like those of many other continents, is in part the product of late Pleistocene or Subrecent extinctions that decimated many lineages of large

vertebrates. The quality of the fossil record of Australasia surpasses that of other southern continents and provides and opportunity to study another evolutionary "experiment" as climatic change and human intervention had their impacts on such late Pleistocene and Subrecent biotas.

The authors of this volume have provided us with both a valuable standing ground and a significant point whose pages soon will acquire the patina characteristic of oft-consulted references. The contributions clearly illustrate the current, rapidly accelerating pace of vertebrate palaeontological research in Australasia and document our colleagues’ research

accomplishments. As a major reference work, it is destined to serve as a starting point for many lines of future research.

W.A. Clemens Berkeley, California August 1990

feng HoAGuL aT fers UNS

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CONTENTS

Preface v

Introduction x1

Acknowledgements xiii

Geologic Time Scale xv BACKGROUND TO THE FOSSIL RECORD

1. Squatters, Priests and Professors: A Brief History of Vertebrate Palaeontology in Terra Australis. P. Vickers-Rich & N. W. Archbold 1

2. Vertebrate Palaeontology in Australia: The American Contribution. R.H. Tedford 45

3. | Musings on New Guinea Fossil Vertebrate Discoveries. M.D. Plane 85

4. Palaeoclimatic Setting and Palaeogeographic Links of Australia in the Phanerozoic. L. A. Frakes & P. Vickers-Rich 111

5. An Introduction to the Literature of Palaeontology with Reference to the Fossil Vertebrates of Australasia. M. Chiba 147

TECHNIQUES AND ANALYSIS OF FOSSILS

6. | Techniques Used in Preparation of Terrestrial Vertebrates. M. Whitelaw & L. Kool 173 7. Predicting the Diet of Fossil Mammals. G. D. Sanson 201 8. The Diet of the Extinct Bandicoot Chaeropus ecaudatus. W. Wright, G. D. Sanson & C. McArthur 229 9. Reconstructing the Natural History of Extinct Animals: Ektopodon as a Case History. N. Pledge 247 10. The Taphonomy of Late Quaternary Cave Localities Yielding Vertebrate Remains in Australia. R. F. Baird 267 11. Preservation of Biomolecular Information in Fossils fron Australia. M. Rowley 311

VERTEBRATE FOSSIL RECORD OF AUSTRALASIA

12. The Long History of Australian Fossil Fishes. J. A. Long 337

13. Palaeozoic Vertebrate Microfossils in Australia. S. Turner 429

14. —_ Australian Mesozoic and Cainozoic Lungfish. A. Kemp 465

15. Chondrichthyans in the Cretaceous and Tertiary of Australia. N.R. Kemp 497 16. Australian Fossil Amphibians. A. Warren 569

17. _ Australian Fossil Frogs. M. J. Tyler 591

12. Fossil Reptiles in Australia. R. E. Molnar 605

19. The Fossil Turtles of Australia. E.S. Gaffney 703

The Mesozoic and Tertiary History of Birds on the Australian Plate.

P. Vickers-Rich 721

The Quaternary Avifauna of Australia. R. F. Baird 809

Fossil Eggs from the Tertiary and Quaternary of Australia. D. L. G. Williams & P. Vickers-Rich 871

The History of Mammals in Terra Australis. T. H. Rich 893

The Pleistocene Megafauna of Australia. P. Murray 1071

The Australasian Marine Vertebrate Record and its Climatic and Geographic Implications. R. E. Fordyce 1165

A New Look at the Fossil Vertebrate Record of New Zealand. R. E. Fordyce 1191 The Quaternary Avifauna of New Zealand. P.R. Millener 1317

Vertebrate Fossil Faunas from Islands in Australasia and the Southwest Pacific. C. W. Meredith 1345

The Fossil Vertebrate Record of New Caledonia. J.C. Balouet 1383 Systematic, Geographic and Geologic Index 1411

Index 1419

INTRODUCTION

Vertebrate Palaeontology of Australasia is the direct outgrowth of an earlier book entitled The Fossil Vertebrate Record of Australasia edited by P. V. Rich and E. M. Thompson and published first in 1982. The original book grew from a series of lectures given in second and third year Earth Sciences and Zoology courses coordinated by P. V. Rich and J. W. Warren at Monash University. These courses involved many of the book's authors, whose participation was underwritten and supported by both the departments of Earth Sciences and Ecology and Evolutionary Biology at Monash University. This initial support was critical to making a reality of both resultant books.

