- ——— .
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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Bsc en 4

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

4-RICH & ARCHBOLD

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

op ArAntl: PREMIER SNS
gee err ~~ TNS
4 Fé

a

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).

6- RICH & ARCHBOLD

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

ian fossil

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8 - RICH & ARCHBOLD

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, Owen also took a keen
interest in other vertebrates as well, and he produced a prodigious number of papers.

Other foreign experts included Darwin's "bulldog," T. H. Huxley (1862), Gervais (1848-
1852), Hochstetter (1859), and R. Lydekker (1887, 1896), among others. None of these
workers, however, published as prolifically on Australian fossil vertebrates as Owen. Because
of his prodigious publication record, a few, but only a few, mistakes crept into his work, such
as his description of an elephant (supposedly a mastodont) from Australia. Many people
(Leichhardt 1855, Falconer 1863) questioned the authenticity of the elephant record in
Australia, and it has been suggested that the specimen probably entered as a trade item. After
the challenge by Falconer, Owen quietly abandoned his claim (Dugan 1980).

Such early collections’ of fossil vertebrates from Australia had some effect on European
science, especially the rich discoveries at Wellington caves. Even though the majority of the
material was described by Owen, who disagreed with Darwin on the mechanism that formed
new species, natural selection, according to Dugan (1980), the Wellington fossils favoured
Darwin's ideas. Dugan suggested that the Law of Succession was formulated in part based on
the types of fossils that occurred in the Wellington collections. This law states that fossil
animals in any particular geographic area are 'succeeded’ by other animals that are closely
related to them, no matter what the environmental conditions. Owen in his own writings
claims to have formulated this law (D. Ride, pers. comm.) and it gave no support to the
special creationists of the time, who had suggested that certain animals were "created" to be
perfectly suited to their environment, and that if such environmental conditions were present,
then certain predictable kinds of animals should be there too. Thus, one would expect the same
kinds of animals in the tropical regions of Australia and Africa. Mitchell noted the effect the
Wellington discoveries had on certain creationists: "I understand Buckland's nose is put
completely out of joint by the bones from Australia, their not being those of lions and hyenas

HISTORY OF AUSTRALIAN VERTEBRATE PALAEONTOLOGY - 9

74
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Figure 5. Plate from T. L. Mitchell (1838) illustrating the first Australian vertebrate fossils reported in a
scientific publication. (Courtesy of E. B. Joyce).

10 - RICH & ARCHBOLD

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Figure 6. Sir Richard Owen as a young man (A) and near the end of his career (B, C). He is often
associated with the New Zealand moas (A, B), a group he described prodigiously, almost single-handedly.
The outstanding comparative anatomist of the day, he became the authority on the fossil vertebrates of New
Zealand and Australia, describing the first material that was collected by Mitchell and Ranken in the
Wellington Valley. (Courtesy of the British Museum (Natural History), London).

HISTORY OF AUSTRALIAN VERTEBRATE PALAEONTOLOGY - 11

is, I find, a fact which is considered in England to entirely upset his theory. And I have now
heard from the best authority that their fossil bones belonging to animals similar to those now
existing has worked a great change in all their learned speculating on such subjects at home."
(Ranken 1916) Buckland, in fact, later did modify, and finally abandon his original ideas on
this issue (Vallance 1975; Ride, pers. comm.).

Despite the dominance of foreign experts during this period, it was also at this early stage
that the first native born (in Parramatta) Australian began to study vertebrate fossils
systematically - the naturalist Edmund Charles Hobson (1814-1848) (Vallance 1978).
Hobson, a founding member of the Tasmanian Society, had studied under Sir Richard Owen
and established a medical practice in 1838 in Hobart, but subsequently, for reasons of health,
moved to the Port Phillip district in 1840 (Gunn 1848a). An early death at the age of 34 ended
a highly active mind that investigated animal and human physiology, geology and vertebrate
palaeontology (Hobson 1841la-b, 1845a-c, 1846a-b, 1847a-f, 1848a-b). His last paper,
published posthumously, was one on a vertebrate fossil, a Diprotodon jaw. Hobson's wife
(nee Adamson) was also a keen naturalist and sketched the specimens figured in the two plates
of Hobson (1845c). Her sketches were used by the accomplished lithographer, Thomas Ham,
to produce the final plates. She also discovered the first trilobite recorded from rocks (Silurian)
of the Melbourne district, identified by Gunn (1848b) as probably Asaphus. Until much later,
she is the only woman who is recorded to have had any association with the field of vertebrate
palaeontology in this country.

GOLD RUSHES, MUSEUMS, AND SYDNEY GENTRY,
BEGINNINGS OF A HOME-BASED SCIENCE

Throughout the latter part of the 19th century and well into the 20th, vertebrate
palaeontology in Australia was in the process of establishing a home base here. The transition
from Basalla's stage 1, where Australia provided only the grist, the data for the European
intellectual mills, to stage 2, where although still colonial in aspect, the local scientific
community was becoming steadily more independent, occurred during this period. It was a
gradual transition, and no clear boundaries can be drawn at any one time.

Certainly the increasing independence must be related to the increasing population size and
its concentration in growing urban centres such as Sydney, Melbourne and Hobart. It also
must be tied to the increasing wealth that accumulated in these communities, especially that
generated by the discovery and mining of gold in several parts of Australia, notably in the area
north of Melbourne. Not only was there an incentive to excavate, which in itself led to the
discovery of fossils, but the economic necessity, which brought about the establishment of the
first state geological survey, whose job it was to document the rock record in the gold bearing
regions. Coincident with this, the settling of the interior led to the discovery of bones on a
number of sheep and cattle runs as wells were excavated or when drovers reported exposed
fossils as they traversed the countryside behind their slow moving herds.

With the opening up of the interior and the mining boom of the mid-19th century, several
members of the European scientific community took up temporary or even permanent residence
in Australia. Two men from this period stand out in the move towards an independent
palaeontological community, Ralph Tate in Adelaide and Robert Etheridge in Sydney. Both
were determined advocates of Australian science standing on its own. Both firmly believed in
cooperation as equals with foreign experts, neither serving them nor foolishly ignoring them.

12 - RICH & ARCHBOLD
GOLD AND BEGINNINGS OF THE STATE SURVEYS

The discovery of gold, mainly in Australia's southeast provided the first major economic
incentive that affected the course of palaeontology. The first of Australia's geological surveys,
the Victorian Colonial Survey was established in 1852, with A. R. C. Selwyn (1824-1902) as
its director (Dunn 1910, Darragh 1987). Selwyn's directive was to document the geology of
the gold-bearing regions, probably in the hope that such work would allow prediction of further
producing goldfields. Selwyn had been trained by some of the most prominent geologists in
Britain and, thus, was well prepared for the task of recording the stratigraphic succession,
almost unknown in Australia at this time. When he assumed his duties as Director the most
authoratative summary of geology of Australia was Jukes’ A Sketch of the Physical Structure
of Australia, So Far As It Is At Present Known published in 1850 by T. & W. Boone in
London. This small text shows how general the knowledge of geology was at the time and
how restricted to the margins of the continent it was. Selwyn and his team of geologists added
much to the detail of southeastern Australian geology by preparing well over three score of
detailed maps of present day Victoria before political controversy managed to bring about the
dissolution of the Survey for a while. In the meantime, Selwyn's survey was able to locally
carry out and publish its own scientific results and to locally train a number of geologists,
who later served to set up other surveys and institutions in Australia. Following the
establishment of the Victorian survey, all other states followed suit by 1890, and these
institutions served to foster collection and storage of fossils, some of which were vertebrates.

REPOSITORIES OF FOSSIL OBJECTS: THE FIRST MUSEUMS

Repositories of fossil objects had been established in Australia even before the state
surveys, and these, together with the surveys served as a growing resource for comparison in
the latter part of the 19th century (Kohistedt 1983). Since the early voyages to Australia by
the French and British, and even before them the Dutch, considerable British and European
interest in Australian natural history specimens had led to a lively trade in Australian
oddments, both for scientific and commercial reasons. Respectable collections of Australian
specimens accumulated in London, as well as in European museums, at first through the
efforts of such men as Sir Joseph Banks and Robert Brown. As a result, these museums,
especially the British Museum (Natural History) came to own many of the type specimens of
the newly discovered natural curiosities.

The pattern began to change, however, with the arrival in Sydney of Alexander Macleay in
1826. Macleay was sent as Colonial Secretary to the Government of New South Wales, a
position he held for the next decade. With him, from Europe, came a fine library and a fine
insect collection, amongst the best known anywhere at the time. Interestingly, Macleay's own
son and his nephew, both with the name of William, were to add to Alexander's collections
and perpetuate the family's support of science in Australia (Fletcher 1920, Stanbury 1975).

William Julian, the nephew, expanded the family's personal museum in the Elizabeth Bay
residence (after both Alexander and William Sharp Macleay had died) in part by importing
specimens from abroad. His wish was to possess a truly international collection, not just a
local one. Macleay went so far as to hire a curator, George Masters, out of his own funds. He
also continued the tradition of the Macleay's for serving as an intellectual hub in the
community by giving "whisky parties" (actually scientific gatherings around the drink) for staff
from the University of Sydney, as well as interested personages - explorers, doctors, visiting
scientists, amongst others. This undoubtedly led to his offering in December 1873 to bequeath

HISTORY OF AUSTRALIAN VERTEBRATE PALAEONTOLOGY - 13

his museum to the University of Sydney including a salary of 6,000 pounds to pay for
curation.

At the time Alexander Macleay stepped ashore in Sydney town a museum of sorts already
existed there. This consisted of a small room in the Colonial Secretary's Office, initially set
up by the colony's first scientific society, the Philosophical Society of Australasia (Branagan
& Townley 1976). In the beginning each of the seven members of the society paid 5 pounds
to have the collection organized and for the purchase of books. Major Goulburn provided the
single room that constituted the museum. Australian specimens were put on display, but
perhaps more important than that, such a museum allowed establishment of official contacts
with foreign museums, which in turn encouraged the exchange of specimens between this and
foreign institutions. The flow of scientific material was now in both directions, not just from
Australia to the colonial powers.

To this infant museum Macleay added his own collections and together with his
enthusiasm, the foundations of the Australian Museum were established. Naturally, this first
museum became a storehouse for antipodean specimens and gradually encouraged more and
more residents to retain material in Australia as well as to be more observant of their native
fauna. It was quite amazing that a museum was able to exist at all at this time. The Sydney
colony was "a convict settlement.....racked with dissention between free immigrants and
emancipists, businessmen and farmers, army and government, colony and colonial office... an
environment conducive [only] to.....activities [such as] those directed to individual survival and
aggrandisement" (Strahan 1979). In this milieu, science, even that part devoted to
palaeontology, had its beginnings.

When the Philosophical Society first set up its museum, Sydney was still small, with a
population approaching 20,000. Yet, by 1837, this tiny museum had public hours each
Tuesday and Friday from 11 am to 4 pm and contained exhibits of native fauna and flora
including more than 300 species of birds (Fletcher 1920). Also present in the collections were
ethnological and geological specimens, and by 1832 the government was providing 200 pounds
annually - a beginning.

The Australian Museum was Australia's first museum, but soon others followed. The
National Museum of Victoria, now the Museum of Victoria, was set up in Melbourne in 1854
(Pescott 1954). Two thousand pounds were set aside by the Victorian government in 1854 for
the fledgling National Museum, and by March 1854 Captain Andrew Clarke, who was
instrumental in the initial stages of this museum, saw to it that two rooms were set aside
above his offices at the old Assay Office in Latrobe Street (Fig. 7), just west of what was to
become a more permanent site. The first staff appointment was made on April 1, 1854,
William Blandowski, whose personality eventually led to considerable conflict with the council
that oversaw the museum and the newly appointed Frederick McCoy as Director (in 1858).
Blandowski energetically mounted a number of expeditions, which greatly expanded the
museum's holdings, and interestingly engaged such people as Gerard Krefft, who was later to
assume a curatorship in the Australian Museum. Blandowski finally resigned, and McCoy
took over the reins of power. It was McCoy who determinedly built up the Museum
collections and expanded the original facilities. He orchestrated and oversaw the removal of the
museum from the Assay office to a new site on the campus of the University of Melbourne
(Fig. 8), which was described in a poem published in the Melbourne Punch (Pescott 1954):

THE RAID ON THE MUSEUM

There was a little man,
And he had a little plan,
The public of their specimens to rob, rob, rob,

14 - RICH & ARCHBOLD

So he got a horse and dray,
And he carted them away,
And chuckled with enjoyment of the job, job, job.

Blandowski's pickled ‘possums

And Mueller's leaves and blossoms,

Bugs, butterflies, and beetles stuck on pins, pins, pins,
Light and heavy, great and small,

He abstracted one and all -

May we never have to answer for such sins, sins, sins.

There were six foot kangaroos,

Native bears and cockatoos

That would make a taxidermist jump for joy, joy, joy.
And if you want to know

Who took them you should go

And should seek information from McCoy, Coy, Coy.

When one's living far away,

Up the country I dare say,

It's very nice to have such things at hand, hand, hand,
Yet it don't become professors,

When they become possessors,

Of property by methods contraband, band, band.

The collections were to remain at the University of Melbourne until 1899, when shortly
after McCoy's death they were moved again to the present site of the Museum of Victoria on
Russell Street (Fig. 9).

Other museums followed as well: the Queensland Museum in Brisbane in 1855 (Mack
1956), the South Australian Museum in Adelaide in 1856 (Hale 1956), and the Tasmanian
Museum (Hobart), the Queen Victoria Museum (Launceston), the Western Australian Museum
(Perth), and finally the Northern Territory Museum (Darwin and Alice Springs). In addition to
these governmentally sponsored institutions, several private museums, such as the Kyancutta
Museum in South Australia, were managed with private funding. Many of the private
collections were later incorporated into state and federal museums, either by direct donation or
purchase. Although not without political intrigue and funding difficulties (Kohlstedt 1983),
museums did continue to expand into the 20th century. But they were no longer simply places
for storage of Australia's heritage, its natural wonders and antiquities. They also became
centres for public education, research and exchange of ideas, all in an Australian setting. The
museums and those associated with them, likewise, became the source of personnel and

HISTORY OF AUSTRALIAN VERTEBRATE PALAEONTOLOGY - 15

funding that sponsored locally based expeditions, such as Macleay organized and financed to
northern Australia and New Guinea in 1875 (Stanbury 1975).

SS

Figure 7. The Crown Lands Building, the original National Museum (now the Museum of Victoria) was
located in the assay section of this building from 1854 to 1856. (From the //lustrated Melbourne News
1858, courtesy of the Museum of Victoria).

BEGINNINGS OF INDEPENDENT TRAINING: THE UNIVERSITIES

Universities generally developed slightly later than the surveys and museums, but they
played a major role in establishing the independence of science in Australia. Frederick McCoy
(1818-1899) (Fig. 9), a major force in the founding of the National Museum of Victoria, also
was the Professor of Natural Science at the University of Melbourne from 1854 to 1899. He
was particularly interested in palaeontology and worked occasionally on vertebrate fossils,
although invertebrates were his main research subjects. During the forty years he led an active
scientific life in Melbourne, he ruled over it with an autocratic air (Branagan & Lim 1984) and
because of this attracted few students. T. S. Hall (1858-1915) (Fig. 10) was an exception to
this rule, an exception partly enhanced by his ability to get along with McCoy, despite the fact
that he well might disagree with him on a number of issues (Robin 1987). Hall had hoped he
would be offered the Chair of Geology when McCoy was gone, but it instead was offered to
John Walter Gregory, an import from Scotland.

J.W. Gregory (1864-1932), a remarkable "international" geologist, succeeded McCoy in
December 1900 as Professor of Geology, carrying on with the interest of the University of
Melbourne in palaeontology (Fig. 11). Gregory, who began his career as a palaeontological
assistant in the British Museum (Anonymous 1932), was interested in popularizing geology
and teaching it as a practical subject, very unlike McCoy. A great organizer, Gregory led a
group of Melbourne University students on camels into the Lake Eyre Basin during the
summer of 1901-1902. On that expedition he discovered a remarkable fossil field, mainly
along Cooper Creek. The account of this early vertebrate palaeontological expedition appears

16 - RICH & ARCHBOLD

8. The National Museum of Victoria with headquarters on the University of Melbourne campus, with

Figure

(Courtesy of the Museum of Victoria).

external and internal views.

HISTORY OF AUSTRALIAN VERTEBRATE PALAEONTOLOGY - 17

Figure 9. The move of the National Museum of Victoria from its university address to that on Russell
Street, where it is located today as the Museum of Victoria. Sir Frederick McCoy (as a young man, and as a

mature scientist) was responsible for the growth of this museum as well as the Department of Geological
Sciences at the University of Melbourne. (Courtesy of the Museum of Victoria).

18 - RICH & ARCHBOLD

Figure 10. Nineteenth and early twentieth century Australian palaeontologists. A, E. C. Stirling,
Honourary Director of the South Australian Museum from 1889 to 1914. He was involved in the excavation
and later monographing of a variety of vertebrate fossils recovered from Lake Callabonna in South Australia.
B, A. H. C. Zietz, Assistant Director of the South Australian Museum from 1888 to 1909. Like Stirling, he
excavated at Lake Callabonna and coauthored many papers with him on the discoveries there. (Courtesy of
the South Australian Museum and N. Pledge). C, T. S. Hall, Frederick McCoy's student at the University of
Melbourne was the first locally-trained palaeontologist in Australia. His specialty was invertebrate
palaeontology, but he found time to work on fossil whales as well. D, Frederick Chapman, primarily
interested in invertebrate palaeontology, mainly of Victoria, published a few papers in the early part of the
twentieth century on fossil vertebrates. (Courtesy of the Museum of Victoria).

HISTORY OF AUSTRALIAN VERTEBRATE PALAEONTOLOGY - 19

Figure 11. J. W. Gregory, as a young man (C, D) and as older explorer (A, B). Gregory was responsible
for the discovery of Pleistocene fossil vertebrates along Cooper Creek in the Lake Eyre Sub-basin, an area
that was later to produce the first concentrations of Tertiary terrestrial vertebrates from the continent. Osmar
White's caricature in The Super Roo of Mungalongaloo (D) illustrates how Gregory's expedition into central
Australia captured the public interest. (Courtesy of the University of Glasgow (A, B), Mrs. C. Gregory, and
J. Hook (D)).

20 - RICH & ARCHBOLD

in his Dead Heart of Australia published in 1906 in London, as well as in a number of local
newspapers including the Melbourne Age (Fig. 12). It was not until half a century later that
the full potential of this area was recognized when R. A. Stirton (see detail in account by R.
H. Tedford, this volume) followed in some of Gregory's footsteps. Stirton not only relocated
many of Gregory's Pleistocene locales, but he discovered the first concentrations of Tertiary
terrestrial vertebrates ever found on this continent.

Gregory was not only associated with the university system but also simultaneously served
as Director of the Geological Survey in the Mines Department of Victoria (from 1901),
demonstrating the often repeated pattern in Australia's early history of geology of a single
person serving concurrently in several institutions (such as museums, surveys, universities).
He also became involved in the Victorian Chamber of Mines and was on the council of the
Australian Institute of Mining Engineers (now the Australasian Institute of Mining and
Metallurgy) as well as an office-holder in the Royal Society of Victoria (Branagan & Lim
1984). Even more than this, he became involved in presenting extension courses, in addition
to his already heavy load of university courses, and was interested in both primary and
secondary education. This was, in part, a reflection of how small the scientific community
was, and how talented people had to carry many different responsibilities. Gregory was an
extremely energetic man, which probably helped immensely to manage two such jobs. George
(1975) described this restless energy: "By repute he [Gregory] could at any one time nurse his
infant on his knee, correct the proofs of one book with his left hand while writing another with
his right, and dominate a polemical discussion on any topic." Gregory, despite seemingly
boundless reserves, finally resigned his post at the university in despair in June of 1904
because of his inability to extract enough funding from the government to operate effectively.
Ironically, only a few months later, in September, the same month he returned to Scotland
(where he took the Chair of Geology at the University of Glasgow), the funding became
available, unfortunately, too late. Gregory continued his expeditionary work in many parts of
the world until he drowned on an expedition to the Amazon Basin in 1932. Appropriately and
ironically Gregory had copied a poem on the fly-leaf of the first of his Peruvian notebooks:

"I wander'd till I died.

Roam on! The light we sought is shining still.
Dost thou ask proof? Our tree yet crowns the hill.
Our Scholar travels yet the loved hill-side."

Alexander M. Thompson, who occupied the Chair of Geology at the University of Sydney
from 1866, had sufficient interest in fossil vertebrates to spend a considerable time in the caves
in the Wellington area. This obsession most likely hastened his early death in the 1870's
shortly after an expedition to the caves with Gerard Krefft (Branagan & Townley 1976).

Another university personage of marked significance to palaeontology in Australia was
Ralph Tate (1840-1901). He arrived from England to take up the foundation chair in geology at
the newly founded University of Adelaide in 1874. Tate taught palaeontology with an
infectious enthusiasm and an open mind and lured his students and volunteers into the field by
packing a keg of beer as part of the field gear (Alderman 1967, Vallance 1978)! His research
was careful, abundantly published and of good quality (e.g. Tate 1893), and his production of
good students was unrivalled at the time. Besides his charisma, Tate also held the view that
the Australian record should be viewed as separate and independent from that elsewhere and "had
little but scorn for those he thought believed all the rules of geology were written in Europe.
"Sir F. McCoy appears to object to any Australian deposits being called Eocene unless the
fossil species are identical with those occurring in the London Clay, Paris Basin, and other
European Eocenes, peculiar Australian species being open to grave suspicion." Tate is
remembered as one of the first Australian palaeontologists, who was not always looking over

HISTORY OF AUSTRALIAN VERTEBRATE PALAEONTOLOGY - 21

140
Sketch map of! the

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Figure 12. Map of the Lake Eyre re di
in the early twentieth century. Gregory discovered many vertebrate fossil-bearing localities along many of
the creeks draining into Lake Eyre. (From Gregory 1906).

22 - RICH & ARCHBOLD

his shoulder for advice from abroad or trying to shove the Australian record into a European
mould (Vallance 1978).

The universities provided permanent, indigenous positions for professional palaeontolgists,
and the first Australian-educated students began to appear. T. S. Hall, who was trained by F.
McCoy and worked with the Victorian Geological Survey under Selwyn, was the first of many.
Although locally-trained vertebrate palaeontologists did not appear until the twentieth century
in Australia, palaeontology by then had its independence. For the sub-discipline of vertebrate
palaeontology, Australian maturity was deferred, mainly due to the lack of a trained
professional base in this country and the absence of financial support for their work.

RAPID COMMUNICATION ON A LOCAL SCALE: THE SCIENTIFIC
SOCIETIES

Yet one other factor which gave independence of scientific endeavour in Australia (Branagan
and Townley 1976) was the establishment of local scientific societies (Prince 1979). Patterned
after groups such as the Royal Society of London, these societies gave the educated gentry a
chance to exchange information at meetings and perhaps, more importantly, to give local
scientists a rapid, local source of publication. One of the earliest of the societial publications
was the Tasmanian Journal of Natural Science, Agriculture, Statistics &c published by the
Tasmanian Society for the Advancement of Natural Science or The Tasmanian Society, as it
came to be known, a society that was founded in 1839 and flourished under the leadership of
Sir John Franklin (Plomley 1969). The Tasmanian Journal, first published in 1841, continued
into the 1840's (Fig. 13).

MEN OF INFLUENCE, PIONEERS IN AUSTRALIAN VERTEBRATE
PALAEONTOLOGY

During the transition period from dependence to independence, Basalla's stages 1 and 3, a
number of scientists, collectors and interested individuals substantially influenced the course of
vertebrate palaeontology in Australia. These included the Rev. William B. Clarke, Alexander
Macleay, Edmund C. Hobson, the Rev. Julian Tenison Woods, Frederick McCoy, Gerard
Krefft, Robert Etheridge Jr., H. Y. L Brown, Charles W. DeVis, E. C. Stirling, A. H. C.
Zietz and J. W. Gregory, to mention but a few.

1839 marked the arrival of the Reverend W. B. Clarke (1798-1878) in Sydney, migrating
with his family to Australia because of his own ill health. Here he assumed the position as
Rector of Willoughby, North Sydney after coming from a background of study at Cambridge
University. He had been very much influenced by Adam Sedgwick, Professor of Geology at
Cambridge University. Clarke maintained a close friendship with Sedgwick throughout his life
(Branagan in Stanbury 1975). On coming to Australia, Clarke was an avid prospector for
fossils and a keen geologist and took every opportunity to show visiting scientists the local
rocks of the Sydney area. He had an ongoing correspondence with many well-known
geologists such as Murchison and Sedgwick in England and Dana in the United States. He
certainly did not work in an isolated atmosphere. He managed to do more than just collect and
serve as a geological tour guide; he also published a number of articles in the local newspapers
and in both Australian and overseas journals, especially on the geological record of the Sydney
environs (Branagan & Townley 1976). He drew together many of his geological and
palaeontological ideas and the results of several survey trips in The Sedimentary Formations of
New South Wales, which appeared in several editions between 1867 and 1878. Despite this

HISTORY OF AUSTRALIAN VERTEBRATE PALAEONTOLOGY - 23

THE

TASMANIAN JOURNAL

Oo;

NATURAL SCIENCE, AGRICULTURE,
STATISTICS, &e.

VOL. I.

TASMANIA:

TAMES BARNARD, GOVERNMENT PRINTER, HOBART.

LONDON:

JOHN MURRAY, ALBEMARLE STREEP.

1842.
Royal SocieTY OF TATMANIA
Figure 13. The front page of the initial issue of the Tasmanian Journal of Natural Science, Agriculture,

Statistics, &c. , an early locally produced scientific journal that made it possible for local, rapid publication
of scientific papers. (Courtesy of N. Kemp).

24 - RICH & ARCHBOLD

effort on his part, however, most of the vertebrate fossil material that he collected or that was
sent to him was forwarded to scientists abroad for final study.

A part of the same Sydney community to which Clarke belonged was Alexander Macleay
(1767-1848), already mentioned above in the context of museum development. His large
house at Elizabeth Bay with its exquisite library, by far the best palaeontological library in
Australia at the time, plus his natural history collections, certainly served as an intellectual
hub of this colonial town. But, although he wrote articles for the local newspapers about
fossils, he published no substantial scientific papers. His contribution was primarily that of a
great resource on which the infant scientific community could build.

It was certainly no easy task to foster or pursue science in a society that was small, nearly
half unlettered convicts and cut off from rapid communication with other centres of scientific
endeavour in Europe and England. Scientific efforts were restricted to a small group of educated
men who had both the training and the means of support to keep themselves going.

Julian Tenison Woods (1832-1889) was another early worker on vertebrate fossils (Fig.
14). A Roman Catholic priest-geologist, he led a varied and somewhat controversial life that
took him from England to Australia to Europe and Asia (including China) and back to
Australia where he died (Press 1979). He spent much of his time in Australia in the area
around Penola, to the southeast of Adelaide. He collected in the mid to late Cainozoic deposits
in this area and wrote a number of scientific papers based on the material that he had collected,
or had collected for him, such as the large bird bones from native wells around Penola, some of
the first dromornithids reported. He maintained an active correspondence with geological
enthusiasts (Player 1983) as well as other palaeontologists and geologists in Australia, such as
William Macleay (Press 1979) and overseas, such as Sir Charles Lyell, one of the most
prominent geologists of the time. He was also an active member, even president (Linnean
Society of New South Wales from 1879-1880), of several scientific societies and a member of
the Board of Trustees of the Australian Museum (1880). He used what few chances there were
for intellectual pursuits within his parish when he stopped for a while on properties such as
that of Samuel Pratt-Winter at Murndel near Hamilton, Victoria. Murndel was ideal for
Woods, as it combined an abundance of Miocene fossils (marine), a superb library and a well
educated and well travelled land-owner (Press 1979). Even though Pratt-Winter was not a
Roman Catholic, he would travel to Tenison Woods presbytery in Penola, when going to Mt
Gambier, and often would bring books for the priest to read in his absence from Murndel.
Woods certainly didn't confine his explorations to southeastern Australia, but made a number
of excursions to several parts of the continent, for both scientific and religious reasons. He
visited Malaysia and China, and, of course, he spent his early life in England. Despite the fact
that Woods described and named many of the fossils that he collected, however, he often
forwarded specimens to Melbourne or to London to the experts for a final decision. He oft
times used such specimens as trade items with other workers such as Macleay in Sydney.

A major influence in South Australia was H. Y. L. Brown (1844-1924), a Nova Scotian
who received his training initially at the Royal School of Mines in London (Fig. 15). He
worked for a time in both Canada and New Zealand. He trained under Selwyn in the Victorian
survey. He spent time in New South Wales before taking a job in 1882 as the government
geologist of South Australia. He held that job for 30 years and used all manner of transport to
cover much of South Australia and the Northern Territory (then a part of South Australia) in
his geological surveys. He was a man of catholic tastes, and his reports and maps (Fig. 16)
detailed not only the geology but also water resources, local environment, distribution of fauna
and flora, and ethnology of each area he visited.

Brown (1892, 1894) collected the remains of Diprotodon, a giant, now extinct marsupial,
and the giant goanna, Megalania, as well as a number of other bones in the area northeast of
Lake Eyre. He noted that the native peoples of this area accounted for the presence of such

HISTORY OF AUSTRALIAN VERTEBRATE PALAEONTOLOGY - 25

B

Figure 14. Julian Tenison (J. E. T.) Woods as a young priest in South Australia (A) and a traveller (B) in
southeast Asia. He was an important early collector and writer on the geology and palaeontology of
southeastem Australia. (From Press 1979).

bones by their being remains of the cadimurka , a large fish that lived in the bottom of the
waterholes in the area. These cadimurka had never been seen alive by anyone, however.

Simultaneous with Brown's work in central Australia was the first major expedition
collecting vertebrate fossils, that at Lake Callabonna carried out by the South Australian
Museum (Figs 17, 18). Brown (1894) reported on this expedition. H. Hurst was originally
sent to Lake Callabonna by the South Australian Museum after an Aboriginal stockman
pointed out the occurrence of giant bones on the lake's surface. Parts of 80 skeletons of large,
extinct vertebrates, mainly Diprotodon, were discovered by Hurst, and later by E. C. Stirling &
A. H. C. Zietz (Fig. 10) and others from the South Australian Museum who took over from
Hurst. Brown's report on the area (dated 27 June 1893) astutely recognized the importance of
this site: "In view of the importance of preserving these relics of a bygone age for the future
scientific exploration I would recommend that the whole area of the lake be reserved for that
purpose, and to prevent the indiscriminate digging up and removal of portions of the
specimens." This recommendation was implemented on 30 November 1901 by the South
Australian government.

In the late 19th century two scientists with palaeontological interests stand out as
independent workers who did not automatically seek foreign expert opinion to give credence to
their own ideas. These were Johann Ludwig Gerhard Krefft (1830-1881) and Robert Etheridge
Jr. (1847-1920) of the Australian Museum, Sydney. Both men strongly believed in their own
ability to make reliable decisions without outside confirmation. Etheridge was an imaginative
and careful scientist and together with Ralph Tate supported the idea that the Australian
stratigraphic sequences might not be a direct reflection of those in the Northern Hemisphere.

26 - RICH & ARCHBOLD

Figure 15. H. Y. L. Brown (A, B younger, and C, older) was responsible for the discovery of much fossil
vertebrate material from the interior of the continent, mainly South Australia and the Northern Territory.
(Courtesy of the South Australian Archives, Adelaide, and N. Pledge).

HISTORY OF AUSTRALIAN VERTEBRATE PALAEONTOLOGY - 27

MAP

TO ACCOMPANY

GOVERNMENT GEOLOGISTS REPORT
On country in the neighbourhood of do

. LAKE EYRE |
a . . fy

“\_L. HOPE

Recent 4 Tertiary on Mesozvce

(Cretaceous 4 Ooltac)

aw)

SLAKE EYRE)

= (North ) :
g

HARRY BORE
(ARTES(AM)

/ Tertary on/Mesozou /
A ‘

f
LAN nigh

an ‘Raiden SCALE ri , Northern oid of the Mairv a Bande
ite eee ES te" (Primary rocks)

AY L.Brown, Govt Geologist. 11 10.92

SURVEYOR CENERALS OFFICE ADELAIDE A Viughan hotel thographer
PP NO /4/.\

Figure 16. Map of the Lake Eyre Sub-basin prepared by H. Y. L. Brown, then the Government Geologist
for South Australia. This area was rich in fossil vertebrates, many of which were discovered by Brown.
(Courtesy of the National Library of Australia and G. F. van Tets).

Krefft is often remembered for his public disagreement with Sir Richard Owen over the
feeding habits of the fossil marsupial Thylacoleo, clearly demonstrating his belief in the value
of his own opinion. Krefft had sent Owen material for study for some time, but he rather
unfortunately stopped this when Owen interpreted the jaws and teeth of Thylacoleo as
belonging to a carnivorous animal. Krefft adamantly and rather unscientifically disagreed,

28 - RICH & ARCHBOLD

claiming that the beast was a plant-eater, something like a giant rat kangaroo. Krefft remained,
throughout his life, an outspoken defender of his own, independent opinions. He is best
known for his work on fossil mammals, but he worked on a number of other vertebrates as
well (e.g. Whitley 1958-1959, 1967-1968; Krefft 1866, 1870, 1873; Archer & Clayton 1984).

Robert Etheridge Jr. first came to Australia to be a part of Selwyn's geological survey.
Although he returned to England after his service in the survey, he was drawn back to Australia
by an abiding interest in this new country, and was important in establishing the science of
vertebrate palaeontology on this continent. He served as a palaeontologist at the Australian
Museum and as well as a palaeontologist with the Geological Survey of New South Wales
(Strahan 1979) and was eventually appointed as Director of the Australian Museum in 1887,
where he remained until 1919. During that time he worked on fossils from all parts of
Australia and had close working links with other geological survey personnel, such as H. Y. L
Brown. He was "aloof, rather dour ...and shared his enthusiasms with few, though so many
profited by them" (Vallance 1978). He had an incredible capacity for work, and his
publication record was impressive (more than 400 papers) (e.g. Etheridge 1878, 1918;
Etheridge & Jack 1882, Jack & Etheridge 1892, Dun & Rainbow 1926). Even more
impressive was the accuracy of his assessments in those papers. His careful scientific work
has stood the test of time. "Etheridge's writings, like Tate's, betray a well-informed sense of
historical scholarship" (Vallance 1978), and together with Tate, Etheridge set the stage for
independence in Australian palaeontology.

E. C. Stirling and A. H. C. Zietz (Fig. 10), based at the South Australian Museum, are
perhaps most remembered in vertebrate palaeontology for their excavations at Lake Callabonna
in South Australia (Figs. 17, 18). Fossil bones were originally discovered at Lake
Callabonna by an Aboriginal stockman (?Jackie Nolan) who reported them to Mr. F. B.
Ragless. Two days later Mr. Ragless visited the site. A few days later the station cook also
visited the site, and knowing that there was a reward posted for the recovery of the feet of
Diprotodon, took the bones to Adelaide to claim the reward. Because of the confusion
concerning just who should receive the reward, no one ever did! At this point the South
Australian Museum dispatched Mr. H. Hurst to investigate the discovery in January 1893.
After four months of field work a considerable amount of material was returned by "buck-
board" buggy by Hurst (Stirling & Zietz 1900). After evaluation of the Hurst work, Stirling
and Zietz decided to return themselves to Callabonna in August of 1893 and Hurst resigned his
appointment upon their arrival. Despite appalling field conditions including bogged camels,
the difficulty of acquiring feed and firewood, rabbit plagues, illness and high temperatures, a
major part of the world's largest collection of Diprotodon skeletons was recovered and
subsequently transported to the South Australian Museum in Adelaide.

One of the first excavations of its kind, where whole animals were being recovered in
numbers, Lake Callabonna gave Australia's small population and that of the world a glimpse
of what the entire skeleton of such animals as the giant marsupial Diprotodon (Fig. 18) and
the massive bird Genyornis really looked like. In other localities known up to that time
skeletons were disarticulated, not associated, because of the jumbling that occurred in caves,
stream channels, and even swamp accumulations. Stirling and Zietz were not simply field
collectors, but also studied and published on what they had found, producing a series of
excellent, large format, well illustrated monographs on a variety of the Callabonna vertebrates
(Stirling & Zietz 1896a, 1896b, 1900, 1905, 1913). Sir Richard Owen, who had toiled so
long and hard on understanding Diprotodon, would have envied such work, or would perhaps
have done the work himself had he the chance. Ironically, Owen died in 1892, the very year
that bones were discovered at Lake Callabonna and never knew what the feet of his treasured
Diprotodon looked like. Callabonna held the secret that Owen would never know.

Another late nineteenth century vertebrate palaeontologist was Charles de Vis (1829-1915)
(De Vis's name is variously spelled DeVis and deVis). He differed from Gregory, Brown,

HISTORY OF AUSTRALIAN VERTEBRATE PALAEONTOLOGY - 29

Figure 17. Lake Callabonna, South Australia (above) and skeleton of a Diprotodon weathering out on the
surface of the lake during the South Australian Museum's expedition to this area in the late nineteenth
century. (From Stirling and Zietz 1913).

30 - RICH & ARCHBOLD

Figure 18. Lake Callabonna, skeleton of a Diprotodon partially excavated (above) and fully prepared and
mounted (below). These specimens were recovered by the South Australian Museum's expedition in the late
nineteenth century led by Hurst initially and then later by Stirling and Zietz.

HISTORY OF AUSTRALIAN VERTEBRATE PALAEONTOLOGY - 31

Stirling and Zietz in that he did little field collecting himself. Instead, he worked on the
collections made by others, such as Brown and Gregory. His origins lay in Manchester and he
worked at many jobs upon coming to Australia, including being a librarian in Rockhampton
for some time. He often published articles in local newspapers, such as The Brisbane Courier,
The Telegraph and The Queenslander) under a pseudonym, Thickthorn (Ingram 1986), Later he
assumed a curatorship at the Queensland Museum, and there he published profusely, naming
many new forms, primarily from late Tertiary and Quaternary deposits. His comparative
collections were exceedingly small, and his communication with the remaining scientific world
was hampered by distance. Primarily because of this, and his lack of understanding of
variability within species, most of the material he described was set up as new, and presently
extinct species. He seems also to have believed that all fossil forms must represent extinct
species. Thus, many of the original deVis names have been found to be invalid, the species
instead representing fossil remains of extant taxa. In fairness to de Vis, however, it is worth
remembering the isolation in which he worked, and the minimal funds and the small
comparative collections with which he dealt. Besides his scientific work, de Vis made a
significant contribution to museums serving as educational institutions based on his experience
in Britain (Kohlstedt ms.).

As well as full-time professionals, a variety of other part-time vertebrate palaeontologists
were important . Robert Broom (1866-1951) serves as an example. Probably best known for
his work on australopithecines in South Africa, he also spent a time as a medical practitioner
in Australia. He arrived in Sydney on 28 May 1892 (Hunt 1974), and spent four years on the
continent. He spent the greatest time in Taralga, New South Wales as the town's doctor, but
found enough time to collect fossil vertebrates from the Wombeyan caves, despite some
resistance from the New South Wales government. Some of this material eventually was
deposited in the Australian Museum, but the vast majority of it followed him overseas when
he returned to Glasgow, perhaps in part because of the resistance of officialdom to his work in
the caves of New South Wales, and perhaps due to a somewhat cool reception that he received
at times from Etheridge.

By the beginning of the 20th century Australian-based vertebrate palaeontologists were
collecting, describing and thinking about Australian fossils. They were no longer
automatically shipping them overseas. But, there was still little funding for this science,
either for collection and study or for the hiring of professional vertebrate palaeontologists in
permanent positions. As a result, students were not being trained in this discipline on this
continent, and little expeditionary work was mounted locally.

AUSTRALIAN INDEPENDENCE AND INTERNATIONAL
COOPERATION

Vertebrate palaeontology in Australia has seen decided expansion during the 20th century,
especially since the 1950's. The greatest activity has occurred in the study of fossil fish and
mammals. Both of these disciplines have provided biostratigraphic information very useful in
establishing rock sequences in the deformed Devonian sediments in eastern Australia as well as
in the flat-lying, monotonous Tertiary carbonate-rich channel deposits that mimic the
underlying Cambrian marine limestones in northern Australia.

32 - RICH & ARCHBOLD

4 eit se Bn ealliaaet as
: se all i . oo

Figure 19. The early 1950's saw the discovery of concentrations of Tertiary marsupials in northem South
Australia. This discovery was the result of joint expeditions of the University of California (Berkeley) and
the South Australian Museum. Three individuals were instrumental in these discoveries: R. A. Stirton
(above, right) from the University of California, P. Lawson (above, left) from the South Australian Museum
arid R. H. Tedford (below) from the University of Califomia. R. H. Tedford presents a detailed account of
this pioneering work in his chapter in this volume. (Courtesy of P. Lawson).

HISTORY OF AUSTRALIAN VERTEBRATE PALAEONTOLOGY - 33

EXPLO
ig 4 ce MUSEU mS oe r

Figure 20. International cooperation of the National Museum of Victoria and the Denver Museum in a joint
expedition to the Nullarbor. Edmund Gill, a vertebrate palaeontologist from the Victorian museum (fourth
from right), was a prominent member of the expedition. (Courtesy of the Museum of Victoria).