Unlike The Fossil Vertebrate Record of Australasia, this book has been published through a commercial publisher. This was possible because of the commitment of this publisher to the principle of making the book, this time twice the length of its predecessor, available at a price that students and scientists alike could afford, both in Australia and overseas. And, this economical production was carried out without sacrificing editorial and production quality. We, as editors, are extremely grateful for the efforts made by Derrick Stone who heads Pioneer Design Studio in this regard, and would encourage more such cooperative efforts along these lines between authors-editors and publishers. This book would, likewise, not have been viable without the generous monetary support provided by the Monash University Publications Committee, who by their injection of funds allowed a larger press run, which, of course, lowered unit costs. To them we give our greatest appreciation.

The purpose of this volume on Australasian vertebrate palacontology is to present a state- of-the-art resumé of the different disciplines that compose this rapidly growing scientific endeavour in Australia, New Zealand, the southwest Pacific and Antarctica. Vertebrate Palaeontology of Australasia is divided into three major sections: Background to the Fossil Record, Techniques and Analysis of Fossils and Vertebrate Fossil Record of Australasia. The first section presents the history of vertebrate palacontology on the Australian continent and in New Guinea; an overview of the geological history and palacoenvironmental setting during the history of vertebrates, with emphasis on Australia; and a final chapter on the literature of vertebrate palaeontology for the Australasian area.

The second section outlines the different kinds of techniques - collecting, preparation, and analytic - that have been applied to Australasian fossil vertebrates, not fundamentally different from those applied anywhere else, but the results of case studies on reconstructing of function based on morphologic form are unique to this biogeographic region, because the animals dealt with are endemic. Many of the chapters in this section are new, such as that by Whitelaw & Kool (Chap. 6) on preparation and collection techniques and that by Rowley (Chap. 11) on biomolecular analyses as applied to vertebrate fossil remains in Australasia. New, too, is a chapter on the interpretation of the dict of a recently extinct bandicoot, Chaeropus, which still had available for study a carcass with stomach contents that could be consulted after interpretations based on dental morphology were drawn (Wright, Sanson & McArthur, Chap. 8), as well as a chapter on taphonomy of vertebrate bone accumulations in caves (Baird, Chap.

he third section deals specifically with the vertebrate fossils that have been recovered from Australasia, and this data has grown considerably since the publication in 1982 of The Fossil Vertebrate Record of Australasia. Much of the added length of this book over its predecessor is a direct reflection of the growth of this data, and, in fact, this may be the last time it is

possible to write such a compendium, unless a multivolume work is produced. The new information accruing is enormous, and with the rapidly growing number of new workers in vertebrate palaeontology dealing with Australasian subjects, this trend is likely to continue and accelerate in the years to come.

All of the original topics covered in The Fossil Vertebrate Record of Australasia have been retained in this new book and updated, but in addition, new chapters on vertebrate microfossils (Turner, Chap. 13), fossil turtles (Gaffney, Chap. 19), fossil eggs (Williams & Rich, Chap. 22), the Quaternary avifauna of Australia (Baird, Chap. 21), the Quaternary megafauna (Murray, Chap. 24), the Quaternary avifauna of New Zealand (Milliner, Chap. 27), the vertebrate fossil faunas of islands of Australasia, including the southwestern Pacific (Meredith, Chap. 28) and the fossil vertebrates of New Caledonia (Balouet, Chap. 29) have been added.

This volume is the end result of 8 years of work to bring the preliminary volume, The Fossil Vertebrate Record of Australasia, up to the standard of a finished version, both updated and polished, and to increase both the quality and quantity of the illustrations. We hope that the book will find use both as a standard reference work for the Australasian area and as a textbook for the beginning student of palaeontology who has special interests in this most intriguing biogeographic area of the Earth.