During this period, advancement has been greatly influenced and enhanced by a few
energetic workers and the discovery of important new fossil fields (see also Tedford (Chap. 2)
and Plane (Chap. 3) this volume), It was also during this period that enough information
began to accumulate to make meaningful summaries possible (Chapman 1914a, 1914b, Hills
1958, Ride 1964, Rich & Thompson 1982, Archer & Clayton 1984).

Names that stand out amongst the many vertebrate palacontologists who worked or are still
working during the twentieth century are E. S. Hills (Aust.), R. A. Stirton (U.S.A.), R. H.
Tedford (U.S.A.) (Fig. 19), M. O. Woodburne (U.S.A.), W.D.L. Ride (Aust.), J.A. Mahoney
(Aust.), E, D. Gill (Aust.) (Fig. 20) (Gill 1953, 1957, 1965a-b, 1968), E. Lundelius (U.S.A.),
W. Turnbull (U.S.A.), M. Archer (Aust.), R. Wells (Aust.), P. Murray (Aust.), T. F.
Flannery (Aust.), J. Hope (Aust.), S. Hand (Aust.), N. Pledge (Aust.), J. A. Long (Aust.), T.
H. Rich (Aust.), A. Ritchie (Aust.), KS, Campbell (Aust.) S. Turner (Aust.), R. Miles
(U.K.), G. Young (Aust.), R. Molnar (Aust.), and T. Thulborn (Aust). All of these scientists
contributed significantly in major field discoveries, prolific description of new taxa and in
novel interpretation of the newly found material. Hill's (1958) review of the entire field of
Australian vertebrate palacontology and Ride's (1964) summary of Australian
palacomammalogy were the first real attempts to draw together the rapidly accumulating data.
Hills’ as well as Long's, Young's, Campbell's, Ritchie's and Miles’ specialty is Devonian fish,
while Stirton, Tedford, Woodburne and their associates finally located the first concentrations
of Tertiary mammals in Australia, in the Lake Eyre Basin where H.Y.L. Brown and J. W.

34 - RICH & ARCHBOLD

Gregroy had trekked with camels. Gill, through his enthusiasm as a collector, and his
publications, also promoted vertebrate palaecontology, especially in Victoria.

Figure 21. Joint British Museum (Natural History) and Western Australian Museum expedition near the
productive sites at Gogo, Wester Australia. (Courtesy of the British Museum (Natural History).

Lundelius and Turnbull together with staff (including Duncan Merrilees) from the Western
Australian Museum developed the Pleistocene record of Western Australia as well as the unique
Pliocene site of Hamilton in Victoria, one of the few radiometrically dated vertebrate sites in
Australia. Stirton and Ride promoted Australian vertebrate palaeontology not only by their
own field and research work, but by training a number of students and collaborating with
scientists around the world on Australian projects. Current researchers, such as T. H. Rich and
M. Archer studied with Stirton and Ride respectively, and they in turn have supervised
additional students, for example T.F. Flannery and J.A. Long. The current field of vertebrate
palaeontology has much expanded over that of the 1950's. The long list of professionals in
this field given in Rich & van Tets (1985) is a reflection of the current level of activity (Fig.
21). Each has made their own unique contribution, from setting up an economically useful
microvertebrate biostratigraphy (S. Turner) to collecting and mounting impressive public
displays (T. Flannery, A. Ritchie) to guiding museums filled with vertebrate fossils (A.
Bartholomai). Activity is by far the highest it has ever been in this field in Australia.

HISTORY OF AUSTRALIAN VERTEBRATE PALAEONTOLOGY - 35

Figure 22. A joint expedition of the British Museum (Natural History) and the Western Australian Museum led
to the discovery in the mid-twentieth century of the rich Devonian Gogo locality in northwestern Wester
Australia. (Courtesy of the British Museum (Natural History).

36 - RICH & ARCHBOLD

Figure 23. Meeting of many of the currently practicing vertebrate palaeontologists in Australia at the
Conference on Vertebrate Evolution and Systematics held in March 1989 at the Australian Geographic
headquarters in Sydney. Front Row (left to right): 1, Groves; 2, Jeanette Muirhead; 3, Rhys Walkley; 4, ?;
5, Corrie Williams; 6, Dietlind Knuth; 7, Coral Gilkeson; 8, Sue Creagh; 9, Anne Warren; 10, Sue Hand.
Back Row (left to right): 1, Tony Thulborn; 2, Michael Loy; 3, Zhang Gue Rui; 4, Gavin Young; 5, Robert
Jones; 6, Tim Hamley; 7, Susan Bergdolt; 8, Bernie Cooke; 9, Brian Mackness, 10, Miranda Gott; 11, Mike
Durant; 12, John Barry; 13, Walter Boles; 14, Neville Pledge; 15, Julie Barry; 16, Arthur White; 17, Alex
Ritchie; 18, Jim Lavarack; 19, Mary White; 20, Sue Lavarack; 21, Pat Rich; 22, ?; 23, Tom Rich; 24, 7; 25,
John Long; 26, Henk Godthelp; 27, Peter Murray; 28, Paul Willis; 29, Michael Archer; 30, John Scanlon.
(Courtesy of Australian Geographic).

Discoveries such as those in the Wellington Valley and at Lake Callabonna stand out as
significant in the nineteenth century, when Australian vertebrate palaeontology was just being
weaned. Several significant finds mark the twentieth century as well. These finds were made
by a variety of people, mainly Americans, Australians and British. Such localities include:
Gogo, a number of rich Devonian fish localities on Gogo Station in Western Australia
discovered and developed by the British Museum (Natural History) and the Western Australian
Museum (Figs 21, 22); the Devonian armoured fish localities in western New South Wales
and southeastern Victoria, primarily developed by personnel from the Australian Museum, the
Museum of Victoria and Monash and Melbourne universities, Australian National University
and the Bureau of Mineral Resources; the Cretaceous terrestrial and marine sequences
containing reptiles in southwestern Queensland and Victoria, the former through the efforts of
the Queensland Museum, Harvard University, and the British Museum (Natural History) and
the latter primarily by the Museum of Victoria and Monash University. The Tertiary
vertebrate-bearing clays and sands of the Great Artesian Basin originally discovered and
developed by R. A. Stirton and his collegues based at the University of California, Berkeley,
together with the South Australian Museum (sce Chap. 2 by R. H. Tedford, this volume, and
in Rich & van Tets 1985), was further explored by a number of American and Australian

HISTORY OF AUSTRALIAN VERTEBRATE PALAEONTOLOGY - 37

groups, and is still a research focus. The mid to late Cainozoic fossiliferous Riverslcigh area
of northwestern Queensland is one of the rare jewels that was discovered during the initial
exploration of the 20th century, but its real potential and significance lay unrecognized until
Michael Archer (University of New South Wales) initiated a detailed survey of the area in the
1980's. And finally, the Quaternary cave deposits along the southern and eastern parts of
Australia, investigated by a number of Australian and American workers from many
institutions (Australian National University (Canberra), Flinders University (Adelaide),
Western Australian Museum, University of Texas, Field Museum of Natural History in
Chicago, the University of California at Berkeley, amongst others) have produced very large
collections of vertebrate material, some quite young, but including many extinct forms.

The discovery by R. A, Stirton (Fig. 19) during the mid-20th century of Tertiary
vertebrates in the Lake Eyre Basin is particularly momentous. It not only found a number of
previously unknown mammals and birds, some quite unusual and quite distinct from any of the
living groups in the Australian extant or Quaternary fauna, but it quite clearly led to a real
acceleration of activity in Australian vertebrate palacontology. Ruben Arthur Stirton had come
to Terra Australis, an explorer in 1952, to find pre-Pleistocene mammals in quantity. He did
what he came to do (Stirton ef al. 1961, Stirton et al. 1968). From those expeditions that he
led, fanned on by his infectious enthusiasm, came students, funding and a momentum that
continues to the present.

By the 1970's and early 1980's Australian vertebrate palacontology had finally come of age.
Certainly by this time, Basalla's stage 3 had been reached. Australian vertebrate
palaeontologists were quite visible and looked for the most part to their overseas colleagues as
equals and co-workers. Indigenous expeditions as well as a number of joint expeditions
involving such groups as the South Australian Museum (Adelaide), the University of
California (Berkeley and Riverside), the Bureau of Mineral Resources (Canberra, especially
with regard to work in the Northern Territory and Papua New Guinea (see Plane, this volume,
Chap. 3), the American Museum of Natural History (New York), the Smithsonian Institution
(Washington, D. C.), the Queensland Museum (Brisbane), the Museum of Victoria
(Melbourne), the British Museum (London), the Field Museum of Natural History (Chicago),
the University of Texas (Austin), the University of New South Wales (Sydney), Monash
University (Melbourne), and the Australian Army. This approach differs vastly from the
colonial days where material was often collected by local collectors and then shipped overseas
for study by people such as Owen. By this time field work and theoretical work alike was
being carried out by both locals and foreign experts, and neither group tried to shove the
unruly Australian record into a European or North American mould.

Today with a number of full time and part time positions filled by vertebrate
palaeontologists in Australia, research and training of personnel locally is ensured. Although
becoming increasingly restricted, funding for field work, research and publication in this area is
available locally from both governmental and private sources. Because of this together with
the lively interest of scientists, and some funding agencies (such as the National Geographic
Society and Earthwatch) and private enterprise, from around the world, in the development of
Australia's unique biota, a lively period of discovery is insured for what remains of the 20th
century. Much of this work will undoubtedly be directed by Australians, but it will be
significantly enriched by interactions on an international level, interactions which should be
encouraged and nurtured. The scientific findings will, in turn, be of no small interest to the
general public (Fig. 24).

38 - RICH & ARCHBOLD

Figure 24. An official crest, of Mr. Rhys Walkley, which incorporates one of Australia's extinct fossil
ventebrates, Diprotodon, here shown with a trunk, (Courtesy of R. Walkley).

HISTORY OF AUSTRALIAN VERTEBRATE PALAEONTOLOGY - 39

ACKNOWLEDGEMENTS

Many people are to be thanked for their assistance in preparation of this paper; D.F.
Branagan and W.D.L. Ride for their reviews; T.H. Rich, M.V. Macdonald, J.R. Macdonald, J.
Monaghan, and C. Williams for their editorial remarks; E. B. Joyce for the lend of his copy of
T.L. Mitchell (1838); several individuals noted in the figure captions for providing illustrative
material; D. Gelt, S. Morton and F. Coffa for illustrative work and material; A. Player for
information on J.E.T. Woods; and Australian Geographic, Dick Smith, S. Hand and J.
Lavarack for providing Fig. 23.

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Rept. Brit. Assoc. Adv. Sci., York. (1844) 14: 223-240

OWEN, R., 1877. Researches on the fossil remains of the extinct mammals of Australia, J, Erxleben,
London.

OWEN, R., 1879a. Memoirs on the extinct wingless birds of New Zealand with an appendix on those in
England, Australia, New foundland, Mauritius and Rodriquez. J. van Voorst, London.

OWEN, R., 1879b. On Dinornis containing a restoration of the skeleton of Dinornis maximus (Owen), with
an appendix on additional evidence of the genus Dromornis in Australia. Trans, Zool. Soc. Lond,
10(3): 147-188.

OWEN, R., 1879c. On Dinornis. Pars. 19: containing a description of a femur indicative of a new genus of
large wingless birds (Dromornis australis) from a post-tertiary deposit in Qld. Trans. Zool. Soc.
London, 8: 381-384.

OWEN, R., 1882. Description of portions of a tusk of a proboscidean mammal (Notelephas australis, Owen).
Phil. Trans. R. Soc. 173: 777-781.

PENTLAND, J.B., 1831. Une communication sur des ossemens trouves dans une breche calcaires sur la
riviere de Hunter, dans le Norde-est de la Nouvelle-Hollande. Bull. Soc. geol. Fr.1: 144-145.

PENTLAND, J.B., 1832. On the fossil bones of Wellington Valley, New Holland or New South Wales.
Edinburgh New Phil. J. 12(24): 301-308.

PENTLAND, J.B., 1833a. Uber die fossilien Knochen vom Wellington-Thale in Newholland (Sud-Wales).
Neu. Jahrb. Min. Geognos. Geol. Petrefaktenkunde. 4: 603-605.

PENTLAND, J.B., 1833b. Observations on a collection of fossil bones sent to Baron Cuvier from New
Holland. Edinburgh New Phil. J. 13: 120-121.

PENTLAND, J.B., 1833c. Beobachtungen iiber eine Sammlung fossiler Knochen, welche aus dem
Wellington-Thale in Neuholland am Baron Cuvier eingesendet worden. Neu. Jahrb. Min. Geognos.
Geol. Petrefaktendunde. 4: 605-606.

PESCOTT, R.T.M., 1954. Collections of a Century. The History of the First Hundred Years of the National
Museum of Victoria . Nat. Mus. Vict., Melbourne.

PLAYER, A.V., 1983. The Archer letters. Argyle Press, Goulbum.

PLOMLEY, N.J.B., 1969. The Tasmanian Joumal of Natural Science. Pap. Proc. R . Soc. Tasmania 103:
13-15.

PRESS, M.M., 1979. Julian Tenison Woods. Catholic Theological Faculty, Sydney.

PRINCE, J.H., 1979. The First One Hundred Years of the Royal Zoological Society of N.S.W. Surrey Beatty
& Sons, Sydney.

RANKEN, C.G., 1916. The Rankens of Bathurst. S.D. Townsend, Sydney.

RICH, P.V., 1979. The Dromorithidae, an extinct family of large ground birds endemic to Australia. Bull.
Bur. Min. Res, 189.

HISTORY OF AUSTRALIAN VERTEBRATE PALAEONTOLOGY - 43

RICH, P.V. & VAN TETS, G. F., eds., 1985. Kadimakara. Extinct Vertebrates of Australia. Pioneer Design
Studio, Lilydale.

RICH, P.V. & THOMPSON, E.M., eds., 1982. The Fossil Vertebrate Record of Australasia. Monash Univ.
Offset Printing Unit, Clayton.

RIDE, W.D.L., 1964. A review of Australian fossil marsupials. J. R. Soc. W. Aust. 47(4): 97-134.

ROBIN, L., 1987. Thomas Sergeant Hall (1858-1915): Scholar and enthusiast. Hist. Rec. Aust. Sci. 6(4):
485-492.

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as observed in several species of fossil shells. Tran. Linn. Soc. London 12: 514-516.

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of fossil struthious bird found at Lake Callabonna, South Australia. Trans. Proc. R. Soc. S. Aust. 20:
171-190.

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genus and species of fossil struthious bird. Mem. R. Soc. S. Aust. 1(2): 41-80.

STIRLING, E.C. & ZEITZ, A.H.C., 1905, Fossil remains of Lake Callabonna. III. Description of the
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STIRLING, E.C. & ZEITZ, A.H.C., 1913. Fossil Remains of Lake Callabonna. IV. Descriptions of some
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paleontology of the Tirari Desert, South Australia. Rec. S. Aust. Mus. 14(1): 19-61.

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VALLANCE, T.G., 1975. Origins of Australian geology. Proc. Linn. Soc. N.S.W. 100: 13-43.

VALLANCE, T.G., 1978. Pioneers and leaders - a record of Australian palaeontology in the nineteenth
century. Alcheringa 2: 243-250.

VALLANCE, T.G., 1981. The fuss about coal. Troubled relations between palaeobotany and geology. In
D. J. & S. G. M. Cart, eds. Plants and Man in Australia. Academic Press, Sydney: 136-176.

WHITLEY, G.P., 1958-1959. The life and work of Gerard Krefft. Proc. R. Zool. Soc. N.'S.W. 59: 21-34.

WHITLEY, G.P., 1967-1968. Gerard Krefft and his bibliography. Proc. R. Zool. Soc. N.S.W. 68: 38.

44 - RICH & ARCHBOLD

Ngapakaldia from Miocene terrestrial sediments of northern South Australia. These sheep-sized
diprotodontids inhabited the Centre when it was much better watered. (From Rich & van Tets
1985, with permission of The Museum of Victoria).

CHAPTER 2

VERTEBRATE
PALAEONTOLOGY IN
AUSTRALIA: THE
AMERICAN CONTRIBUTION

Richard H. Tedford!
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1 Department of Vertebrate Paleontology, American Museum of Natural History, New York, New York 10024,
U.S.A.

46 - TEDFORD

INTRODUCTION

It may seem presumptous in the context of the history of vertebrate palaeontology in
Australia to focus on an American contribution if it were not for its conspicuous presence in
the years after the Second World War. American financed and directed work in the early post-
war years demonstrated the necessity of direct exploration for Australia's fossil vertebrates.
This was not a foreign invasion directed from the outside with all the spoils going overseas,
but a cooperative venture between palaeontologists of the United States and Australia. The
most important fruits (type specimens) of this international effort remained in or were returned
to Australia to stimulate research in the country of origin. This cooperative approach to the
development of the fossil record is largely responsible for the post-war burgeoning of the field
in Australia, as has already been documented in historical reviews by Rich (in Rich &
Thompson 1982 and Rich & van Tets 1985), Archer & Hand (in Archer & Clayton 1984) and
in this volume.

To place the American contribution in context, I briefly examine the state of vertebrate
palaeontology in Australia before and just after World War II, and then detail the post-war
American presence during four decades, each of which is characterized by different levels of
involvement.

BETWEEN THE WARS

Australian vertebrate palaeontology was about as isolated from foreign involvement as its
unique vertebrate fauna during the years between WWI and WWII. The field, however, held a
number of prominent and active enthusiasts who continued to describe the fauna while carrying
out official duties in other areas. Three were museum directors, C.A. Anderson (Australian
Museum), L, Glauert (Western Australian Museum) and H. Longman (Queensland Museum),
and two were academics, E.S. Hills (Univ. Melbourne) and the Rev. R.T. Wade from Sydney.
Between these men there were studies of Devonian and Mesozoic fishes, Cretaceous dinosaurs
and marine reptiles, Cainozoic fishes, reptiles and mammals. Discoveries in the science
depended mostly on contacts with interested laymen and their initiative to send material to the
museum. Such field work as was carried out was often done within the investigators’ private
means. Nevertheless, important discoveries were made and promptly reported in the scientific
literature. Prominent among these were Hills’ studies on the largely unknown Devonian
fishes; Longman's description of the first important remains of Cretaceous dinosaurs, and
Anderson's work on the first Tertiary (Pliocene) marsupials from New Guinea.

Toward the end of this period Edmund Gill, palacontologist of the National Museum of
Victoria (now Museum of Victoria), began a geochemical investigation of the provenance of a
number of vertebrate fossils in that collection. Comparing the flourine content of museum
specimens with material obtained in situ of the purported sites, Gill (1957) was able to
demonstrate the Tertiary age of other remains from Victoria and Tasmania (including
verification of the provenance of the marsupial Wynyardia). This work came to fruition at the
time of the first American effort to find Tertiary mammals, and Stirton and Gill's collaboration
(1957) gave the first hint of the nature of the Neogene fossil record of marsupials.

THE 50'S, THE FULBRIGHT YEARS

There were a number of factors that brought the first American contingent of vertebrate
palaeontologists to Australia. Perhaps the most important was the "world-view" of the United
States in the heady days of outreaching international influence that gripped the nation at war's

AMERICAN CONTRIBUTION TO VERTEBRATE PALAEONTOLOGY - 47

end. This was manifested by such programs as the Marshall Plan, and most significantly for
academics, by the Fulbright scholarship program that eventually sent hundreds of American
graduate students and professors overseas.

The lack of documentation from the fossil record of the history of vertebrates in Australia
has long been noted by biologists. This large and intriguing biological problem focused
attention on the continent as a science frontier. In the years after World War II state and federal
geological surveys were increasingly active in mapping the areal geology of Australia, so that
the distribution and character of the outcropping rocks was becoming sufficiently well known
for exploratory work. This made it possible to actually find rocks of specific types and ages
and to plan field work to explore for a fossil record rather than wait for reported discoveries.
This combination of international science, an important biological problem, data to initiate a

ie and time and adequate funds to pursue it, all came together in the first decade after the end
of the war.

Figure 1. Four vertebrate palacontologists who worked in northem South Australia in 1950's: left to right,
Alden Miller, Richard Tedford, Paul Lawson and R.A. Stirton. (Courtesy of Paul Lawson).

Ruben Arther Stirton (Fig. 1), Professor of Paleontology at the University of California,
Berkeley, and his graduate student Richard H. Tedford, both obtained Fulbright Scholarships in

48 - TEDFORD

1953 and spent nine months in Australia searching for a Tertiary mammal record. They had
important allies in the Geology Department of the University of Adelaide, in the South
Australian Museum and the Mines Department of South Australia, all of whom generously
contributed important knowledge and support, including the help of G. Davidson Woodard, then
a University of Adelaide graduate student, and Paul F. Lawson of the Museum, Agreements
were forged with the South Australian Museum over the disposition of collections and many
other matters that have served as a basis for continued cooperation with that institution
extending to the present day. Lawson, in particular, continued to serve the joint project for
many active years and into retirement.

, a

Figure 2. The initial excavation of the Woodard Quarry, Lake Palankarinna, South Australia, July 1953.
The holotype lower jaw of the diprotodont Meniscolophus mawsoni Stirton, 1955 is in the centre foregound
between the brushes.

AMERICAN CONTRIBUTION TO VERTEBRATE PALAEONTOLOGY ~- 49

Figure 3. Looking southeast across the bluffs at the Tedford Locality on the western shore of Lake
Palankarinna, South Australia. White deposits are dolomitic mudstone within the Etadunna Formation
(Middle Miocene) overlain by darker sandstones of the Tirari Formation (Plio-Pleistocene),

The historical development of this work has been detailed elsewhere by Tedford (in Rich & van
Tets 1985). In brief, the Fulbright program made possible an extended reconnaissance
necessary to give the explorers time to become acquainted with the special conditions of the
Australian nonmarine Cainozoic. The serendipitous discovery of Lake Palankarinna (Figs 2-3),
east of Lake Eyre, in northern South Australia saved the 1953 effort from near failure and
served as a focal point for a number of successful field excursions in later years. During the
50's Fulbright and University of California intramural funds along with funds from the
collaborating Australian institutions enabled Stirton to send three additional expeditions (1954,
1957, 1958) to the Lake Eyre Basin. Personnel! on these expeditions varied, but Lawson (Fig.

50 - TEDFORD

4) served as liaison officer in all of them. Stirton was joined by his student Leslie F. Marcus
in 1954, and Marcus remained in Australia into early 1955 to work on the Bingara Pleistocene
fauna at the Australian Museum. William Ricdel (1954) and Brian Daily (1957) (Fig. 5) in
their capacity as Curator of Fossils at the South Australian Museum were also contributors.
Daily, in particular, set the framework for the Neogene stratigraphy of the Lake Eyre Basin.

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Figure 4. Paul F. Lawson at Lake Pitikanta in 1958 excavating a small marsupial skeleton from the Etadunna
Formation .

Tedford was the only American member in the 1957 field party from the South Australian
Museum that reaffirmed the presence of an important late Cainozoic record east of Lake Eyre.
This work recovered four superposed fossil vertebrate faunas of Miocene through Pleistocene
age in central Australia (Figs 6-15). All the taxa in the older assemblages were new to science,
considerably expanded the diversity of some families of living marsupials and established the
presence of extinct groups vindicating a long held belief that Australia's fauna had a lengthy
and complex history.

Ernest L. Lundelius Jr. (University of Texas) and William A. Turnbull (Field Museum of
Natural History) had also been intrigued as graduate students at the University of Chicago with
the lack of a record of mammals in Australia before the Quaternary. Lundelius was encouraged
to apply for a Fulbright by visiting lecturer in Zoology, A. R. Main, of the University of

AMERICAN CONTRIBUTION TO VERTEBRATE PALAEONTOLOGY - 51

Figure 5. Brian Daily plotting the expeditions course, lower Strzelecki Creek, South Australia, 1957.

52 - TEDFORD

Figure 6. Lower jaws and bones of the wallaby Prionotemnus palankarinnicus Stirton, 1955 in situ in the
Mampuwordu Sand, Lawson Quarry, Lake Palankarinna, South Australia, 1961.

Western Australia, and he went to that institution for his scholarship year in 1954-1955.
Lundelius was also intrigued by the research of Claude Hibbard of the University of Michigan
on the response of American mammal faunas to environmental change during the Quaternary.
When he saw the rich, undescribed Quaternary collections in the Western Australian Museum
and the potential of field work in this area, he recognized that material was at hand for similar
studies in Australia, Thus, his objective shifted to Quaternary and Holocene faunal sequences,
resulting in his first contribution in this field in 1960. This has remained a major focus of his
Australian research to this date.

AMERICAN CONTRIBUTION TO VERTEBRATE PALAEONTOLOGY - 53

THE 60'S, THE NATIONAL SCIENCE FOUNDATION
GRANTEES

In the 1960's support for work in Australia became increasingly costly, and the American
investigators sought funds from the National Science Foundation. Stirton's work eventually
involved earth moving equipment as well as the expensive field vehicles.

Field parties remained small, principally involving himself, Tedford and Lawson. Alden H.
Miller, University of California palaeornithologist, joined in 1961 and Stirton's graduate
student Michael 0. Woodburne in 1962 (Figs 16-21).

Figure 7. Looking south along the bluffs on the north-westem side of Lake Palankarinna, South Australia.
Vehicle on the right is below the Lawson Quarry; the Woodard Quarry is on the flats at the foot of the bluffs
just to the right of the farthest bluff, 1962.

54 - TEDFORD

Figure 8, Looking west across Lake Kanunka, South Australia. Bluffs expose fossiliferous Tirari and
Etadunna formations; Stirton Quarry lies behind the hill at the right hand end of the outcrop, 1957.

A new Miocene faunal level was discovered in the Lake Eyre sequence in 1962 and an
exploratory trip to the upper Sandover River north of Alice Springs in Northern Territory was
made by Stirton, Tedford and Woodburne to investigate a Late Miocene fossil site discovered by
the Bureau of Mineral Resources geologists. This site, on Alcoota Station, was so promising
that Woodburne returned in 1963 with U. C, graduate student John Mawby and BMR
personnel, to open a large excavation (Woodburne 1967) (Fig. 22). At the same time Tedford
with Alan Lloyd of the Bureau of Mineral Resources undertook a reconnaisance of Tertiary
deposits in southern Northern Territory and western Queensland finding a few Miocene
vertebrates at Kangaroo Well, south of Alice Springs (Deep Well Station), and reaffirmed the

AMERICAN CONTRIBUTION TO VERTEBRATE PALAEONTOLOGY - 55

Figure 9. Green Bluff, Warburton River, South Australia, looking downstream. The white limestone band is
in the Etadunna Formation claystones and is overlain by fossiliferous Pleistocene deposits, 1957.

occurrence of Miocene vertebrates from the Carl Creek Limestone on Riversleigh Station of
western Queensland ( Tedford 1967, Lloyd 1967).

In 1962 Stirton visited the Wau mining district in Papua-New Guinea to examine newly
collected Pliocene mammal remains from the gold-bearing Otibanda Formation (see Plane, this
volume, Chap. 3). These were the same rocks that yielded the first Tertiary mammals collected
from New Guinea described by Anderson in 1937, Geoffrey Woodard had been sent to Wau in
1955 by Stirton, but most of the small collection obtained was lost in shipping. The new
effort was spurred by the discoveries of Mike Plane of the Bureau of Mineral Resources
working out of Wau. Plane (1967a-b) subsequently studied all the material in Berkeley, where
he described it as the Awe Fauna.

56 - TEDFORD

Figure 10. R.A. Stirton ("Stirt") in 1958 at Stirton Quarry, Lake Kanunka, South Australia. The holotype
jaw of the extinct kangaroo Troposodon kentii Campbell, 1973 is in situ in the Tirari Formation at his nght

hand.

AMERICAN CONTRIBUTION TO VERTEBRATE PALAEONTOLOGY - 57

This decade saw two reviews of the developing Tertiary faunal sequence for Australia. In
1961, Stirton, Tedford & Miller presented the biostratigraphic framework for the Lake Eyre
Basin. The four faunas were augmented with a fifth Miocene assemblage in 1967 by Stirton,
Tedford and Woodburne. A further review combining all Australian (including the newly
discovered Miocene Bullock Creek Fauna, Plane & Gatehouse 1968) and New Guinea data into
a synthesis of a continent-wide faunal succession was published in 1968 by Stirton, Tedford &
Woodburne, two years after Stirton's fatal heart attack.

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Figure 11. Moming tea at Lake Ngapakaldi, South Australia in early August 1961 during early work at
Ngapakaldi Quarry. From left, R.A. Stirton, H.J. Bowshall, Alden and Virginia Miller.

Ermest Lundelius Jr. (now at University of Texas) returned to Australia in 1963-4 with
William Turnbull of the Field Museum, Chicago. Their NSF supported field work involved
both a search for Tertiary mammals in Victoria and Tasmania and continued work on
Quaternary and Holocene sites in Western Australia, particularly the caves of the Nullarbor
Plain. This work resulted in their obtaining a definitive Pliocene fauna, the first to be
radiometrically dated, from the site at Hamilton in western Victoria discovered by Gill in the
early 50's (Turnbull and Lundelius 1970). They also exploited the fossiliferous deposits in
Madura Cave on the Nullarbor Plain of Western Australia working out a !4C-dated Holocene
and Late Pleistocene succession (not yet completely reported in a series of papers of Lundelius

58 - TEDFORD

& Turnbull 1973, 1975, 1978, 1981, 1982, 1984, 1989). These sites were further exploited
by Lundelius and Turnbull during seven weeks in 1966-7 with funding from the University of
Texas and the Field Museum.

Figure 12. Removing overburden at the Stirton Quarry, Lake Kanunka, South Australia in 1961. Alden
Miller is at the scoop, while Tedford and Lawson mind the winch. (Courtesy of R. A. Stirton).

In 1960 Charles Camp of the University of California followed up a lead to Triassic
vertebrates in the Blina Shale of the Fitzroy Basin, northwestern Western Australia. Bureau of
Mineral Resources geologists had made the discovery in the early 50's, and the material had
been seen by Stirton in Canberra in 1953. Camp's work was conducted with the Western
Australian Museum, and his graduate student John Cosgriff was included in the field party.
This work, which recovered the first diverse Early Triassic vertebrate fauna from Australia, was
followed by further investigations by the Western Australian Museum in 1963 and 1964 (E. H.
Colbert of the American Museum of Natural History participated in 1964). In 1965 Jim
Warren and John Cosgriff revisited the sites for Monash University. Cosgriff (1965, 1969, and
with Garbutt, 1972) described the labyrinthodont amphibians from the Blina Shale, but the
other elements of this vertebraie fauna remain to be described.

AMERICAN CONTRIBUTION TO VERTEBRATE PALAEONTOLOGY - 59

*

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Figure 13. Two partial skeletons (both headless) of the small diprotodont Ngapakaldia tedfordi Stirton,
1967 in Ngapakaldi Quarry, Lake Ngapakaldi, South Australia, 1962. Skeleton in upper right of view lies on
its back, front feet to the right; skeleton at lower left lies on its right side, facing left.

In 1964 John Cosgriff was appointed a Senior Research Fellow in the Department of
Geology of the University of Tasmania, where he remained until 1967. During this period he
continued his studies of Triassic labyrinthodonts, culminating in his review of the Tasmanian
forms published in 1974 after his return to Wayne State University.

In 1961 the first American vertebrate palaeontologist appointed to an academic post in
Australia arrived at the newly formed Monash University. Dr, James Warren soon became
Professor of the Department of Zoology of that institution, and he initiated the first graduate
studies in vertebrate palacontology in eastern Australia. W. D. L. Ride, Director of the
Western Australian Museum held a joint appointment with the University of Western Australia
in 1957-1974, which included graduate studies in vertebrate palaeontology. _—_ His first Ph.D.

60 - TEDFORD

Figure 14. Lawson Quarry, Lake Palankarinna, South Australia, near the end of excavations in 1961.
Kangaroo long-bones on either side of geology hammer are in the base of the Mampuwordu Sand. Top of the
Mampuwordu Sand is about level with the bottle tops, vertical at the face is the Tirari Formation.

AMERICAN CONTRIBUTION TO VERTEBRATE PALAEONTOLOGY - 61

student in that field was the Fulbright scholar Michael Archer, who came from Princeton
University in 1967.

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Figure 15. Backhoe in operation at Lawson Quarry, Lake Palankarinna, South Australia in 1962. Lawson
operates the hoe, M.O. Woodbume collects material left exposed the previous year.

Toward the end of the decade the Geology Section of the Australian and New Zealand
Association for the Advancement of Science held the first symposium on vertebrate
palaeontology at their annual meeting in Melbourne in 1967. The meeting was dominated by
Americans (Stirton, Tedford, Woodburne, Lundelius, Turnbull, Cosgriff, and Warren) but
included reports by Michael Waldman, Warren's first Ph.D. student, and Michael Plane of the
BMR, newly returned from thesis work under Stirton at Berkeley.

62 - TEDFORD

Figure 16. R. A. Stirton (left) and R.H. Tedford at Leaf Locality excavation, Lake Ngapakaldi, South
Australia in 1962. Fossil leaf-bearing shales have been stripped off the top of the ancient stream channel
by the backhoe, and the collectors are working on the sands, conglomerates and clays that fill the deeper
channel where fossil vertebrates occur. (Courtesy of M. O. Woodbume),

THE 70'S CAREERS FOR AMERICAN VERTEBRATE
PALAEONTOLOGISTS IN AUSTRALIA

The early part of the decade proved to be a favourable time for American vertebrate
palacontologists particularly interested in research in their field in Australia to actually find

AMERICAN CONTRIBUTION TO VERTEBRATE PALAEONTOLOGY - 63

Figure 17. M. O. Woodbume picking out small bones and teeth from Leaf Locality pebbly sands that
have been washed through screen wire in ground-water pools dug at Lake Ngapakaldi, South Australia.

academic or museum positions there. The discoverics of the 1950's and 1960's had
demonstrated that an expanding range of evidence was available, including chapters in the
history of vertebrates heretofore unwritten. At the same time the field was also attracting
Australian students, so that by 1971 when the Geology Section of ANZAAS held its second
symposium on vertebrate palacontology there were only three speakers from the United States
(Cosgriff, Romer and Tedford) and twelve Australians reporting diverse researches from
Devonian fishes to Pleistocene marsupials. Most of the Australian speakers held academic or
museum positions. There were few students sufficiently advanced to step into the jobs that
were being offered by administrators aware of the intellectual promise of this developing field.
An exception was Michael Archer, who returned to his native land (he was born in Sydney) as

64 - TEDFORD

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Figures 18-21. Field notes and drawings
Australia. (Courtesy of the Museum of Paleontology, University of

Locality, Lake Ngapakaldi, northem South
California, Berkeley).

AMERICAN CONTRIBUTION TO VERTEBRATE PALAEONTOLOGY - 65

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66 - TEDFORD

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AMERICAN CONTRIBUTION TO VERTEBRATE PALAEONTOLOGY - 67

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68 - TEDFORD

Figure 22. M.O. Woodbume excavating at Paine Quarry, Alcoota Station, Northem Territory, in the initial
work in 1962. Bones can be seen drying after shellacing in the left foreground and in the excavation to the

tight of the whiskbroom.

AMERICAN CONTRIBUTION TO VERTEBRATE PALAEONTOLOGY - 69

a Fulbright scholar in 1967, obtained his degree at the University of Western Australia in 1972
and was appointed Curator of Mammals of the Queensland Museum the same year. He
remained in Queensland until 1978, when he accepted an appointment in the School of
Zoology of the University of New South Wales where he remains to this date. Archer's move
to an academic position at the University of New South Wales, and his exploitation of new and
very rich Miocene sites at Riversleigh in northwestern Queensland has resulted in a flood of
interest in vertebrate palaeontology and in turn attracted a number of graduate students, At
present count (Archer, pers. comm. 1988) fourteen students are enrolled in higher degree
programs under Archer's guidance. In addition to this academic interest, Archer's program has
promoted the involvement of interested nonprofessionals, many of whom are making
important contributions to the field and laboratory work at Riversleigh.

During the early 1970's another American student to study vertebrate palaeontology in
Australia was Larry Marshall, who completed an M.Sc. at Monash in 1974 under Jim Warren.
Marshall's work (1973) was concerned with the Plio-Pleistocene mammal faunas from sites on
the Murray River region in the tri-state area that would have been inundated by construction of
the Chowilla Dam. His work revealed an important faunal succession recently calibrated
palaeomagnetically. Marshall returned to the United States in 1974 to continue studies of New
World marsupials at the University of California. His subsequent research has continued to
emphasize this field.

Thomas Rich and his wife, Patricia Rich, were introduced to Australia as members of
Tedford's 1971 field party that worked Miocene sites in the Lake Frome area. The couple
immigrated to Australia in 1973, first with Fulbright support for Pat, and later both took up
positions at the Museum of Victoria (Tom) and the Earth Sciences and Zoology Departments
of Monash University (Pat) where they remain to date. T hey have spent much time in
wideranging reconnaissance for fossil vertebrate sites throughout Australia - most recently, and
successfully, in the Early Cretaceous deposits of coastal Victoria where a remarkable high-
latitude fauna of dinosaurs and other vertebrates is turning up in the joint work of the Museum
of Victoria and Monash University (Rich et al. 1988, Rich & Rich 1989). Pat Rich's Ph.D.
thesis (Columbia University) reviewed all available fossil bird remains from Australia,
emphasizing the dromornithid birds, especially the newly available Tertiary forms. She
continues her paleornithological studies at Monash, and she and Tom have guided M.Sc. &
Ph.D, students in that field, including both Australians (e.g. Tim Flannery, John Long,
Charles Meredith), as well as Americans (e.g. Robert Baird), amongst others.

In the late 70's two other American academics came to Australia. Ralph Molnar initially
took a post in the Department of Anatomy, University of New South Wales but in 1979
became Curator of Mammals and Fossil Vertebrates at the Queensland Museum. His reviews
of Australian lower tetrapods and studies of Crocodilia, theropods and Australia's first
pterosaurs are important contributions to the Mesozoic fauna. Peter Murray, University of
Chicago physical anthropologist, joined the Tasmanian Museum staff, and very quickly
became involved in determining the late Quaternary mammal succession in the island State.
Important new data were acquired during exploration of caves in northern and eastern Tasmania
with collegues at the University of Tasmania. Murray described the vertebrate remains and
reviewed the record of the giant echidna Zaglossus in Australia. In 1981 he moved to the
Northern Territory Museum, first to Darwin and later to Alice Springs, to the post of Curator
of Fossil Vertebrates and has pursued studies on Tertiary mammals there.

The 70's saw a renewal of field work in Australia by Stirton's students Woodburne and
Tedford, who joined with other American and Australian colleagues to continue exploitation of
the Lake Eyre Basin sites and to explore for new sites in Queensland and South Australia. The
National Science Foundation provided most of the support for these expeditions, but important
contributions were made by the cooperating institutions, namely the South Australian and
Queensland museums.

70 - TEDFORD

Figure 23. Excavations at Lake Callabonna, South Australia in 1983 by the Museum of Victoria, Monash
University and a contingent of the Australian Army led by John Wild and Tom Rich. A, excavation of one of
five Diprotodon skeletons found during the 10 day expedition; B, preserving the bones before plaster-
jacketing them for removal; C-D, plaster jacketing the bones for protection during their transport back to
Melboume; E, removal of the plaster-jacketed specimen to a waiting Army vehicle; F, John Wild, commander
of the Australian Army unit in charge of the Callabonna expedition, holding a Scotch Whiskey jug left by the
Stirling and Zietz expeditions in the late 19th century, Discovery of this relic allowed location of the
campsite that Stirling and Zietz and Hurst had used during their early excavations (see Rich & Archbold, Chap.
1, this volume). (Courtesy of the Australian Army).

AMERICAN CONTRIBUTION TO VERTEBRATE PALAEONTOLOGY - 71

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Australia in 1971 indicating the first discovery of pre-Pleistocene platypus (Obdurodon, the tooth illustrated)
and porpoise (a petrosal) remains in central Australia. (Courtesy of P. V. Rich).

72 - TEDFORD

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AMERICAN CONTRIBUTION TO VERTEBRATE PALAEONTOLOGY - 73

In 1970, Tedford and Columbia University graduate student Bob Emry, joined with the
Smithsonian's Clayton Ray and Frank Pearce and South Australian Museum's Neville Pledge
and Paul Lawson to reinvestigate the famous Pleistocene sites at Lake Callabonna (Fig. 23) in
South Australia. This was the first large-scaled work there since the original expedition
sponsored by the South Australian Museum in 1893 and Stirton's visit in the 50's. The fauna
was enlarged by finding taxa other than Diprotodon. Diprotodon trackways (Tedford 1973)
were discovered, and the local geology was outlined for the first time and fit into the Cainozoic
sequence developed for the nearby Lake Frome area. Near the close of this field work the South
Australian Museum hosted a field conference led by Tedford at Lake Callabonna and in the
eastern Lake Eyre Basin. This trip lasted 7 days and included David Ride and Harry Butler
(Western Australian Museum); Alan Bartholomai (Queensland Museum); Jim Warren (Monash
University), Grant Inglis, Neville Pledge and Paul Lawson (South Australian Museum); Mike
Plane (Bureau of Mineral Resources); and Mines Department South Australia and Bureau of
Mineral Rresources geologists,

Roger Callen, indefatigable geologist of the Mines Department, South Australia, discovered
Tertiary bone fragments southeast of Lake Frome in 1970, and these became the harbinger of
the Miocene fauna discovered by Tedford's party in 1971 (Figs 24, 25, 26). That year the field
party included Tom and Pat Rich then graduate students at Columbia and Rod Wells, a graduate
student at the University of Adelaide. This party made the first collection of vertebrates from
that area, including many taxa new to the Australian Miocene. Moving into Queensland, Alan
Bartholomai of the Queensland Museum joined the party. The group went on to explore
(without success) deposits east of the Dividing Range in Queensland, then moved to the
Winton area in central Queensland where a few fragmentary dinosaur bones and a spectacular
trackway site (Lark Quarry) were located in the Winton Formation with the assistance of a
local grazier, Peter Knowles. No fossil vertebrate remains were found in the overlying Tertiary
deposits.At the close of this trip, after Bartholomai returned to Brisbane, the party returned to
South Australia and briefly joined Woodburne's group working at Lake Palankarinna east of
Lake Eyre.