ACKNOWLEDGEMENTS

Vertebrate Palaeontology of Australasia would not exist if it were not for much hard work and dedication of a considerable number of individuals. Four stand out from the rest because of the massive time and effort put into this long term project: Corrie Williams, Mary Lee Macdonald, Elizabeth Thompson and Mary Walters. They were involved in such activities as the detailed editing, proof-reading, paste-ups, letter writing to authors and reviewers, photocopying and pursuing numerous jobs related to production of everything from initial manuscripts to final camera-ready copy. Corrie was also involved in compiling the systematic appendix. Much of the work they did was unpaid, which makes their dedication all the more appreciated.

Absolutely critical to completion of this project, too, was draftswoman Draga Gelt (Earth Sciences Department, Monash University), who provided most of the artwork in this volume, most of which was originally prepared as parts of research papers or for teaching purposes, but most had to be slightly modified for inclusion in this book. Steve Morton (Physics Department, Monash University), photographer extraordinaire, produced most of the photographic illustrations and provided copies of material so that backup was available in case of loss during production of the final book.

Much of the writing and manuscript production, especially of the final camera-ready copy, was carried out using Microsoft Word (both versions 3.0 and 4.0) on an Apple Macintosh SE and an Apple Laserwriter Plus for printing. Professor Gordon Lister was critical in convincing PVR to use the Macintosh system, and thus we are grateful to him not only for that but for providing half of the funds needed to purchase the Mac SE. We are also grateful to the Earth Sciences Department at Monash University for use of the Laser Writer and to the Ecology and Evolutionary Biology Department for providing the funds to purchase paper and ink cartridges for the Laser Writer. We are also grateful to Monash University, especially the Earth Sciences Department, for providing the atmosphere in which such a book could develop, as a direct result of an intensive, in depth series of courses dealing with vertebrate fossils and evolution, over a period of years. Francis de Souza provided invaluable ("life saving") computer assistance in compilation of the index

During the final stages of this project Computer Knowledge in Melbourne provided us with a second Mac SE needed for editorial work, for which we are most grateful. We wish to especially thank Michael Smart and Bernie Hogan of Computer Knowledge for their help with hardware and software throughout this project.

Many other people are also due our gratitude: Frank Knight, Derrick Stone of Pioneer Design Studio and the Museum of Victoria for the use of reconstructions by Frank Knight of Australian fossil vertebrates from Kadimakara. Extinct Vertebrates of Australia, Frank Coffa (Museum of Victoria, Department of Photography) for providing photographs of Australian fossil vertebrates; Simon Lai for translation of magnetic media into a usable form; J. R. Macdonald and Rhys Walkley for their reviews of each chapter; L. Kool, I. Brailey, N. Schroeder for assistance in editing and gathering research materials; R. K. Johns and the Department of Mines and Energy, South Australia for providing illustrations of H.Y.L. Brown, M. Beckers for her help in typing two of the chapters; P. Hermansen and Francis de Souza for assistance with cranky computers and software; A. Carle, D. McCarry and G. Royce for help with the financial aspects of the project. Also important in allowing this book to develop were fellow members of the Monash University staff, who through discussion, and in some cases

provision of illustrative material and financial support, aided in development of ideas and final production of this book, especially Ray Cas, Joe Monaghan, Ian Nicholls, Neil Archbold, Bob Gregory and Jim Warren. Patricia Komarower is especially thanked for carrying a heavy load of teaching during two years of co-teaching with PVR, which allowed editorial work to proceed, when otherwise it certainly would have faltered. Many other individuals provided illustrative material, and they are thanked in captions for the figures throughout the text.