In 1971 Woodburne had combined with William A. Clemens of the University of
California to set up the first large-scale screen washing of fossiliferous matrix from Miocene
rocks in the Tirari Desert east of Lake Eyre. They were joined by Colin Campbell, Australian
graduate student at California, Mike Archer, just finishing at the University of Western
Australia, and Neville Pledge (Fig. 26). The washing technique proved to be highly successful
with many new taxa coming to light. This was an inspiration for Tedford's work in 1973
when the washing technique was applied to the Namba Formation east of Lake Frome.
Tedford's 1973 party included Alan Bartholomai, Mike Archer (newly appointed at the
Queensland Museum), Rod Wells (then at Monash University), Neville Pledge and Mike Plane
and Dick Brown of the Bureau of Mineral Resources, Canberra. Most of the field season was
spent exploiting the Miocene Namba Formation southeast of Lake Frome where a large and
very successful washing program was set up. A few weeks were also devoted to completing
the reconnaissance of the Palaeogene Eyre Formation outcrops in northeastern South Australia
and adjacent western Queensland. This work failed to find any animal remains in these well-
exposed deposits.

Woodburne spent his sabbatical year in Australia in 1972 during which he again joined
with Australian colleagues Mike Plane, Mike Archer, Paul Lawson and Winston Head of the
South Australian Museum in the field at Lake Palankarinna (Fig. 27) with further success in
washing the Etadunna Formation there.

In 1976 Lundelius and Turnbull (Figs 28, 29) returned to Western Australia to continue
work on Quaternary faunas and to explore the Pilbara district for Tertiary mammal occurrences.
They all visited Hamilton in Victoria. Funding for this work again came from the Fulbright
Program (for Lundelius) along with University of Texas, National Geographic Society and

74 - TEDFORD

Figure 26 A, expeditions into the Lake Eyre Basin in the early 1970's. Above is the field party to Lake
Palankarinna, South Australia in 1971. Left to right: Colin Campbell, Mike Woodbume, Tom Rich (a
inember of Tedford’s party), Rod Wells (a member of Tedford's party), Mike Archer, Dick Tedford, and Bill
Clemens. Picture is taken at Stirton’s old camp site. B, R.H. Tedford's field party that discovered the new
fossil field in the Lake Frome area in 1971: from the left, Tom Rich, Dick Tedford, and Rod Wells

(Courtesy of P. V. Rich).

AMERICAN CONTRIBUTION TO VERTEBRATE PALAEONTOLOGY - 75

Field Museum contributions. No Tertiary vertebrates were found in the Oakover beds, but
studies of Quatemary mammals were advanced.

Figure 27. A, R. A. Stirton (right) and Paul Lawson on early reconnnissance trip in the Lake Eyre Basin
during the 1950's. B, Excavation in the Lake Frome area or South Australia (Lake Tarkarooloo) in 1976 by a
party from the Museum of Victoria; sediments are of Miocene age and related to those at Lake Palankarinna;
P. Morrison (foreground) and M. Vickers. C, Stirton's favourite camping spot at Lake Palankarinna, behind
the Cannuwalkaninna sanddune. Several expeditions mounted by the BMR, the Museum of Victoria and
Monash University, the Queensland and South Australian Museums and the University of New South Wales
retumed to exploit Stirton's old sites and locate new ones in the 1970's and first half of the 1980's. (B and
C courtesy of P. V. Rich).

76 - TEDFORD

This decade saw an enormous increase in field work undertaken by Australian workers using
private, university, and federal funds (Australian Research Grants Scheme, Australian Research
Council) and funds from cooperative projects with other institutions. Notable were Archer's
privately supported (Dr. R. E. Lemley (Fig. 30), Rapid City, South Dakota) work in
Queensland which included exploiting the discovery of a Pliocene site near Charters Towers
(Bluff Downs), and his realization of the enormous potential of the Riversleigh Miocene site in
the northwest of the state. Joint projects between Queensland and Victoria museums to the
Frome and Lake Eyre Miocene sites in South Australia and the Miocene at Alcoota (Fig. 22)
in the Northern Territory provided comparative material and new taxa from the sites discovered
by the American parties,

Palaeozoic fishes were collected from eastern Australia by the Australian Museum (Alex
Ritchie, who was also carrying on field work in Antarctica), Monash University (Jim Warren
and John Long), Australian National University (Ken Campbell), Bureau of Mineral Resources
(Gavin Young) and Queensland Museum (Susan Turner and Anne Kemp). The spectacular
Devonian Gogo site in the Kimberley district of Western Australia was discovered and worked
initially in the 60's by joint British Museum (Natural History) and Western Australian
Museum teams and later by parties from the Australian National University, the Bureau of
Mineral Resources and the Western Australian Museum.

An important new site for Triassic tetrapods was found in southern Queensland and worked
by Anne Warren and Latrobe University parties and Tony Thulborn of the University of
Queensland. Cretaceous vertebrates were collected from the Eromanga Basin in Queensland by
joint British Museum (Natural History) and Queensland Museum group. Systematic
prospecting of the Early Cretaceous in Queensland, New South Wales and especially Victoria
by Ralph Molnar (University of New South Wales, Queensland Museum) and Pat and Tom
Rich (Monash University, Museum of Victoria) has improved the record of vertebrates from
these rocks.

Quaternary vertebrate records were also improved, spectacularly, with the discovery of rich
deposits in part of the Victoria Cave in the Naracoorte district of South Australia by Rod Wells
(Flinders University) and the South Australian Caving Group. Neville Pledge added a new
Miocene level to the sequence at Lake Palankarinna during a joint field trip of Australian
vertebrate palaeontologists and Ernest Lundelius following the International Geological
Congress in Sydney in August of 1976 (Fig. 30).

THE 80'S, AUSTRALIAN VERTEBRATE PALAEONTOLOGY
COMES OF AGE

At the beginning of the present decade vertebrate palaeontology was being addressed in most
universities and state museums throughout Australia, and in some federal departments as well.
The Queensland Museum has Alan Bartholomai, Director, Ralph Molnar and Mary Wade,
curators, and Susan Turner and Anne Kemp Research Fellows and Tony Thulborn in the
University of Queensland. In New South Wales, Mike Archer is at the University of New
South Wales, his former graduate students Tim Flannery is at the Australian Museum
(Mammalogy) and Suzanne Hand is a Research Fellow at the University. Alex Ritchie is
Curator of Fossils at the Australian Museum, These individuals also offer or cooperate in
graduate programs in the Sydney area universities. Jeannette Hope (Fig. 28),
palacomammalogist, employed by the New South Wales Parks Department, is also based in
Sydney. Canberra has palaeoichthyologist Ken Campbell at ANU, Mike Plane (now retired)
and Gavin Young at the BMR, and vertebrate palaeontologists David Horton and John Gorter
are associated with the Institute of Aboriginal Studies. In the Melbourne area Tom Rich is
Curator of Fossils at the Museum of Victoria, Pat Rich (Earth Sciences and Botany/Zoology)

AMERICAN CONTRIBUTION TO VERTEBRATE PALAEONTOLOGY - 77

(Botany/Zoology) are at Monash University and Anne Warren is at Latrobe University. Former
Curator of Fossils at the Museum of Victoria, is fossil wombat specialist Eric Wilkinson,
geologist for the Department of Mines, Victoria in Ballarat. In Tasmania, Museum Director
Don Gregg and Curator of Fossils Noel Kemp are vertebrate palaeontologists, and
palaeoichthyologist John Long is Research Fellow in Geology at the University of Tasmania.

Figure 28. Vertebrate palaeontologists who attended the International Geological Congress held in Sydney
in 1976. This group attended a field conference that visited sites in westem New South Wales, including the
Wellington Caves: left to right, Dr. Lehman, M. Vickers, E. Ritchie, T. Rich, ?, B. Ritchie, P. Rich, A,
Ritchie, S. Ritchie, L. Dawson, M. Archer, J. Hope, ?, E. Lundelius, J. Lundelius, N. Pledge. (Courtesy of P.

V. Rich).

Honours and graduate studies in vertebrate palaeontology are conducted in South Australia
by Rod Wells at Flinders University. Neville Pledge is Curator of Fossils at the South
Australian Museum. Both coordinate a number of amateurs in the state and work with local
caving groups to the great benefit of the science. The Northern Territory Museum has
employed Peter Murray as Curator of Vertebrates, the first vertebrate palacontologist in that
important region. Only Western Australia at present lacks a permanent academic or museum
position in vertebrate palacontology. Alex Baynes, Research Associate at the Western
Australian Museum, is the only vertebrate paleontologist in the state, but Ken McNamara,
Curator of Fossils of the Western Australian Museum has more than a passing interest in the

subject.

78 - TEDFORD

Figure 29. Localities producing vertebrate fossils in Australia worked during the 1970's and 1980's. A-C,
Hamilton, a Pliocene site dated radiometrically by the basalt that overlies the fossiliferous sediments. C,
Tom Rich in the foreground and Tim Flannery. The Museum of Victoria and Monash University carried out
major excavations and sieving operations on this site that had originally been discovered by Edmund Gill
and later worked by Emest Lundelius and William Tumbull. D-F, Lord Howe Island, South Pacific,
Pleistocene: D, partial specimen of a procellarid bird still in the matrix; E, Alex Ritchie (foreground) and
Steve Barghom excavating Meiolania , the Lord Howe Island Horned Turtle; F, Gene Gaffney puttin
plaster jacket on a Horned Turtle specimen. (Courtesy of P. V. Rich).

AMERICAN CONTRIBUTION TO VERTEBRATE PALAEONTOLOGY - 79

This roster lists only those actively engaged in the field as professionals. There are many more
students and a growing number of amateurs attracted to the science. As mentioned, Wells and
Pledge work closely with vigorous amateurs and Archer has incorporated a corps of amateur
enthusiasts into a "Riversleigh Society” to help transform the mountain of limestone from
these deposits into research specimens. The field by its very nature continues to capture public
attention, and this has been important in developing private as well as public funding necessary
for sustenance of field and laboratory work and the growing cost of publication of scientific
papers.

Figure 30. A, Australian Army support on expedition into the Lake Eyre Subbasin with T. H, Rich in the
mid-1970's; B, Ray Lemley, who provided finances for several of M. Archer's expeditions into central and
northem Australia; C, Michael Archer; D, mid-Tertiary northern Australian limestone country at Bullock
Creek, Northern Territory, a site of similar nature to the rich Riversleigh deposits of northwestern
Queensland. (Courtesy of P. V. Rich).

80 - TEDFORD

American contributions to field studies have continued into the 80's. Both Tedford and
Woodburne pursued field projects begun in the 50's in attempts to tie-up loose ends of the Lake
Eyre Basin faunal sequence. Woodburne spent 14 months in Australia in 1984-5 during which
he conducted two field seasons of work at Miocene sites in the eastern Lake Eyre Basin with
his students Judd Case and Mark Springer. Bruce MacFadden of the University of Florida,
working with the Woodburne party, initiated magnetostratigraphic studies on the Etadunna
Formation. This detailed biostratigraphic work has resolved the "Ngapakaldi Fauna” of
Surton, Tedford and Miller (1961) into a six-fold faunal succession that spans sufficient time
so that evolutionary change is visible in most lineages that extend through the succession.
Bruce MacFadden's student, Mick Whitelaw, an honours graduate of Monash University, is
continuing to apply magnetostratigraphic techniques to the Plio-Pleistocene successions.

In 1980 and 1983 Tedford joined with Rod Wells, Paul Lawson and Well's student Dominic
Williams plus a number of other willing workers from the Flinders University staff and
Students to work in the Lake Eyre Basin. Steven Barghoorn of New York a member of the
party in 1983 conducted a magnetostratigraphic study of the Tirari Formation. This work was
jointly funded by ARGS and the National Geographic and was designed to clarify the Pliocene
and Pleistocene stratigraphy and vertebrate palacontology of the deposits east of Lake Eyre.
The results were reported at the de Vis Symposium and will be published in those proceedings,
but a preliminary statement has been published (Tedford, Williams & Wells 1986), The
"Katipiri Sands" of Stirton, Tedford & Miller (1961) was resolved into two formational units
representing penultimate and ultimate glacial ages, and new local faunas were discovered in the
Late Pliocene Tirari Formation and the overlying Pleistocene units.

In 1980 Eugene Gaffney (Fig. 29) of the American Museum of Natural History revitalized
the search for the homed tortoise (Meiolania) on Lord Howe Island while holding a Visiting
Curatorship at the Australian Museum. He was assisted by then Columbia University graduate
students Steve Barghoorn and Paul Sereno in cooperation with Alex Ritchie and Bob Jones of
the Australian Museum. This work met with gratifying success, and Gaffney returned to Lord
Howe Island in 1982 supported by funds from the American Museum and NSF. Barghoorn and
Columbia graduate student Ann Burke completed the party, A further trip in 1987 supported
by the American Museum employed local help on Lord Howe Island, This work enabled
Gaffney to completely describe the osteology and relationships of the Lord Howe Island species
of Meiolania, the most completely known representative of the genus. In the course of these
studies Gaffney also reviewed the fossil record of Australian turtles and made a number of
contributions to the morphology of Australian taxa present in museum collections there and in
the United States,

High levels of activity sustained by this growing population of vertebrate palaeontologists
has increased the rate of discovery of important links in the chain of vertebrate history in
Australia over the past decade. Highlights of this activity would certainly include the
description of well preserved moulds of the Ordovician vertebrates, Australia's oldest (Ritchie
& Gilbert Tomlinson 1977); the continuing yield of new and spectacularly preserved Devonian
fishes from Gogo, Western Australia (Long 1987); the Triassic fauna at Rewan, Queensland
including the first Australian therapsid (Thulborn 1983); the increasingly diverse and peculiar
Early Cretaceous dinosaurs (Rich et al. 1988, Rich & Rich 1989); the oldest Australian
mammal, a platypus (Archer et al. 1985); one of the oldest Australian birds (Molnar 1986) the
diverse array of vertebrates from the Miocene of Riversleigh (Archer et al. 1989), a rich
undescribed Late Miocene or earliest Pliocene fauna from a fissure fill in South Australia
(Pledge pers. comm.) and the spectacular Victoria Cave deposit, South Australia now shown to
be of penultimate glacial age (Wells et al. 1984). Much of this research was summarized and
new work reported at the de Vis Symposium held in at the Queensland Museum, Brisbane, in
1987. Thirty-five Australian and six foreign contributors discussed aspects of the entire
vertebrate record. Notable at this third major meeting of vertebrate palaeontologists (after a gap

AMERICAN CONTRIBUTION TO VERTEBRATE PALAEONTOLOGY - 81

of 17 years) was the number of students reporting thesis research and the professional reports of
two amateurs. Clearly the field has reached maturity in Australia.

CONCLUSIONS

The historical review presented above indicates that the American contribution to vertebrate
palaeontology in Australia has been most direct in initiating and sustaining for over thirty
years a level of field work that has directly revealed aspects of the history of the fossil
vertebrates of that continent. The American effort has consistently been directed toward
exploration for new sites and their exploitation. From the beginning this work has been
conducted with Australian colleagues, few at first, but later with growing numbers of persons
as the field generated interest among Australians. By the 1970's most of the field work was
being conducted from within Australia funded by local institutions, the National Geographic
Society and the ARGS. Some investigators also found significant private sources or obtained
the assistance of the Australian armed forces for larger-scale logistical support. Much of this
work exploited localities previously discovered in order to increase knowledge and to gain
Australian reference materials from the more important sites. At this point the American effort
focused more on completion of long-term studies in areas initially discovered or worked by
them, and such work continues at diminishing scale to the present day.

Certainly the greatest contribution to come from the American presence is the excitement
the field has generated in the minds of Australians as is so amply testified by the number of
Australians fully involved professionally in this field and the growing number of students
entering it. This situation took only a couple of decades to develop into the mature state the
field now enjoys in Australia.

Much has been done, but much more remains to fully realize Australia's potential in
documenting its vertebrate history. Although there are several Devonian fish sites scattered
across the continent, the other Palaeozoic periods are poorly represented or unrepresented.
Nevertheless, the Palaeozoic is present in thick geoclinal rock sequences of untapped potential
in contrast to the cratonic Mesozoic and Cainozoic where shallow basins and long exposure to
weathering and erosion limit the possibility for a record considerably. Despite this, the
Triassic and Early Cretaceous have yielded much, although the Jurassic is an almost complete
blank. The thicker rift-filling Cretaceous of Victoria has special significance because of its
polar position, and indeed most of Australia was at high latitude during the Mesozoic. The
Palaeogene is nearly a total blank except for a few turtles and penguins; the Tertiary record is
limited at present to the latest Palaeogene (except perhaps for one early Tertiary site recently
discovered in Queensland) and Neogene when carbonate sedimentary environments favorable to
the preservation of bone appear in the continental record. Filling this "ghastly blank" in
vertebrate history will require a new approach to prospecting, perhaps focusing on Palaeogene
continental carbonates either fissure fills or clastic carbonate fluviatile or lacustrine deposits
(the Miocene Carl Creek Limestone is a model). Large blanks still remain in the Neogene and
Quaternary but favourable geological situations are still awaiting exploration. Continued work
at the levels applied in the present decade promise a revolution in our knowledge of the
vertebrate history of Australia by the next century.

ACKNOWLEDGEMENTS

These historical notes were put together from personal records and the literature. Through
the years many people have helped to flesh-out this recent phase of the history of vertebrate
palaeontology in Australia. I have called on Ernest Lundelius Jr., Mike Woodbume and Ralph

82 - TEDFORD

Molnar to help check aspects of the story, and to them and all my colleagues in the ever
exciting task of determining Australia's vertebrate history, I express my deepest gratitude.

REFERENCES

ANDERSON, C., 1937. Palaeontological notes. No. IV. Fossil mammals from New Guinea. Rec. Aust.
Mus. 20(2): 73-76.

ARCHER, M. & CLAYTON, G., 1984. Vertebrate Zoogeography & Evolution in Australasia. Hesperian
Press, Carlislet.

ARCHER, M, FLANNERY, T.F., RITCHIE, A. & MOLNAR, R.E., 1985. First Mesozoic mammal from

Australia - an early Cretaceous monotreme. Nature 318: 363-366.

ARCHER, M., GODTHELP, H, HAND, S. & MEGIRIAN, D., 1989 Fossil mammals of Riversleigh,
northwestern Queensland: preliminary overview of biostratigraphy, correlation and environmental
change. Aust. Zool .25(2): 29-65.

COSGRIFF, J.W., 1965. A new genus of Temnospondylii from the Triassic of Wester Australia.J. Proc. R.
Soc. West. Aust. 48: 65-90.

COSGRIFF, J.W., 1969. Blinasaurus, a brachyopid genus from Westem Australia and New SouthWales. J.
Proc. R. Soc. West. Aust. 52: 65-88.

COSGRIFF, J.W. & GARBUTT, N.K., 1972. Erythrobatrachus noonkanbahensis, a trematosaurid species

from the Blina Shale. J. Proc. R. Soc. West. Aust. 55: 5-18.
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Mem. natn. Mus. Vict. 21: 135-203.

LLOYD, A., 1967. An outline of the Tertiary geology of northem Australia. Bur. Min. Res. Bull. 80.

LONG, J.A., 1987. Late Devonian fishes from the Gogo Formation, Westem Australia - new discoveries.
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LUNDELIUS, E.L., IR. & TURNBULL, W.D., 1973. The mammalian fauna of Madura Cave, Western
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LUNDELIUS, E.L., JR. & TURNBULL, W.D., 1975. The mammalian fauna of Madura Cave, Western
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LUNDELIUS, E.L., JR. & TURNBULL, W.D., 1978. The mammalian fauna of Madura Cave, Western
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LUNDELIUS, E.L., JR. & TURNBULL, W.D., 1981. The mammalian fauna of Madura Cave, Western
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LUNDELIUS, E.L., IR. & TURNBULL, W.D., 1982. The mammalian fauna of Madura Cave, Western
Australia, Part V. Diprotodonta (Part). Fieldiana Geol. n.s. 11; 1-32.

LUNDELIUS, E.L., JR. & TURNBULL, W.D., 1984. The mammalian fauna of Madura Cave, Western

Australia, Part VI: Macropodidae: Potoroinae. Fieldiana Geol. n.s. 14: 1-63.

LUNDELIUS, E.L., JR. & TURNBULL, W.D., 1989. The mammalian fauna of Madura Cave, Western
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of the fauna. Fieldiana Geol. n.s. 17: 1-71.

MARSHALL, L., 1973. Fossil vertebrate faunas from the Lake Victoria region, southwest New South Wales,
Australia. Mem. natn. Mus. Vict. 34: 151-171.

MOLNAR, R., 1986. An enantiomithine bird from the Lower Cretaceous of Queensland, Australia. Nature
322(6081) : 736-738.

PLANE, M. D., 1967a. Stratigraphy and vertebrate fauna of the Otibanda Formation, New Guinea. Bur. Min.
Res. Bull. 86: 1-64.

PLANE, M.D., 1967b. Two new diprotodontids from the Pliocene Otibanda Formation, New Guinea. Bur.
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PLANE, M.D. & GATEHOUSE, C., 1968. A new vertebrate fauna from the Tertiary of northern Australia.
Aust. J. Sct.30(7): 272-273.

RICH, P.V., RICH, T.H., WAGSTAFF, B.E., MCEWEN MASON, J., DOUTHITT, C.G., GREGORY, R.T. &
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RICH, T.H. & RICH, P.V., 1989. Polar dinosaurs and biotas of the Early Cretaceous of southeastem
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RICH, P.V. & THOMPSON, E.M., 1982. The Fossil Vertebrate Record of Australasia. Monash University
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AMERICAN CONTRIBUTION TO VERTEBRATE PALAEONTOLOGY - 83

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STIRTON, R.A., 1957. Tertiary marsupials from Victoria, Australia. Mem. natn. Mus.Vict. 21: 121-134.

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STIRTON, R.A., TEDFORD, R.H. & WOODBURNE, M.O., 1967. A new Tertiary formation and fauna from
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STIRTON, R.A., TEDFORD, R.H. & WOODBURNE, M.O., 1968. Australian Tertiary deposits containing
terrestrial mammals. Univ. Calif. Publs. geol. Sci. 77: 1-30.

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sediments and fossil vertebrates. In The Lake Eyre Basin Cainozoic Sediments, Fossil Vertebrates and
Plants, Landforms, Silcretes and Climatic Implications, R. T. Wells & R. A. Callen, eds., Australasian
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THULBORN, R.A., 1983. A mammal-like reptile from Australia. Nature 303: 330-331.

TURNBULL, W.D. & LUNDELIUS, E.L., 1970. The Hamilton fauna: a late Pliocene mammalian fauna from
Grange Bum, Victoria, Australia. Fieldiana Geol. 19: 1-163.

WELLS, R.T., MORIARTY, K. & WILLIAMS, D.L.G., 1984. The fossil vertebrate deposits of Victoria Fossil
Cave, Naracoorte: An introduction to the geology and fauna. Aust. Zool. 21(4): 305-333.

WOODBURNE, M.O., 1967. The Alcoota fauna, central Australia: An integrated paleontological and
geological study. Bur. Min. Res. Bull. 87: 1-187.

84 - TEDFORD

Large leaf-eating stenurine kangaroos. In the background Sthenurus tindalei, in the foreground
the largest, and as yet undescribed, species from the Pleistocene deposits at Lake Callabonna in
South Australia.

CHAPTER 3

MUSINGS ON NEW GUINEA
FOSSIL VERTEBRATE
DISCOVERIES

Michael D. Plane!
Introduction .............cccccceeccecceeceseceucesccenceescens 86
The First Fossil Vertebrate Remains.................. 86
The European Discoveries.......c..cceccescceeeceesees 86
The Stirton Years........ccecscssssssssseseseressseees 87
Bureau of Mineral Resources Involvement.......... 90
Further Excavations in the 1960's ..................00. 90
Stirton's First Visit to New Guinea

and a Second Period of BMR

INVOIVEMENL...........ccecccesccessccceceuscceeececeece 91
Further Work in the Late 1960's to Present......... 95
WieS ta UTIANe 8 es ie Mate he ee eo a 97
Conclusions? c05: 1b .5h coset Rvcvedt coteeM ce 97
References... sis. cBikoteccss odeegts uedstvadselelealecd 97
PIAS codes ache cd edbtie cede: hele dutht te Ics be ds 98

1 "Allsun” via Gundaroo, New South Wales 2620, Australia.

86 - PLANE

INTRODUCTION

The discovery of fossil vertebrates in Papua New Guinea has progressed in a rather
haphazard, but not unexpected fashion. This discussion is a highly personal and anecdotal
account of that progression over the last fifty years. It is far from complete, as I am not au fait
with all of the work done by the myriads of archaeologists who have excavated human
habitation sites, and who have discovered fossil vertebrate remains in their digs (e.g. Bulmer &
Bulmer 1964, White et al. 1970, White 1972).

THE FIRST FOSSIL VERTEBRATE REMAINS

Natural scientists have long realized that the living faunas of Australia and New Guinea
have strong similarities and mutual genera on both sides of Torres Strait. It was correctly
assumed in the 1930's, when Europeans were just starting to explore the hinterland of this
great island that in due course some of Australia's extinct forms would also be found in New
Guinea. With the commencement of alluvial mining in areas of Tertiary and Pleistocene
sedimentary rocks that is just what happened.

We shall never really know who first saw fossil vertebrate remains, but given the
indigenous populations’ cultural preoccupation with natural objects, it would be surprising
indeed if someone had not discovered fossil bones well before Europeans came to New Guinea.
Unfortunately, there is no recorded history, so we shall have to start with what has been
written down.

THE EUROPEAN DISCOVERIES

The first fossil bones came to the attention of the scientific world through the offices of
two stalwarts of New Guinea geology, G. A. V. Stanley and N. H. Fisher, neither of whom
was a palaeontologist. In the mid-1930's, Stanley, then working as a geologist for Oil Search
Ltd., visited Fisher, the Government Geologist for the Territory of New Guinea, at his Wau
office, where he was shown two "mandibular rami" that had been brought in from the gold
workings in the Watut Valley. Fisher forwarded these fossils, together with additional
material, which he obtained after visiting the Watut workings, to the Australian Museum in
Sydney. On a subsequent visit to the Museum, whilst on leave, Fisher provided Charles
Anderson, the Director, with observations on the geology and a geological sketch map.
Anderson, who was a mineralogist and crystallographer, had a keen interest in palaeontology
and, fortunately, realised the importance of these fossils. He published an account of the
geology, based on Fisher's observations and map, and described Nototherium watutense, the
first fossil mammal published about from Papua New Guinea (Anderson 1937).

The alluvial gold workings in the Watut and Bulolo valleys (Fig. 1) continued to produce
fossils, both before and after World War II. Some of the material went to the Australian
Museum, one specimen ended up at the British Museum of Natural History in London, and
much was kept as local curiosities. Immediately pre-war, L. C. Noakes, Assistant
Government Geologist, wrote accounts of the geology of the Upper Watut and Bitoi/Black Cat
areas. Both covered areas within which fossil vertebrates had been found, but unfortunately
both remained unpublished (Noakes 1938a, 1938b). Towards the end of the war, Fisher's
outline of the geology of the Morobe Gold Field appeared (Fisher 1944).

NEW GUINEA FOSSIL VERTEBRATE DISCOVERIES - 87

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Figure 1. Vertebrate fossil bearing sites in New Guinea, excluding archaeological materials.
THE STIRTON YEARS

In 1953, R. A. Stirton, an American from the Berkeley campus of the University of
California, came to Australia on a Fulbright Fellowship (see also Tedford, this volume, Chap.
2). Stirton was preceded by a reputation for being a tenacious and persistant seeker of fossils
in inhospitable and out of the way places. Amongst the many things that Stirton
accomplished during his first year in Australia, was a thorough survey of the Tertiary fossil
material in Australian museums. He was intrigued by the material from New Guinea and made
a note to look into it further when time and opportunity allowed. His notes of 14 October
1953 (Fig. 2) state:

"I squeezed in time to sketch Figs. 115-116, the macropodid from New Guinea
which looks more like the Palankarinna form than any-thing I have seen yet.
This New Guinea material (Figs. 111, 112, 113, 114, 115, 116) is of unusual
interest. It has the feel of early Pliocene or late Miocene. Later I hope to spot
these localities on a map - much of it comes from the Watut area. Dr. Norman
Fisher asked me about N. watutense when I was in Canberra. He probably can
give us more dope on the geology of the area (also see C. Anderson, 1937). I
hope it can be arranged to get some field work done in that area."

88 - PLANE

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(Courtesy of the Museum of Paleontology, University of California, Berkeley).

fossils of New Guinea.

NEW GUINEA FOSSIL VERTEBRATE DISCOVERIES - 89

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90 - PLANE

By 1955 Stirton had, indeed, organised funds which allowed G. D. Woodard, an Australian
from Adelaide, who was Stirt's student at the University of California, to go to New Guinea
and search the Bulolo-Watut area for fossil mammals. Woodard made his base at Bulolo, the
mining headquarters of the Bulolo Gold Dredging Co. He was befriended and assisted by Keith
Collins-Rubie, a mining foreman who had an intimate knowledge of the geology at the
"Sunshine" and "Widubash" operations, and who displayed a keen interest and understanding of
the fossils. A sizeable collection was made, and the bulk of the material was crated and
transported by road to the port of Lae for shipping to San Francisco. It was never seen again.
Fortunately, Woodard had decided to carry the best specimens back to the United States by
hand, These were incorporated into the Museum of Paleontology collections and eventually
worked on in 1964-1965.

BUREAU OF MINERAL RESOURCES INVOLVEMENT

It seems clear that neither Collins-Rubie nor Woodard made contact with the geologist, on
secondment from the Australian Bureau of Mineral Resources to the Territory of New Guinea
Administration, who had his office just 17 kilometers by all weather road to the south at Wau.
Had they done so, much of mutual benefit might have resulted.

J.G. Best was the Resident Geologist at that time, and he, guided by Horrace Clissold of
New Guinea Goldfields Ltd., the leasee of the Korange Open Cut Mine near Wau, had
photographed and partly removed the skull of a juvenile diprotodontid, from the Otibanda
Formation exposed in Koranga Creek. I say "partly removed," because from a photograph,
one can discern that the specimen was at one time in excellent condition. Very little of it was
eventually to find its way into the Commonwealth Palaeontological Collection, much being
lost during excavation.

Some further spasmodic collecting was carried out in the Watut area during the years of
1956 to 1962, but none of the specimens reached scientific institutions.

FURTHER EXCAVATIONS IN THE 1960'S

During 1960 in the highlands, near Chuave, archaeologist Susan Bulmer excavated the left
half of a mandible of Thylacinus. This exciting find demonstrated once again the close ties
between Australia, Tasmania and New Guinea and the tremendous geographic range of
Thylacinus (Van Deusen 1963).

By 1962, Stirton's curiosity, egged on by one or two tantalising Woodard specimens of the
genus Protemnodon, which he had under study at the time, roused him to organise, in
conjunction with Alden Miller, an ornithologist and palaeornithologist, and Bill Lidicker, a
mammalogist, both from the Museum of Vertebrate Zoology at Berkeley, an expedition to
New Guinea, By this time Stirton had been in touch with the Bureau of Mineral Resources
(BMR) in Canberra. Keith Rochow, a BMR geologist working with Alan Newsome,
mammalogist from the CSIRO (Commonwealth Scientific and Industrial Research
Organization), had visited and realised the worth and potential of the fossil deposits at Alcoota,
northeast of Alice Springs. Here N. H. (Doc) Fisher again enters the story, He was by now
Chief Geologist with the BMR (later to become its Director). He was impressed, and who
could fail to be, with Stirton's enthusiasm for fossil mammals and their utility in the
correlation of non-marine rocks. The BMR at that time had responsibility for Northern
Territory geology, and Fisher agreed to co-sponsor some work at Alcoota. Stirton, of course,
knew of Fisher's early association with the New Guinea fossils and mentioned his impending

NEW GUINEA FOSSIL VERTEBRATE DISCOVERIES - 91

visit to the Watut-Bulolo area. Fisher then wrote to the Resident Geologist at Wau giving
instructions that every possible assistance was to be given to Stirton and his party.

STIRTON'S FIRST VISIT TO NEW GUINEA AND A SECOND
PERIOD OF BMR INVOLVEMENT

I was the recipient of that letter, and frankly, I was nonplussed. I had no formal training in
palaeontology, it being taught at the University of Cape Town when I was an undergraduate as
a throw away subject. Fourteen lectures out of Wood's Palaeontology, a book so dry that it
was in danger of spontaneously combusting and one which reduced a potentially fascinating
subject to abject boredom, was my sum total of experience with the subject. The charming
lady who taught the subject was a brilliant petrologist, but her interest in fossils must have
rated no higher than a one on a scale of ten. What on earth was I supposed to do with this
unexpected visitor? Well, it didn't take me long to realise that here was someone who exuded
enthusiasm and had a real love for his subject. I couldn't resist and very soon became involved
in all aspects of the work.

Stirton, again helped by Keith Collins-Rubie, concentrated on the sites at "Sunshine",
where Woodard had had success in finding fossils. He, too, unearthed bones, but the sites
proved to be rather thinly populated. Three incidents stay in my mind from that time, and
they demonstrate the cameraderie, zest and sense of fun which so many vertebrate
palaeontologists bring to their work.

In order to get to the fossil site at "Sunshine", one had to ford a river, which ran rather
swiftly at about waist high on a person of average height. The bed of the river was rather
coarse boulders. To see Stirton, who was himself rather short, negotiating the muddy waters,
stripped and with his clothes, collecting gear and hat perched on his head gave us many a
laugh. The bobbing cork managed it, day in, day out, and our predictions about him losing
the lot were certainly unfounded.

My field assistant, Timbu Tasong and Stirton achieved instant rapport, Tasong clearly
feeling a sense of responsibility towards this visitor to his country, tried to mother him while
holding him in great awe for his prowess with the shotgun. On the day that Stirton put up
two quail and calmly shot both, one with each barrel, Tasong's admiration turned to adulation.

On another occasion, Stirton had us all in stitches when he shot a duck, which fell
inconveniently into the middle of a 400 metre wide tailings dam. Not a breath of wind to
carry it to shore, no dog to retrieve it - off came the clothes and in went our naked professor to
retrieve the prize in his mouth in a manner never bettered by any gundog!

"Sunshine" having proved a little disappointing, we turned our attention to the Watut area,
where we found some fossils and located tuff beds that subsequently proved suitable for
potassium/argon dating.

All too soon, however, Stirton, Miller and Lidicker had to return to the United States. We
parted having agreed that we would continue joint work on the fossils, stratigraphy and
sedimentology. Timbu Tasong went to work prospecting, particularly in the Watut. He
found many sites (Fig. 3, Pls 1, 2), and we developed a number of them into productive
quarries. The fossil material was sent to the University of California Museum of
Paleontology (U.C.M.P.), where I eventually ended up during 1964-1965, ostensibly to work
on the stratigraphy of the Otibanda Formation, and to learn something about vertebrate
palaeontology. Stirton, in his kind and disarming fashion, handed over all the fossil material
from New Guinea, so I had, in the nicest way, been thrown in at the deep end.

92 - PLANE

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(After Plane 1967a).

NEW GUINEA FOSSIL VERTEBRATE DISCOVERIES - 93

Figure 4. A, Wallabia rufogrisea, right pex x1/3, Recent; B, Protemnodon otibandus, right pes with left
calcaneum drawn in reverse and restored parts indicated with broken lines, x1/2; C, left calcaneum, ventral
view, x1/2; D, left calcaneum, dorsal view. x1/2; E, metatarsal IV, side view, x1/2; F, metatarsal IV, dorsal
view, x1/2. (From Plane 1967a).

94 - PLANE

Figure 5. Fossils from the Otibanda Formation, Pliocene. A-C, right P? of Protemnodon otibandus; a,
occlusal view; B, posterior view; C, lingual view; tooth length 17.8mm; D-E, Crocodilus cf. porosus, viewed
from two sides. F, right proximal tarsometatarsus, anterior vies of Casuarius sp; maximum length 64.5mm.
(From Hoch & Holm 1986). G-I, Thylacinus sp., left P2; G, lingual view; H, labial view; I, posterior view.
(From Plane 1976).

NEW GUINEA FOSSIL VERTEBRATE DISCOVERIES - 95

FURTHER WORK IN THE LATE 1960'S TO PRESENT

At the University of California I spent much of my time studying and writing up the
recently found New Guinea material. This proved to be a most productive time for me, with
two publications resulting in short order (Plane 1967a, 1967b) (Fig. 4). I returned to New
Guinea in 1967 to wet screen some of the Watut sites for small vertebrates. The locality I
worked was ideal for this type of operation, with good, flowing water, plenty of willing
helpers and good sunshine for drying. The results, however, were quite disappointing, but I
was at least able to confirm the presence of rodents during the Pliocene, albeit indeterminate

with regard to species.

Some field work has been carried out in the Watut area after my work there - by Pat
Wooley, a mammalogist from La Trobe University and Ella Hoch, a Danish palaeontologist
from Copenhagen (Hoch & Holm 1986) (Fig. 5, Pl. 3).

“OS NEW IRELAND

NEW BRITAIN |”

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BOUGAINVILLE
PAPUA NEW GUINEA ISLAND.

25 Measured strike and dip
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Figure 6. Geology and location of the Pureni site, which has produced a Pleistocene fauna. (From Rich ef

al. 1988).

Figure 7. Hulitherium tomasettiit, a panda-like diprotodontid from Pureni (artist P. Schouten, in Flannery &
Plane 1986).

Besides the fossils found in the Wau Valley, additional material, incduding unidentifiable
rodent remains, were recovered at Pureni (Fig. 6) in the Eastern Highlands. A Late Pleistocene

NEW GUINEA FOSSIL VERTEBRATE DISCOVERIES - 97

site was discovered there during improvements to the mission airstrip during 1967. The local
tribesmen, digging with handtools, discovered a layer rich in bones and plant material.
Although nominally Christians, the people were both excited and frightened, bones being
associated with ancestors. The result of this was that some of these tabu objects were broken
by the nervous poking and proding. Luckily, the late Father Bernard Tomasettii realised the
scientific worth of the material and salvaged it.

In 1969, I visited the site together with Paul Williams, a post-doctoral fellow from the
Australian National University. We investigated the stratigraphy, made further fossil
collections and put down an auger hole at the vertebrate site (Williams ef al. 1972). A
fascinating diprotodontid, Hulitherium tomasettii, named for its discoverers and salvager, was
described in 1986 (Flannery & Plane 1986)(Fig. 7, Pls 6-10), as later was a pygmy
cassowary, Casuarius lydekkeri (Rich, Plane & Schroeder 1988) (Pls 4, 5). Tim Flannery, a
mammalogist at the Australian Museum, in addition to co-studying Hulitherium has made
further investigations in the area and has published on new macropodid species from other Late
Pleistocene sites in New Guinea (Flannery ef al. 1983).

Rod Wells of Flinders University carried out field work in New Guinea, in 1975. I had
been asked if I would be interested in going to excavate some fossil bones discovered by
English speleologists deep in caves in the Western Highlands karst country. Being a total
coward and not at all fond of deep, dank, humid holes in the ground, I immediately offered the
opportunity to Wells, who revels in those conditions. He went, collected the fossils of a
sirenian but to this day claims that I tried to get him killed. On leaving the cave, he was
almost overtaken and drowned by rapidly rising flood waters caused by a thunderstorm at the
surface.

WEST IRIAN

No mention has been made of the western half of island New Guinea, and to date only
limited vertebrates have been described from there (Zygomaturus; Hardjasamita 1985). They do
exist, and material from cave deposits has been found. There are tantalising hints at a rich new
Pleistocene fauna, which includes small diprotodontids, possibly related to Hulitherium.

CONCLUSIONS

Given the difficult nature of field work in New Guinea, the unravelling of its prehistoric
faunal history is likely to be slow and erratic. That its vertebrate fauna during the late Tertiary
and Pleistocene is closely related to forms in Australia is undoubted. The specialisations
displayed by the diprotodontids, however, such as Kolopsoides from the Watut and
Hulitherium from Pureni, demonstrate that new and exciting discoveries lie in wait for those
who continue the search for the ancestors of New Guinea's fauna.

REFERENCES

ANDERSON, C., 1937. Palacontological notes. No. IV: Fossil marsupials from New Guinea. Rec. Aust,
Mus. 20(2): 73-76.

BULMER, S. & BULMER, R., 1964. The prehistory of the Australian New Guinea Highlands. Am. Anthrop.
66(4): 39-76.

FISHER, N.H., 1944. Outline of the geology of the Morobe Goldfield. Proc.R. Soc. Qd. 55(4): 51-58.

FLANNERY, T.F., MOUNTAIN, M.J. & APLIN, K., 1983. Quatemary kangaroos (Macropodidae, Marsupialia)
from Nombe rock shelter, Papua New Guinea, with comments on the nature of megafaunal extinction in
the New Guinea highlands. Proc. Linn. Soc. NS.W. 107(2): 75-98.

FLANNERY, T.F. & PLANE, M., 1986. A new late Pleistocene diprotodontid (Marsupialia) from Pureni,
Southern Highlands Province, Papua New Guinea. Bur. Min. Res. J. Aust. Geol. Geophys. 10: 65-76.

98 - PLANE

HARDJASAMITA, H.S., 1985. Fosil diprotodontidi Zygomaturus Owen 1859 Dari Nimboran, Irian Jaya.
Psekmnan Ilmiah Arkeologi 3, Jakartai, PPAN: 999-1004.

HOCH, E. & HOLM, P.M., 1986. New K/Ar age determinations of the Awe Fauna gangue, Papua New Guinea:
Consequences for Papuaustralian late Cenozoic biostratigraphy. Modern Geol. 10: 181-195.

ies it L.C., 1938A. Preliminary geological report on the Upper Watut area. New Guinea Admin. Rep.
unpubl.).

Ee L.C., 1938B. Geological report on the Upper Bitoi-Black Cat area. New Guinea Admin. Rep.
unpubl.).

PLANE, M.D., 1967a. Stratigraphy and vertebrate fauna of the Otibanda Formation, New Guinea. Bur. Min.
Res. Bull. 86: 1-64.

PLANE, M.D., 1967b. Two new diprotodontids from the Pliocene Otibanda Formation, New Guinea. Bur.
Min. Res. Bull. 85: 105-128.

PLANE, M., 1976. The occurrence of Thylacinus in Tertiary rocks from Papua New Guinea. Bur. Min. Res.
J. Geol. Geophys. 1(1): 78-79.

RICH, P.V., PLANE, M. & SCHROEDER, N., 1988. A pygmy cassowary (Casuarius lydekkeri) from late
Pleistocene bog deposits at Pureni, Papua New Guinea. Bur. Min. Res. Aust. Geol. Geophys. 10:
377-389.

VAN DEUSEN, H.M., 1963. First New Guinea record of Thylacinus. J. Mammal. 44: 279-280.

WHITE, J.P., 1972. Ol tumbuna: archaeological excavations in the Eastern Highlands, Papua New Guinea.
Terra Australis, Aust. Nat. Univ., Canberra.

WHITE, J.P., CROOK, K.A.W. & RUXTON, B.P., 1970. Kosipe: a late Pleistocene site in the Papua
Highlands. Proc. Prehist. Soc. 36: 152-170.

WILLIAMS, P.W., MCDOUGALL, I. & POWELL, J.M., 1972. Aspects of the Quatemary geology of the Tari-
Koroba area, Papua. J. geol. Soc.Aust. 18: 333-347.

PLATES

Plate 1. A, Stream channel in the Otibanda Fm, Koranga Creek, near Bulolo, Papua New Guinea; B, Awe
Fauna type locality on top of cliff near the umbrella. Looking south, Ekuti Range in back-ground, beneath
clouds (from Plane 1967a).

Plate 2. Typical exposures in upper Watut Valley, looking west. High ridge in background is Ekuti Range
(from Plane 1967a).

Plate 3. Right mandible of Protemnodon buloloensis, The sectorial P3 more elongate than any molar,
and all teeth wear simultaneously. Otibanda Fm, Upper Watut area (photo by B. Bang from Hoch & Holm
1986).

Plate 4. Casuarius lydekkeri. Femora: right, CPC26605b, A, internal view, D, proximal view; left,
CPC26605c, B, internal view, C, proximal view. Tibiotarsi: right, AMF50094, E, posterior view, G, distal
view, L, anterior view, M, internal view; right, CPC26605d, F, posterior view; H, distal view, J, anterior
view, N, intemal view; left, CPC2660Se, K, internal view, L, proximal view. Scale bar, 1 cm. All
specimens from Pureni, except AMF50094, whose locality is unknown.

Plate 5. Casuarius lydekkeri. Tibiotarsi, right, AMF50094, A, external view; right, CPC26605d, B,
extemal view. Tarsometatarsi, right, CPC26605f, C, anterior view, E, proximal view; left, CPC26605g, D,
proximal view; left, CPC26605h, F, anterior view, G, distal view. Phalanges: Casuarius phalanx 2 digit II:
C. benneitti, left, H, proximal view, L, distal view, O, lateral view, C. casuarius, right, I, proximal view, M,
distal view, P, internal view; C. sp. from Awe, P.N.G., UCMP70129, left, J, proximal view, N, distal view,
Q, lateral view; UCMP 70129, right, K, proximal view, R, intemal view. Scale bar, 1 cm. All C. lydekkeri
specimens except AMF50094 are from Pureni.

Plate 6. A, lateral view, and B1, B2, stereo occlusal view of partial cranium of holotype of Hulitherium
tomasettii from Pureni (after Flannery & Plane 1986).

Plate 7. Stereo dorsal view of cranium of Hulitherium tomasettii (from Flannery & Plane 1986).

NEW GUINEA FOSSIL VERTEBRATE DISCOVERIES - 99

Plate 8. Holotype of Hulitherium tomasetti (from Flannery & Plane 1986). A, C, stereo occlusal view of
partial left M3.5, xl; B, D, stereo occlusal view of right M3 and Ms, xl; E, G, stereo occlusal view of
posterointernal comer of right P3, x2; F, H, stereo occlusal view of partial right M34, xl; I, K, stereo
anterior view of anterior face of protolophid left M2, x2; J, L, stereo view of central portion of buccal face
of right P3, x2 (after Flannery & Plane 1986).

Plate 9. A, B, stereo view of posterior face and D, E, anterior face of proximal left femur fragment; C,
anterior face, and F, posterior face of right tibia of Hulitherium tomasettii (after Flannery & Plane 1986).

Plate 10. A, B, stereo view of anterior face, and C, D, posterior face of right proximal humerus fragment,

E, posterior face, and F, anterior face of distal right humerus fragment of Hulitherium tomasettii (from
Flannery & Plane 1986).

100 - PLANE PLATE 1

PLATE 2 NEW GUINEA FOSSIL VERTEBRATE DISCOVERIES - 101

102 - PLANE PLATE 3

PLATE 4 NEW GUINEA FOSSIL VERTEBRATE DISCOVERIES - 103

104 - PLANE RLATES

PLATE 6 NEW GUINEA FOSSIL VERTEBRATE DISCOVERIES - 105

106 - PLANE PLATE 7

PLATE 8 NEW GUINEA FOSSIL VERTEBRATE DISCOVERIES - 107

108 - PLANE PLATE 9

PLATE 10

NEW GUINEA FOSSIL VERTEBRATE DISCOVERIES

- 109

ee

4
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110 - PLANE

rb “ i ce * Kull
BAGH. oe e ee eat
82 ee ek ok ta ie. # a

Many species of flamingoes or related birds are known from the now arid Centre of Australia.
This reconstruction is of central Australia 15 million yBP. In the foreground Phoeniconotius
eyrensis cares for a juvenile. the more gracile Phoenicopterus novaehollandiae feeds in the mid
distances, while the white-coloured palaelodids stand in the mid-background, (From Rich & van
Tets 1985, with permission of The Museum of Victoria).

CHAPTER 4

PALAEOCLIMATIC
SETTING AND
PALAEOGEOGRAPHIC
LINKS OF AUSTRALIA IN
THE PHANEROZOIC

Lawrence A. Frakes! and Patricia Vickers-Rich2

TA OGUCH ONE sis dee eis ont ce hen b dee oh ag on debBiBsleles oa trees 112
A Brief History of Global Climates.................. 113
Phanerozoic Climates of Australia................04. 116
Cambrian Period .............cccsceecscceeeeneeeeeeees 116
Ordovician Period...........cccccccceeececeneeeeeeeees 117
Silurian Period ...........cccceceeecececececeeeeeeeeees 118
Devonian Period ...........cececcessceceeeeeeeeeseeens 119
Carboniferous Period ..............cccececeeeeeeeeees 120
Permian Period ............ccccececeeeceeeeeeeeeeeees 122
Triassic Period..........ccccccssesececessceceeeeerenees 123
Jurassic Period.........cccccsessseccscceceseceeeeeeees 124
Cretaceous Period.........cccecccsssceceseeceeeeeeees 125
CAIN OZ OICHERA sce ton obs. 0rvee cutee cetess se vea' tecths 127
Geographic Links and Barriers: Determinants
of Biotic Distributions...............ccceceeeeeeeees 130
CONCLUSIONS. ..0..t.csscie itis ces nddevebiedeertiessesecedadane 137
REPETCTICES Foe. wce:siere streets lo XG- oho Sars ajinn oh faglobeaneibon selves soe 143

a

1 Department of Geology & Geophysics, University of Adelaide, South Australia 5001, Australia.
2 Earth Sciences and Botany/Zoology Departments, Monash University, Clayton, Victoria 3168, Australia.

112 - FRAKES & RICH

INTRODUCTION

A look at the distribution of climates on the modern globe reveals that elements such as
mean annual temperature, rainfall, etc. are controlled largely by latitude. That is, similar
climates tend to occur in latitudinal ("zonal") bands, unless strongly influenced by topographic
irregularities. This banding arises from the equator-to-pole gradient in heat energy received
from the sun, but the effect is imperfect because of geographic controls on transport of heat
toward the poles in both atmosphere and oceans (e.g. the Gulf Strearn). Seasonal variations in
distribution of sunlight owing to the tilt of the earth cause climate bands to shift their
latitudinal positions over the year.

The relative elevation of continents, and their locations, also have strong influence on
regional climate. Climates in the Alps are markedly colder and wetter than lowland climates at
the same latitude, and on a global scale, the east coasts of continents in mid-latitude are
characterized by more humid climates than are west coasts. Additionally, the location of a
continent is important because land masses serve as barriers to circulation in the air and in the
oceans and thus affect their own climates. Also, since land reflects more sunlight than does
water at the surface of the earth, concentration of land at low latitudes means that less heat will
be retained in the system, and overall, climaies of the earth will be cooler. Thus, in
considering palaeoclimates it is vital to know not only the latitudes of a continent but its
elevation (especially mountains), its relationship to oceanic circulation patterns and the global
distribution of land and sea. Climatic changes through the Phanerozoic, as well as the
changing geography, have affected the options available to the biota for dispersal and
distribution.

There is abundant evidence, in the form of widespread tillites, evaporites, reef limestones,
etc. as well as in palaeotemperatures inferred from measurement of oxygen isotopes, that the
thermal state of the climate system has varied greatly over earth history (Frakes 1979). The
climate of a continental land mass, therefore, varies not only as a function of its position and
elevation but also as a consequence of the global thermal state. For example, parts of Australia
were glaciated during the late Palaeozoic, but the continent remained nearly, if not entirely free
of permanent ice during the Mesozoic, although the continent lay at approximately the same
latitudes during both intervals. The first half of the Permian was a time of great cooling over
most of the earth, whereas the Mesozoic featured exceptionally warm climates, except in polar
areas.

Several factors combine to make interpretation of palaeoclimatology difficult. First, the
climate system is complicated by a network of positive and negative feedback mechanisms,
which hinder determination of cause and effect relationships. Second, geologic materials, or
proxy evidence as they are referred to by Lamb (1977), must be correctly and precisely dated in
order to establish the sequence of events in the time scale. This almost never seems to be the
case. Third, closcly spaced and synchronous data points are required if the extent of a particular
type of climate is to be found, as would be important, for example, if one were trying to
estimate the difference between the climatic state at the time in question versus that at present
or at some other time. Although data are never sufficient for these purposes, progress is being
made, at least to the point when an attempt can be made to sketch the climatic history of
continents.

This chapter is only slightly modified from that of Frakes & Rich (1982). A useful
companion to this summary is Frakes, McGowran & Bowler (1987). Reference to Fig. 1 will
aid in location of major depositional areas discussed throughout the chapter.

PALAEOCLIMATE & PALAEOGEOGRAPHY OF AUSTRALIA - 113

A BRIEF HISTORY OF GLOBAL CLIMATES

The earliest known sedimentary rocks (as xenoliths in the Ishua Group, Greenland) include
fragments of banded iron formation older than about 3,800 myBP. While these themselves are
not diagnostic of climates of the time, somewhat younger rocks (~3,200 - 2,600 myBP)
elsewhere are characterized by an abundance of carbonates, suggestive of warm climates.
However, until latitudes of deposition are known for these strata, it is not possible to say
whether the global situation was warm or otherwise. Although unlikely, the case may be that
all carbonates of this interval were laid down in relatively low latitudes. Some oxygen isotope
work on cherts also suggests quite warm climates (Knauth & Epstein 1976), but again at
unspecified latitudes.

The earth's first glacial episode took place at around 2,500 - 2,300 million years ago
(myBP), with large tracts of glacial debris being deposited in Canada (the Huronian sequence),
South Africa (Witwatersrand Group, etc.) and possibly in Western Australia (Wyloo Group).
In Canada the glaciation was multiple in that at least three glaciations are recorded. Following
this and until the next known major glaciation beginning at about 1,000 myBP, climates once
more seem to have been characterized by warmth and equability. The late Precambrian
glaciations appear to have lasted about 400 my (from ~1,000 to 600 million years ago) and are
known to have affected all continents with the exception of Antarctica, where geologic
information is sparse. Interestingly, many tillite localities of this age were in relatively low
latitudes, based on palacomagnetic data. It is likely that the late Precambrian was the coldest
time in the history of the globe.

By the beginning of Phanerozoic time (Fig. 2), climates had warmed and glaciers had
disappeared. What evidence is available suggests that all land masses were then at latitudes of
less than 60°. The highest latitudes to which these warm climates extended is not known;
however, warm-water indicators in shallow seas at 55° latitude imply a comparatively warm
global situation. This is further suggested by a general lack of evidence of cold climates. The
Ordovician Period (Fig. 3) seems to have been a time of fairly rapid oscillation of climate as
suggested by variations in carbonate distribution and in eustatic sea level changes. Near the
end of the Ordovician a major phase of glaciation affected Africa and marginal effects are seen
in adjoining continents, including Europe and North America. This cold episode ended early in
the Silurian (Fig. 4), and was followed by generally warm conditions.

The Devonian (Fig. 5) was a time of very great warmth on the globe, although there may
have been limited glaciation in near-polar areas of South America. Late Devonian evaporites
extend to about 40° palaeolatitude, at least 5° beyond their present area of formation. The
aridity of the Devonian gave way to very humid conditions during the Carboniferous (Fig. 6)
as deduced from the abundance of coals in North America, Europe and Asia. This global high
humidity probably contributed to glaciation, which began in the middle Carboniferous,
inasmuch as abundant precipitation on high-latitude land masses eventually led to the
formation of glacier ice. The second half of the Carboniferous and the first quarter of the
Permian (Fig. 7) saw glaciation spread over a large part of Gondwana. For unknown reasons
this abruptly halted, and the globe again became relatively warm by the mid-Permian.

The earth apparently reached its thermal maximum during the Mesozoic Era (Figs 8, 9). In
the Late Cretaceous, oxygen isotopes define two peaks of warmth, Albian and Coniacian-
Santonian, the latter synchronous with the highest peak on the Phanerozoic sea level curve of
Vail et al. (1977). The latest Cretaceous began the long and intermittent cooling which
resulted ultimately in the Pleistocene glaciations (Figs 10, 11). Large temperature drops

Figure 1. Major tectonic and depositional features of the Australian continent during the Phanerozoic.
(Derived from the Bur. Min. Res. Earth Science Atlas of Australia, Canberra, 1979).

114

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occurred in the oceans at the end of the Eocene and in the Middle Miocene, the latter
apparently being the time during which most Antarctic ice accumulated. The last two
million years has been characterized by advances and retreats of ice sheets in both hemispheres.
Cooling of the oceans over the last 700,000 years operated on a cyclicity of about 100,000
years. The last interval of significant ice growth ended about 18,000 years ago.

PHANEROZOIC CLIMATES OF AUSTRALIA

The climates of Australia over the last 570 million years, during the history of vertebrates
on the continent, are a reflection of two factors. Firstly, they reflect the variable
palaeolatitudinal position of the continent and secondly the global climatic state. Climates of
the continent at any given time can be expected to be varied owing to the great size of the land
mass. However, in many cases geologic information on climates is limited to certain regions
or to widely separated areas and generalizations about the whole continent accordingly are weak.
Since dating of diagnostic sedimentary deposits commonly is imprecise, detailed description of
short time-intervals is not yet possible. These limitations place constraints on interpretations
of Australia's climatic history.

CAMBRIAN PERIOD

Climatically significant rocks of Cambrian age in Australia include only evaporites.
Archaeocyathid limestones, oolitic or pisolitic limestones and red beds embrace warm
palacoclimates amongst their possible depositional environments. For example, red beds
probably represent deposition under warm conditions, at least when associated with evaporites,
but they are known to form and to be diagenetically altered, under a variety of conditions.
Accordingly, red beds and non-reefal limestones are interpreted with reservations herein.

Evaporites are known from the Ord Basin (Fig. 1) of northwest Australia and the Georgina
Basin. These appear to be related to a widespread transgression of the sea, which began in the
early Middle Cambrian and lasted into the Late Cambrian. Limestones and dolomites are
common deposits resulting from this transgression and oolitic varieties possibly represent
deposits of warm shallow seas (Bonaparte Gulf and Georgina basins, Adelaide Geosyncline).
Red beds of Cambrian age include the Ayers Rock Arkose of the Northern Territory.

Palaeogeographic reconstructions vary in positioning Australia at this time (and other
times) (e.g. Irving, 1964). However, all available ones place Australia in relatively low
latitudes; the reconstruction in Fig. 2 shows Australia lying between about 5° and 40°N
latitude. The east coast in this scheme should have been exposed to comparatively cold oceanic
waters traversing down from high latitudes, but if so, any evidence is masked by the dominance
of clastic sedimentation at this time. Localities at lower latitudes are characterized by warm
water indicators, such as evaporites in the Northern Territory, South Australia and Western
Australia, For areas where Cambrian strata are unknown, it can be predicted that most of the
western half of the continent experienced tropical climates with abundant rainfall in the
Cambrian and the southern areas, being more isolated from the sea, may have seen the
develoment of desert climates. Strata of the Middle to Late Cambrian Dundas Group offer few
clues to climatic conditions as most were deposited in deep water.

Finally, the Middle Cambrian may have been warmer than either the early or late parts of
the period. This interpretation is based on the idea that the eustatic sea level rise in the Middle
Cambrian corresponds to a slight global warming. However, the associated transgression of
the sea, which occurs on other continents as well, may instead reflect increased tectonic activity
in the oceans and a consequent decrease in the volume of ocean basins (Hallam, 1977).

PALAEOCLIMATE & PALAEOGEOGRAPHY OF AUSTRALIA - 117

CAMBRIAN (Late) 510 my.

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Figure 2. Palaeogeography and palaeoclimatic indicators for the Cambrian Period. Black triangles, reefs;
black squares, evaporites, and black circles, coals; symbols apply to Figs 1-10. (Modified from Scotese et
al. 1979).

ORDOVICIAN PERIOD

Ordovician epeiric seas occupied a broad belt across the continent from the Canning Basin
southeastward to the Lachlan Fold Belt. However, sediments in these areas are mostly of Early
and Middle Ordovician age. This is in keeping with the global sea-level scheme for the
Ordovician proposed by Spjeldnaes (1961). Important deposits occur in the Canning and
Georgina basins, the Lachlan Fold Belt and western Tasmania.

Most Ordovician deposits of Australia consist of dark shales and turbidites laid down in
deep water, and these have little climatic significance. The so-called "shelly" facies of the
Ordovician has a restricted distribution and consists of both shelf-type limestones and local
bioherms. The bioherms are known only from two areas in Tasmania and New South Wales
(Cliefden Caves Limestone, near Orange). Solitary corals occur in several places, but their
temperature tolerances are unknown. Oolitic limestones, suggestive of warm shallow seas,
occur in western Tasmania and the Georgina Basin. The other main occurrences of carbonates
are scattered throughout the Lachlan Fold Belt and in the Amadeus and Canning basins and in
western Tasmania.

Arid regimes are indicated by traces of evaporites in the Canning Basin (anhydrite),
Amadeus Basin (gypsum, halite casts) and the Georgina Basin (gypsum). Also, there is a
suggestion of aridity in fanglomerates of the Junee Group in western Tasmania. There is no

118 - FRAKES & RICH

trace in Australia of the glaciation which affected the African sector of Gondwana; the only
suggestion of Late Ordovician cooling comes from the apparent regression of the sea at that
time.

The distribution of climatic indicators as shown in Fig. 3 suggests relatively warm
conditions over most of the continent. This is in keeping with the position of Australia
between 20°S and 20°N during the Middle Ordovician. Dry easterly equatorial winds from over
the main land mass of Gondwana may have contributed to these conditions. There is, thus,
little observable change in the climates of Australia between the Cambrian and Ordovician
periods.

MIDDLE ORDOVICIAN 465 my.

Figure 3. Palacogeography and palaeoclimatic indicators for the Ordovician Period. See caption Fig. 1.

SILURIAN PERIOD

The reconstruction for the Silurian (Fig. 4) shows Australia positioned between the equator
and about 35°S latitude. The continent was considerably less inundated by the sea than during
the Ordovician, and extensive outcrops are limited to the Lachlan Fold Belt. Marine strata are
abundant there and include biohermal, biostromal, and oolitic and detrital limestones and some
dolomite. Additional occurrences of marine rocks are those of the Carnarvon Basin and
scattered areas in northern Queensland. Dating of many Silurian non-marine rocks is difficult
owing to a lack of fossils; several evaporite deposits accordingly are referred to as Siluro-
Devonian (Wells 1980).

In the Lachlan Fold Belt, limestones were deposited on shelf areas and local "highs",
particularly in the south of New South Wales. Many of these deposits contain solitary coral

PALAEOCLIMATE & PALAEOGEOGRAPHY OF AUSTRALIA - 119

genera of uncertain climatic significance. Similar deposits occur in the New England area, in
western Tasmania, and along the Chillagoe Shelf in northern Queensland. What appear to be
bioherms, suggesting warm conditions, are limited to northern Queensland (Wairuna
Formation) and the Cobar region (New South Wales). Oolitic limestones have also been
reported from the Wairuna Formation (White 1961).

MIDDLE SILURIAN 415 m.y.

Figure 4. Palaeogeography and palaeoclimatic indicators for the Silurian Period. See caption Fig. 1.

The only known Silurian evaporites are substantial deposits in the Carnarvon Basin, the
Dirk Hartog Formation of Wenlockian age. Deposition of anhydrite and halite in the Dirk
Hartog Formation may correspond to a global marine transgression determined by Vail et al.
(1977). Aeolian, possibly desert, conditions are suggested by cross-stratified units in the
Mereenie Sandstone of the Amadeus Basin. Evaporitic units of "Siluro-Devonian" age include
strata in the Canning Basin (Carribuddy Formation).

From the above evidence, Australian Silurian climates can be seen to be characterized by
warmth and aridity, at least in the west. Probably warm and arid conditions existed over the
interior of the continent where evidence is lacking. From configurations on Fig. 4, warmth
along the eastern seaboard can be explained by southward deflection by the land mass, of warm
westward-moving equatorial currents in the adjacent ocean. Although indicators are lacking,
wet climates would also be expected here. Western and northwestern parts of Australia,
although at slightly higher latitudes, remained warm and arid, probably because the region was
positioned beneath the subtropical low-pressure zones, as are modern deserts. There was little
high-latitude ocean in the Southern Hemisphere to cool the continent at this time. The African
glaciation, which began in the Late Ordovician, terminated in the Early Silurian; its effects are
not recorded in the Silurian rocks of Australia. Late Silurian strata appear to be less abundant

120 - FRAKES & RICH

than expected in Australia, perhaps reflecting a large global regression of the sea as indicated in
the sea-level curve of Vail et al. (1977).

DEVONIAN PERIOD

The position of Australia on the globe (Fig. 5) is essentially the same as during the
Silurian, and the distribution of climate indicators, therefore, could be expected to be similar.
There are striking differences, however, probably the result of a marked global warming in the
Devonian. Local effects, no doubt, were introduced as a consequence of widespread tectonism at
this time in Australia.

EARLY DEVONIAN 390 m.y.

GONDWANA

a

Figure 5. Palaeogeography and palacoclimatic indicators for the Devonian Period. See caption Fig. 1.

Evaporites and reefal limestones are abundant in Devonian sediments of Australia.
Evaporites occur in the Adavale, Arckaringa, Bonaparte Gulf and Canning basins, and
commonly are associated with red clastic sediments. Most apparently were deposited during the
Middle Devonian, although there are suggestions that some are of Early or Late Devonian age.
Anhydrite is most common, but halite is more abundant than at any other time, and gypsum
also occurs in the Adavale Basin. This flourishing of evaporite deposits in Australia parallels
the global total, in which Devonian evaporites are abundant (Gordon, 1975). Further, the
concentration in the Middle Devonian is in agreement with a marked global transgression at
about this time (Vail e¢ al. 1977, Haq et al. 1988). It is likely that transgressions led to
evaporite deposition through the flooding of shallow marginal basins in low areas near the
shoreline. Interestingly, although coals make their first appearance in the Devonian with the
advent of land plants, Devonian coals are unknown in Australia, and indeed in Gondwana. This

PALAEOCLIMATE & PALAEOGEOGRAPHY OF AUSTRALIA - 121

perhaps results from the geographic isolation of Gondwana (Fig. 5), rather than from
particularly harsh climates for plant growth (at any rate, the genus Leptophloeum is well
known in the Late Devonian of eastern Australia.)

As in earlier periods, the accumulation of shelf carbonates was widespread during the
Devonian, They are known from the Lachlan Fold Belt (Victoria, New South Wales) and the
basins of northern Queensland; in both areas Early to Middle Devonian reef structures have
been identified, and they also occur in the Canning Basin where they are of Late Devonian age.
Oolitic limestones are known from the Wee Jasper area, New South Wales. It is notable that
marine invertebrate faunas of the Devonian show great diversity, another feature of warm
climates.

It is difficult to explain the occurrence of evaporites in the Adavale Basin, in view of the
fact that the east coast of the continent lay exposed to warm, equatorial ocean currents. They
could represent strictly local evaporative conditions. The lack of Devonian coaly deposits in
Australia can perhaps be attributed to a scarcity of peat-forming plants due to geographic
isolation from an area of radiation in the Northern Hemisphere. The warm and highly
evaporitive climates of the west are predictable because of their slightly higher latitudes and
their location away from equatorial sources of moisture for precipitation.

CARBONIFEROUS PERIOD

In the Carboniferous, Australia was located at much higher latitudes than earlier (Fig. 6),
and as a consequence, glaciation was initiated. However, sizeable bodies of ice which could
leave a record in the rocks, were not generated until the Westphalian interval, about halfway
through the Carboniferous. The pre-Westphalian interval saw the deposition of limestone,

LATE CARBONIFEROUS 290 m.y.

Figure 6. Palaeogeography and palaeoclimatic indicators for the Carboniferous Period. See caption Fig. 1.

122 - FRAKES & RICH

although not as extensively as in earlier periods, and these include reefs and oolitic limestone
in the Yarrol Trough and oolitic limestone in the Tamworth Trough and the Carnarvon Basin.
Early Carboniferous coaly layers also are known from the Tamworth Trough. Anhydrite
occurs in the Anderson Formation of the Canning Basin, the only known example of warmth
and/or aridity on the continent during the Late Carboniferous. In Australia, evaporites show a
continuous decrease in abundance since the Middle Devonian; increasing humidity is indicated.

In contrast to the generally warm conditions of the Early Carboniferous, the later half of the
period shows marked cooling. The earliest evidence for this is in glacial deposits of
Westphalian to Stephanian age in the Kempsey region of New South Wales; deposits in
Tasmania and near Heathcote in Victoria may be as old. Most Australian late Palaeozoic
glacial deposits span the Carboniferous-Permian boundary, and some can only be referred to as
"Permo-Carboniferous" in age owing to poor resolution in dating. However, Carboniferous ice
was much less extensive than Permian ice. It appears that the former occurred primarily
(between New England and the Bowen Basin) as alpine glaciers on highlands and islands along
and near the eastern seaboard (Crowell & Frakes 1971). Climates were cold here, in part
because of elevation effects, but the remainder of the continent, the western two thirds, was
somewhat warmer then and became progressively cooler through the period and into the
Permian. Glacials of the Grant Formation in the Canning Basin are also of Stephanian to
Sakmarian age. True ice-sheet conditions apparently did not exist on the continent until the
earliest Permian.

Australia lay between about 40° and 70°S latitude during the Carboniferous (Fig. 6),
following a rapid change of palaeolatitude. This migration brought Western Australia into
higher latitudes than the east, but glaciation in the west lagged, possibly because the region
was isolated from sources of warm ocean waters to provide a moisture source to build glaciers.
On the other hand, eastern Australia (and Victoria Land in Antarctica) lay where subtropical
Pacific currents, deflected southward by Coriolis acceleration, supplied warm surface waters for
evaporation and hence precipitation. This situation led eventually to the construction of
glaciers and ice-sheets. These trends are strongly supported by the occurrence of coal in the east
and evaporites in the west during the Early Carboniferous.

PERMIAN PERIOD

Australia continued to occupy a high-latitude position during the Permian (Fig. 7).
Glaciation expanded to such an extent that a major ice-sheet occupied much of the southeastern
half of the continent and large ice bodies were in place on the western half (Crowell & Frakes
1971). However, glaciation began to wane at the end of the Sakmarian Stage and the ice-sheet
was restricted to parts of the east (Bowen Basin and Tasmania) by the Artinskian. All traces of
glaciation were gone by the end of the Kazanian (early Late Permian). Over most of the
continent, post-glacial sediments include coaly strata. The ages of these coals vary from place
to place, as does the age of the termination of local glaciations.

Extreme refrigeration affected all of Australia in the Early Permian, as it did most of
Gondwana (Frakes 1979). The major southeastern ice-sheet appears to have originated partly in
adjacent parts of Antarctica and spread northward and north-westward into New South Wales,
Victoria, South Australia and the Northern Territory. Meanwhile, ice masses grew in areas
adjacent to the Bowen, Officer, Perth and Carnarvon basins and both to the southwest and the
northeast of the Canning Basin. Melting of Australian ice in coastal areas led to the deposition
of glacial marine sediments in widely separated areas in Tasmania and Queensland during the
Artinskian, Kungurian and Kazanian stages. Although morainal tillites of this age are not
known in Australia, there is no doubt that terrestrial ice was present as parent material for

PALAEOCLIMATE & PALAEOGEOGRAPHY OF AUSTRALIA - 123

icebergs. Over other parts of Gondwana all ice had disappeared before the Kungurian - this
signals a global warming the effects of which over-shadowed the continued high latitude
position of the Australia-Antarctica sector of Gondwana. Glaciation ceased, although land
masses still existed in the polar zone.

Growth of ice in Australia again is attributed to a source of warm surface waters diverted
from subtropical seas. During the postglacial Permian, precipitation was abundant, at least
intermittently, but this did not lead to the formation of new ice or glaciers, probably simply
because the earth-atmosphere system was warmer than earlier. Abundant precipitation did lead,
however, to widespread accumulations of vegetative debris and coal formation at times in the
Late Permian (Bowen, Sydney, Canning, Perth and Galilee basins and in Tasmania). As in
earlier periods, evaporites are restricted to the west (gypsum, Perth Basin). These lie between
the glacial Nangetty Formation and the Irwin River Coal Measures and are restricted in
distribution, suggesting a local climatic sequence which changed from cold to dry to moist over
the interval Sakmarian to Kazanian.

LATE PERMIAN 240 m.y.

SIBERIA SS

Figure 7. Palaeogeography and palaeoclimatic indicators for the Permian Period See caption Fig. 1.

TRIASSIC PERIOD

The opening of the Mesozoic Era saw a major regression of the sea from the Australian
continent. As a consequence, Triassic sedimentary rocks are rare, and non-marine types
predominate. Marine strata appear to be limited to marginal areas of the present continent - in
the Canning, Carnarvon and Perth basins and southeastern Queensland. In the Triassic,
Australia's palaeolatitudinal position was little different from that in the Permian, Despite

124 - FRAKES & RICH

being located between about 40° and 65°S latitude, Australian Triassic climates appear to have
been warm and with seasonal rainfall. These conditions generally reflect the global Triassic
picture.

Rocks diagnostic of climate are poorly represented in the Australian Triassic, and fossil
floras and faunas similarly are scarce. Even where present, fossils as yet are of little value in
evaluating climate. Coals are most important; they are known from east-central South
Australia (Leigh Creek), Tasmania, and coastal basins in New South Wales and southern
Queensland. Of interest is the fact that most of these coals seem to be Middle or Late Triassic
in age, suggesting an increase in precipitation since the Early Triassic. Middle Triassic red
beds were deposited in the Fitzroy Graben (Canning Basin), and Early Triassic ones were laid
down in the Bowen, Galilee and Sydney basins, and near Hawker in South Australia. At the
latter locality, gypsum occurs in shales. Interpretations of red beds as indicating climates in
the source area are now known to be unreliable, first because in some cases the pigmentation is
formed during diagenesis and second because colour may result from oxidizing conditions at
any place during transport or deposition, or in the weathering cycle. The significance of
Australian Triassic red beds has not yet been determined through detailed petrologic-
geochemical studies. However, the lack of Early Triassic coal deposits suggests that at least
this interval was characterized by relatively dry climates.

The scarcity of Triassic evaporites can be taken as evidence for generally humid climates
throughout the period. However, on a global basis, evaporites of the Triassic are among the
most abundant for any period. It is much more likely that Australian evaporites were not
deposited because large marine embayments were unavailable, than because of an increase in
humidity. Most Australian Triassic non-marine sediments appear to have been laid down in
fluvial environments; a few were formed in swamps.

JURASSIC PERIOD

Australia was again positioned between about 35° and 65°S latitude (Fig. 8) in the Jurassic
and was the site of continued non-marine sedimentation. However, the situation changed in the
Late Jurassic when transgressions of the sea began to occupy marginal areas of the continent
near the New South Wales-Queensland border and the Perth, Carnarvon and Canning basins of
the west. This corresponds in time to a known global warming of moderate magnitude.

Coal is the dominant climatic indicator in the Australian Jurassic, being known from the
east, south and west (Laura, Carpentaria, Eromanga, Surat, Ipswich-Moreton [east], Polda
[south], and Perth basins [west]). Concentration of coals in the eastern basins is again
explained by the proximity of the region to a moisture source - southward-moving subtropical
currents in the Pacific. Western regions still had little access to such currents and, therefore,
tended to remain drier. Early rifting along the Australia-Antarctica join may have provided a
moisture source for Polda Basin coals.

Most Australian Jurassic coals are of Middle Jurassic age; Early Jurassic ones appear to be
of minor thickness and extent. On this basis one can suggest a gross climate trend from fairly
humid climates in the Early Jurassic, in the west only, to quite high precipitation in the middle
of the period, to less precipitation in the Late Jurassic. This scenario does not compare well
with the global trend as based on evaporite abundances - a relatively wet Early Jurassic and
increasingly arid conditions through the remainder of the period. It is likely that local
tectonism and other factors, such as relatively slight transgressions, played a part in
determining the distribution of coals. Again, relatively more widespread precipitation is
indicated for the east than for the west, and this is taken as reflecting the presence of an
evaporative source in the southwestern Pacific.

PALAEOCLIMATE & PALAEOGEOGRAPHY OF AUSTRALIA - 125

LATE JURASSIC 135 m.y.

Figure 8. Palaeogeography and palaeoclimatic indicators for the Jurassic Period. See caption Fig. 1.

CRETACEOUS PERIOD

Fig. 9 shows late Maastrichtian palaeogeography, but as in previous diagrams, climatic
indicators are shown for the whole period. Again, Australia maintains its high latitude
position ( 35° to 70°S latitude). Embleton (1972) and Veevers (1984) show a clockwise
rotation of the continent for the Mesozoic relative to the late Paleozoic; this suggests that
western Australia (at 45° - 60°S latitude) might have experienced somewhat warmer climates in
the Mesozoic than earlier. This is certainly the case, indeed for the entire continent, but overall
there is little indication that the west was warmer than the east (45° - 85°S latitude) during the
Mesozoic. Such generalizations are particularly difficult for the Cretaceous because of the
complexity of sedimentation.

The global record of Cretaceous climates shows warming late in the first half and cooling
late in the second half of the period. The warmest times, possibly averaging 10°C warmer than
at present, probably occurred in the Albian and the Coniacian-Santonian stages. These
intervals coincide with times of high relative sea level (Haq et al. 1988). On a global basis,
the Cretaceous is notable for containing plants reminiscent of modern subtropical types to
latitudes as high as 55°; these lately have been reinterpreted as indicating strong seasonality,
including formation of winter ice.

In the Early Cretaceous a major seaway probably crossed the continent from the Canning
Basin through the Officer and into the Great Artesian and Eucla basins. At times, the Great
Artesian Basin, occupying much of the centre of the continent, probably also had eastward
connections to the sea by way of the Surat, Eromanga and Clarence-Moreton basins and

126 - PRAKES & RICHI

northeastward ones via the Carpentaria and/or Laura basins, Other marine depocentres included
the Murray, Carnarvon and southern Perth basins, and non-maring sediments accumulated in
the Otway and Gippsland basins, The Late Cretaceous saw intermittent marine sedimentation
inthe Canning, Carnarvon, Perth and Otway basins, and in the region around Darwin,
Northern Terntory, while non-maring accumulations took place over part of the centre of the
continent, “The Neoconian, Apuian and Albian all were times of regionally important marine
transgressions; Cenomanimn and Turonian transgressions were of lesser extent. The sea
withdrew for the last tame from interior Australia probably at the end of the Albian (Frakes ef
al, 1987)

LATE CRETACEOUS (Maastricohtian) G5 my

Pigure 9% Palacogeography and palacocliumatio indicators for the Cretaceous Ponod, See caption Pig. |

Climate imdicators are not abundant im the Australian Cretaccous, Limestones occur
sporadically, but reefs and oolites are lacking, Evaporites and red bed sequences are unknown
except for local gypsum (probably of secondary origin) in the Wallumbilla Formation of the
Hromanga Basin (Smart & Semor 1980), However, coal is widespread in Lower Cretaceous
units ino many of the eastern Australian basins, although it is very scarce in the Upper
Cretaceous sequences, Marine invertebrate faunas suggest fairly warm climes (Scheibnerova
1971, Hany 1979), while elements of Cretaccous floras suggest temperate climates with
increasing precipitation through the period (Dettmann & Playford 1969, Douglas & Williams
1982), Recent work on the palacobiota (Rich ef a/, 1988, Rich & Rich 1989), isotope
geochemistry (Gregory ef al, 1989) and scdimentology (Frakes & Francis 1988) suggest that in
the southern and central parts of Australia temperatures may have dropped below O8 C, at
times (particularly during the long polar winters), and scasonal ice may have formed during the
Early Cretaceous (Frakes & Francis 1988), ‘The evidence trom coal distributions (decreasing

PALAEOCLIMATE & PALAEOGEOGRAPHY OF AUSTRALIA - 127

precipitation) may thus be at variance with that from analyses of floras, but it is likely that
coal abundance results merely from the extent of ponding due to local and regional isostatic and
eustatic effects. Distribution of coals along the eastern side of the continent again reflects the
availability of warm surface waters in the Pacific as a moisture source, and their concentration
in the early half of the Cretaceous suggests that Australian coals are coincident in time with the
major transgressions which are known to have affected the continent. Late Cretaceous climates
on a world scale, are the warmest known in earth history.

Seafloor spreading between Australia and Antarctica was initiated in the Cretaceous prior to
86 myBP (Cande & Mutter 1982), perhaps as long ago as 95 myBP (Veevers 1986a), and this
was to have a marked effect on oceanic circulation and climate throughout the Cainozoic.

CAINOZOIC ERA

Climates of the last 65 million years are highly variable but are well documented by an
abundance of information. In this section, climates of the Cainozoic will be discussed in
general terms and reference will be to two figures only, Figs 10 and 11, which refer to the mid-
Oligocene (30 m.y.) and to the time when the last glaciation was at its peak (18,000 years ago)
respectively. Climatic data on Fig. 10 represent an epoch in the Tertiary Period.

In response to seafloor spreading, Australia separated from Antarctica in the Late Cretaceous
to early Tertiary and moved slowly northward relative to the South Pole beginning in the Late
Palaeocene (Embleton 1973, McElhinny et al. 1974, Cande & Mutter 1982). As a result,
Australia presents some interesting examples of climate change resulting from movement
across lines of latitude, but to evaluate these, the global cooling trends of the Cainozoic must
first be accounted for.

OLIGOCENE 35 my.

Figure 10. Palaeogeography and palacoclimatic indicators for the Oligocene Epoch. See caption Fig. 1.

128 - FRAKES & RICH

The earth as a whole underwent a progressive cooling in the Tertiary; this was an extension
of a major global cooling which began in the Late Cretaceous. Significant temperature
declines took place at the end of the Eocene and in the Middle Miocene. Prominent reversals of
this trend occurred in the Early Eocene, the early Late Eocene, and the Early to Middle Miocene
(see oxygen isotope record in Shackleton & Kennett, 1975). The end of the Pliocene saw the
development of marked short-term fluctuations, and, at about 800,000 yBP these developed into
cycles of about 100,000 years (Bowler 1976, 1982). There was a large increase in global
precipitation in parallel with the global cooling, and the largely arid climates of the Mesozoic
gave way to more humid conditions, particularly in low and mid latitudes. The first positive
evidence for this is seen in fossil land plant assemblages of Palaeocene age. In general, from
the Miocene onwards, climates were drier during cold intervals than during warm times.