Each chapter was assessed by at least two reviewers, some who have remained anonymous, and their help is gratefully acknowledged: A.K Behrensmeyer (National Museum of Natural History, Washington, D.C.), H. Olson (National Museum of Natural History, Washington, D.C.), W. Boles (Australian Museum, Sydney), C. Mourer-Chauviré (University Claude Bernard, Villeurbanne Cedex, France), C.W. Meredith (Australian Biological Research Group, Melbourne), D. W. Steadman (New York State Museum, Albany, New York), Dianne Clifford (Golden Grove, Western Australia), F. Whitmore (National Museum of Natural History, Washington, D.C.), P.R. Millener (National Museum of New Zealand, Wellington), J. Bowler (Museum of Victoria, Melbourne), J.A. Long (Western Australian Museum, Perth), N.A. Pledge (South Australian Museum, Adelaide), G. C. Young (Bureau of Mineral Resources, Canberra), A. Ritchie (Australian Museum, Sydney), T. F. Flannery (Australian Museum, Sydney), J. Hope (Australian National Parks and Wildlife Service, Sydney), R. Wells (Flinders University, Adelaide), G. D. Sanson (Monash University, Melbourne), G. F. van Tets (C.S.I.R.O., Canberra), D. F. Brannagan (University of Sydney, Sydney), W. D. L. Ride (Australian National University, Canberra), M. O. Woodburne (University of California, Riverside), G. Lowenstein (University of California, San Francisco), J. Ramshaw (C.S.LR.O., Melbourne), E. L. Lundelius (University of Texas, Austin), P. Janvier (Université Paris VI), M. Davies (University of Adelaide), R. Estes (San Diego State University), K. Kelly (Museum of Victoria, Melbourne), I. Norton (Queen Victoria Museum and Art Gallery, Launceston), and K. F. Hirsch (University of Colorado, Boulder).

A great debt of gratitude is due many funding agencies, which, either through direct support for this project or indirect support for research programmes that yielded the information in this book, have been critical to its completion: the National Geographic Society, the Australian Research Council, Monash University, Computer Knowledge, the Museum of Victoria, Earthwatch, the Ingram Trust, the Danks Trust, the Ian Potter Foundation, Western Mining, International Chemical Industries, Safeway Australia, the Australia-China Council, the Australian Academy of Sciences, the Australian National Parks and Wildlife Service, the Australian-American Educational Foundation, Friends of the Museum of Victoria, Sunshine Foundation, the Australian Army, Atlas Copco, Ingersoll-Rand, Shell, Mobil Oil, the Victorian Police, the Surf and Life Saving Association and last, but certainly not least, the Publications Committee of Monash University.

This book is dedicated to four people who have been especially significant in nurturing, in some way, most of the current crop of vertebrate palacontologists active in Australia today as well as significantly pushing ahead the frontiers of this science in the 20th century: Dr. R. A. Stirton ("Stirt"), now deceased, formerly of the University of California, Berkeley; Dr. W. D. L. Ride (Department of Geology, Australian National University, Canberra); Dr. R. H. Tedford (Department of Vertebrate Paleontology, the American Museum of Natural History, New York) and Mr. Paul Lawson (formerly of the South Australian Museum, Adelaide). Their stories are told in the pages of this book. Without their infectious enthusiasm, their uncanny ability to find bones and inspire others, vertebrate palaeontology in Australasia would most certainly not be the vibrant science that it is today.

——

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CHAPTER 1

SQUATTERS, PRIESTS AND PROFESSORS: A BRIEF HISTORY OF VERTEBRATE PALAEONTOLOGY IN TERRA AUSTRALIS

Patricia Vickers-Rich! and Neil W. Archbold?

PEE CHOTE ta) cea ee gat enlntess cantare if. reatey tees 1 Antipodean Discoveries, Grist for the Bir Peay MAT ectsicsresteansnnts yevete eos thinas agen 1 Barkly XcOdStaleSutve ys nccte.sslunesecte ss eae 1 Ei PAnGl: SUVS YS Bote iiss cliccedetealeds (ahd etotnenb carts 4 NBOLOLET PO RDIOKIS, Ad.Te acl vrcdt svapaaplre an hie tehe wie 8 Gold Rushes, Museum and Sydney Gentry Beginnings of a Home Based Science....... 11 Gold and Beginnings of the State Surveys...... 12 Repositories of Fossil Objects: the First VIS SUTIES 6 alent satis tp oh aoe anne cdoeen slew taete {2 Beginnings of Independent Training: THE TWMEVOTSHUES At betas cium Aletta ene eles 15 Rapid Communication on a Local Scale: the Scieheirio- SOCMGS 56.0555 005 oe te are 22 Men of Influence, Pioneers in Australian Vertebrate Palacontolo PY ins ccescescnsenttovsnt 22 Australian Independence and International COGPCTANOM YEA MR Ti ee ah A 31 PRORSHOW LOGS OMVCAIS c02 50 tis vcnu decd ste dnedcuctics subciest de 39 IREIEIENCES «ioh sig tobv aided Pada eeete a ei dca bE 39

1 Earth Sciences and Ecology/Evolutionary Biology Departments, Monash University, Clayton, Victoria 3168, Australia. 2 Department of Geology, University of Melbourne, Parkville, Victoria 3052, Australia.