Indicators of climate in the Australian Cainozoic include coals and lignites (Gippsland,
Bass, Otway, Murray, St. Vincent's Gulf and Eucla basins, and in local areas in New South
Wales and Queensland.) Most Cainozoic Australian coals are dated in the interval Eocene to
Miocene, Their concentration in the east is remindful of present conditions (humid east,
relatively arid west). Reefs occur in the Eucla Basin, but they are small and limited to the
Early Miocene. Evaporites are not uncommon in Quaternary sediments of the fresh-water
basins of the Centre, but Tertiary ones appear to be limited to the Late Palaeocene of the
Canning Basin and the Early Miocene of the Murray Basin. Limestones make their first
appearance in abundance since the Palaeozoic in many marginal basins.

LAST GLACIAL MAXIMUM 18,000 YEARS

eS

ak: a

Figure 11. Palaeogeography for the Pleistocene Epoch showing in heavy stipple the extent of the ice caps
during the last glacial maximum. (Modified after Smith & Briden 1977 and Rich 1975.)

PALAEOCLIMATE & PALAEOGEOGRAPHY OF AUSTRALIA - 129

Fossil floras are important indicators of climate in the Australian Cainozoic (Kemp 1978,
Kemp & Harris 1982). The picture which emerges from Kemp's and Kemp and Harris’ reviews
is one of humid climates from the Palaeocene through the Miocene. However, indications of
temperature are variable and inconclusive. In the Eocene, climates appear to have been humid
over nearly the whole continent, but there was restriction of humid (rainforest) conditions to
the southeast after the Middle Eocene. Continent-wide conclusions are difficult for the
Oligocene and later epochs because of a scarcity of data points, but a marked change to
grasslands occurred in the interior by Middle Miocene time. More recent work suggests that in
Victoria and New South Wales the onset of aridity was first felt at the end of the Miocene.

In basins marginal to the continent, marine fossils are of value in determining climate.
This is especially so in the case of the larger foraminifera (McGowran 1978). The occurrence
of these forms has long been known in the Gippsland and Bass basins (Middle Miocene) and
the Carnarvon Basin (Eocene through Pliocene), but they are now thought to have made large
excursions from the tropics to higher latitude sites in Australia during the Early Eocene, Late
Eocene, Late Oligocene and the Early and Middle Miocene. Accordingly, these intervals may
represent times of significant warming, as they seem to be on a global basis.

Another indication of humid conditions in the Australian Tertiary record is in the abundance
of weathered profiles signifying deep chemical weathering. McGowran (1981) assigns these,
without discussion, to several intervals as follows: Early Palaeocene, Late Palaeocene, Early
Eocene, Middle Eocene, Late Eocene, Late Oligocene-Early Miocene, Middle Miocene and
Early Pliocene. However, the only weathering profiles with comparatively narrow age ranges
are two levels from the Eromanga Basin - Maastrictian to Early Eocene, and Late Oligocene
(30 + 15 my). These are dated by palaeomagnetic methods (Idnurm & Senior 1978). The
latter date may also apply to profiles in Western Australia and South Australia, and weathering
at this time (mid-Tertiary) is considered by some to reflect intensely humid climates.

The northward migration of Australia carried it into progressively lower latitudes and
warmer climatic zones. Yet during the early Tertiary the continent experienced successively
cooler climates, as a result of pronounced global cooling. The proximity of Antarctica with its
expanding ice cap did not lead to Tertiary glaciation in Australia, and, in fact, carbonates and
other indicators of warmth were deposited in southern coastal basins at times, Like other
continents in lower latitudes, early Tertiary Australia experienced a wet phase, which probably
began in the Palaeocene. It was not until at least late in the Miocene that the continent
experienced widespread climates characterized by moderate rainfall, owing to its position
beneath the subtropical high pressure zone (sinking dry air from aloft). Northern areas first felt
these effects of migration, although at least moderate aridity had featured in the far west since
the Cretaceous. Quite likely, the zone of drier climates has moved south over time, but data
are too sparse to document or disprove this. Regarding surface temperature, the decline which
began in the Cretaceous affected eastern and southern Australia most markedly, but again, this
was intermittently punctuated by warm intervals in the Eocene, Oligocene and Miocene, as
well as by late Quaternary events. It is not yet possible to sort out the relative strengths of
these warming phases, although, on some evidence, the Middle Miocene one seems most
pronounced, both regionally and globally.

The Quaternary was the time of extensive drying out of the continental interior and no
doubt some coastal areas in the west. Bowler (1976, 1982) suggests that widespread dune
construction in the interior coincided with very dry periods, which first began at some time
prior to 300,000 years ago. Analyses of core material from Lake George, New South Wales,
have given a fine record of climate changes for the Southern Tablelands (Singh et al. 1981).
From about the end of the Miocene until about 940,000 years ago, the lake was full,
suggesting moist climates. The following interval up to the present is characterized by four
cycles probably related to glacial/interglacial stages but in an irregular fashion, and the record
for the last 400,000 years correlates with the 100,000 year cycles detected in variations in

130 - FRAKES & RICH

oceanic 018/916 ratios. Quaternary sea levels of eastern South Australia have been correlated
with the cycles as well (Idnurm & Cook 1980), but this gives no direct evidence bearing on the
climates of the region. A dramatic event in the Quaternary history of Australia was the
initiation of glaciation in the highland centre of Tasmania. Two ice advances are recorded, the
last probably corresponding to the last global glacial maximum of the time scale (~18,000
years ago).

GEOGRAPHIC LINKS AND BARRIERS:
DETERMINANTS OF BIOTIC DISTRIBUTION

In addition to the location and extent of climatic "zones", the physical continuity or lack of
continuity of land masses (or oceans in the case of marine organisms) can strongly influence
faunal and floral distributions in times past (Simpson 1940, amongst many others). Thus, for
each of the periods (e.g. Ordovician, Silurian, etc.) discussed in this book during which
vertebrates have left a record in Australia, maps are presented (Figs 1-11) that have been
modified after Scotese et al. (1979, for the Palaeozoic) or Smith & Briden (1977, and Smith,
Hurley & Briden (1981 for the Mesozoic and Cainozoic). Such maps clearly show the
proximity of different parts of the world's continents of today as well as point out where lands
and seas existed during the period discussed. The distribution of specific types of sedimentary
environments are also summarized at the end of this chapter in a series of maps (Figs 12-20)
modified from those of Truswell & Wilford in Rich & van Tets (1982) and Rich et al (1989).

ORDOVICIAN

Figure 12. Australian Ordovician palaeogeography (modified from Truswell & Willford in Rich & van Tets
1982). Slight differences in continental plots for Australia between Figs 2-11 and Figs 12-20 are due in part
to different opinions of authors and in part to the plots representing slightly different times in a single
period. See Fig. 20 for legend. In Figs 12-20 the maximum extent of different depositional environments
for each period are indicated.

PALAEOCLIMATE & PALAEOGEOGRAPHY OF AUSTRALIA - 131

What is clear from these palaeocontinental reconstructions, as well as those of Ballance
(1980), Powell et al. (1980) Weissel et al. (1977) and many others, is that Australia was
closely linked with the Gondwana continents through the Palaeozoic and into part of the
Mesozoic. What is interesting is that during part of this time Australia was also closely
associated with parts of China and Southeast Asia from the Cambrian until at least the
Devonian, and both terrestrial (Young 1987, among others) and marine (several papers in Cox
1974, Gray & Boucot 1979) faunas reflect this near proximity in being markedly similar.
After the Devonian, Gondwana and the present day Laurasia developed continuity, and
apparently terrestrial vertebrates at this time could have moved back and forth across this super-
continent hindered only by climate and mountain barriers, etc., but not by marine conditions.
This geographic state remained in effect into the Mesozoic and probably accounts for the
cosmopolitan nature of the dinosaur faunas that characterize these times. By Early Cretaceous
times, however, terrestrial biotas were beginning to exhibit endemism (Rich & Rich 1989).

DEVONIAN

Figure 13. Australian Devonian palaeogeography. See caption Fig. 12.

During the Mesozoic the Gondwana supercontinent began to break apart with continental
fragments often developing their own separate histories, and with epicontinental seaways often
providing barriers even on continuous tectonic plates. Australia remained physically connected
to Antarctica until some time in the early Cainozoic (Eocene), and then, about 55 million years
ago, completely severed connections as it drifted from a relatively high southern latitude
(Tedford 1974, Rich 1975, Cande & Mutter 1982, Veevers 1986a) to near its present low
southerly latitude by the Miocene. After its break with the Antarctic, and further west with
South America (to which it was probably attached by an archipelago during part of the

132 - FRAKES & RICH

Mesozoic), Australia was extremely isolated from all other continental areas. Australia became
both a Noah's Ark and a Viking

LATE CARBONIFEROUS
Figure 14. Australian Late Carboniferous palaeogeography. See caption Fig. 12.

PERMIAN
Figure 15. Australian Permian palaeogeography. See caption Fig. 12.

PALAEOCLIMATE & PALAEOGEOGRAPHY OF AUSTRALIA - 133

TRIASSIC

Figure 16. Australian Triassic palaeogeography. See caption Fig. 12.

JURASSIC
Figure 17. Australian Jurassic palaeogeography. See caption Fig. 12.

134 - FRAKES & RICH

CRETACEOUS

Figure 18. Australian Cretaceous palacogeography. See caption Fig. 12.

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Mo Mo RY 7

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EARLY TERTIARY

Figure 19. Australian early Tertiary palaeogeography. See caption Fig. 12.

PALAEOCLIMATE & PALAEOGEOGRAPHY OF AUSTRALIA - 135

MID TERTIARY

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Australian middle Tertiary palaeogeography. See caption Fig. 12.

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136 - FRAKES & RICH

both a Noah's Ark and a Viking Funeral Ship (McKenna 1974), that is, it carried on it a fauna
and flora that was derived from a Gondwana "population", and when it moved northwards, this
biota developed independently, then was superimposed on the "foreign" biota of the Oriental
region as Asia and Australia neared one another in the Late Cainozoic. The intermixing of
these two faunas has produced the complex biogeographic situation that characterizes the Indo-
Malaysian area with its numerous lines (Wallace's Line, Weber's Line, etc.) denoting the
divisions between the Oriental and the Australasian biogeographic realms (Simpson 1977,
Rich & van Tets 1982). Australia is also a Viking Funeral Ship that carries with it
Gondwanan fossils that will be superimposed alongside those of Laurasia when the collision of
Asia and Australia is complete in 50 million years time (Fig. 21).

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THE FUTURE, 50my from Present

Figure 21. Possible Australian palaeogeography in the future, 50 million years from now. (Modified
from a map in Scientific American, 1970, vol. 233(4): 39.)

The northward drift of Australia in the Tertiary led to the continent moving across several
degrees of latitude and through many different climatic zones. The development of aridity
during this period is reflected by the marked changes, particularly during the Late Miocene to
Pleistocene, in the fauna and flora (Hope 1982, Truswell & Harris 1982) (Fig. 22). The
grasslands and their associated grazers and fast runners expanded and diversified at the expense of
forests with their associated fauna of browsers and silvophiles. At the same time invaders from
the north, such as the varanid lizards, Acacia and murid rodents (Keast 1981, Archer &
Clayton 1984) had their impact on the endemic biota. The final affront by at least 40,000
years ago was that of man and his entourage of domestic associates. This accentuated trends
already set in motion by climatic change. The impact of climate, man and the inevitable
intermingling of Asian and Australian biotas in the future will precipitate massive extinctions
of the Australian endemic biota. The final result of these changes will have to be left to future
observers, but it will most certainly be tied to the perigrinations of the continents themselves.

PALAEOCLIMATE & PALAEOGEOGRAPHY OF AUSTRALIA - 137
CONCLUSIONS

Australia and its biotic composition has not only been affected by changing associations
with other continents, by severe isolation, and changing world climates, but has also been
affected by its own changing latitude. As the palaeogeographic maps here illustrate, Australia
was in an equatorial position and rotated 90° to its present orientation during the Ordovician,
when the first vertebrates are known on this continent. Subsequently in the Silurian and
Devonian, it moved south into higher, but still subtropical and temperate latitudes, and rotated
more towards its present day orientation. By the Carboniferous and Permian, it had moved
decidedly further south into temperate and even polar latitudes. During the Mesozoic, Australia
remained in much the same southerly position, even when New Zealand broke away in the late
Mesozoic, and it wasn't until the late Mesozoic to early Tertiary that once again the continent
changed latitudes, moving northwards to near its current position by the Miocene, after
crossing several degrees of latitude. As the initial spreading rate between Australia and
Antarctica is regarded to have been slow (Cande & Mutter 1982), interchange of terrestrial biota
between the two continents may have been possible long after the onset of breakup.

All of these factors need to be kept in mind when considering Australian vertebrate
biogeography and evolutionary patterns throughout the Phanerozoic (the last 600 million
years). All of them certainly must have had their effects and have markedly complicated
paleogeographic interpretations of the origin and evolution of the biota made 20 years ago
when biogeographers used the present day arrangement of continents.

Figure 22. Palaeontological and geological data bearing on the development of aridity in Australia.
(Derived primarily from Bowler 1982, Hope 1982, Truswell & Harris 1982).

138 - FRAKES & RICH

CHRONO-
STRATIGRAPHY

MAGNETO: GEOGRAPHIC
CARREY POSITION avin. TEMPERATURES

EUSTATIC CURVES

ak! ’
M

SERIES.

MAGNETIC
ANOMALIES
POLARITY

Antarctic ice at
nearly modern
extent

MIDDLE

MIOCENE

Warmer - foramin-
iferans indicate

warm surface H,O
Southern Australia

018/018 indicate
steep temperature
drop

OLIGOCENE

ind
ws
a
a
3

Decline of temp-
erature from
previous highs,
018/018 fluctuating
temperatures

MIDDLE

No ice at poles,
018/0' indicate
temperatures
warmer than
present

PALAEOCENE

{MINOR REVERSED POLARITY EVENT

[EE orm pouanry

REVERSED POLARITY

Haq et al. 1988

Truswell and
Harris 1982;
Hope 1982;

Bowler 1982.

Veevers 1986a,b

PALAEOCLIMATE & PALAEOGEOGRAPHY OF AUSTRALIA - 139

CHRONO
STRATIGRAPHY

SERIES.

L

UPPER

MIDOLE

MIOCENE
a

LOWER

| OLIGOCENE
Y=
[UPPER LOWER UPPER

MIDOLE

EOCENE

| LOWER

UPPE

PAL AE OCENE

LOWER

i
‘of

SEL AEELaEn EERE ER aR

40+

018/016 °C | WEATHERING SEDIMENTS Pein

oT
} Se ae

i

— NZ (Wellington)

++ Aust.(Victoria)

Devereoux 1967;
Dorman 1966;
Frakes & Kemp
1973

Shackleton &
Kennett 1975.

Possible
seasonal
dryness

Possible
seasonal
dryness

Silcretes
indicate
seasonal
aridity?

Deep
7, weathering
profiles,
humid
regime?
Idnurm &
Senior 1978
Quilty 1984
Truswell &
Harris 1982:
Wopfner, Callen &
Harris 1974.

—>

van)
n
=)
<x
oO
Ns
t=
c
®
(o)
£
Yn
o
oD)
w
=
w
©
ne}
fe)
®
is)
o
a
©
>
oO
<

Dolomites form-
ing in central
Australia

Palygorskite-
rich sediments

Drying up

of alkaline
lakes, semi-arid
weathering

Lack of
carbonaceous
pollen- bearing
seds. in
Central
Australia

Carbonaceous,
pollen —- bearing
seds. in many
parts of Central
Australia

Coarse
clastics,
flooding

Lignites, peat — |}

forming swamps
in Central
Aust.

Coarse clastics,
flooding

Quilty 1984
Truswell &
Harris 1982;
Van de Graaff
1977.

Central,
northern,

NW Aust.

with increasing
dry, anti -
cyclonic air
circulation

Continued
steepening of
temp. gradients,
intensified
circulation

Steepening of
temp. gradients
equator to pole;
increasing
intensity of
circulation

Temp. gradients,
pole to equator
low

Circulation
sluggish

Bowler, 1976;
Bowler, 1982;
Frakes &
Kemp 1973
Truswell &
Harris 1982.

140 - FRAKES & RICH

CHRONO-
STRATIGRAPHY

SERIES

MIOCENE

LOWER

OLIGOCENE
T

LOWER

| UPPER

EOCENE
MADOLE

LOWER

|

PALAE OCENE

{

30+

40

50fF

POLLEN & MACROPLANT MAJOR BOTANICAL |MAJOR ARID ZONE
RECORD EVENTS PLANT LOCS.

>__/ Open vegetation dominant

in arid regions
Chenopods, Eucalyptus, |; .i%j Brief expansion of rain-| Eyre Peninsula,S.A
; iss

an Arid floras become
established Lake Frome

a few rainforest forms forest in Cent. Aust. Western/N.S.W.

Nothofagus absent Decrease in diversity
Dacrydium suggests A and abundance of
some moisture aa Nothofagus

&
ro

oka

Acacia, Sie Ne
Casuarina A »* i
we TJ

Nothofagus, podocarps
abundant

Grass pollen in %'s

suggesting grasslands present
in interfluves

Lake Eyre and
Tarkarooloo
subbasins

Early radiation of
Eucalyptus
Leptospermaceae

(Etadunna and
Namba Fm)
Woomera,
S.A.

First Acacia
pollen (northern
migrant )

Rainforest trees

Reduction of

still abundant, 4 diversity of geile Kalina
but less diverse : 2 F asin,
: ie 8 rainforest SA
than in Eocene rr e, | A.
we l species
Wu Sa |

Glenn Florrie,
W. Aust.

More palaeobotanical
sites than any other
part of Cainozoic Wu

First grass

-?high rainfall? pollen Hale River, N.T.
Trees of rainforest arapheny, N.T.
affinities abundant; Marked increase in A Pan m.,

Nothofagus Nothofagus diversity.

abundant

Goat Paddock,
W. Aust.

First Myrtaceous
pollen

Gymnosperms dominant,
related to temperate
rainforest forms

(e.g. Dacrydium)
Angiosperms present, whose
relatives now in vale Hee ce
high rainfall a eas @iiaiii
(e.g. Proteaceae, § a te tgs
Winteraceae) “hz

Ayers Rock, N.T.
Lake Eyre Basin
(Eyre Fm.),S.A.

Bint, 1981; Truswell & Harris
Lange 1978, 1982; 1982; Wells &
Martin 1973, 1978 Callen 1986.
Truswell &

Harris 1982.

PALAEOCLIMATE & PALAEOGEOGRAPHY OF AUSTRALIA - 141

VERTEBRATE FOSSIL RECORD
in present arid zone)

\\=
in

(uolseqqns oojooue ye 1) sulydjog prejsopqeyy

—

(aepuny)
s}uspoy

Spl}Seyosojed

|
Splyucpo}odig

splodoioeyw

SPp1010}0d

Genetic Diversity
of some Marsupial
Families

Familial
Diversity of
Marsupials

eepiyouAyoyyUO

— sooBiuiels

——————S SBD Oo goa aoacs

AHdVYDILVELS
-ONOYHD

ca co [ nD | es [ comes 9 ees 9 a IC — oS eS — i
no
2
Splu}!UIOWO01G oO
OLE
cq fo)
D>OH
Odvnw
o Sd|I|POI0ID
qQocoocojcontac 7f—Cc SS. SS ee

[ee [cos as fee 9 ces 9 es ee I ee Ol ees 9 ees 9 eee Bee 9 egy f cee)

ysijHun | yuopoyeiey

at Hi) n st (eve seaenas ews n n ae axes
° " 2 6 S g 8 8 $ ° 8 8 g 2
7 7
° | n | 1 W3ddN TIOGIW y3M07 Wd3ddN yw3MO1 | ¥3ddN | JIGGIN | 307 Y3dd/N Ww3M01
w | | 1 Lava

a 4

: S| 3Na9 AN3Z00IN 3N3909110 3N3900 43 3N300 3V Wd
O

Archer & Clayton 1984
Patterson & Rich 1987;

Rich, et al. 1987;

Rich & Thompson 1982;
Wells & Callen 1986.

142 - FRAKES & RICH

CHRONO
STRATIGRAPHY

CENE

MIOCENE

OLIGOCENE

AL AE OCE NE

MIDOLE | UPPER jejyu

LOWER

UPPER

MIDDLE

an |

WE

urren | LO"

LOWER

30+

50+

MAJOR ARID ZONE TERR. {MAJOR ARID ZONE ECOLOGICAL
VERT. INDICATORS VERTEBRATELOCS SUMMATION

Lake Eyre and
Tarkarooloo
subbasins.

Intense episodes of
aridity

Increased aridity &
increased seasonality
with oscillations

Fluctuating conditions
Humid at beginning &
middle of Epoch
Increasing aridity in
younging direction
Central Aust. lakes
occasionally drying out
Loss of Nothofagus,
rainforest elements
Expansion of grasslands ,
xeromorphic vegetation

Extinction of megafauna,
flamingoes, palaelodids ; local
loss of lungfish & crocodiles;

grazing forms dominate.

|

Lungfish, crocodiles,
a variety of
waterbirds, water
dwelling and arboreal
mammals (mainly
marsupials),
browsing

Alcoota, N.T.

Bullock Creek, N.T.
Riversleigh, Qld.
Lake Eyre and
Tarkarooloo
subbasins

Decrease in precipitation
Rainforest still dominant
Lack of carbonaceous seds.
indicate increasing aridity
Cooling at beginning

of Epoch

Humid over most of
Australia

Warmer than at present,
but cooling from beginning
to end of Epoch.

Some arid pockets in
Central Australia,
indicated by
grasslands

Humid, well watered
environments

Peat-forming swamps
abundant

Eucalyptus [f:] Browsers No long term aridity

Grazers

Chenopods

Myrtaceae
Nothofagus
Acacia

Grasses “RS Ferns

PALAEOCLIMATE & PALAEOGEOGRAPHY OF AUSTRALIA - 143

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HAIG, D.W., 1979. Global distribution patterns for mid-Cretaceous foraminiferids. J. Foram. Res. 9: 29-40.

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IDNURM, M. & COOK, P.J., 1980. Palaeomagnetism of beach ridges in South Australia and the Milakovitch
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KEAST, A., 1981. Ecological biogeography of Australia. Junk, The Hague.

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PALAEOCLIMATE & PALAEOGEOGRAPHY OF AUSTRALIA - 145

VEEVERS, J.J., 1984. Phanerozoic Earth History of Australia. Clarendon Press, Oxford.

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WOPENER, H., CALLEN, R. & HARRIS, W.K., 1974. The lower Tertiary Eyre Formation of the southwestem
Great Artesian Basin. J. geol. Soc. Aust.21: 17-52.

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146 - FRAKES & RICH

> EAH

Hipposideros (Brachipposideros) nooraleebus (left) and the less common megadermatid, a False Vampire
Bat (right) are forms found at the rich Oligo-Miocene sites on Riversleigh Homestead in northwestern
Queensland. (From Rich & van Tets 1985, with permission of The Museum of Victoria.)

CHAPTER 5

AN INTRODUCTION TO THE
LITERATURE OF
PALAEONTOLOGY WITH
REFERENCE TO THE
FOSSIL VERTEBRATES OF
AUSTRALASIA.

Marta Chiba!

IN(ROGUCHON:, oc hs elie Anat Dude old oh tee cease gnaveetane 148

A Brief History of the Literature of
Palacontology..ic...t20. hoch aceeeseel nag sigeeeee 148

An Outline of the History and Growth of
an Australian Literature of

PalacOntOlO Py: ns sasies.cvueeedhspahay Veblen sit 150
The Literature of Fossil Vertebrates of
AUSULalaSiQ...... cee ecceeeeeeeee ceeeeeeeeeeeeeees 151
Searching the Older Literature of
Palacontology........sceseeeseeeeeeeseeeenee sense 153
Biblio sraphiesys ctx: seasiens ehty of Sivaghea baa enanes sae dees 153
The Use and Structure of Subject
VAtEPAtre sco... .tsige teecve teseeneatyonpes Da geeblane 155
Literature of Palacontology .........ceceeeeesee eee ee ees 157
Primary Publications in
Pala OntOlOPy...osicecscesssss odaadeel cava 'aes voe'es 157
Secondary Publications in
PalacOntOlogy -. on ccieveeleessiden dedesevaeeeteles 159
Tertiary Publications in
Palaeontology isis. csstvscnecavwsesas leeeenetetds 163
Hints on Searching the Literature for
AATOFMALOM ss... cancer sea dnae Seay teas Sasaeet tenets 165
Computerized Information Services
Covering Palacontology.......ceeeee 168
RGICRNCES Ts. di snhanhs agehlceseg rales torec edb veyetele sone: 169

1 Hargrave Library, Monash University, Clayton, Victoria 3168, Australia.

148 - CHIBA

INTRODUCTION

This chapter gives a brief outline of the history, structure and use of the literature of
palacontology, with reference to information sources on the fossil vertebrates of Australasia.
The aim is to assist the reader in searching for a specific item of information or embarking on
a comprehensive review of the literature on the subject.

The volume of literature on palaeontology grows more slowly than other scientific
literatures. Unlike Physical Sciences or Technology, where information dates quickly and
publications are superceded by more recent ones within five to ten years, information on fossils
remains relevant for a long time. The literature of palaeontology is characterized on the one
hand by the large, cumulative reference works, and on the other, by the multitude of reports
published by geological surveys, government organizations, museums, research institutes,
commercial firms and just about anyone interested in fossils. The problem facing the student
or researcher is how to find an efficient approach to retrieve relevant information from the
large, cumulative, international and fragmented literature on the subject. An understanding of
the nature of the discipline and how it exerts an influence on the structure of the literature and
on the pattern of publications should be helpful to the reader in the choice of reference or serial
publications.

During the last decade, an increasing number of computerized information services appeared
with a generally selective coverage of palaeontology. These computerized information services
offer easy, online access to the recent literature of palaeontology. Some of the databases are
national, others are intemational in their scope and coverage of scientific publications, however
none are devoted exclusively to palaeontology.

Despite the lack of a specialist information database on palaeontology, many Zoology,
Earth and Life Sciences information databases are useful for identifying key publications,
important authors, conferences or institutions that carry on and report research on fossils.
Computerized information services having a substantial coverage of palaeontology, especially
literature on vertebrate palaecontology of Australasia, will be reviewed.

A BRIEF HISTORY OF THE LITERATURE OF
PALAEONTOLOGY

It is not sensible to separate the history of the literature of a subject from the history of its
development. The literature represents the record of achievement in a field of study. Progress
made in the state of the art is indicated by the time span and the volume of literature on the
subject,

One of the daunting prospects facing the researcher is having to decide on the retrospective
coverage of the older literature on the subject. Whilst many of the important older
publications are cited in recent works, the serious researcher cannot afford to miss older
publications relevant to the problem, but overlooked in recent works.

The development of palacontology as a separate branch of science with its own subject
literature dates back to the first half of the nineteenth century. It was preceded by three
centuries of debate on fossils, arguing whether they were the remains of plants and animals,
minerals, or the illusions of nature. The literature of natural sciences could be regarded as the
parent discipline of palaeontology. Leonardo da Vinci (1452-1519), working as an engineer on
the construction of canals in Italy, noted fossil remains in rocks and suggested that they were
marine organisms that had once lived there. Robert Hooke (1635-1703), the English physicist
and mathematician was the first to suggest using fossils as a record of the Earth's history.

AUSTRALIAN VERTEBRATE PALAEONTOLOGY LITERATURE - 149

According to Zittel (1901), a well known palaeontologist and historian of the discipline, the
first famous masterpiece of palaeontological literature was Brocchi's Conchyliologia fossile
subapeninnium published in Milan in 1814, The work contained an accurate description of the
occurance and distribution of Tertiary Mollusca in Italy and a chapter on land mammals, whales
and fish. Another notable early contribution to palaeontology was William Smith's unfinished
work on Strata Identification By Organized Fossils (1816-19).

Sarjeant & Harvey (1973), described the early history of palaeontological literature and
regarded the works of Lamark (1744-1829) on vertebrates, Brongiart (1801-1879) on fossil
plants, and Ehrenbert (1795-1876) on microscopic fossils as pioneering works, laying the
foundation for the development of palaeontology as a separate discipline with its own specialist
publications. A landmark in the study of fossil vertebrates was Sir Richard Owen's
Palaeontology (1860). The work provided an excellent general survey of vertebrates, but
treated invertebrates less fully. The early literature of palacontology was often influenced by
the home discipline or the special interest of the author. Thus, palaeontology was treated
variously as part of botany, zoology, comparative anatomy or frequently as a branch of
geology.

The study of fossils was given the name palaeontology by two eminent authors: Ducrotay
de Blainville and Fisher von Waldheim in 1834 (Zittel 1901). The term was rapidly adopted in
France and in England; in Germany, however, the older name of petrefaktenkunde or
petretaktologie was used for awhile. The first German university to establish separate chairs in
palaeontology and geology was in Munich in 1843.

An all important event marking the beginning of the publication of specialist serials
devoted to palaeontology was the establishment of the Palaeontological Society in London in
1847. The Society's aim was to describe the complete stratigraphical series of British fossils.
To promote its aim, the Society immediately embarked on the publication of a monograph
series. The first volume of the monograph was published in 1847, and it carried a list of
authors and titles of works on British fossils, to be published in later monographs between
1847-1861. Many eminent nineteenth century palacontologists published the results of their
research in the monographs of the Palaeontological Society: authors like Sir Richard Owen,
H. Milne-Edwards, E. Forbes, T. Davidson and H. Woodward. The papers published in the
monographs were carefully researched. The first paper published in the series on Tertiary Crag
Mollusca from Britain, carried a bibliography of 137 references. Many of the papers cited were
from the publications of learned societies, museums, academies, and magazines of natural
history. Papers published in the Journal of the Linnean Society, the Zoological Record of the
Zoological Society of London and in the Philosophical Transactions of the Royal Society of
London were among the most frequently cited references, and these journals could be regarded
with some justification as important information sources in the early literature of
palaeontology. Publications from other countries, particularly from France, Germany and
Belgium were also represented among the list of references cited in the early volumes of the
Palaeontological Society's monographs.

By the end of the nineteenth and the early part of the twentieth century, societies with a
special interest in palaeontology were formed in other European countries and in the United
States. Their purpose was to promote the publication and exchange of information on
palaeontology. These societies performed the function of an invisible college for
palaeontologists, furthering advances in the new discipline.

The first periodical offering a comprehensive coverage of palacontology was
Palaeontolgraphica, edited by W. Dunker and H. von Meyer, published by Cassel in Germany
for the first time in 1846. Other periodicals devoted to palaeontology started to appear in the
late nineteenth and the early twentieth century. The Bulletin of American Paleontology
commenced publication in 1895, Annales de Palaeontologie (Paris) in 1906 and Journal of
Paleontology (Tulsa, Ok.) in 1927. Alcheringa, the Journal of the Association of Australasian

150 - CHIBA

Palaeontologists of the Geological Society of Australia, commenced publication in 1975,
much later than similar periodicals in Europe or North America. The majority of the specialist
periodical literature devoted to the coverage of palaeontology, or one of its subfields like
micropalaeontology or foraminifera, commenced publication in the second half of the twentieth
century. Despite the existence of a well defined specialist periodical literature of
palaeontology, many papers on fossils continue to be published in journals covering related or
broader fields than palacontology, reflecting the wide professional interest in the subject.

The nineteenth century saw the rise of palaeontology as a new discipline, and specialization
within the field of study. The theoretical foundation of the new discipline was laid by the
pioneer works of eminent scientists such as Lamark, Brongiart, Ehrenberg and Sir Richard
Owen. The literature of palacontology emerged as a specialist literature in the middle of the
nineteenth century, its development hastened by the formation of learned societies, museums of
natural history and by the establishment of separate chairs in palacontology at some
universities.

AN OUTLINE OF THE HISTORY AND GROWTH OF AN
AUSTRALIAN LITERATURE OF PALAEONTOLOGY.

The development of palacontology in Australia, and the growth of an Australian literature
on fossils was late in comparison to other countries. The history of palacontological studies
in Australia had a typically colonial beginning, showing a strong dependence on the work of
European, particularly English palaeontologists (Rich et al. 1982, Vallance 1978). Fossils
collected by the French (Nicholas Baudin, 1800-1804) and the English (Matthew Flinders
1800-1804) expeditions to Australia, and later by the various explorers of the interior of the
continent, were sent to England for identification and description by eminent palaeontologists
such as Sir Richard Owen. In this pioneer stage of Australian palaeontology, there were only
isolated examples of work on Australian fossils by resident palacontologists published in
Australian journals such as the Tasmanian Journal of Natural Sciences, or the Proceedings of
the Royal Society, Hobart, and even in some obscure Sydney newspapers. The mainstream of
literature on Australian fossils was published in England up until the late nineteenth century.

The infrastructure needed for the establishment of an Australian base for palacontology and
for the development of an Australian literature on the subject, was created in the latter part of
the nineteenth century by the formation of scientific societies (the Royal Society in Hobart
founded in 1841, followed by the Royal Societies of New South Wales, Victoria, South
Australia and others), by the establishment of museums (the National Museum of Victoria, the
Queensland Museum in Brisbane, the South Australian Muscum in Adelaide, etc.) and by the
setting up of the state geological surveys. These societies and institutions had a firm interest
in the study of Australian habitat and resources. To publicize their activities they issued serials
and pamphlets giving local scientists an accessible outlet for publication. The proceedings and
transactions of the local scientific societies such as the Proceedings of the Royal Society of
Victoria quickly became, and to this date remains, an important forum for the publishing of
papers on Australian fossils. The museums were more than a repository for the fossil fauna
and flora of Australia. They had scientifically trained staff eager to study and ready to
contribute, generally to publications issued by their museums, invariably named: records,
circulars, or memoirs. An outstanding example of an early Australian publication on
palaeontology was Frederick McCoy's (1817-1899) Prodromus of the Palaeontology of
Victoria, issued in seven parts between 1874 and 1882 by the National Museum of Victoria.
McCoy, in addition to his involvement with the National Museum of Victoria, also held the
position of Professor of Natural Sciences at the University of Melbourne, between 1854 and
1899. The state geological surveys were especially important in the growth of an Australian

AUSTRALIAN VERTEBRATE PALAEONTOLOGY LITERATURE - 151

literature of palaeontology, as they accepted the responsibility for the systematic study,
recording and publishing of information on the state's natural resources in their bulletins,
occasional papers and special publications, an activity which they continue to this date. Brown
(1946) regards the mid-nineteenth century as the turning point in the history of Australian
palaeontology which culminated in the publication of the Geology and Palaeontology of
Queensland and New Guinea by Jack and Etheridge in 1892.

University-based research in palaeontology by scientists like Frederic McCoy, Ralph Tate
(1840-1901), who was appointed to a foundation chair at the University of Adelaide in 1874,
and A.M. Thomson, who held the chair of geology at Sydney University from 1896, gave an
additional impetus to the development of the subject and to the growth of an Australian
literature of palaeontology.

An outstanding palaeontologist who made a significant contribution to the early literature
on Australian fossils was Robert Etheridge Jr. (1847-1926), who was appointed
palaeontologist to the N.S.W. Geological Survey and became the director of the Australian
Museum, Sydney in 1895, a position which he held until his death, By the end of the
nineteenth century, Robert Etheridge Jr. and Professor Ralph Tate together, through their
extensive studies of Australian fossils and their numerous publications, brought palaeontology
in Australia out of its pioneer stage and helped to establish an Australian literature of
palaeontology.

By the beginning of the twentieth century there were many Australian publications, mainly
serials, that carried information on Australian fossils. The first Australian textbook of
palaeontology was F, Chapman's Australian Fossils, published in Melbourne by G. Robertson
in 1914, The twentieth century saw the growth and proliferation of Australian publications in
palaeontology and the increasing concentration of literature on Australian fossils in Australian
publications. Whilst Australian palaeontologists, like palaeontologists in other countries,
may publish the results of their research overseas, the core literature, i.e. most important, on
Australian fossils is concentrated in Australian publications. A quantitative analysis of titles
included in the Annotated Bibliography of the Palaeontology of Western Australia, 1814-1974,
compiled by P. Quilty and published by the Geological Survey of Western Australia in 1975,
confirmed the continuing importance to palaeontology of serials published by the state
geological surveys, the Bureau of Mineral Resources and the various royal societies in each
State.

THE LITERATURE OF FOSSIL VERTEBRATES OF
AUSTRALASIA

In scientific disciplines, English language writers read, cite and contribute to English
language publications. The generalization holds particularly well for palaeontology. A casual
perusal of references cited by English language papers on palaeontology reveals that over
ninety percent of the literature cited is in the English language, with preference for local and
national publications. Despite the relative youth of an Australian indigenous literature on
vertebrate palaeontology, the core literature on the subject is published in Australia, especially
in serials. To test the assertion, the Australian Earth Sciences Information Database (AESIS)
was searched online for papers published between 1980 and 1985 on the fossil vertebrates of
Australasia. The database covers both Australian and overseas publications. The search
retrieved two hundred papers on the subject. Of the two hundred papers retrieved, twenty-seven
or thirteen point five percent were overseas publications, all but one published in English.
Serial publications accounted for sixty-one percent of the total. Australian serials yielded forty-
eight percent of the papers retrieved. The following serials published papers on fossil

152 - CHIBA

vertebrates of Australasia between 1980 and 1985. The serial titles are ranked in order of the
number of papers published on the subject during the period covered by the search (Table 1).

See

Table 1. Journals that Published Papers on Australasian Fossil Vertebrates Between 1980-
1985. (Source: AESIS Search 1980-1985).

Title Number of Pages

Alcheringa 25

Queensland Museum. Memoirs 20

Victoria. National Museum. Memoirs 10

Journal of Paleontology

Palaeontology

Royal Society of Western Australia. Journal

Nature

Queen Victoria Museum. (Launceston, Tasmania) Records

Australia. Bureau of Mineral Resources, Geology and Geophysics.
B.M.R. Journal of Australian Geology and Geophysics

Australia. Bureau of Mineral Resources, Geology and Geophysics.
Bulletin

Royal Society of Tasmania. Papers and Proceedings

Search

South Australia. Museum. Records

Australian Natural History

Australian Museum (Sydney). Records

Australia. Bureau of Mineral Resources, Geology and Geophysics.
Records

Fieldiana: Geology

Geological Society of Australia. Journal

Linnean Society of New South Wales. Proceedings

Royal Society of South Australia. Transactions

American Museum of Natural History. Bulletin

Postilla

Journal of Biogeography

Journal of Vertebrate Paleontology

Nomen Nudum

Palaeogeography, Palaeoclimatology, Palaeoecology

Paleaontological Association of London. Special Papers in
Palaeontology

Royal Society of New South Wales. Journal & Proceedings

Royal Society of Victoria. Proceedings

Science

Scottish Journal of Geology

South Australian Naturalist

Western Australia. Geological Survey. Records

Western Australian Museum. Records

WwW PAHMAN

NNWW WwW

RRR RPE NNN N ND bd

— eR

Total Number of Journal Articles 122

Twelve monographs yielded sixty-two titles representing thirty-one percent of the total number
of papers retrieved. Fifty of the monographic papers come from just two Australian books:

AUSTRALIAN VERTEBRATE PALAEONTOLOGY LITERATURE - 153

Archer, M. & Clayton, G., eds., Vertebrate Zoogeography and Evolution
in Australasia. Hesperian Press, Carlisle, Western Australia. 1984. 1203 pp.
figures, tables, references.

Rich, P.V. & Thompson, E.M., eds., The Fossil Vertebrate Record of
Australasia. Monash University Offset Printing Unit, Clayton, Victoria. 1982.
xxiii, 759 pp. appendices, figures, maps, references, tables.

Only one of the twelve monographs was published overseas, the rest in Australia. The
remaining sixteen papers consisted of six theses and ten unpublished reports.
The online search of AESIS for the period 1980-1985 confirmed the concentration of the
literature on fossil vertebrates of Australasia in Australian publications and the predominance
of journal articles. A subsequent search of AESIS on the same topic covering 1988-89 did not
reveal a significant change in the pattern of dispersal of journal articles in Australian and
overseas serials. Alcheringa was the most productive serial, followed by the Queensland
Museum Memoirs and the Western Australian Museum Records

SEARCHING THE OLDER LITERATURE OF
PALAEONTOLOGY.

There is no easy approach to searching for titles published before the twentieth century. A
major problem encountered is the lack of authoritative works providing a comprehensive
coverage and an adequate subject approach to the older literature of palaeontology.

There are a number of reference publications with substantial coverage of older publications
on palaeontology. Readers wishing to locate a list of reference publications useful for
searching the older literature are advised to consult the guide books to the literature of earth
sciences (e.g. Wood 1973) or to geology (e.g. Pearl 1961) and to read the chapters describing
bibliographic access to the older literature of palaeontology. Encyclopaedic works such as
scholarly treatise designed to summarize the documented knowledge on a subject are also useful
sources for locating titles published before the twentieth century, There are a number of
scholarly treatise covering palacontology. They are described later in this chapter, as they offer
organized bibliographic access to the recent as well as to the older literature.

In the following section, reference works offering substantial coverage of the older literature
of palaeontology are described briefly.

BIBLIOGRAPHIES

Bibliographies offer an important organized approach to older publications. The name
bibliography was first used to describe a list of books on a subject. Modern usage of the term
describes reference works designed to provide an organized approach by authors, titles or
subjects to the literature. Bibliographies appear in many forms. Some are published as books,
others as special issues in journals or as part of review articles published in serials. Generally,
major bibliographic works are published in several volumes, over a period of time.

One of the oldest, most important bibliographic works covering the older literature of
palaeontology is the Repetorium Commentatorium a Societatibus Litteraris Editarum,
compiled by D.J. Ruess and published by Dietrich in sixteen volumes covering papers
published by scientific societies between 1665 and 1800. It is useful for locating articles on
palaeontology published in the journals of the learned societies such as the Royal Society of
London and the Linnean Society.

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The most important bibliography covering the nineteenth century scientific literature,
including palaeontology, is the Catalogue of Scientific Papers, 1800-1900, compiled by the
Royal Society, London and published in four series between 1867-1925. The catalogue is an
author index to scientific papers published mostly in Europe, during the nineteenth century.
The Catalogue covers articles which appeared in some fifteen hundred journals between 1800
and 1900. It offers no subject approach to palaeontology. The researcher must know the
names of authors before embarking on a search for nineteenth century papers on fossils.

A continuation of the Royal Society's Catalogue is the International Catalogue of
Scientific Literature 1900-1914, published by the Royal Society of London on behalf of the
International Council of Scientific Workers. The Catalogue is arranged under broad subjects,
and Section K is Palaeontology. There are also annual author and subject indexes to this work.

One of the most important encyclopaedic reference works that carry substantial
bibliographies of older literature is the Dictionary of Scientific Biography. published in New
York by Scribner in 1974 in sixteen volumes. This is an authoritative work which was
sponsored by the American Council of Learned Societies. The Dictionary carries articles on
the life and work of eminent scientists, including palaeontologists, from the beginning of time
to the early part of the twentieth century. Each article concludes with a bibliography of works
by, and a bibliography of works about, the life and contribution of the particular scientist.
Volume sixteen is an index, offering a detailed subject approach. For example, under
palaeontology the index lists works chronologically from the Renaissance to the early
twentieth century, in addition to works arranged under aspects or subfields of palaeontology.
This is a useful work for discovering key papers, particularly on the early history of
palaeontology.

J.C. Poggendorf's Bibliographisch-Literarisches Handworterbuch zur Geschichte der Exacten
Wissenschaften, is a multi-volume Bio-bibliographic reference work on the life and work of
scientists from early time to 1953. For each author listed, the biographical entry is followed
by a brief bibliography of their more important works. The Handworterbuch was published in
four series, each covering a different period of time. The first series was published in Leipzig
by Barth in 1863. The Handworterbuch is useful for tracing early contributions to
palaeontology, particularly those written in foreign languages.

The reader searching for papers on the development of palaeontology in general or in a
specific country, is well advised to consult W.A.S. Sarjeant's Geologists and the History of
Geology: An International Bibliography from the Origins to 1978 (New York, Arno Press,
1978). The first volume of this five volume bibliography carries references to papers on the
history of palaeontology and its subfields, in different countries, including Australia.

There are many special bibliographies of palaeontology. Most include references to early
contributions to the subject. For example, the Bibliography of Fossil Vertebrates Exclusive of
North America, 1509-1927 (New York, Geological Society of America, 1962) offers an author
and a subject approach to the world literature on fossil vertebrates. Early contributions to
vertebrate palaeontology in the United States can be located in the Bibliography and Catalogue
of Fossil Vertebrates of North America, (1902) published as Bulletin 179 by the United States
Geological Survey. The more recent literature on fossil vertebrates can be searched in CL.
Camp et al. Bibliography of Fossil Vertebrates, a quinquennial bibliography of the world
literature on the subject from 1928, published in New York by the Geological Society of
America between 1938 and 1973, as special issues to the GSA Special Paper and the GSA
Memoir series. The publication of the Bibliography of Fossil Vertebrates was continued on an
annual basis from 1978 by the American Geological Institute, the Society of Vertebrate
Palaeontology, and by the Museum of Palaeontology of the University of California at
Berkeley. A single, retrospective volume covers the literature published between 1973 and
1977.

There are a number of special bibliographies of palaeontology covering the Australian

AUSTRALIAN VERTEBRATE PALAEONTOLOGY LITERATURE - 155

literature. Generally, they are published by the state geological surveys in their bulletins or
reports or by the Bureau of Mineral Resources in its Bulletin. The first comprehensive
bibliography of Western Australian palacontology compiled by Patrick Quilty was published
as Report number three (1975) by the Western Australian Geological Survey. The
Bibliography covered publications on Wester Australian fossils from 1853 to 1973.

Special bibliographies offering access to the carly literature of palacontology sometimes
appears as journal articles or as chapters in books. I. Crespin's A Bibliography of Australian
Foraminifera (1975) was published in the journal Micropalaeontology (volume one. pp. 172-
186), whilst a chapter on the Literature References to the Fossil Terrestrial Mammals of
Australia and New Guinea is included in this volume.

Bibliographies on palacontology may be cited by abstracting or indexing services and in
reference books, discussed later under secondary and tertiary publications. The only abstracting
service going back far enough to cover the older literature of a palacontology is Zoological
Record, published by the Zoological Society of London since 1864. Zoological Record offers
an author, subject, and a systematic approach to ninctcenth century publications on fossil
fauna, including Australian fossils under the name of the genera or subgencra.

A comprehensive coverage of the older literature of palacontology requires patience, an
appreciation of the history of the discipline and above all, a careful planning of search strategy.
Knowledge of important people and epoch making papers will help to provide the key to this
early literature. Important early contributions are cited in modern works and in citation
indexes.

The method of approach to searching the carly literature of palacontology should be guided
by the purpose of the search and by the available access to bibliographic resources. The writer
of an introductory paper may be excused for not paying adequate attention to the history of the
subject. The researcher, however, cannot afford to overlook important early contributions,
which could influence the approach to the problem or the interpretation of the findings. When
searching for early contributions on a subject, patience and perseverence are essential to ensure
success.

THE USE AND STRUCTURE OF SUBJECT LITERATURES

The literature of any subject may be divided into three broad categories on the basis of the
function and the type of information contained in the publications making up these categories.

Primary publications carry the latest published information on a subject. They represent a
vast pool of up-to-date information, growing at an exponential rate. The main type of
publications belonging to this category are journals, reports, conference papers, the varied
forms of contributions by institutions, socicties, organizations and government bodies and
theses. The function of primary publications is to report with the minimum of delay, research
work and new findings.

Secondary publications represent the first attempt to provide a systematic access to the
literature under authors, titles and subjects. Secondary publications provide bibliographic
access to different types of publications, such as journal articles, conference papers, research
reports and books. Secondary publications are represented by abstracting and indexing journals,
by review serials, bibliographies and current awarencss services. As the catalogue of a library
is the key to its collection, secondary publications serve a similar function for subject
literatures, as they offer a bibliographic approach to searching the literature for a specific tile
or author or for publications on a topic. In secondary publications, bibliographic entries are
sometimes accompanied by a brief indicative or information summary or abstract of the subject
matter covered in the item of literature cited by the service.

Abstracting journals assist the reader to identify relevant publications from the literature.
Review serials survey the advances made in a field of study during a stated period of time.

156 - CHIBA

Review articles are followed by references to important contributions. Articles in review
serials often evaluate contributions made to the subject.

Bibliographies present the literature on a subject, topic or country, or author in an
organized form. Their scope and coverage generally includes all types of publications in
different languages. Current awareness services carry contents pages of journals. Their
function is to alert the reader of the existence of the most recent papers on the subject
published in serials.

Generally, the function of secondary publications is to assist the reader in the search of the
literature in order to identify, select and locate the relevant information resources.

Tertiary publications are books, including reference books. Information in tertiary
publications is not as up-to-date as in the primary or secondary publications, but it is organized
and summarized to cater for different levels of understanding, ranging from the introductory to
the scholarly, informative level. Tertiary publications include general books, textbooks,
research monographs and reference books such as handbooks, dictionaries, encyclopaedias and
treatise. The function of tertiary publications is to introduce, explain and to summarize
knowledge.

The essential difference between general books, textbooks and research monographs is in
the treatment of the subject. General books describe even the most complex topic using terms
which a layperson can understand, Texbooks are designed to teach the student of the subject.
Research monographs are written by specialists for specialists. The treatment of the subject in
research monographs assumes formal education in the field of study. Books are written for a
well defined audience in mind. The reader is advised to search for books on the subject designed
to suit the desired level of understanding of terms, concepts and ideas.

Reference books serve to orient the reader and to introduce a topic. They are not meant to
be read from cover to cover. Information in reference works is organized in a systematic way
to facilitate the looking up of a definition, a formula, a concept, or is an introductory article on
a topic.

Reference works may be general or special, depending on whether they cover the whole field
of knowledge or one of its subsets, such as a discipline or a single subject. Some reference
works are intended for the interested layperson, whilst others serve the sophisticated
information need of the subject specialist. Handbooks for example, exist to provide in a
compact form, principles, data and tables for professional practice. The arrangement of
information in handbooks generally follows a careful classification of the field of coverage.
Handbooks are more important in the applied than in the pure sciences. Their function is to
furnish facts and principles. Dictionaries and encyclopaedias can be general or special. A
general dictionary gives the meaning, usage, spelling, etymology and the pronunciation of
words. Special dictionarics on the other hand, define and explain concepts, theories and terms.
General dictionaries are used in every-day communication, while special dictionaries are used in
scientific, technical or professional communication. Similarly, general encyclopaedias cover
the whole field of knowledge. Special encyclopaedias cover a branch of knowledge or a single
subject. The narrow field of coverage in specialist reference works allows the treatment of the
subject in considerable depth. Articles in special encyclopaedias present an overview of the
field, including its history, development and current status with illustrative examples, graphic
material and a short bibliography of supporting literature on the subject. General or special, an
encyclopaedic article is intended to augment knowledge. A typical encyclopaedic article starts
with a definition of the topic, indicating its relationship to other, related fields of study. A
good encyclopaedic article also contains the main key-words which describe the subject, to
assist the reader to define the topic and thus facilitate the search of the literature for additional
information.

Treatise are scholarly reference works that provide an authoritative and carefully documented
summary of the literature on a subject. The function of a treatise is to present a summary of

AUSTRALIAN VERTEBRATE PALAEONTOLOGY LITERATURE - 157

all the known facts, theories and principles, together with a complete bibliography of
important contributions to the field of study. The field of coverage in treatise type
publications is narrow, to allow the complete, in-depth treatment of topics by recognised
authorities working in well defined specialities. Information in treatise articles should be
adequate to evaluate scholarly contributions to the field at a given date. Treatise are
indispensable reference works to scholarly research.

Tertiary publications offer either a broad or an in-depth summary of the literature, and are
useful to introduce the reader to an unfamiliar topic or to provide background information for
serious research.

Fig. 1 illustrates the use and the structure of subject literatures. It shows how with the
passage of time the fast growing, up-to-date primary publications, are included in bibliographic
services or secondary publications, and finally in tertiary publications, which may introduce,
define, explain or summarize the documented literature on a subject.

THE LITERATURE OF PALAEONTOLOGY.

In this section, only key publications in palaeontology are described. Readers wishing to
gain a more detailed knowledge of publications in palaeontology are advised to refer to the
guidebooks to subject literature, including those listed in the bibliography, at the end of this
chapter.

PRIMARY PUBLICATIONS IN PALAEONTOLOGY.

The importance of the different types of primary publications varies with the differences in
social organization and communication patterns in the various disciplines. The most
important forms of primary publications in palaeontology are journals and reports.
Palaeontological studies are generally carried out by institutions, museums, government bodies
and commercial organizations applying palaeontological techniques in exploration work.
Among the government organizations, geological surveys and departments of mines make
significant contributions to the study of fossils.

Accounts of palaeontological investigations are usually written up in journal articles or in
papers presented at professional meetings. A substantial part of palaeontological
investigations is published in the report literature. Reports, particularly the ones by private
companies, sometimes remain internal reports and can be identified only in the secondary
services covering the literature of palacontology. The researcher generally relies on the
‘invisible college’ (colleagues working in the same ficld) for information about unpublished
reports and research in progress.

The report literature on palaeontology is vast and fragmented. There are so many reports
issued by such a wide spectrum of sources, that it would be unwise to single out some and not
mention others. Published reports can be identified in the secondary services such as
abstracting and indexing journals, under the subject of the paper, or the name of the author.
Unfortunately, bibliographic access to unpublished reports is limited. Annual reports of
organizations and house organs (publications designed to publicize the activities of firms and
organizations) are valuable sources of information on internal reports and on research in
progress. Similarly, newsletters issued by learned societies and research organizations can be
useful to identify internal reports and unpublished conference papers.

158 - CHIBA

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AUSTRALIAN VERTEBRATE PALAEONTOLOGY LITERATURE - 159

The periodical literature of palacontology is compact in comparison to other disciplines.
There are a relatively small number of specialist periodicals devoted to the coverage of
palaeontology. However, articles on palacontological topics also appear in journals covering
related fields in the earth and life sciences and in journals issued by scientific socicties. The
27th edition of Ulrich's International Periodicals Directory (New York, Bowker Co., 1988-
1989), a worldwide directory of current periodicals, listed ninety-nine thousand four hundred and
fifty journal titles under broad subject headings. Under 'Palacontology' there were ninety-three
specialist periodicals devoted to the subject with a further fifty-six references to related serials.
The majority of current journals devoted to palaecontology were published in the English
CNBUagS. Foreign language journals were well represented, however, by French, Russian and

erman.

In 1980, the oldest primary journal still current was Palaeontographica (v.1- ; 1846-)
published by the Schweizerbartsche Verlagsbuchhandlung in Stuttgart. The Australian journal
of palaeontology, Alcheringa, (v.1- ; 1975-), is one of the most recent primary periodicals in
palaeontology and the only recent journal with a general coverage of the field. Most of the
recent journals in palaeontology are specialist journals devoted to the coverage of a subfield of
palacontology. Prestigious older journals listed included Journal of Paleontology (v.1- ;
1927-) issued by the Society of Economic Palaeontologists and Mineralogists in Tulsa,
Oklahoma, Palaeontologische Zeitschrift v.1- ; 1914-) issued by the Schweizerbartsche
Verlagsbuchhandlung in Stuttgart, West Germany, and the Bulletins of American Paleontology
(v.1- ; 1895-) issued by the Palaeontological Research Institution in Ithaca, New York.
Generally, primary journals issued by national academies, leamed socicti¢s and by international
publishing houses are the most cited and prestigious journals in palacontology.

Papers presented at professional meetings when published, appear in a collected form. Such
publications are generally called Proceedings, Colloquium, Conference, Symposium, etc.
followed by the title of the meeting. Some papers presented at meetings of professional and
learned societies are not published in proceedings form. They may be identified in the relevant
abstracting and indexing journals as conference papers under subject headings and the name of
the author. Preprints of unpublished conference papers can be obtained from the sponsoring
body of the meeting or from the author of the paper. There are fewer conference papers than
journal articles or reports in the primary literature of palaeontology.

Theses are also primary literature, as they report original research, carried out under
supervision al tertiary institutions. Dissertations Abstracts International (v.1- ; 1969-)
published by the University Microfilm Corporation in Ann Arbor, Michigan offers a good
coverage of doctoral theses in palacontology. Section B: Sciences and Engineering and Section
C: European Abstracts of the Dissertations Abstracts are relevant to palacontological research.
Theses listed in this service are available in microform or as a paper copy from the University
Microfilms Corporation.

The primary literature of palaeontology is characterized by a steadily growing number of
specialist journals devoted to a subfield of palacontology, and by the vast, fragmented,
international and rapidly growing report literature, which presents considerable problems for the
researcher trying to discover the existence of relevant reports.

SECONDARY PUBLICATIONS IN PALAEONTOLOGY

The reader wishing to scan the literature for current publications on palacontology, or to
carry out a search, going back some years covering the literature of the subject, has a choice of
using specialist secondary publications devoted exclusively to palacontology or secondary
services with a broad subject coverage including palacontology.

The most comprehensive English language index to earth sciences publications with an

160 - CHIBA

excellent world-wide coverage of palaeontological publications is the Bibliography and Index of
Geology, published in twelve monthly issues with annual cumulations, since 1969. The
Bibliography succeeded the Bibliography of North American Geology, a U.S. Geological
Survey publication, which covers literature from 1785 to 1970, and the Bibliography and Index
of Geology Exclusive of North America, published by the Geological Society of America
between 1933 and 1968. Between 1969 and 1978 the Bibliography and Index of Geology was
published by the Geological Society of America and from 1979 to date by the American
Geological Institute.

The monthly issues of the Bibliography are divided into four sections: Serials, Fields of
Interest, Subject Index and Author Index. The main section in the monthly issues is the
‘Fields of Interest’, which provide a subject approach to the literature. Under each subject
category, citations are grouped into books, meetings, theses, maps and papers. The indexes
refer to the citations through the consecutive entry numbers. The ‘Field of Interest’ 08:
Palaeontology, general (studies on fossil plants and animals, concepts, life origin,
applications, methods etc.) is most useful for identifying information on fossil vertebrates.

For each citation listed under ‘Fields of Interest’, there are an average, 3.4 entries in the
Subject Index. The Author Index contains the names of personal authors, corporate authors and
editors; the Serials section contains information on serials cited in the Bibliography.

The annual cumulation of the Bibliography, parts 1 and 2, is an alphabetical listing under
authors of bibliographic entries included in the monthly issues. There is no cumulation by
Fields of Interest. The cumulative index, parts 3 and 4 consists of the cumulation of the
monthly subject indexes.

The following sample entry shows the information provided in the Cumulative
Bibliography under authors:

Smith, Meredith J. Small fossil vertebrates from Victoria
Cave, Naracoorte, South Australia: II., Peramelidac.
Thylacinidae and Dasyuridae (Marsupialia): R. Soc. S.
Aust., Trans., Vol. 96, Part 3, p. 125-137, illus., 1972.

Dec 16 E72-39583

The publication was indexed in the Cumulative Index under the following headings:

South Australia — paleontology

Mammalia, Pleistocene, Naracoorte, Victoria Cave
(Smith, Meredith J.) Dec 16 E72-39583

Mammalia Marsupialia

Pleistocene, South Australia, Naracoorte, Victoria
Cave, bones, morphology, taxonomy,
Peramelidae, Thylacinidae, Dasyuridae
(Smith, Meredith J.) Dec 16 E72-39583

The monthly issues of the bibliography fulfill the role of a current awareness service. The
annual cumulations provide bibliographic access to the literature under indexing terms which

AUSTRALIAN VERTEBRATE PALAEONTOLOGY LITERATURE - 161

describe the subject matter of documents and by authors, and there is a User Guide to the
Bibliography and Index of Geology, edited by J. Mulvihill which was issued by the American
Geological Institute in 1982. The American Geological Institute published two special
bibliographies for palacontologists from its Bibliography. The Bibliography of Fossil
Vertebrates is issued annually from 1978- in cooperation with vertebrate palaeontologists
working at the Museum of Palacontology of the University of California at Berkeley.

The occasional use of foreign language abstracting and indexing services is necessary as no
English language service provides a worldwide coverage of the palaeontology literature. The
following foreign language abstracting and indexing journals are useful to palaeontologists:

Bulletin Signaletique - Bibliography des Sciences de la Terre: Section 227, Cahier H:
Paleontologie (Bureau des Recherches Geologiques et Minieres et Centre national de la
Recherche Scientific, Paris), This is a monthly publication with short abstracts arranged
according to a special subject classification scheme. Abstracts are in French, but titles are
given in their original language. There is a monthly subject index and annual author and
subject indexes. The emphasis is on French and continental European, including Russian
language publications. Bulletin Signaletique came into existence in 1956 as a continuation of

the Bulletin Analytique.

For German language publications in palacontology the specialist service is the Zentralblatt
fur Geologie Teil 2: Palaeontologie an irregular (about seven issues per year) publication
issued since 1950 by the Schweizerbartsche Verlagsbuchhandlung in Stuttgart.

For Russian and Eastern European publications in palaeontology, the most comprehensive
abstracting service is Referativnyi Zhurnal: Geologiya (Akademiya Nauk SSSR, Moscow,

1956-). In the monthly issues of this service, abstracts are arranged in a subject sequence with
an author index. Although the abstracts are in Russian, the full bibliographic details are given
in the original language of the paper. There are annual author (avtorski) and subject
(predmetnyi) indexes to the abstracts. The subject index is in Russian Cyrillic alphabet, but
the author index is in two sequences, a Cyrillic and a Roman alphabet, according to the
language of the paper.

There are a number of English language abstracting and indexing journals useful to
palacontologists. Zoological Record (Zoological Society of London, 1864-) and Biological
Abstracts (Biosciences Information Service and Biological Abstracts, 1927-) have a good
coverage of papers of palacontological interest. Science Citation Index (1955-) published in
Philadelphia by the Institute for Scientific Inforraation has a worldwide coverage of scientific
publications. The service cites papers on earth sciences, and also palaeontology. Provided the
researcher knows the bibliographic details of a key paper, the citation index permits the
retrieval of other papers cited by the key (source) paper. Science Citation Index consists of a
Citation Index, a Source Index and Permuterm Subject Index. The Source Index gives
bibliographic details of papers published in the journals scanned by the service. The Citation
Index consists of references cited in source items in alphabetical order, under the name of the
first author, The Permuterm Subject Index is an index using the words contained in the titles
of the papers listed in the Source Index,

The Australian literature of palacontology is covered by two secondary services published in
Australia. The Australian Earth Sciences Information System (1976-) AESIS is a national
information service which covers the Australian literature including published and unpublished
material in the earth sciences, including palacontology. The service appears quarterly in paper
form and cumulates annually, and later covering a number of years in a single alphabetical
sequence in microform, In the quarterly issues of AESIS, annotated bibliographic entries are
arranged under broad subject categories. Section 1430 is Palaeontology. Each quarterly issue
contains an Author, Subject, Locality Name, Stratigraphic name and Map Reference Index.
The following is a sample entry from AESIS quarterly, reproduced to illustrate the elements of
the bibliographic citation and the extent of indexing used by the service.

162 - CHIBA

Q12-4595 The Devonian dipnoan Ho/odipterus: dental form variation and remodelling growth mechanisms.
Campbell, K S W Smith, MM
Australian Museum. Record 39(3) September, p131-167; | appx, 24 fig, 17 ref (1987)
Vertebrate palaeontology/ Lung fish/ Morphology/ Devonian/ Canning Basin/ Gogo Formation/ Western Australia/

AESIS Quarterly Vol. 12 No. 4 December 1987

The Annual Cumulative Index to AESIS is published in the December issue each year. It
consists of Author, Subject, Locality, Map, Mine/Deposit/Well-name and Stratigraphic-name
indexes. There are also five year cumulative indexes on microfiche. AESIS is an excellent
secondary service that offers in depth subject analysis of the Australian Earth Sciences
literature. AESIS scans many internal reports missed by other abstracting and indexing
journals, and is also available as a machine readable database, which is used to generate special
bibliographies on specific fields of interest.

Australian Science Index (1957-1982) was intended to cover articles in Australian scientific
and technical publications. Entries are arranged under broad subject headings. Papers on
palaeontology are listed under the subject heading 'Earth Sciences - Palaeontology’. The
annual cumulation of Australian Science Index carries subject and author indexes.

As the primary literature continues to grow at an exponential rate, it is increasingly
difficult to keep abreast of developments in one's own field and in related fields of interest.
Review articles are extremely useful to the reader wishing to survey the current literature on a
subject. There are a number of review serials in earth sciences, none devoted exclusively to
palaeontology. Earth Sciences Reviews (Elsevier Scientific Publishing, Amsterdam, 1966-) is
a specialist review serial which frequently carries survey articles on progress made in
palaeontology. The review articles in this service are always informative, often evaluative and
sometimes critical. In all cases, the articles are followed by a long list of references to current
literature on the subject.

Review articles may appear in specialist review serials generally entitled Advances in...,
Progress in..., Reviews of... etc., and in primary journals or in books. Most abstracting
services, including the ones described earlier in this section, will include review articles. The
inclusion, however, of review articles is not reliable in any of the secondary services covering
palaeontology. /ndex to Scientific Reviews (1974-), a semiannual hardcover publication by the
Institute for Scientific Information, Philadelphia, is a special indexing service devoted to the
coverage of review articles collected from all types of publications. Most of the information
input to this service is based on the papers covered by Science Citation Index. Index to
Scientific Reviews has a Source Index, a Permuterm Subject Index, a Corporate index and a
Research Front Specialty Index. The Source index is a cross referenced author index to the
current review literature, giving a complete bibliographic description of each review article
(source item) followed by numbers representing research front specialities treated in the articles,
The Research Front Specialty Index provides a classified subject approach to research
specialities surveyed in the review articles.

Current awareness services are mostly title announcement services of papers published in
journals. The following current awareness publications are useful to palaeontologists. Current
Contents: Physical, Chemical and Earth Sciences (1957-) is a weekly publication by the
Institute of Scientific Information in Philadelphia. Current Contents consist of content pages
of journals regularly scanned by the service. Each issue has an author index and directory to
facilitate the acquisition of reprints of journal articles. Geotitles Weekly (London,
Geosystems, 1969-) is a weekly publication in three parts, the main part consists of a
classified list which gives details of authors, titles, and sources of approximately fifty to sixty
thousand items per year. The classification is based on the Geosystem subject classification.
There is an author index and a ‘key to coded sources’ in each issue.

AUSTRALIAN VERTEBRATE PALAEONTOLOGY LITERATURE - 163

TERTIARY PUBLICATIONS IN PALAEONTOLOGY

Palaeontology is a descriptive and a cumulative science. For this reason, books,
particularly reference books, are important to the student of palaeontology, while treatise and
fossil catalogues are indispensable to the researcher.

English language books on palaeontology currently in print can be identified in Books in
Print, (New York and London, Bowker Company) an annual author, title and subject index to
books in print. Scientific and Technical Books and Serials in Print, 1988 (New York) Bowker
Company, in volume 1, which is a subject index, lists several books on palaeontology under
the subject and also by country e.g. Palaeontology - Asia. Books are listed under the heading:
Vertebrates, Fossil. Palacontological publications no longer in print can be identified in special
bibliographies and guidebooks to the literature, in library catalogues and in guides to reference
material such as E.P. Shechy's Guide to Reference Books (10th edition, American Library
Association, Chicago, 1986) and A.J. Walford's Guide to Reference Material, 3rd edition, The
Library Association, London, 1973). Pure and Applied Science Books 1876-1982 (New York,
Bowker Co.) is a bibliography of some 220,000 titles in a six volume set. Inclusion is not
limited to books published in the United States, but also covers overseas publications which
were distributed in the United States. The retrospective Bowker Bibliography lists several
hundred titles on palaecontology under the general subject as well as by country, type of
publication, and speciality e.g. Vertebrates, Fossils - Bibliography. In the following, only a
brief reference will be made to examples of the different types of reference works in
palaeontology. For more detailed or complete listings of reference books, the reader is advised
to consult the guidebooks to the literature of Earth Sciences.

For a definition of a topic or for keywords to formulate a search question on an unfamiliar
topic, special dictionaries and encyclopaedias are particularly useful. The difference between the
two is blurred, as some dictionaries go beyond the usual single volume presentation, whilst
some encyclopaedias cover such a narrow field of speciality that a single volume may allow the
encyclopaedic treatment of topics falling within the field of coverage. An unusual example of
a popular but informative presentation of information in a dictionary form, covering a narrow
field of speciality is D.F. Glut's The Dinosaur Dictionary (Secaucus, New Jersey, Citadel
press, 1972), There are many books written on dinosaurs, the difference between the Dinosaur
Dictionary and similar books on the subject is in the dictionary type presentation, of
information by genus of dinosaurs. This special dictionary is useful for the interested
layperson and to the student of palaeontology. The dictionary has a bibliography of sources
consulted and recommended for further reading.

The Encyclopaedia of Paleontology (Stroudsburg, Pa., Dowden, Hutchinson and Ross,
1979) edited by R.W. Fairbridge and D. Jablonski, is an example of an encyclopaedic
dictionary of palaecontology. The Encyclopaedia carries fairly long articles, followed in each
case by a bibliography. The McGraw-Hill Encyclopaedia of Science and Technology isa
multi-volume encyclopaedia which includes many brief entries on topics of palaeontological
interest. The articles in this work are written by specialists, often illustrated and generally well
documented. A separate subject index to this work allows the survey of information on the
various aspects of the same topic. A single volume encyclopaedia published by the same
company which could be useful to the student of palaeontology is the McGraw-Hill
Encyclopedia of the Geological Sciences (New York and Sydney, McGraw-Hill, 1978). There
are a number of encyclopaedic dictionary-type reference works in the earth and biological
sciences, These works can be identified in bibliographies, guidebooks and library catalogues,
generally under the name of the subject followed by a subheading indicating the form of
publication e.g. Palaeontology- Dictionaries and Encyclopaedias.

Handbooks are useful reference publications for looking up special techniques, application

164 - CHIBA

data and principles. There are a number of handbook-type reference works in palaeontology.
Handbooks on palacontology or on related subjects can be located in the same way as suggested
for dictionaries and encyclopaedias, under the name of the subject e.g. Palaeontology-
Handbooks; Zoology-Handbooks. It should be noted that some reference publications listed as
handbooks are not handbooks in the strict sense of the definition, but are guidebooks and
realise covering narrow specialitics. The most comprehensive handbook on palaeontological
techniques is the Handbook of Palaeontological Techniques (San Francisco and London, W.
Freeman, 1965) a multiauthored volume work edited by B. Kummel and D. Raup. The
Handbook was prepared under the auspices of the Palacontological Society and it is in five
parts covering I. General procedures and techniques applicable to major fossil groups; IT.
Description of specific techniques; HI. Techniques in palynology; IV. Bibliography of
paleontological techniques and V. Compilation of bibliographies of use to palaeontologists and
stratigraphers. The Handbook is a source book for palaeontological techniques, with
illustrations, photographs, drawings and tables.

As palacontology is a cumulative science, scholarly treatise are essential tools for research.
In palaeontology there are numerous smaller works called treatise, which cover a narrow
specialty as well as multi-volume treatise that summarize the state of knowledge embracing the
whole field or a substantial part of palaeontology. Treatise-type palaeontological publications
are not necessarily identified by their title as treatise. It is often necessary to look beyond the
title of the work to ascertain that the treatment of the subject meets the criteria for a treatise.

Traite de Paleontologie (Paris, Masson, 1962-69) is a multi-volume treatise edited by J.
Pireteau, which presents in considerable detail a survey of documented knowledge of fossils
with a special emphasis on fossil fauna and vertebrate palacontology. The first volume of the
treatise commences with a summary of knowledge on primitive fossils, and the last volume of
the treatise covers primates. Volume one also contains a brief history of palaecontology with
portraits of palaeontologists, and a chapter on the process of fossilization and methods of fossil
studies. All contributions in this treatise are thoroughly documented, the papers carry
illustrations, photographs, maps and lengthy bibliographies.

A scholarly treatise covering the whole ficld of palacontology is a Russian work translated
into English by the Israel Program for Scientific Translations, entitled Fundamentals of
Palaeontology (Osnovy Paleontologiee) edited by A. Orlov and originally published in
Moscow by the State Scientific and Technical Press (Gosudartsvennoe Nauchno Technicheskoe
Izdatelsvo). This is a fifteen volume treatise-type manual for palacontologists and geologists
in the USSR. It is a carefully documented work, presenting a systematic summary of
knowledge in separate chapters dealing with different fossil groups. The treatise is illustrated,
and the chapters are accompanied by a bibliography in two sequences, the first, listing
publications in Russian, the second, listing publications in other languages. Volume fifteen
carries an alphabetical index of species, gencra, subgenera, etc. to facilitate the looking up of
specific information.

Fossil catalogues and fossil indexes are widely used in the identification, classification,
naming and description of fossils. Fossil catalogues are sometimes based on a collection of
fossils held by a muscum or an institution. There are numerous examples of fossil catalogues.
Fossilium Catalogus (Berlin, Junk, The Hague, Feller, 1913-) is a fossil catalogue published
in over one hundred parts to serve as a comprehensive index to the published generic and
specific names of fossils. The work is in two series: Series One, Animalia, and series two,
Plantae. Each part in the series consists of a catalogue of known species and an index and
bibliography to the class covered.

Examples of fossil catalogues based on collections of fossils are provided by the fossil
catalogues published by the American Museum of Natural History, which are substantial
works, and by the British Museum of Natural History, that are pamphlet type publications.

AUSTRALIAN VERTEBRATE PALAEONTOLOGY LITERATURE - 165

HINTS ON SEARCHING THE LITERATURE FOR
INFORMATION

It is important to adopt a flexible but systematic approach to the literature when searching
for information. The method of approach should be guided by the purpose of the search, the
volume of the literature, the scholarly level of the information required and the availability of
information resources. The most important thing in a literature search is to have an open mind
and a flexible approach to the problem. It is well to remember that documents and people
together make up the information resources on a subject. Private communication, professional
advice and suggestions from colleagues can save hours and put the searcher on the right path,
when documents on the subject appear inadequate to solve the problem. Librarians are
information specialists trained to search for documents and to access computerized information
services. In their daily professional practice they often use a wide range of information
resources. They search for information in books, in primary or secondary publications, in
audio-visual sources, microform and in computerized information services.

When embarking on a search it is important to have the right keywords to retrieve relevant
information. There are different ways to ensure that important keywords have not been
overlooked. The use of correct terminology can save time and frustration of having to start
again after a faise start or even worse, missing important information on the subject.

If the purpose of the search is to find documents held in a library, then the library catalogue
is the key to the collection. The keywords in this case are the authorized subject headings used
by the library to describe the subject contents of documents in the collection, the names of
authors and the titles of publications. Journals and serials are entered under the first word of
their title and under the subject. Most libraries in English speaking countries use the Library
of Congress Subject Headings. It is helpful to look up indexing terms in the Library of
Congress Subject Headings, before embarking on a search of the subject catalogues. If no
information is found under the correct subject heading in the catalogue the reader is advised to
check the entries under related or broader headings. The following is a sample entry showing
the range of indexing terms entered under palaeontology in the 9th edition of the Library of
Congress Subject Headings: paleontology

Includes both general and zoologi<l pa-
leontology.

sa Animal remains (Archaeology)

Bioherms

Classification—Books—Paleontology

Coprolites

Extinct animals

Footprints, Fossil

Forests, Submerged

Geodes

Living fossils

Micropaleontology

Paleobiogeography

Paleobiology

Paleobotany

Paleopathology

Sedimentary structures

Thanatocoenoses

Thin sections (Geology)

Trace fossils

Birds, Fossil; Brachiopoda, Fossil;

166 - CHIBA

Plankton, Fossil; Vertebrates, Fossil;
and similar headings
x Animals, Fossil
Antediluvian animals
Fauna, Prehistoric
Fossils
Paleontology, Zoological
Paleozoology
Prehistoric fauna
xx Extinct animals
Geology
Geology, Stratigraphic
Natural history
Paleobiology
Phylogeny
Science

Zoology

See also references refer the reader to related headings and see references to headings used for
the subject. It should be remembered that books may treat more than a single subject and can
be indexed under several subject headings.

Different types of publications are listed in the subject catalogue under the name of the
subject heading, followed by a subdivision indicating the type of publication, e.g.

Paleontology - Bibliography
Paleontology - Congresses
Paleontology - Dictionaries
Paleontology - Handbooks, Manuals
Paleontology - Periodicals

The classification used by the library offers an additional subject approach to the collection.
Most libraries use classification to arrange documents to facilitate browsing by readers. The
function of the classification is to keep documents on the same subject together, and
publications on related subjects in close proximity. The Dewey Decimal Classification is
among the most widely used classification in libraries.

The following is a sample entry from the 19th edition of the Dewey Classified Schedules,
used for palaeontology:

560 Paleontology Paleozoology

1 Philosophy and theory

AT Stratigraphic paleontology, paleobotany, paleozoology
Class specific fossils or groups of fossils in 561-569

A71 Archeozoic and Proterozoic (Precambrian)
paleontology

172 Paleozoic paleontology

1723 Cambrian period

172 4 Ordovician period

172 Silurian period

5
172 6 Devonian period
7 Mississippian period
Class here Carboniferous periods

For Pennsylvanian period, see 560.1728

AUSTRALIAN VERTEBRATE PALAEONTOLOGY LITERATURE - 167

STEP 1: ORIENTATION

USE REFERENCE BOOKS

SUBJECT OR SPECIAL DICTIONARIES, ENCYCLOPAEDIAS
GLOSSARIES, HANDBOOKS

STEP 2: SURVEY THE LITERATURE

USE REVIEW ARTICLES, BIBLIOGRAPHIES

REVIEW SERIALS; REVIEW PAPERS IN PRIMARY JOURNALS, BOOKS, SUBJECT

BIBLIOGRAPHIES, STATE OF ART REVIEWS AND SUMMARIES OF LITERATURE IN
TREATISES

STEP 3: IN-DEPTH READING

USE BOOKS TO SUIT A LEVEL OF UNDERSTANDING
GENERAL BOOKS, POPULAR INTRODUCTIONS, TEXTBOOKS, MONOGRAPHS,
CONFERENCE PROCEEDINGS

STEP 4: UPDATE INFORMATION

USE PRIMARY PUBLICATIONS

IDENTIFY RELEVANT PAPERS IN ABSTRACTING, INDEXING, CITATION JOURNALS
DATA BASES AND CD-ROM SERVICES

STEP 5: IDENTIFY RESEARCH IN PROGRESS

USE RESEARCH DIRECTORIES

SUBJECT, PERSONAL AND CORPORATE RESEARCH GUIDES, DIRECTORIES

OF R&D ORGANIZATIONS, ASSOCIATIONS, ACADEMIC AND RESEARCH
INSTITUTIONS, INFORMATION DATA BASES

Figure 2. Model of a comprehensive approach to a literature search.

168 - CHIBA

There is an alphabetical index to the Dewey Decimal Classification to guide the user to the
relevant classification numbers.

When embarking on a search for information in abstracting and indexing journals, the
enquirer should check the indexing terminology used by the service. There are special guides to
indexing terms used by secondary services generally called Thesaurus of Indexing Terms (a list
of authorized subject headings) used for an in-depth subject analysis. Indexing terms used in
secondary services are more specific than the list of subject headings used by libraries. An
outline of the system of classification used by a secondary service is also helpful to locate
relevant information in the current issues of the service.

Fig. 2 is a model of a comprchensive approach to a literature search showing steps
taken and the type of publications used. Only a small percentage of enquiries require a
comprehensive search for all available information resources on the subject.

COMPUTERIZED INFORMATION SERVICES COVERING
PALAEONTOLOGY

Computerized information services were developed in the 1970's as a by-product of the
application of computers to the coding, storage and retrieval of information for printed
secondary services, such as abstracting and indexing journals. Computers used in the
production of the printed abstracting and indexing journals replaced the paper copy of the
journal by a machine readable database which offers online access to the information that
appears in the paper copy of the service.

The advantages of using computerized information services over manual (paper or
microform) information services are many, Computerized information services provide
flexibility of approach to searching through the use of boolean operators which combine
indexing terms, with each other as well as with subject categories, titles of publications and
names of authors, to retrieve relevant information in minimum time with maximum ease. The
use of natural language (free text) rather than authorized indexing terms for searches is a feature
offered only by the computerized information services. Finally, as information is updated first
in machine readable form, the database equivalent of a printed abstracting service is more up-to-
date than its printed counterpart. Advances in telecommunication and computer technologies
brought the cost of using computerized information services within the means of most people.

There are a number of computerized information services with a coverage of palaeontology.

GEOREF is a computer database produced by the American Geological Institute. The
database covers references included in the following publications:

Bibliography and Index of North American Geology (1961-1970)
Bibliography and Thesis in Geology (1965-1966)

Geophysical Abstracts (1966-1971)

Bibliography and Index of Geology Exclusive of North America (1967-1968)
Bibliography and Index of Geology (1969 to present)

we whe

GEOREF covers technical literature on geology, geophysics and palaeontology on a
worldwide basis. The database indexes journal articles selected from over 4,500 serials, books,
conference papers, government publications, theses, reports and maps. From 1967 the
coverage of literature is international. An estimated 40% of the publications indexed by the
service originate in the United States. Information in the database is updated monthly,

GEOARCHIVE is produced by Geosystems in London. It is a comprehensive database
covering the geosciences, including palacontology. GEOARCHIVE provides an international
coverage of the literature. It scans over 5,000 serials, books from over a thousand publishers,

AUSTRALIAN VERTEBRATE PALAEONTOLOGY LITERATURE - 169

theses, technical reports, maps and conference papers. The database covers geoscience literature
from 1974 to date. The service is updated monthly. Approximately 5,000 citations are added
per update to the database.

AES] is the machine readable equivalent of the paper and microform version of AESIS, the
Australian Earth Sciences Information System. The coverage in the database is restricted to the
Australian literature of earth sciences and palacontology. The database can be searched online
from 1976 to date.

AUSTRALIS is the collective name for the Australian Scientific and Technical Information
Database marketed by CSIRO: the Commonwealth Scientific and Industrial Research
Organization in Australia. AUSTRALIS databases cover research reports, conference papers,
journal articles, books, theses and research in progress. The database service is supported by
AUSTRADOC, a document delivery service offering online ordering of full text documents
retrieved from AUSTRALIS databases.