2- RICH & ARCHBOLD

INTRODUCTION

In his study on 'The Spread of Western Science' George Basalla (1967) presented a model of how Western science has characteristically developed and grown outside of Europe, often in three major stages: (1) a stage when the newly discovered territory serves as a source of new data for European science; (2) a stage still primarily colonial in aspect, but during which the local scientists accept fuller responsibility for investigation and interpretation of the data themselves; and (3) a stage when the indigenous scientists attain, or make efforts to attain, an independent scientific tradition constructing self-supporting institutions, receive scientific training in their own country, develop independent societies, and "formulate indigenous scientific attitudes and goals" (Moyal 1976). Certainly, this has been much the path taken by vertebrate palaeontology in Australia, and it has only recently entered the final of Basalla's stages. It is still a science limited to a small band of professionals with a growing support of associated non-professionals. It is still a science with unfathomed areas in need of exploration, still very much in a pioneering era of discovery (Vallance 1975, 1978, Rich & Thompson 1982).

The following paper is a brief overview of vertebrate palaeontological work in Australia, starting with its beginnings in the early 19th century and continuing to 1989. It is organized utilizing Basalla's developmental divisions, even though there are often no clearcut boundaries separating each of these stages.

ANTIPODEAN DISCOVERIES, GRIST FOR THE EUROPEAN MILL

Prior to discoveries by Europeans, Aboriginal legends existed, which perhaps had stemmed from an acquaintance with prehistoric bones or even living prehistoric animals themselves (Fig. 1). Tribes in eastern Australia were quite fearful of the bunyip (Barrett 1946), sometimes described as a monstrous animal that supposedly inhabited deep waterholes and roamed the billabongs at night. When confronted with the remains of some of the now extinct Australian marsupials, Aborigines would often identify them as the bunyip (Barrett 1946, Dugan 1980). Rich (1979 and in Rich & van Tets 1985) has noted legends about the mihirung paringmal of western Victorian Aborigines, which may allude to the currently extinct giant birds, the Dromornithidae. Some of the legends describing such creatures led to the discovery of rich vertebrate fossil fields, such as those at Lake Callabonna in South Australia in the late 19th century (Hale 1956).

It was not these legends, however, that led to a detailed understanding of the past veretebrate faunas of Australia, but European and later indiginous exploration that produced the fossils and formed the basis for the recognition of a succession of Australian vertebrate faunas spanning almost the last 500 million years.

EARLY COASTAL SURVEYS

The first European expeditions in the early part of the 19th century did not produce the remains of fossil vertebrates, but did, instead, locate invertebrate and plant fossils and even the living remnants of some vertebrate groups now extinct. The Matthew Flinders Expedition of 1801-1805 (Flinders 1814) and the French Nicholas Baudin Expedition of 1800-1804 (Fig. 2) were two such enterprises. The Baudin Expedition, splendidly outfitted with both equipment

HISTORY OF AUSTRALIAN VERTEBRATE PALAEONTOLOGY - 3

and scientists, returned fossil and modern natural history material to Europe. This material included living specimens of the now extinct, dwarf King Island Emu (Dromaius baudinus), which were kept alive in France for some time after their arrival. J. C. Bailly, a mineralogist attached to the Baudin expedition, reported fossil ferns in shales near Parramatta and fossil plants collected from Tasmania, which were assigned to what would now be called the Carboniferous by Leopold von Buch (1814), after examining the specimens in Paris.

Figure 1. Aboriginal art in the Cape York Peninsula caves and elsewhere in Australia depict animals that may now be extinct. This cave art in the Quinkan Gallery, site B(5) recorded in Trezise (1971) maybe one of the extinct dromomithids, perhaps Genyornis.

Also included in the collections from Terra Australis was the clam Trigonia brought up in a dredge haul off King Island in Bass Strait. Jean Baptiste Lamarck, famous for his evolutionary theories but also a highly respected and influential invertebrate zoologist of this period, was struck by the resemblance of the living Trigonia to forms known only as fossils in Europe and elsewhere in the world. The concept that Australia was somehow a haven, a refuge, for organisms that could no longer survive elsewhere had its origins in these early discoveries. Australia was viewed as a land of living fossils, and this was further reinforced as exploration continued inland later in the 19th century.