A search of AUSTRALIS databases for information on Vertebrate Palaeontology retrieved
relevant documents from a number of AUSTRALIS databases. The most productive
AUSTRALIS database on the subject was CSIRO Index (CSX), which lists over 51,000
documents issued by CSIRO from 1969 onwards. The search also identified no less than
thirty-three Australian institutions, state geological surveys, museums, tertiary institutions and
research organizations with an active interest in palacontology.

CD-ROMS AND VERTEBRATE PALAEONTOLOGY. Initially CD-Rom technology
was developed as an alternative to online access to expand the market for information services
in remote areas without telecommunication facilities. Compact discs have large storage
facilities; a single CD-Rom can store the equivalent of 200,000 printed pages of text. For the
user, the CD-Rom version of a database eliminates the need for bearing the cost of search and
telecommunication services. The International Guide to CD-Roms in Print 1988-89 (Emard
1988-1989) lists over two hundred reference and information databases available on CD-Rom.
In the 1988-1989 guide there are a number of GeoScience Information databases available on
CD-Rom, but none so far that would be expecially useful for identifying information resources
on vertebrate palaeontology.

In recent years the pheonmenal increase in the usage of micro- and personal computers, the
emergence of laser optical discs for data storage and the availability of user friendly software,
has led to an explosive growth in the number of GeoScience databases. The Directory of
Government GeoScience Databases in Australia lists fifty-seven reference and one hundred and
ninety-six factual databases produced by thirty-seven organizations, The development of Open
Suystem Interconnections (OSI) will allow in the foreseeable future, direct, online access to
GeoScience databases in Australia and a demand for individually packaged, integrated databases
specified by the end user of the systems.

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Geotitles. 1969- (formerly Geotitles Weekly) 1969- . London: Geosystems.

GINSBURG, R.D., 1977. State of the Art Survey of Reference Sources in Geology. Reference Services
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CHAPTER 6

TECHNIQUES USED IN
PREPARATION OF

TERRESTRIAL
VERTEBRATES.

Michael Whitelaw! and Lesley Kool?

IMNOMUCTION 0.0... eee ecceeeceeceeeereesccneceees 174 Acid Supply..........ccccceseceeeeeeeees 185
Chemical Techniques - Acid Etching....... 174 Moving Blocks Around the
Detailed Methodology...............00006+ 175 Laboratory........e.c.eceeeeeeeeeee 185
Preparation of the Block Prior to Fluid Movement around the
Etching.......cceeecccccccesecceseeee 175 Laboratory ...........cccseeeeeeeeees 186
Orientation and Mounting Preparation Materials................. 186
Of BIOCK..........ceeeeeeeeee 175 Non-Consumables................ 186
Paperwork ............cccccesseceeesecees 176 Consumables ................2.2005 186
Acid Choice and Pouring............ 176 FOAMS ...........cececeeececeeeeeeeee 187
Acid Types and GCS. 5 se5 steele teiletywwes odes 187
Concentrations............... 176 Hardenesrs............cccececceeeeeees 187
Pouring Acid .................0005 179 Preservatives ...........cececee eens 187
Duration of Reaction Time....179 Physical Preparation Techniques............. 188
Post-Reaction Processing........... 180 Collecting the Specimen.................. 188
Emptying Acid Tubs............ 180 Preparation Methods ...............:...0008 188
Block-Inspection and Laboratory Safety ...............ceeseeeeeees 189
Cleaning..............cc ce eeeee 180 Recommended Tools..............0000ee0e 190
Mineral Salt Removal ......... 180 Field Techniques ...............ccccecececsseeeees 190
Washing and Drying............. 181 Screen Washing ............cscceeseeeeeeeee 190
Final Acid Neutralization ...... 181 Plaster Jacketing ..............ccceceeeeee eee 192
Preparation of Partially Etched Casting and Moulding Techniques.......... 193
BOMES.,......... ee eeeeeeeeeceee es 181 Preparation of Specimens for
Hardening Vertebrate Moulding 2.00.0... cecec ee eeeeeeeeese ees 193
SPeCiMENS.........e eee 182 Latex Peel ...........ececccecececeeeee ees 196
Acid-Resistant Lithologies .... 182 Silicone Rubbet.................000000 197
Laboratory Safety...........c cs eeeeeeeeees 183 Plaster of Paris-Mould Supports .. 197
Laboratory Design and Casting from Moulds..................0008 198
Materials...............:.:eeeeeeeeee 183 Plaster of Paris...............c.c020e0e 198
Laboratory Design ...............00608 184 Polyurethane Foam .................4. 198
Acid. Tubs ...........:. ce eeeeeee cree ee ees 184 Epoxy ReSin...............cecceeeeee eee 199
Acknowledgements .............0ceececeeveseeees 199
RELELOHCES tile ccc. tiga deetah isi wer Avatars 200

1 University of Texas at El Paso, El Paso, Texas 79968-0555, U.S.A.
2 Department of Earth Sciences, Monash University, Clayton, Victoria 3168, Australia.

174 - WHITELAW & KOOL

INTRODUCTION

The quality of research in palaeontology is directly related to the quality of the material
available for study. Therefore, the aim of fossil preparation is to make the specimen amenable
to study whilst preserving, as closely as possible, its natural form. This chapter will deal
specifically with the preparation of vertebrate fossils from three very different, although
common. lithologies: a calcium carbonate matrix that typifies the mid-Tertiary Bullock Creek
locality m the Northern Territory; the sandstones and mudstones characteristic of the Early
Cretaceous fluviatile and lacustrine facies of southern Victoria; and the unconsolidated fluviatile
sands and silts typical of the Tertiary sites of Central Australia. Fossils obtained from these
lithologies have been prepared by using acid etching, physical, and screen washing techniques,
respectively, and will be described in detail. These techniques are the ones most frequently
utilized in preparing a large proportion of the vertebrate fossils of Australasia and should give
the beginner a basic idea about how to proceed. This chapter is not intended to be a
comprehensive review of the preparation methods available. If a broad coverage of techniques
is required, Kummel & Raup (1965) and Rixon (1949, 1976) are suggested as further reading.

This chapter will discuss how to prepare moulds and casts of fossils. In many situations
casts may be used as substitutes for valuable specimens, thus ensuring the safety of the
original. Such replicas are commonly used in research, for exchange and for displays. The
production of accurate, detailed copies of unique, irreplaceable specimens is, therefore, a vital
skill for any preparator.

CHEMICAL TECHNIQUES - ACID ETCHING

Variation in the chemical composition, between bone and the surrounding matrix, allows
the acid etching method to be employed in the extraction of vertebrate material. Calcium
carbonate (CaCO3) is susceptible to attack by dilute acetic acid (CH3COOH), whilst bone,
which consists of collagen hardened by mineral salts, largely calcium phosphates
(Cas5(PO4)3(OH,F,Cl), which have often altered to apatite in fossilized specimens, is not.
This method has been used to prepare a variety of fossils including Devonian placoderms from
the Gogo Fauna of Western Australia, varied marine invertebrate faunas and, as discussed in
this chapter, terrestrial vertebrate fossils such as those of the Bullock Creek Local Fauna
(Miocene, Northern Territory).

The essentials of the acid etching method used is well described in Rixon (1949, 1976),
and Kummel & Raup (1965). This report seeks to give some idea of the methods, problems
and costs of setting up an acid etching laboratory and to give practical examples, based on the
Bullock Creek work.

Acid etching involves the dissolution of carbonate matrix from a bone aggregate or
specimen, by submersion in dilute acetic acid. The general form of the chemical reaction
involved is CaCO3 + 2CH3COOH = Ca2t+ + 2CH3COO- + CO2 + H20. As the fossil is
gradually exposed during this process, it must be periodically strengthened and hardened. This
is achieved by impregnation of the bone with acid resistant plastic solutions and glues between
acid treatments (Fig. 1).

The carbonate lithologies from the several localities that comprise the Bullock Creek
Local Fauna vary quite widely, ranging from a relatively pure carbonate matrix, to clay and
sand rich matrices, to cemented pebble/bone/nodule breccias. This lithologic variation has a
strong influence on the etching characteristics of each matrix block and, therefore, on the
processing method employed. The type of bone elements and their preservational requirements
are also controlling factors in the etching process. The range and combination of these factors

PREPARATION TECHNIQUES - 175

necessitates the treatment of each block on an individual basis, with regard to acid volume and
concentration, immersion period and type and quantity of preservatives used.

DETAILED METHODOLOGY
Preparation of Block Prior to Etching

Before the block is immersed in acid, the condition of exposed bones should be carefully
checked. If the bones are cracked or crushed, some form of consolidation or support will be
needed. The hardening agent used is a 3-5% solution of Synocryl 9122x (a polybutyl-
methacrylate) in acetone, which may be painted on the bone surface. This solution has a high
penetration and should be applied, in a series of coats until the bone is hardened and
consolidated. If necessary up to a 30% solution of Synocryl may be applied, but at these
concentrations it has low penetration and will only form a skin on the bone surface.

In many limestones, the rock is harder than the bone, leading to the clean breakage of
bone elements along matching block faces. A close examination of the bone surface will often
reveal that the interstices have been filled with either carbonate matrix, calcite crystals or both.
For these bones, strengthening is not possible, as hardenin g agents are incapable of penetrating
the carbonate filled interstices to any practical degree. These blocks must first be put in acid
before they can undergo additional consolidation.

If two blocks with matching faces have bones exposed, every attempt should be made to
glue the two faces together so as to etch the two as one block. Once etching begins, it is
difficult to prevent skeletal elements from chipping along exposed edges and this prevents good
"joins" when attempts are made to repair bones. Naturally, this is limited by the size and
weight of the blocks that can be comfortably handled during preparation.

Orientation and Mounting of Block

After all exposed bones have been treated, the orientation of the block in the acid tub
should be considered. If possible, the block should rest on a face which has no bones exposed
exposed and/or with the most important specimen to be etched facing up. This will facilitate
access during preparation. Most small blocks can be dealt with in this manner.

Blocks weighing in excess of a few kilograms require a different approach. As the block
is etched, bones are exposed on all surfaces, and those exposed on the base will be crushed by
the weight of the overlying rock. The application of an acid proof base will alleviate this
problem. After the optimal etching orientation for the block is chosen, it is inverted, and a
waterproof polyurethane foam base is applied. This foam is chemically inert (acid-proof) and
available as a two part mixture consisting of a resin and an activator. Once mixed and applied,
the resin foams with a considerable increase in volume and then rapidly cures. The block
should be placed in its tub whilst the foam is still active allowing the foam to mould itself to
the space between the block and the tub bottom. This seals the bottom of the block and
provides it with a stable base which will prevent the block from rolling over and crushing
exposed bones as etching proceeds.

The use of the foam base requires this important caution. Since the size of the block is
reduced with each acid pour, the strong positive buoyancy of the foam will cause the block to
become unstable and eventually capsize. As etching continues gradual reduction of the foam
base, by shaving it with a knife, will alleviate this problem.

176 - WHITELAW & KOOL
Paperwork

The importance of maintaining accurate and up-to-date records during processing can not
be over emphasized. Each etching tub should be given a number, letter, name or some form of
identification, and the locality information of the block within it must be recorded. Along with
this information, a list of any exposed or identifiable benes should be kept. We found it
a to use a table on which to record processing details, including the following (see Fig.

):

(a) date, concentration and volume of acid pour

(b) state of the block (in acid, soaking, or in preparation)

(c) any notes on preservational details - glues used, problem bones, etc.
(d) a list of any bone elements or specimens recovered.

Acid Choice and Pouring

Acid Types and Concentrations

Once the block has been prepared and placed in its tub, acid may be added. For most
calcium carbonate matrices, acetic acid (CH3COOH) is the preferred agent, although formic acid
(HCOOH) may be used. Acetic acid is less expensive and poses less of a physiological hazard
to the user (Rixon 1976, p.111).

Experiments with acid concentration led to a 10-12% solution being adopted as the most
efficient and effective etching agent. Higher and lower concentrations were rejected for several
reasons. Rixon (1976) experimented with very high acid concentrations and found that acid
solutions stronger that 33% could not effectively ionize and would not dissolve carbonates.
Concentrations between 20% and 33% not only etch calcium carbonates, but also calcium
phosphates (bone). Concentrations in excess of 15% produce large amounts of salts in
solution from the dissolved matrix which tend to buffer the acid. This causes activity states to
tail-off exponentially, resulting in incomplete acid consumption, even over quite long
immersion periods. Concentrations over 15% also tend to create strong effervescence, through
the rapid production of CO, which creates excessive turbulence that may damage fragile bone
elements.

Concentrations below those recommended are not directly detrimental to bone elements
but necessitate more frequent replacement of the spent acid. This requires extra handling, which
unnecessarily exposes the specimen to damage.

Figure 1. Flow chart outlining the steps involved in acetic acid preparation of vertebrate fossils contained
in a limestone matrix.

PREPARATION TECHNIQUES

AND BROUGHT TO LAB.

| LIMESTONE BLOCKS ea

INSPECTION

BONES O.K.

SIZE SORTING

SMALL < 4kg

BONES DAMAGED

CONSOLIDATION
AND REPAIR

LARGE > 4kg

POLYURETHANE
BASE

PLACED IN SMALL PLACED IN LARGE
202TUB 75tTUB

ACID ETCHING
ROCK SUBMERGED IN 10-12%
CONC. ACETIC ACID
ACID EXHAUSTED IN
5-14 DAY PERIOD
DRAIN TUB
INSPECT BLOCK AND
SCRUB CLEAN
FURTHER
DISSOLUTION
REQUIRED
PREPARATION ?
REPAIR ?

YES
TUB CONTENT —

= J IN WATER

SALTS
FORM
BLOCK DRIED

NO SALTS

FINISHED ELEMENTS AND EXPOSED BONES HARDENED
SEDIMENTS RINSED &

AND REPAIRED
SIEVED RESPECTIVELY

|

FINISHED MATERIAL
SOAKED IN 5% AMMONIA

;