The unfortunate Flinders Expedition, unlike the highly successful Baudin enterprise, ended in shipwreck and the loss of most specimens, except for a few that the shipboard botanist, Robert Brown (1773-1858), had surreptitiously taken ashore with him in Sydney when he left the expedition. Although not mentioned in his catalogues, of which he kept a duplicate set when Flinders sailed away, fossil invertebrates and plants were returned to Europe and England. Brown returned to England in 1805 with three cases of "minerals," a part of his possessions that passed through Customs (Vallance 1978). Some of these fossils were definitely presented to the Rev. William Buckland at Oxford, who concluded (1821) that the Australian coal was comparable to that of the Carboniferous of England and that the marine fossil invertebrates

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were similar to those of the Mountain Limestone of Derbyshire. James Sowerby (1818a, 1818b) had previously described morphological details of the invertebrates. Brown's name comes up several times in new species being named by palaeontologists in both England and

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Figure 2. Nicholas Baudin and his ships used on the first major scientific expedition to Australia at the beginning of the nineteenth century. Bavdin's expedition was splendidly outfitted and returned to France with an array of new forms, both fossil and recent, that greatly expanded the European knowledge about Australia. (Courtesy of the Museum d'Histoire Naturelle, Paris).

Europe describing Australian material, e.g. Glossopteris browniana, a fossil seed fern described by Adolphe Brongniart (1828) based on fossils from New South Wales passed onto him by William Buckland. Buckland had evidently received it originally from Brown. Charles Konig (also spelled Koenig, see Archbold 1986) was also to describe some of the invertebrate fossils, such as the brachiopod, Trigonotreta stokesii (Konig 1825, Brown 1946). Both of these forms are now known to be classic Permian Gondwanan species, reflecting a very different arrangement of the continents of the world than characterise the present.

INLAND SURVEYS

Other coastal surveys followed, and visitors to Australia returned collections of fossils to Europe. Plants and invertebrates were mentioned (see Vallance 1981 and Archbold 1986 for details), but no vertebrate fossils of note were found. This was to change dramatically with the inland explorations carried out by Major (later Sir) Thomas Livingstone Mitchell, Surveyor- General of New South Wales from 1828 until his death in 1855 (Foster 1985).

T. L. Mitchell was to map in detail and procure many specimens of bones from the Wellington Valley caves (Fig. 3) of New South Wales (Mitchell 1838). He first visited the caves on 26th June 1830 with a local colonist, George Ranken. Ranken had previously

HISTORY OF AUSTRALIAN VERTEBRATE PALAEONTOLOGY - 5

Figure 3. A younger (left) and an older Sir Thomas Mitchell, who led an expedition into interior New South Wales in 1830. His recovery of fossil vertebrates from caves in the Wellington Valley (below) led to the first extinct vertebrates from Australia being described in a scientific paper. (Courtesy of E. B. Joyce from Mitchell 1838).

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discovered some bone fragments (Ranken 1916) and had taken them to Sydney in order to send them to Professor Robert Jameson of the University of Edinburgh (Anderson 1933).

Ranken's discovery of fossil bones was announced in the Sydney Gazette of 25th May 1830 in an anonymous letter (signed L.) by the Rev. Dr John Dunmore Lang. Lang left Sydney on the 14th August 1830 with Ranken's specimens, his own Sydney Gazette letter and a short manuscript by Mitchell on the Wellington caves. By early 1831 all were in the hands of Jameson, the two notes being published in the Edinburgh New Philosophical Journal. Both notes were credited to Lang, but in the subsequent volume Mitchell's note was correctly attributed to him (see Lang 1831, Mitchell 1831a).

Mitchell revisited the caves on the 3rd July 1830 and collected further specimens. These were apparently sent to the Geological Society of London with a letter dated the 14th October 1830 (read at the Geological Society of London meeting of 13th April 1831 - see Mitchell 1831b).

Various specimens collected by Ranken and possibly Mitchell were examined by William Clift, Conservator of the Hunterian Museum (College of Surgeons), who identified dasyurids, wombats and kangaroos (Clift 1831). Joseph Barclay Pentland (see footnote by T. G. Vallance in Dugan 1980) living in Paris, commented extensively on material sent to Paris from England and also independent information on the Wellington caves from Peter Cunningham, author of the 1827 book Two Years in New South Wales (see Pentland 1831, 1832 and Jameson 1831b). Jameson also offered editorial comment on William Clift's conclusions (Jameson 1831a - see Dugan 1980 on the importance of this for challenging Baron Georges Cuvier's contemporary catastrophist theories). William Buckland (1831) considered that some bones might represent either rhinoceros or hippopotamus, and Baron Cuvier (see Pentland 1833b, 1833c) also examined specimens.

Such was the interest in Europe on the Wellington caves discoveries, that many of the notes and letters discussed above were translated and published in contemporary German journals (Jameson 1832a, 1832b, Mitchell 1832a, 1832b, Pentland 1833a, 1833c ).

Mitchell's records on the discovery of vertebrate fossils at the Wellington caves (Fig. 4) are not without humor, for as he noted in his diary:

"The pit (Breccia Cave) had been first entered only a short time before I examined it, by Mr. Ranken, to whose assistance in the researches, I am much indebted. He went down, by means of a rope, to one landing place, and then fixing the rope to what seemed a projecting portion of rock, he let himself down another stage, where he discovered, on the fragment [a giant bird femur, probably from a member of the family Dromornithidae] giving way, that the rope had been fastened to a very large bone, and thus these fossils were discovered" (Mitchell 1838: 362)."

The bone which Mr. Ranken misjudged was the "lower end, mutilated, and encrusted with the red stalagmite of the cave ...." of a femur that was identified by Sir Richard Owen as belonging to a large bird, previously unknown. It was figured in Mitchell's (1838) publication (Fig. 5), but was subsequently lost, perhaps during the bombing of London during World War Il.

Mitchell's discoveries of fossil bones had aroused the interest of overseas scientists in extinct Australian vertebrates, and there followed many years of European and Australian alike collecting fossil remains. Most all of this material was sent from the shores of Terra Australis for description and study by foreign experts, as the needed expertise and comparative collections did not exist in Australia. It was not until the latter part of the 19th century, that indigenous workers began to study the local fossils in any serious way, even though several

HISTORY OF AUSTRALIAN VERTEBRATE PALAEONTOLOGY - 7

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residents, such as Leichhardt and Stutchbury suggested local material should remain in Australia (Branagan, pers. comm.).

"FOREIGN EXPERTS"

It was Sir Richard Owen (1804-1892), renowned British comparative anatomist, who described much of the new fossil vertebrate material from Australia (and New Zealand) as it came to light when inland exploration and settlement expanded (e.g. Owen 1843, 1845, 1877, 1879a, 1879b, 1879c, 1882) (Fig. 6). In an appendix to Mitchell's volumes on his three expeditions into interior Australia published in 1838 (Fig. 5), Owen identified some of the fossils as gigantic marsupials, Nototherium and Diprotodon . Although Mitchell's and Ranken's material from Wellington caves was examined and reported on by Cuvier and Pentland, and Darwin was aware of it as well when he visited Australia in 1836, it was Owen who undertook the tedious job of description and study. He, like the French before him, suggested that "it was necessary to search Britain's secondary (oolitic) [Mesozoic] formations to find specimens analogous to Australia's recent marsupial fossil forms" (Moyal 1975, 1976).

Owen, over the next 40 years, made Australian and New Zealand vertebrate palaeontology his own domain. In this he was aided by many resident Australians who sent him material. Friedrich Wilhelm Ludwig Leichhardt provided and helped describe bones from southern Queensland in 1844. W. B. Clarke and S. Stutchbury recovered bones in their northern surveys from the Darling Downs of Queensland as well as closer to home near Sydney. F. McCoy and G. Krefft, likewise, provided specimens that came to their attention as the officials in charge of the National Museum of Victoria and the Australian Museum. Local pastoralists in digging wells or in surveying property came upon and then sent material to the youthful Australian Museum, and often these treasures eventually made their way to Owen's desk. Although much of his Australian work centred on fossil marsupials,