FINISHED MATERIAL
SOAKED IN WATER

SALTS
FORM
~~~{__ MATERIAL DRIED ee
Ne ji ae
BONE ELEMENTS FINAL REMAINING SEDIMENTS

PREPARATION —- HARDENING PICKED FOR
VIA VACCUUM IMPREGNATION MICROFOSSILS

177

178 - WHITELAW & KOOL

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Figure 2. Sample of a typical record sheet, used for detailing processing stages, of a limestone block from
the Miocene Bullock Creek site, N. T. (Courtesy of S. Morton and Monash University, Melbourne).

PREPARATION TECHNIQUES - 179

Pouring Acid

In practice, the easiest method of obtaining the right acid concentration for the etching
procedure is to simply fill the etching tub with a measured volume of water until the block to
be etched is submerged. Concentrated acid (usually either 90% or glacial) can then be slowly
added to make up the solution to 10-12%. This avoids the necessity of having to allow for
block displacement in concentration calculations. If the rock volume needs to be considered,
calculations to within tolerable limits may be obtained by dividing the weight of the rock
(grams) by the specific gravity of calcite (2.71 g/cm3).

After several pours the volume, and, therefore, the displacement of the rock will be
considerably reduced; thus, the volume of concentrated acid used will have to be adjusted
accordingly. Most blocks tested exhibited some tolerance to concentration variation.
Solutions in the range 10 + 4% do not damage emergent bone, and there is minimal waste of
acid or etching time.

Duration of Reaction Time

The duration of the acid reaction is a function of the lithology of the block, and the
concentration, volume and temperature of the acid solution. The rate of carbonate dissolution
decays exponentially as a result of acid consumption, a salt solution buffering effect and the
build-up of noncarbonate sediments on the block. The reaction normally reaches a practical
working minimum within 7-14 days, although continued reactions of over 3 weeks have been
observed. As long as the reaction continues, CO? gas will form as a by-product. As it rises to
the surface, it serves to keep fine sediments in suspension by generating turbulence or by
buoying up individual grains. Useful etching may be considered over when CO? production
has ceased, as evidenced by a lack of sedimentary particles in suspension.

Reaction times may be decreased significantly if the acid solution is maintained at
temperatures of 30-35°C. In small volume tubs, this may be achieved by the use of electric
heating elements, such as those used in aquaria. Heating of the acid solution in large volume
tubs was found to be impractical, but an inexpensive partial solution is to use hot water when
they are initially filled. Large volume tubs will hold heat for up to 24 hours, thus increasing
the etching rate. Care should be exercised with this procedure, as hot solutions produce violent
initial reactions (strong effervescence), which may damage fragile bone.

In some cases, the use of large volumes of acid, relative to the size of the block, will
reduce the salt buffering effect, thus allowing more carbonate dissolution per etching cycle.
The larger acid volume normally means that each etching cycle will take longer to complete.
However, the rate of removal pays dividends by markedly lowering the number of treatments
necessary to fully etch a specimen. The efficiency of this method is largely dependent on block
lithology. Clean carbonate will etch rapidly whilst clay rich matrix will develop an insoluble
skin that will inhibit etching.

After the acid has been exhausted, some workers simply reactivate the solution by adding
fresh acid. This practice is permissible for most invertebrate and some well preserved vertebrate
specimens, but is not recommended. Excessive immersion periods, without hardening
treatments, will damage bone elements, no matter what acid concentration is used. Complete
acid changes also allow regular inspection of exposed elements, a necessary task if reasonable
preservation of all bones in a block is to be attained. Simple tub reactivation will eventually
result in a substantial waste of acid and time as it allows a cumulative increase of mineral salts
in solution. These will act as a basic (high pH) buffer, which will inhibit further etching.

180 - WHITELAW & KOOL
Post-reaction Processing

Emptying Acid Tubs

Once etching has been completed, the used acid must be removed, At the Monash
University laboratory the fluid is siphoned or pumped directly into a sink. If the etching tubs
can be placed above the drainage point the siphon is preferred. The slow removal of spent acid
by this method allows gradual draining of fluid from bone cavities, thereby minimizing the
stress on exposed elements. Nylon garden hose was found to work quite well, and it is cheap,
easily available, and can handle high sediment loads without blocking. Acid-proof pumps are
available, but they are expensive and rarely capable of handling any form of sediment load
without modification. By necessity, a pump is used for some of the tubs in the Monash
laboratory. It is described in the laboratory apparatus section.

The tub should not be totally emptied if the block is to be immediately etched again.
Leaving 5 cm of fluid in the bottom will help prevent the accidental loss of small elements
(i.e. small teeth) by removal of the tub residue. Also, a small amount of fluid left in the tub
acts as a buffer Jeppson et al. 1985).

At Monash, the Melbourne Metropolitan Board of Works (MMBW) regulations require
that the spent acid has a pH range of 6-10 before disposal. Each tub, therefore, is tested with
litmus paper and, if necessary, a small amount of ammonia (approx. 100 ml. of 75% strength)
is added to neutralize any remaining acid.

Block Inspection and Cleaning

Once the tub has been emptied, bones should be inspected for signs of deterioration and
the tub bottom checked for broken elements, which may match pieces still embedded in the
block. If the exposed specimens do not require hardening, and the sediment/bone residue is not
to be collected, the block may be cleaned and another etching cycle initiated.

The cleaning process is important, as clay-rich matrices develop thick "mud-skins"
which inhibit etching. The block should be carefully scrubbed down with a fine bristle brush
and a soft water spray. Water pressure and direction is best controlled by using a length of
flexible rubber laboratory hose attached to the water outlet. Hard bristled or nylon brushes (old
tooth brushes), when used with discretion, are also effective on stubborn matrix.

Under no circumstances should the block be allowed to dry between pours. Drying of
the surface causes dissolved salts to exsolve and crystallize within bone interstices. The growth
of salts exert large internal stresses that may lead to damage and/or collapse of fragile elements.
If bone elements require repair or consolidation, the mineral salts will need to be removed.

Mineral Salt Removal

If the exposed bone elements are in need of hardening or repair, the whole block must
undergo a pre-preparation soak in water. This is necessary for ALL blocks, bones and
sediments before any preparation can be undertaken. Stopping the etching activity of remnant
acid and preventing the growth of insoluble salts on or within bone elements is vitally
important. If a halt for preparation can be anticipated, before the last pour in a series of acid
treatments, salt removal from the block can be expedited by halving the acid concentration.
The block should be soaked in water until salts do not appear as it begins to dry.

The duration of the soaking process for the Bullock Creek blocks ranged from four days
to two weeks and is largely dependent on block lithology, with soaking times increasing
through the series: pure carbonate, sand rich limestone, clay rich limestone and brecciated and
conglomeratic limestone. Times for individual blocks may be significantly decreased if the

PREPARATION TECHNIQUES - 181

water is replaced every 2-3 days, and reduced to a minimum if a constant water exchange is
maintained. The latter is only recommended when time is the important factor, as water
consumption is enormous. Sediments and bones that have dropped off the block during
processing should not be disturbed until the salt removal process is completed. Missing bone
elements are easier to locate and repair when the residue is left in situ.

Washing and Drying

When salt removal is complete, the block should be taken from the tub and rinsed, if it
can be moved without damaging semi-prepared specimens. Also note the location of broken-
off bone fragments in the tub, to expedite their return to the matching section still in the
block, after the individual fragments have been hardened. Rinsing is best carried out on an
open mesh grid placed over a sink. Rixon (1976) suggests the use of diffusion grids from
overhead lights, for this purpose. They are adequate for the lighter blocks but of little use for
the heavy blocks. Bread crates, such as those used by most bakeries, prove to be excellent
alternatives, being capable of supporting blocks up to 40 kgs in weight.

Recovery of microvertebrate elements and sediments from the block is accomplished
using a fine mesh screen, such as fibreglass fly-wire, placed between the block and the grid.
Flywire is inexpensive, acid proof and has a mesh size which allows passage of fine sands and
clays without loss of the smaller vertebrate material. After the block has been scrubbed down,
it should be removed and allowed to dry. The remaining tub residue may then be washed from
the tub and onto the mesh. This concentrate may then be rinsed by holding each corner of the
mesh in one hand whilst running water down the outside until all the material has been washed
into the centre.

The duration of the drying process is a function of several factors. Blocks that are too
heavy or those that contain specimens too fragile to move must be dried in their tubs. The
resultant lack of air circulation slows down the drying time. Clay-rich and brecciated blocks
which absorb large volumes of water also take a long time to dry. In several blocks, drainage
was inhibited by the sealing effect of the polyurethane base. Drainage may be augmented by
the removal of foam from part of the base or by tilting the block at a steep angle to the base,
whilst it dries.

To keep track of blocks, prepared bones and residues it is vitally important to label each
item with its tub number, locality information and, preferably, the date. In a large operation
material is easily misplaced, and the value of a specimen is severely reduced by poor or suspect
locality data.

Final Acid Neutralization

To ensure complete acid removal, both the fully etched bone elements and the sieved
residue are soaked in a 3% ammonia solution. The residue is immersed in the ammonia for
approximately an hour, then in water for 5 minutes and allowed to dry. Large bone elements
are immersed in ammonia overnight and then soaked in water for an equivalent period of time.
The water rinse removes and prevents growth of ammonia salts, which like the acid salts, can
cause damage to bone. If salts begin to form when the bone dries, it should be immediately
rinsed again.

Preparation of Partially Etched Bones

Once a block has been dried, broken bones must be repaired. Acid and water insoluable
glues, preferably plastic ethyl methyl methacrylate-based types (see Glues in "Preparation
Materials” section) are recommended.

182 - WHITELAW & KOOL

Exposed bones must be hardened with a preservative capable of penetrating deep into the
bone. This is accomplished by dissolving a plastic glue in a volatile solvent, usually acetone
or ethanol, by applying it to the bone. The solvent will penetrate the bone, carrying the glue
with it in solution, but rapidly evaporates, leaving the glue behind to harden. At Monash
University, hardening and consolidation of exposed elements is accomplished by application of
coats of 3-5% solutions of a polybutylmethacrylate based glue, in acetone. A light, fine-
haired brush is recommended for this job, but for very fragile bones, and the insides of hollow
elements, a pasteur pipette (or eye-dropper) may also be used. Synocryl 9122X is the glue
currently used for hardening applications.

Fragile, delicate or crushed specimens sometimes require additional structural support.
Elongate bones may be reinfored using temporary splints, which may be made by gluing a thin
piece of wood or plastic along its length. A methacrylate glue, which can later be removed
with acetone, is recommended for this job. Thin, wide bones, such as cranial elements pose a
continuing problem, as hardening treatments strong enough to hold them together tend to
develop a "skin" which prevents any additional treatment of the emergent bone. The acid also
tends to permeate under these skins and reaction with the underlying carbonates produces CO2
gas which separates the support layer from the bone. Presently, such elements are treated in
several ways, on a case-by-case basis. The bone may be structurally supported by the
application of glue-impregnated layers of open-fibered tissue paper or heavily coated with dilute
glue and slowly etched in a 5% acid solution. In some cases such elements have been placed in
the tub so that the endangered bone remains above acid level as long as possible. This will
tend to limit the damage caused by the etching process. When exposed elements are considered
adequately consolidated and supported, the block may be returned to the tub for continued
etching.

Hardening Vertebrate Specimens
Once dry, the sieved sediments should be picked and microfaunal elements mounted or
stored in separate containers. If necessary, these elements may be strengthened with a 5%
solution of Synocryl (see Glue section). Larger elements are hardened, by vacuum
impregnation for 15 minutes or by overnight immersion, in a 10% polyvinyl acetate (P.V.A.)
solution. A 10% solution of a 4:1 ratio of Mowoiil 144 (styrene/acrylic vinyl acetate) and
Mowotil 7001 (plastic/acrylic vinyl acetate) is recommended. A discussion of glues is given at
the end of the acid preparation section.
A separate storage for bone elements and residue from each tub should be kept until each
block is entirely dissolved. Many of these elements may ultimately be repaired by careful
examination of the remnants of a fully etched block.

Acid Resistant Lithologies

Acetic acid will not dissolve a variety of minerals commonly encountered in limestones,
including crystalline calcite, which frequently occur in bone hollows and interstices,
magnesium carbonates (dolomites) and silcretes. Other workers have had some success with
resistant forms of calcium carbonate by using dilute solutions of formic acid (HCCOOH)
(Rixon, 1949). This acid was tried on some "problem" Bullock Creek blocks with little
success. Silcrete is resistant to all acids, except hydrofluoric acid,which will also dissolve
bone. Similarly, calcite is susceptible to hydrochloric acid, as is bone. At present bone
embedded in these matrices must be prepared mechanically or be preferentially etched from the
matrix, so that casts of the bone can be made from the resulting moulds.

PREPARATION TECHNIQUES - 183

LABORATORY SAFETY

The nature of the acids and solvents used demand that some comments on safety be
made. A laboratory coat, rubber gloves, adequate footwear (gum boots) and a face mask, fitted
with a breathing filter suitable for acid fumes, should be worn whenever handling the acid (Fig.
3). Concentrated acetic acid will burn, and its fumes may damage lung tissue. The dilute acid
(10%) may cause skin irritation and/or breathing problems, to some people. Work should be
carried out in fume cupboards whenever possible, but if the operation is too large, in a well
ventilated laboratory. After a pour, the acid fumes may also be reduced by fitting each tub with
a cover or lid. At peak operating capacity the Monash facility will use 200 litres of 90% acetic
acid in a single pour, thereby producing a considerable volume of acid fumes. To avoid
extended exposure to these conditions pours are preferentially carried out on Fridays so that the
laboratory has a full weekend to ventilate.

Fumes and vapours produced by preparation and hardening agents are also a problem.
Those produced by the polyurethane foam are toxic, whilst the plastic glues are nearly all
diluted with acetone and/or other carcinogenic ketones. Preparation should be carried out in a
fume cupboard or a breathing mask should be worn.

Figure 3. Pouring acetic acid into calibrated bucket (A) and then into processing tubs (B). Standard safety
equipment is being wom. Acid-proof footwear is highly recommended, either rubberized shoes or gumboots
will suffice. (Courtesy of S. Morton and Monash University, Melboume).

LABORATORY DESIGN AND MATERIALS

This section describes the lab equipment and materials used at the Monash University acid
etching facility. All equipment and supplies were/are obtained in Melbourne, and specific
name-brand supplies may not be applicable interstate. Costs are those for original purchases
during 1984 or from ongoing consumables as of April 1985. Prices have been updated where
possible.

184 - WHITELAW & KOOL

Laboratory Design

The size and design of an acid etching laboratory is controlled by the anticipated scale of
the program. The current laboratory at Monash University is a portable building, measuring
approximately 3x6x2 m, and designed to operate 30 large and 20 small tubs at peak capacity.
The building was purchased in 1986 from Accom Portables Australia, 8 Fraser St., Airport
West, Victoria at a cost of $5,290.

Two laminated benches, running the length of each side of the room, can accommodate
ten large (75 1) tubs each, whilst a similar number of tubs are located beneath each bench. The
floor tubs are mounted on trolleys, designed by the Earth Sciences Department workshop,
which allow block movement around the laboratory as required. In practice, a large amount of
bench space is required for fossil preparation and storage; therefore, only one bench is utilized
for the large tubs.

The far end of the room is equipped with a stainless stecl sink (100 | capacity and 0.75
m2 work space) where the bulk of the block cleaning and preparation is carried out. The sink
is fitted with double taps and a hose long enough to reach the full length of the laboratory.

A sediment settling tank (3600 | capacity) is installed behind the laboratory. To
conform to Melbourne Metropolitan Board of Works sewer regulations the tank was lined with
150 mm thick, reinforced concrete and measures 2400x1500x900 mm. Waste water and
sediment from the laboratory sink drains to this tank to allow the removal of particulate waste
before it enters the sewer system. Specifications for waste management from acid etching
facilities may vary in different areas, so be aware of possible requirements before a laboratory is
built. In a small operation, sediment may be collected by pumping or washing waste water
into a tub, which is allowed to overflow into a drain or sink. This form of sediment trap will
successfully remove the large particulate matter that is commonly responsible for sink
blockages.

Room ventilation is maintaincd by a WFS Fume Scrubber, manufactured by
Conditionaire International Pty Lid, Moorcbank, New South Wales. This scrubber is suitable
for water soluble gases and particulate matter and may be used in either vertical or horizontal
ducting.

Even with good ventilation, corrosion by acetic acid and its fumes remains a problem,
Apart from stainless steel, acetic acid will corrode most metals. It de-polymerizes many types
of plastics and greatly increases the rate of wood rot. These problems are facts of life in an acid
etching laboratory and can not be avoided. Damage to work surfaces and floors may be reduced
by covering them with vinyl or a similiar protective material. Corrosion of tools may be
minimized by using stainless stecl or resistant plastics; however, the cost of quality stainless
steel items is often prohibitive, and it may be cheaper, in the long run, to buy lower quality
products and replace them more frequently.

Acid Tubs

The only tubs that are guaranteed corrosion-proof are those made of stainless steel or
special purpose plastics, for which prices may be well over $100 each. Manufacturers
specifications of less exotic plastics often indicate that their products may de-polymerize, with
resultant development of brittleness and loss of structural integrity, when exposed to acetic
acid. This problem has not been encountered during the five years that the current tubs, all
made of common plastics such as ethylene and propylene, have been in use. Fibreglass sieving
mesh and nylon hose have also been used for extended periods with no noticeable deterioration.

Three types of acid tubs are employed in the Monash laboratory: (a) for small blocks
(<4 kg), 20 1 tubs (30x40x18 cm) made of polyethylene are used. These were available in

PREPARATION TECHNIQUES - 185

several hardware shops around Melbourne advertised as "Viking Handy Boxes" and cost $9.75
for the tub plus lid; (b) 75 1 tubs (65x42x39 cm) made of polyethylene are used for etching the
larger blocks, which range up to 30 kg. Tubs of the same size may frequently be seen in fresh
food shops and are often called "butcher's tubs". These tubs are very expensive, if bought from
food container suppliers, due to health and quality regulations. However, identical tubs classed
only as "General Storage Containers" are much cheaper and were obtained from Nally Plastics
Pty. Ltd., Blackburn, at $16.80/tub and $4.40/lid; (c) several 100 1 tubs were created by
cutting empty 200 | acid drums in half with an angle grinder. As the acetic acid is sometimes
delivered in non-returnable, plastic lined, drums, no cost is involved. This is also a useful way
of disposing of the empties. The high volume of these tubs allows a rapid rate of carbonate
dissolution and preparation of oversized blocks that cannot be moved once etching is begun.

Acid Supply

Acid consumption is a function of the carbonate/clay ratio, the exposed surface area, acid
concentration and the ambient acid temperature. In the Bullock Creek material, consumption
varies from 0.4-1.2 kg of limestone dissolved per litre of acid, with an average of
approximately 0.85 kg/l. This figure may be of some use in projecting likely acid
consumption in other etching programs.

Acid purity is not a factor in the etching process, so the lowest (cheapest) grade, known
as "Technical Grade", should be used. Ajax Chemicals Pty. Ltd., is the major wholesale
supplier of acetic acid in Melbourne charging $275.00 for 200 | (44 gallons) of 90% technical
grade acid in a non-returnable drum. The acid is also supplied in volumes of 20 1 at $42.00.
Another supplier is I.C.I.dmperial Chemical Industries) Australia, which charges $1.75 per kg
plus a $50.00 deposit (as of November 1988) on the returnable drum.

Although formic acid is rarely used, its cost for 20 litres is $41.00 (Technical Grade;
Ajax Chemicals).

Moving Blocks Around the Laboratory

Most blocks are moved around the laboratory by good, old-fashioned, manual labour.
For the larger blocks, a portable hydraulic floor-crane manufactured by Fleetweld Engineering,
51-61 Maffra St., Coolaroo, Victoria has been used. The crane is effective, but its usefulness
is limited by size and a lack of maneouverability. A problem remains with how to deal with
the super-heavy blocks. In terms of field collection, and preparation, the larger the block, the
greater the chance of obtaining an undamaged specimen. Consequently, during the last field
trip to Bullock Creek, blocks in the 100-150 kg range were collected. Retrieval and transport
of these blocks was carried out in the following manner. A hole was drilled into the top of
each block, and a Ramset rock bolt was then hammered into it. A cable was then attached to
the bolt, and each block was simply winched off the outcrop and packed into a 44 gallon drum
for shipping. These rock bolts were extremely reliable and did not fail during the winching
operations.
Currently, one of the super heavy blocks is being etched by suspending approximately
7/8 of its length in a used 200 1 acid drum. The block is supported by a cable attached to the
original Ramset bolt, which is kept out of the acid to avoid corrosion and weakening. When
enough material has been removed, the block will be inverted and etched in a normal sized tub.
The only other method of handling such large blocks is by installing an overhead rail crane in
the laboratory. Rough estimates by crane specialists suggest that the cost for such a system
would be in the vicinity of $3,500.

186 - WHITELAW & KOOL

Fluid Movement Around the Laboratory

Major movement of fluids around the laboratory include filling the tubs, with both water
and acid, and emptying the exhausted solution after etching is finished. At Monash, this
involves the transfer of 7000 1 of water, acid, and exhausted solution during an etching cycle.
All tubs are filled through lengths of nylon garden hose from taps at the sink.

The acetic acid is obtained from a 200 | drum which has been tilted to a horizontal
position, and mounted on a drum rack. Flow rate is controlled by an acid proof nylon spigot
mounted in place of the 2.25 inch drum plug. Required volumes of acid are poured directly
into a 91 plastic bucket (domestic household type) and then transferred to each tub (Figs. 3A,
B).

A siphon system, again using garden hose, is used to remove spent acid from the tubs
on the bench. A pump, which is both acid proof and capable of handling light sediment loads is
used to empty the floor level tubs. The pump chosen was a Nikkiso 7 Centrifugal Pump,
costing $250.00. It employs a sealed magnetic drive to turn an acid proof plastic (nitrile)
impeller in a centrifugal action. This allows it to handle debris up to 5 mm in diameter. The
non-mechanical power train ensures that the motor will not burn out if larger particles jam the
impeller. The pump has a flow rate of approximately 35 l/min. and has been converted to a
self priming unit by the addition of a 5.0 1 header tank. The header tank has the added
advantage of acting as a sediment trap and prevents large sediment particles from being drawn
into the pump. The system was purchased from a firm called the “Pump Doctor” in
Moorabbin, Victoria. Pumps with higher flow rates which do not require priming, are
available and will probably be recommended, by many salesmen. They are not recommended
because they are rarely able to handle debris without incurring serious damage. Also, the high
rate of fluid removal does not allow blocks to drain slowly enough, and exposed bones are
subjected to excessive stress from water left behind in bone interstices.

Preparation Materials

Non-Consumables

The discussion of preparation materials will be divided into non-consumables and
consumables. Most of the non-consumable items are low cost items such as sieving fly-wire
mesh for sieving, brushes, scrapers, probes, and plastic containers. These are common in
laboratories or easily obtained from hardware suppliers and will not be discussed further. In the
same manner, common chemicals such as ammonia and the solvents, acetone and ethanol, need
no further mention.

Consumables

This section will list the important consumables, namely the foams, glues and hardeners
employed during processing. These preservatives are also used in the physical preparation
techniques that will be discussed later in this chapter. So, this review will serve for both
sections. The only addition to the list of glues previously discussed will be the cyanocrylate
super-glues. These are fast drying and very strong glues which are commonly used in physical
preparation techniques. They de-polymerize in acetic acid, and so, should not be used in
chemical preparation treatments. Many of the glues discussed are off-the-shelf items that may
be purchased in hardware stores. Others were found after consultation with Melbourne
suppliers of specialist plastics and glues. Firms which were interested in our preparation
problems sometimes supplied samples of their products to test. The industrial-sized samples,
commonly donated free of charge, have been enough to keep us operating for much of the past
five years.

PREPARATION TECHNIQUES - 187

Foams

(a) Polyurethane foams. These are available in 4 | cans, as a two part, foam and activator,
which can produce up to 270 | of foam. The volume of foam produced is strongly dependent
on the mixture ratio used. Acctone is the solvent when the foam is liquid, but it is insoluble
once cured. They are supplied by Daystar Pty Ltd, 1 Varman Crt., Nunawading, Victoria and
cost $42.00 per 4 1.

(b) Fomofil Ramset pressure pack. This is a "convenience" product for use when only
small amounts of polyurethane foam are required. Nozzle blockages and difficult operation
made this product inconvenient. It is also expensive at $13.00/30 1. Acetone is the solvent
es liquid, but it is insoluble when cured. It is supplied by Daystar Pty Ltd, Nunawading,

ictoria.

Glues

(a) Plastic glues. There are many "plastic" based glues suitable for bone repair. These
include the ethyl acetate and the ethyl methyl methacrylate glues used in the Monash
University laboratory. Tarzan’s Grip, an ethyl acctate glue, is commonly used and is available
in hardware shops at $2.50/tube. It dries clear, and acetone is the standard solvent. Tarzan’s
Grip appears to have a limited life span (approximately 10-15 years), after which time
shrinkage may ensue, thereby damaging bone (I. Stewart, pers comm.)

(b) Acrifix 92. This is a very strong plastic glue used for large bones or important repairs
in preference to Tarzan‘s Grip. It is an ethyl methyl methacrylate (see also Synocryl and
Vinalak below) which uses acetone as a solvent. Supplier is Daystar Pty Ltd, Nunawading at
$4.95/100 ml tube.

(c) Hotstuff. This is one of the cyanocrylate based superglues used to hold loose bone
fragments during physical preparation. It was purchased from Toy World, Swanston St.,
Melbourne at $10.50/ 30 ml bottle and must be applied via a teflon tube. The glue dries
within seconds, but may be dissolved with acetone. It is manufactured by HS-7, Satellite City,
California.

(d) Locktite Super Glue 3. This is a cyanocrylate superglue which is commonly available
in hardware stores. Acetone is the solvent. It costs approximately $2.00/ 3 ml bottle but,
unlike Hot Stuff, may be applied straight from the tube.

Hardeners

(a) Butvar. This is a polyvinyl butyrol glue, available in powder form. It has good bone
penetration when used in 3-10% solutions in ethanol and leaves a matt finish. Butvar,
although not as strong as the methlacrylate hardeners (described below), is preferred for health
reasons, since ethanol can be used as a solvent instead of acetone. Application is via a brush or
an eye dropper.

(b) Synocryl 9122X. This is a co-polymer (methyl butyl) methlacrylate, which may be
employed as a glue (30% solution) or as a hardener (3-10% solution) in acetone. It is very
strong and gives a matt finish at low concentrations, although higher concentrations will leave
a surface gloss. Synocryl is supplied by Cray Valley Products, Great Britian, and is the
recommended replacement for the widely used Bedacryl, which C.V.P. has taken off the market.

(c) Vinalak 63-513. This is a methyl butyl methacrylate which functions as a good
hardening agent at 5% concentrations in acetone. At higher concentrations, Vinalak has low
penetration and forms a surface skin. It was supplied by A.C. Hatrick Chemical Pty Ltd, 1612
Centre Road, Springvale, Victoria.

Preservatives

(a) Mowilith 144 and Mowilith 7001. These are uscd in a 4:1 ratio in a 10% solution,
with water for the final hardening of etched bone elements. Mowilith 144 is a styrene/acrylic
vinyl acetate, whilst Mowilith 7001 is a plastic/acrylic vinyl acetate. Bones are treated by

188 - WHITELAW & KOOL

either vacuum impregnation or overnight soaking in the solution. The combination of the two
products gives penetration (Mowilith 144) and strength (Mowilith 7001). With its high pH,
the Mowilith 144/7001 combination is used in preference to the standard P.V.A. (polyvinyl
acetate) glues, such as Aquadere. The more acidic P.V.A. glues will cause bone deterioration
over a long period of time (C. Cleeland, pers. comm.). Both Mowiliths were supplied by
Hoechst Australia Ltd, 606 St. Kilda Road, Melbourne, Victoria.

PHYSICAL PREPARATION TECHNIQUES

One of the most common forms of fossilization is petrification, a process in which bone
chemistry is altered, by silica replacement, whilst the structure remains unchanged. If a
silicified fossil occurs in silicate matrix, chemical preparation methods will not work.
Consequently, physical preparation methods, involving some method of mechanical matrix
removal, must be employed.

Mechanical methods involve the use of a tool to remove the matrix with the shape, size
and condition of the specimen controlling the tool choice. The range of tools varies from the
hammer and chisel, for coarse matrix removal, to the use of a fine needle or dental pick and
low-powered microscope, for the delicate teasing of individual grains from around a tiny bone.
There are also a large number of hand-held electric tools including, vibro-tools, electric "dental"
drills, rotary grinders and air-scribes, some of which will be described in this section.

The techniques described are those used by the Museum of Victoria and Monash
University (Melbourne) in the preparation of small vertebrate fossils from the Cretaceous
Dinosaur Cove locality, Victoria. Arkosic sandstones and mudstones are the typical lithologies
at this locality. These sediment types are common in many fossil deposits, and the preparation
techniques described in this section will be applicable or readily adaptable to a great variety of
other localities.

COLLECTING THE SPECIMEN

Fossils are recovered from the sandstones and mudstones of Dinosaur Cove by the
systematic breaking down of blocks with a hammer and chisel. Unfortunately, this usually
means that bones are broken into two or more pieces during discovery. When this occurs the
broken bone surfaces must be treated with a preservative to harden and consolidate the fractured
specimen (see hardeners described in the chemical preparation section). It is imperative that all
the bone fragments, and the surrounding matrix be collected and labelled, so that they may be
reassembled in the laboratory. Most fossils from Dinosaur Cove are relatively small and
protected by the natural mould of the surrounding matrix. Therefore, it is not necessary to
encase the fossil in plaster bandages, a common method of protecting larger specimens, Careful
wrapping, first in tissue or toilet paper, and then in newspaper will protect the specimen during
transport. Details of locality, specimen number and date should be recorded on the outside of
each package, and in a field book, to aid in specimen identification when laboratory preparation
begins.

PREPARATION METHODS

In the laboratory the specimen is unwrapped, and the bone and surrounding matrix
fragments are matched back together. This helps determine if any pieces of the specimen have
been lost and gives a general idea of the bone orientation within the surrounding matrix.
Excess matrix is then trimmed off with a rock-saw to reduce the block to a manageable size.

PREPARATION TECHNIQUES - 189

After cutting, the sawn surfaces should be carefully checked, for signs of any new bones
exposed by this work, and labelled accordingly.

The Dinosaur Cove material is much softer than its surrounding matrix and extreme care
is required during preparation. Bone must be continuously strengthened and hardened as matrix
removal proceeds. The hardening agents described in the chemical preparation section are
appropriate for this task. However, for the bonding of broken fragments, quick drying
cyanocrylate super glues are preferred over the slower drying methacrylate based plastic glues.
These are also discussed in the Glues and Hardeners section of the chemical preparation
techniques.

Matrix is normally removed with a vibro-engraver, but an air-scribe or rotary grinders
have also been used. The vibro-engraver recommended is a Burgess model 74, which removes
matrix with a variable intensity vibrating needle point. A low intensity vibration setting is
used for delicate work and allows gentle removal of matrix close to the specimen. Higher rates
of vibration can be used to remove larger volumes of matrix at a fast rate but should not be
used in close proximity to the specimen. High vibration rates, used too close to exposed
bones, will tend to damage or fragment the specimen. The tool should be held so that the
needle is perpendicular to the surface to be cleaned. Slow, gentle pressure on the point will
remove the matrix most efficiently. Heavy pressure will impede vibration and actually slows
the process. The tool can be adapted to hold a variety of points ranging from gramophone
needles to fine sewing needles. Gramophone needles are recommended, as they are made of a
harder steel than sewing needles and last longer.

This process creates a large amount of dust, which may obscure the specimen, and an
adaption described by R. Robison (Kummel & Raup 1965) is worth noting. Robison describes
the attachment of a flexible air hose to the vibro-tool to keep the specimen free of dust. This
hose is then attached to a small aquarium aerator or simply blown by mouth.

The rotary grinder used in preparation is an Arlec Super Tool, which may be fitted with a
variety of grinding heads. It is slower than the vibro-engraver and lacks the precision of a fine
needle point but has the advantage of reducing the specimen vibration to a minimum.

Some bones are too small or delicate to withstand mechanical vibration and may not be
prepared by motorised hand tools. In these cases, matrix must be removed by teasing it loose
with a hand-held needle or dental pick. This is a slow process, but necessary to achieve
satisfactory results.

Direct lighting, to highlight the surface of the exposed bone, is important - a small
flexible desklamp which allows light to be focused is recommended. Some bones are so small
that surface details are difficult to recognize with the naked eye. The use of an illuminated
magnifier (Magi-lamp) or low powered microscope (6-15x) is essential in these cases.

LABORATORY SAFETY

There are a number of safety aspects to consider with this type of preparation. Vibro-
engravers and air-scribes produce noise loud enough to be detrimental to hearing if used for
prolonged periods. Flying rock fragments produced by operation of these tools are also a
hazard. Therefore, ear-muffs and safety glasses are strongly recommended. Noise may also be
muffled by placing the specimen on a small, sand-filled cloth bag (Fig. 4). Rock dust from the
silicate matrix surrounding specimens can cause irreparable damage to lungs (silicosis) when
inhaled. Therefore, during sawing or grinding of matrix a dust-filtering face mask should also
be worn. There is a wide variety of suitable masks on the market, with many available at local
hardware stores.

190 - WHITELAW & KOOL

Figure 4. Removal of hard sandstone matrix from around dinosaur bone collected from Dinosaur Cove,
Victoria. Operator is using a vibrotool. (Courtesy of S. Morton and Monash University, Melbourne).

RECOMMENDED TOOLS

(a) Vibro-engraver Model 74. This tool is manufactured by Burgess-Illinois, 730
Waverley Road, Chadstone, Victoria.

(b) Air-scribe. The modcl used in our preparation was a Chicago pneumatic. Available
from suppliers of pneumatic tools and accessories.

(c) Rotary grinder. The Arlec Super Tool, which has a variety of grinding heads is
available at major hardware stores.

(d) Portable Dental Drill. This unit (motor, drill piece and flexible drive) is supplied by
Victorian Dental Supplies, Bourke St., Melbourne. A variety of drills, sawing blades and
grinders are available from the same supplicr.

FIELD TECHNIQUES.

SCREEN WASHING

Screen washing is an excellent method of recovering vertebrate fossils from
unconsolidated sediments. In particular, small, isolated mammal tecth are difficult to see in
unprocessed matrix and are often overlooked. This method employs a sieve screen to remove
the fine fraction matrix, thereby concentrating the size fraction which includes the vertebrate
material. The concentrate may then be taken back to the laboratory where it can be picked for
vertebrate remains. This method performs the dual functions of reducing the volume of matrix
that must be transported and increasing the concentration or visibility of any fossils that may
be present. Screen sieving may be donc dry, or in water, and has been successfully used to

PREPARATION TECHNIQUES - 191

recover material from a variety of sites ranging from ocean cliff exposures at Portland,

ane where previously, only macro-fossils had been known, to the dry playa lakes of South
ustralia.

ye oe: - = ——S I
A> L
Soi \, ff °

fy

45cm —
Figure 5. Schematic diagram of a screen-box used in processing unconsolidated matrix (from McKenna in
Kummel & Raup, 1965).

Sa

The amount of moisture, and the lithology of the matrix, will control the sieve method
to be adopted and the volume reduction achieved. If the matrix is dry and unconsolidated, dry
sieving often serves to reduce the sediment volume by at least 50%. If the sediment is
compacted or semi-indurated, dry sieving will have little effect, and a wet sieving process is
required. Ideally, a permanent, small stream is a perfect medium for washing, as the water is
continually replaced. However, depending on the locality, improvisation is sometimes
necessary to find a water source. Sieving has been successfully conducted thigh-deep in ocean
water, or by filling a trailer with water from a bore and using it as a mobile sieve tank. Always
be mindful of not polluting stockwater and check with the appropriate landowner before using
any available water supply.

Wet sieving of clay lithologies is often very successful, and volume reductions of up to
95% have been achieved. However, sediments may often need more than one processing cycle
to reduce volumes by this amount. If matrix can be placed on plastic sheeting and allowed to
dry, between cycles, the rate of reduction will be greatly increased. Some clay mineral
lithologies may resist the reduction process, because of static bonding between mineral grains.
This problem may be alleviated by drying the matrix after the first wash, and then washing it
in kerosene, drying it and sieving it again. Kerosene acts as a deflocculent, and breaks up
aggregated clay particles, which impede the breakdown process.

The amount of sediment placed in the sieve box should also be considered. Too much
sediment in the box takes longer to wash, longer to dry and is hard on your back!

The sieve boxes used are based on the same design as the washing box described by
M.C. McKenna in Kummel & Raup (1965) (Fig. 5). The box is 45x45x30 cm in size and

192 - WHITELAW & KOOL

has a wooden handle attached diagonally to its open top. Aluminium fly-wire is stapled to the

bottom of the box and supported by a single piece of heavy gauge 1 cm galvanised iron mesh.

Two opposite sides of the box measure only 20 cm deep, leaving a 10 cm gap, which is also

covered with the flywire and mesh. This allows water to flow freely in and out of the box as it

ee from side to side and allows for better drainage when the boxes are stacked for
ing.

A mechanised screen sieve has also been successfully employed by one of the authors
(Whitelaw). This system is based on a rotating drum sieve, tilted at a shallow angle, which is
continually sprayed by a series of water jets. Matrix is fed into the top end of the machine and
tumbled in the water spray until it emerges as concentrate at the other end. The method is
rough on bone material and should only be used for recovery of strong and already disarticulated
fossils where large volumes of matrix must be processed. It was successfully used to process
80 tonnes of matrix, from which the Dog Rocks Local Fauna, Geelong, was collected. This
fauna is mostly represented by single teeth and small bone fragments which had undergone a
violent transportation history before being deposited as part of a channel fill sequence. The
material was very hard and showed no sign of damage from being processed by this machine.

PLASTER JACKETING

When collecting vertebrate fossils it is often necessary to protect the exposed bone(s)
with a plaster jacket (Macdonald 1983). Fragile, broken or articulated bones need stabilization
and protection during transport and a plaster jacket is ideal for this purpose. The materials
necessary to make a plaster jacket are:

(a) A strong, open weave material e.g. hessian sacks torn into strips approximately 7 cms
wide and long enough to cover the height and width of the specimen.

(b) Plaster of Paris. The preferred material is dental plaster, a fine quality and quick setting
plaster, but Plaster of Paris of any quality is acceptable.

(c) Enough water to mix with the Plaster of Paris.

(d) A container large enough to hold the Plaster of Paris solution, generally a plastic wash
basin or bucket, will suffice.

The bone(s) to be jacketed should be exposed, as much as possible, to determine
orientation, size, shape and depth of burial, The exposed bone should then be hardened with a
preservative (see Chemical Preparation section). If the specimen is damp, plastic based
preservatives will not pentrate the bone surface and a water soluble glue such as Aquadere
(P.V.A.) should be used.

A trench should then be dug around the specimen to a depth several centimetres below its
base. Initially, a wide margin of matrix should be left around the specimen. This will prevent
any dislodging of the fossil during trenchwork and can be trimmed back when the trench is
finished.

After the trench is completed the specimen should be moistened and covered with wet
paper (newspaper, tissue paper or toilet paper). The layer of wet paper acts as a barrier between
exposed bone and the plaster jacket, thereby stopping plaster from adhering to the specimen.
Any depressions, hollows or narrow spaces between bones must also be packed with paper to
prevent clots of plaster from forming in these places.

The next step is to prepare the material strips and the Plaster of Paris solution. The
strips of open weave material should be soaked in water. Dry strips will tend to draw water out
of the plaster solution and may retard setting and proper hardening (Macdonald 1983). The
Plaster of Paris solution is best prepared by slowly adding plaster to a bucket of water until it
reaches a consistency of soft, thick, pea soup. The material strips are then soaked in the
plaster, one at a time. As they are removed from the plaster, they should be pulled between
two fingers to remove excess plaster. The first strip is placed in the middle of the specimen,

PREPARATION TECHNIQUES - 193

smoothed down and pressed snugly into depressions. Additional strips are each placed so that
they overlap the previous one by about 2 cm, and so that they work towards each end of the
specimen whilst also extending down to the base of the trench. A second layer of overlapping
strips is placed at right angles to the first. If additional structural support is necessary, lengths
of wood can be placed between the two layers of plaster or further layers of plaster strips may
be added. After this is complete the jacket should be left for approximately an hour to set.

Once the plaster has set, the base of the trench is undercut until the specimen is standing
on a pedestal. Undercutting should then continue until the block breaks loose from its base.
With large specimens a tunnel should be cut underneath the specimen, and plaster strips should
be passed through this and bound to the plaster jacket to provide extra support. Extreme care
should be taken when turning over the block. If the jacket does not form a snug fit, the block
may fall out, and a valuable specimen may be damaged or lost.

Once overturned, any excess matrix and plaster can be trimmed to reduce weight. The
underside is then moistened and plastered in the same way as the upper jacket to entirely seal
the specimen. It may then be transported to the laboratory where the jacket may be cut open
and the fossil prepared.

CASTING AND MOULDING TECHNIQUES

The casting of specimens, although not new, has only become a science in its own right
in the past forty years. This is largely due to breakthroughs in the development of polymers
and plastics. These materials have allowed faithful reproductions of original, and often rare,
specimens which preserve details that may only be seen with an electron microscope. Casts
may be used as alternatives to original specimens for display purposes, as teaching aids or for
research, thus protecting the original specimens from possible damage.

Before a cast can be made, it is necessary to make a mould of the original specimen.
The mould type and complexity will depend upon the size and shape of the fossil and can vary
from a simple latex peel to a multi-piece mould.

PREPARATION OF SPECIMENS FOR MOULDING

When preparing a specimen for moulding, all loose material such as dirt, dried glue and
loose matrix, must be removed from its surface. Cleaning aids such as acetone, alcohol, dilute
acetic acid or soapy water may be used. Once clean, the specimen may need to be hardened and
any large cracks or missing sections filled. A substance that will not react with the
moulding medium must be used. Water soluble putty, employing a Polyethylene Glycol 4000

Figure 6. Preparation of moulds and casts for production of replicas of original fossil material. A, emplacing
plasticene wall around the mid-line of the bone; B, brushing vaseline on silicone mould to prevent top mould
from sticking to bottom mould; c, pouring the urethane foam into the prepared mould; d, expanding foam
escaping from mould; E, removing silicone mould from epoxy cast. (Courtesy of F, Coffa and the Museum of
Victoria, Melboume).

194 - WHITELAW & KOOL

A

PREPARATION TECHNIQUES - 195

196 - WHITELAW & KOOL

E

base, is ideal for filling such voids, and will not react with silicone rubber moulding mediums.
Hardeners, such as Butvar B98 or Vinalak, should be used to impregnate and seal porous
specimens. This will prevent the moulding medium fro: adhering to the bone as it sets.
Three different moulding materials and their relevant preparation techniques are discussed below.

Latex Peel

Latex peels are useful for making copies of fine surface detail exposed on bones.
Essentially, they are two dimensional moulds, but quality peels often show more surface detail
than can be seen on the original. Peels are made from latex rubber by the following procedure.

First, the exposed areas of the fossil have to be prepared so that the latex will not stick
to the bone surface. Butvar B98, which has been previously mentioned, is excellent for sealing
surfaces and, thus, preventing sticking, but any consolidant which is not water soluble will
suffice.

Once the specimen's surface has been consolidated and sealed, a test area should be
moistened with a solution of water and 5% ammonium hydroxide. If the solution turns red, the
bone has not been completely sealed, and more preparation of the surface is indicated. If no
reaction occurs, the whole specimen should be covered in a thin film of the ammonia solution.
Several drops of latex are then applied to the bone and spread across the surface by blowing the
latex with a straw. This procedure ensures that the latex fills all the fine surface structure and
avoids introducing air bubbles into the latex. It should be repeated until the entire specimen is
covered by several coats of latex. Each layer should be applied before the previous one has set,
so that a cohesive rubber surface is built up. The final layer should consist of a combined latex
reinforced by cotton wool or gauze, which is added and allowed to dry. This final layer adds

PREPARATION TECHNIQUES - 197

some rigidity to the mould and helps prevent excessive shrinkage. When the latex is
completely dry, carefully peel it off the specimen to produce a high-fidelity mould or latex peel.
Latex peels have a limited life-span, but provided they are kept away from sunlight and

moisture, they will last several years. The major disadvantage of latex peels is distortion.
Original dimensions will alter, generally shrinking, and it is recommended that casts be made
within the first six months of the peel's life. If required, latex peels may be temporarily
expanded or enlarged, to help emphasize fine surface details, by soaking them in paraffin oil or
kerosene.

Two brands of latex rubber are readily available in Victoria. They are:

(a) Betatex, available from Beta Chemicals Pty Ltd, 121-125 Northern Road, West
Heidelberg, Victoria.

(b) Revultex, available from Revertex Industries, 161 Westall Road, Clayton, Victoria.

Silicone Rubber

Silicone rubber is more expensive than latex rubber but is the more popular medium as
it undergoes little distortion or deterioration with age. It also possesses high tear strength, will
not adhere to bone and has a high pattern complexity which allows duplication of very fine
surface detail. It is available as a two component medium (base and catalyst) and its only
disadvantage is the short shelf life, of approximately six months, of the two unmixed
components (approximately six months).

The first step in making a silicone mould is to build a plasticine wall or dam around the
specimen to contain the moulding medium until it sets (Fig. 6A). To make a cast of an entire
specimen requires at least two moulds, one upper and one lower. Therefore, the plasticine wall
is built up to only the midline of the specimen. The silicone rubber is then prepared by
mixing the base with the catalyst in a ratio that can vary between 3% and 8%, giving the
preparator a range of working times in which to apply the rubber. The rubber is then poured
onto the specimen until the top of the dam wall is reached. Once the silicone rubber has set
(approximately 24 hours: Fig. 6B), the first mould is turned upside down and the exposed
rubber is brushed with petroleum jelly to prevent it from sticking to the second application.
Again, a plasticine wall is necessary to contain the silicone while it sets. When the second
half has set the two mould halves may be pulled apart and the specimen removed. At this
point a small opening must be cut into one, or both of the moulds. This will be the pour
hole which will allow the introduction of the casting material into the mould.

Dow Corning produce and import a wide range of rubbers in Australia that are suitable
for moulding. These are available through retail outlets in the eastern states such as Daystar
Pty. Ltd.:

(a) 1 Varman Crt., Nunawading, Victoria.

(b) 396 Princes H'way., Rockdale, New South Wales.

(c) 239 Brisbane Rd., Labrador, Queensland.

As of November 1988, the Q3-3321 rubber, which was the preferred type, is no longer
available. However, Daystar Pty. Ltd. have supplied a replacement, Q3- 3481, and early results
indicate that it is a viable alternative. A 5 kg tub costs $227.90 (trade price).

Plaster of Paris - Mould Supports

Silicone rubber moulds are not very rigid, and need to be supported in a plaster jacket
when casting. The jacket prevents mould distortion and ensures a perfect fit when the two half
moulds are joined. The two halves of the plaster jacket may be made in the following manner.
The original specimen should be enclosed in its two half moulds and placed on the preparation
bench. A plasticine dam, similiar to that made for the silicone mould, should be built to a

198 - WHITELAW & KOOL

height of half the total mould and a diameter slightly greater than that required for the plaster
jacket which will enclose it. The mould pour hole should be protected at this stage so that it
too is not filled in. The dam may then be filled with plaster which can be further reinforced by
the addition of sisal fibre, canvas or metal supports whilst it is still wet. After the plaster has
set, the mould can be inverted and the second half of the support can be made in the same
manner. A certain amount of heat is generated by setting Plaster of Paris, but this will cause
no damage to the enclosed mould or fossil. When the plaster has set and is completely cold,
the specimen can be removed and safely stored, leaving the mould ready to make casts.

CASTING FROM MOULDS

Casting involves filling a mould with a liquid which will take on the shape of the mould
as it hardens, thereby creating a copy of the original specimen. A wide variety of casting
materials are available including Plaster of Paris, polyurethane, epoxy resins, polyesters, low-
melt metals, flow-moulded vinyls and, virtually any other substance that will set hard from a
liquid. The first three are the most widely used materials in our laboratory and they will be
described in this section.

Plaster of Paris

Plaster of Paris is a versatile medium, that is durable, inexpensive and readily available.
Dental plaster, a high quality and quick drying variety, is recommended as it produces tough,
hard casts with good preservation of detail. It is probably the simplest of all casting materials
to use, and is prepared by simply adding it to water until a thick, viscous fluid is produced.
The mould is prepared by enclosing the two halves in the mould support and by binding the
assembly with elastic bands or masking tape.. The liquid dental plaster is then introduced into
the mould cavity, slowly to avoid air-bubbles, through the pour hole and allowed to set. The
cast will be ready to remove from the mould in approximately half an hour.

The disadvantages of plaster casts are that they can be chipped or broken relatively easily
and that large casts are heavy and difficult to mount. Because of this, another medium,
polyurethane foam, is becoming increasingly popular for display purposes.

Polyurethane Foam

Polyurethane foam is a relatively new casting medium which was first used on a large
scale by the Museum of Victoria in 1981. The museum was given the opportunity to cast
complete skeletons of two Chinese dinosaurs, Mamenchisaurus and Tsintosaurus and the
preparators chose polyurethane foam as the casting medium because of its light weight, solid
form and ease with which casts could be made (Kelly 1983), (Figs. 6C, D). The foam selected
was an I.C.I. (Imperial Chemicals Industries.) product - Daltolac SW6\Suprasec 5005 rigid
urethane.

For casting, the mould is prepared in the same manner as that described for plaster
casting. Silicone rubber moulds are recommended for this work as polyurethane foam exerts a
high tear stress as it expands. The urethane foam, supplied as two inert liquid components, is
mixed in equal parts, by weight and poured into the mould before the onset of foaming. This
allows the urethane to mix, expand, and distribute evenly throughout the mould as it forms.
Apart from the volume of mixture used the amount of foam required to fill a mould is a
function of the amount of mixing and the ambient temperature. Over estimates of the amount
of material used, and consequent waste of foam, may be avoided by incrementally filling the
mould with a series of small pours.

PREPARATION TECHNIQUES - 199

Epoxy Resin

Because of its strength and durability epoxy resin is a popular casting medium. Again
the mould is prepared in the same manner as the plaster cast mould. The medium, which is
usually available as a two part resin and hardener combination, is then mixed in a 5:1 ratio, by
weight. If desired, coloured dye may be added at this stage, to give authenticity to the cast.
Once mixed, some of the epoxy is introduced into the mould and the mould is centrifuged to
create a smooth film of epoxy over the mould surface. This eliminates air-bubbles that will
detract from the quality of the cast and enhances the filling of small crevices that preserve the
fine details of the specimen. Once centrifuging is complete the remaining epoxy may be
poured into the mould and allowed to cure. If two half moulds are to be used to cast a
specimen, each half should be filled and then joined together after approximately two hours, or
when the epoxy has become tacky. The mould or moulds may be removed after approximately
24 hours when the epoxy should be totally cured (Fig. 6E).

ACKNOWLEDGMENTS

Thanks go to Craig Cleeland for his major contribution to the section on casting and
moulding and to Ian Stewart for his advice on latex moulds. Jenny Monaghan gave help and
support in the preparation of both the Bullock Creek and Dinosaur Cove material, and the
gentlemen in the Monash University Earth Sciences Department workshop assisted greatly in
setting up the Acid Lab. Guy Royce, Laboratory Manager in the Earth Sciences Department
at Monash University, has graciously dealt with the many logistical problems in the running
of the Acid Lab. Many thanks are also due to Julie Whitelaw for her hard work in reviewing
the manuscript. The technical work discussed in this chapter is financed by grants from the
Australian Research Council, the National Geographic Society and the Sunshine Foundation.
Finally, many thanks to Pat and Tom Rich, Jenny Monaghan and Bob Baird for their patience
and understanding in editing this manuscript.

ADDENDA

Two other techniques not discussed in the text need to be briefly mentioned. One is the air-
abrasive technique and the second is the use of hydrochloric acid in total removal of bone. The
air-abrasive tool projects a stream of air-borne particles under pressure at a given target. The
particles can vary in size from the very finest powder, such as sodium bicarbonate, to a coarse
compound like aluminum oxide, depending on the result required (see G.F. Stucker, M.J.
Galusha & M.C. McKenna’s article "Removing matrix from fossils by miniature sandblasting"
in Kummel & Raup 1965). The hose and nozzle through which the particle-laden pressurized
gas is fed should be sealed in a glass-topped chamber. The stream of particles can be played
onto the surface of the matrix in a sweeping action, effectively removing the most stubborn
matrix. It is advisable to practise with the tool before using it on important specimens, as
incorrect-sized particles or too high a gas pressure can quickly ruin a specimen.

Hydrocholoric acid can be usefully employed when matrix surrounding a fossil is so hard
and/or acid resistant that it makes preparation impossible, or in cases where the fossil is poorly
preserved. Dilute HCl can remove the bone completely leaving behind a natural mold of the
original material, This mold can then be filled with latex to produce an accurate, durable
replica of the original, which can then be studied (see J.K. Rigby & D.L. Clark's article
"Casting and Molding" in Kummel & Raup 1965).

200 - WHITELAW & KOOL

REFERENCES

BEHRENSMEYER, A.K., 1987. Vertebrate preservation in fluvial channels. Palaeogeo., Palaeoclim.,
Palaeoecol., 63: 183-199.

FELDMANN, R.M., CHAPMAN, R.E., HANNIBAL, J.T. , eds., 1989. Paleotechniques. Paleontol. Soc. Sp.
Publ. 4, Univ. Tennessee, Knoxville.

JEPPSON,L., FREDHOLM, D. & MATTIASSON, B., 1985. Acetic Acid and Phosphatic Fossils - a warning. J.
Paleo. 59: 952-956.

KELLY, K.A., 1983. Workshop Manual - The Moulding and Casting of Dinosaurs. Mus. Victoria, Melbourne.

KUMMEL, B. & RAUP, D., 1965. Handbook of Palaeontological Techniques. W.H. Freeman & Co. San
Francisco & London.

MACDONALD, J.R., 1983. The Fossil Collectors Handbook. A Paleontology Fleld Guide. Prentice-Hall, Inc.,
New Jersey.

RICH, T.H. & RICH, P.V., 1989. Polar dinosaurs and biotas of the Early Cretaceous of southeastern
Australia. Nat. Geog. Res. 5(1): 15-53.

RIXON, A., 1949. The use of acetic acid and formic acids in the preparation of fossil vertebrates. Mus. J.
London 49: 116-117.

RIXON, A., 1976. Fossil Animal Remains, Their Preparation and Conservation. Athone Press, London.

CHAPTER 7

PREDICTING THE DIET OF
FOSSIL MAMMALS

Gordon D. Sanson !

Introduction’. wosie42 058 ics scecd dee. 202
Teeth as Adequate Predictors of Diet ................. 203
Physical Properties of Food, and Tooth

Form and Function ............ccccesceceeeeeees 205
The Interaction of Mechanical and

Chemical Digestion ........0......ccccceeeeee ee 207
Body Size, Metabolism and Energy

REQUITEMENtS 2.8: cee lee eee eee 208
BehaviOurs oss ce. cbss ces ie Peine oivonc deen sa eee 209

Tooth Wear and Striation Pattern and Structure...210
Thylacoleo: Predicting the Diet of a Unique

Fossil Without Living Relatives.....00......... 210
Propleopus: Predicting the Diet of a Unique
Fossil with Living Relatives... 213

Fossil Sthenurinae and Macropodinae:
Predicting the Diet of Fossils with

Living Relatives .........c.cccccecesceecseceseceeenens 215
CONCIUSION Pets teetest od idve tees odsthaticw esse eases: 22]
Acknowledgement...........ccceccesecccesecceneeceeeces 224
RELCTENCES Ho. vocis sSesaisee nob ne seoahente SetUbeeseloctios ese ss 225

1 Department of Zoology, Monash University, Clayton, Victoria 3168, Australia.

202 - SANSON

INTRODUCTION

Traditionally, hypotheses about the diet of extinct vertebrates have flowed from the
principle that animals with similar digestive tract morphology, which includes teeth, are
assumed to have similar diets. With the reservations and conditions discussed in this paper,
this approach is the most fruitful, being based on analogy. Convergence of feeding and
digestive system structure is common and occurs across diverse groups, which strongly
supports the idea that important constraints operate and must be met for a particular resource to
be effectively utilised. Predictions of diet based on relationship and homology of the digestive
tract are weaker, as there are numerous examples of closely related forms which differ in their
dictary preferences. Unique forms, by definition, have no analogues. This immediately raises
serious questions about the validity of even attempting to predict the diet of unique fossil
forms. The hypotheses, like any prediction about fossils, can never be tested, except in those
very rare cases where stomach contents are preserved. There is always the danger of producing
ad hoc hypotheses in biology (Gould 1977, Lewontin 1978, Gould & Lewontin 1979), and this
is particularly true about fossils. Kay & Cartmill (1977) and Kay & Covert (1984) address
these issues particularly well and propose criteria which must be satisfied before adaptive
meaning can be ascribed to a particular adaptive trait.

Since indications of the soft parts of the digestive tract are rarely preserved, it is usually the
teeth, and to a lesser extent jaw structure, that are relied upon to infer diet. It is generally
assumed that teeth and jaws do reflect diet very closely. Kay & Hylander (1978) contend that
since dental structure is influenced largely by selection for optimal designs for acquiring and
breaking down food, differences in morphology most often can be related to the physical
properties of the foods. Following this argument, detailed knowledge of functional tooth
morphology should allow accurate prediction of dict.

The reasons for expecting that minor differences in tooth morphology should reflect the
physical properties of the diet will be briefly explored in this paper. The limits to the dietary
information which dental systems can supply will also be examined. It will be argued that
while many good correlations between diet and dentition exist, it does not follow that diet can
always be confidently predicted from teeth.

The class of vertebrate will affect the capacity to make dietary inferences. Fish, amphibians
and reptiles, being ectotherms, have a lower basal metabolism than the endothermic birds and
mammals. In addition, it appears that being herbivorous places different constraints on
digestive systems than being carnivorous. Carnivores essentially consume items with the
same biochemical components as themselves (Mellett 1982). Animals which consume plant
cell wall must cope with a material which is abundant and foreign to them in terms of its
physico-chemical properties, and for which they do not make an enzyme. In this context the
plant cell wall provides a barrier to digestion by animals. Once the cell wall is digested or
penetrated, the cell contents are much more similar to animal tissue and provide no major
impediment to digestion. For cell wall digestion, however, a symbiotic relationship with
microorganisms needs to be established and maintained. This latter capacity is best developed
in mammals but also occurs in a few birds, reptiles and fish.

Fish demonstrate a wide range of jaw and dental adaptations. Some of these can be related
to diet. Amphibians and reptiles show less dental variation, and there is little information
comparable to that available for mammals. Most birds, of course, have no teeth, but beak
morphology may indicate diet to some extent. There is no reason to suppose that in all these
forms gut morphology is not also an important part of the total digestive process. However,
the data is much better in mammals, and only mammals will be discussed further in this paper.

PREDICTING FOSSIL MAMMAL DIETS - 203

TEETH AS ADEQUATE PREDICTORS OF DIET

Kay & Hylander (1978) point out that different species, with different heritages may have
analogous, but not homologous, morphologies that process particular foods. So, species
which have evolved to eat foods requiring large amounts of shearing during mastication need
not necessarily emphasise the same molar shearing blades, but will have shearing capacity.
They argue that teeth do reflect diet.

Eisenberg (1978) has challenged these assumptions, citing Vorontsov (1962) and Carleton
(1973) as indicating that adaptations of the gut for increased microbial activity do not
necessarily proceed in step with the evolution of dental modifications for the mastication of
plant material. Vorontsov concludes that some organs of the digestive system in a species can
be at essentially different levels of specialization. He argues that in the Gerbillinae there is
wide variation of molar structure, while stomach structure exhibits little variation. In the
Microtinae, on the other hand, the variability of stomach structure is wider than that of the
molars, illustrating the phenomenon of the compensation of one organ in a system by another
organ. Eisenberg (1978) concludes that since gut morphology cannot easily be deduced from
fossilized hard parts, and since gut modifications do not necessarily evolve in step with dental
modifications, the reconstruction of the evolutionary history of diet is fraught with difficulties.
This follows from the clear understanding that it is the total digestive system, gut and
masticatory apparatus, which constrains or influences diet (Mellett 1982, Sanson 1985, and for
an excellent holistic treatment see Fortelius 1985).

Kay & Hylander (1978) argue that Vorontsov's conclusions should be viewed with caution
at three levels. First, many of his statements cannot be checked, because he presents no dietary
information on the species he studied. Second, at least for gerbils, increased molar complexity
is correlated with increases in the length of the caecum. Third, Kay & Hylander (1978) argue
that, with respect to Vorontsov's conclusions, it is fundamentally important to remember that
dentitions are mechanical food processing devices, while the gut is an assimilation, chemical
processing and elimination system. They point out that the dentition is responding
evolutionarily to the physical properties of foods, while the gut is responding to the chemical
properties of ingested food. These arguments are important, because Kay & Hylander (1978)
are attempting to establish that dental morphology is an adequate and true indicator of dict.
They seek to demonstrate that minor dental variations correspond remarkably well with what is
known of dietary preferences of species in the wild. There is a great deal of work which tacitly
or implicitly relies on these kinds of conclusions.

Kay & Hylander make some important and relevant points, but it is not clear that
Eisenberg's concerns, based on Vorontsov and Carleton's work, should be dismissed too readily.
Kay & Hylander's first criticism of Vorontsov can and should be assessed by further analysis.
It is true that Vorontsov does not give dietary information about the particular species he has
examined. Rather, he discusses the species in the context of a broad shift from “high-caloric
but hard-to-get food (seeds to small invertebrates) to little-caloric but easy-to-get food
(vegetative parts of plants)" (p. 360), and merely lists the species along morphoclines relating
to his notion of the dietary shift. However, if Vorontsov is correct, and gut and molar
morphology do not necessarily correlate, then, without diet information, it is difficult to know
whether different animals, with similar dentitions but different guts, are on different diets or
gain different benefits from the same dict. Alternatively, it might be inferred that different
animals, with similar dentitions and guts, are on the same diet, or derive different benefits from
different diets. This is particularly likely if gut morphology is adaptive. Therefore, the lack of
data on diet is not critical to Eisenberg's point.

Vorontsov's data on teeth is difficult to interpret. The clearest statement is that in the
gerbils, for example, there is a trend in increased lophodonty and hypsodonty in changing to

204 - SANSON

rough "cellular" food, “trom Gerbillus and Monodia through Meriones and Psammomys to
Khombomys" (p. 363), Unfortunately, Vorontsov's Fig. 6, illustrating the variation of the
gerbil stomach, does not seem to be referred to in the text, nor is there a discussion of the hind
gut in the text, However, he does pive figures of the length of the small, and large, intestine,
and caccum as a percentage of the whole put. These figures are graphically portrayed in Fig. 1.
The perbil species are arranged not as in Vorontsov's ‘Table 1, but in his sequence of changing
molar structure relating to an increasingly vepetative plant diet. Vorontsov gives no means, or
standard errors, and, since all theee Components must add up to 100%, there are corollorics
involved, Statistical treatment of his data is, therefore, difficult. However, it is not clear from
Vorontsov's data, represented in Fig. 1, how Kay & Hylander (1978) could conclude that

\\
P.o R.o

Gd Bp Ms Mp M.t Me ML M.v Mim M.u M.e M.t
Species of Gerbillinae

AN smattint | l/large int lif caecum

Figure ft. Graphical representation of gut parameters of the Gerbillinae presented in Vorontsov (1962),
Ihe data is extracted from ‘Table 1, Vorontsov (1962), and indicates the relative lengths of the small intestine,
large intestine and cacoum, of vartous gerbil species, expressed as a percentage of total gut length, The
gerbil species are arranged from left to right in Vorontsov's sequence of increasing hypsodonty of the molars,
Le. “Gerbillus and Monodia, through Mertones and Pyammomys to hypsodont Rhombomys" (p, 363).
Vorontsov gives no data for Monodia, Vorontsov does not specify where Brachiones and Tatera fit on his
hypsodont meorphooling but they are plotted in Fig, loin the relative position suggested by the data im
Ellerman (1940), ‘The Meriones species are plotted in the sequence in Vorontsov's ‘Table 1. Even if the
position of Tafera and Brachiones are altered, it does not alter the pattem very much, Vorontsov appears to
have plotted the gerbils in order of decreasing relative length of the small intestine, and not taxonomy or
caceal length, Abbreviations: Tit, Tatera indica, Gp, Gerbillus pyramidum, Gad, Gr dasyurus, Bop, Brachiones
preewalvkui, Mis, Meriones shaws, Mop, M. persicus, Mit, M tamarivcinus; Moe, M. erythrourus; M.l, M
Libyous; Mov, Mo vinogradovi; Mam, M. meridianus, Muu, M. waguiculatus; Mic, M. crassus; Mat, M. tristrami;
Po, Prammnomys obexus, Roo, Rhombomys Opimus,

Percentage of total intestine length
Bo

6O

20

\

O

“increasing molar Complexity (as expected tor increased dietary fibre) is correlated with increases
in the length of the caecum, suggesting that, contrary to his [Vorontsov's] interpretation, these
two systems are indeed in step with each other" (p. 174). On the contrary, caecum length does
not seem to vary with dental complexity, supporting Vorontsov. The point is more marked
when Tufera is considered, Ellerman (1940) notes that the genus has molars which are
originally cuspidate, simpler than Gerbillus and less hypsodont than Meriones and other

PREDICTING FOSSIL MAMMAL DIETS - 205

gerbils. Yet Tatera has a caccum longer than Rhombomys, the gerbil with rootless cheekteeth,
Similarly Psammomys though having strongly hypsodont molars, and a molar pattern like that
of Meriones, has relatively the longest caecum,

Kay & Hylander's third point is interesting. It is probably true, as they argue, that the gut
is responding to the chemical properties of ingested food in it. However, the level of chemical
Tesponse will be affected by exposure of adhesion sites to bacteria, and cellular walls and
contents to enzymes. This exposure occurs by physical processing. There is evidence that the
two can be tightly linked (see Sanson 1985 and in press), but also, as Vorontsov argued, that
they can compensate for each other.

Dentitions wear, and as they wear they may change their efficiency (Gipps & Sanson 1984,
Lanyon & Sanson 1986) and may affect diet selection (McArthur & Sanson 1988). Yet
animals manifestly continue to live and reproduce and do not necessarily alter their diet with
age. In many cases the differences between an unworn and a worn tooth may exceed
interspecific differences between unworn teeth. It must be concluded that animals can
compensate for ontogenetic changes in the dental system. The gut can change its form
depending on the quality of the diet, even over short periods of time (Gross ef al. 1985). Many
species change their diet quite markedly on a seasonal basis depending on food availability (e.g.
Leuthold 1977). The degree of compensation may be affected by differences in body size and
the associated changes in metabolic requirements, relative spatial and temporal availability of
preferred dietary items, and behaviour. The latter factor is particularly difficult to predict and
may confound attempts to accurately infer diet. It could be argued that Kay & Hylander's third
concern about Vorontsov's conclusions do not materially weaken and if anything strengthen
Eisenberg's reservations.

Eisenberg's (1978) concerns that one should look with caution on dietary inferences based
solely on the morphology of a single system, and that such inferences should be checked
against other systems, seem to be justified. Since it is not possible to confirm the diet when
fossils are studied, unequivocal interpretations of feeding may be impossible. Kay &
Hylander's general conclusion that minor dental variations correlate remarkably well with what
is known of dietary preferences are reasonable. However, it is not necessarily always so,
because dental morphology can be compensated for, and with fossils we will never know when
a compensation is operating.

PHYSICAL PROPERTIES OF FOOD, AND TOOTH FORM AND
FUNCTION

In mammals with heterodont dentitions, ingestion is mostly performed by the front tecth,
the incisors and canines, which can act to grasp and hold dictary items. Sometimes, aided by
the premolars, these teeth can begin preparation for digestion. They tend to grasp and kill prey
animals, tear and scrape bark and exudates and open fruits and seeds. The anterior teeth are
more directly correlated with the gross structure of foods and may more directly reflect the
ecological adaptations of a species (Maicr 1984).

In addition teeth, more usually the posterior molars, may process the food by reducing it in
size. Therefore, molar structure is often considered to be related to the dict, generally in broad
terms. However, Maier (1984) argues that we should avoid using such generalisations as
omnivory and folivory; instead we should ask what is happening physically between teeth
when specific foodstuffs are being masticated. This has been generally considered to occur ina
series of actions defined as puncture-crushing (Crompton & Hiiemae 1970) and crushing,
grinding, and shearing (Rensberger 1973). Terms such as grinding and shearing, are gross
terms which apply to the relative movement of certain cusps or regions of tecth. The
definitions of these terms refer particularly to the relative movement of teeth, not so much to

206 - SANSON

the consequent effect on food, although, of course, there is a relationship, and certain inferences
are usually drawn.

These definitions and descriptions of tooth action are also too general. While they have
focused attention on the detailed action of the tooth, it is not really clear, for example, why
shearing fibrous composites is better, in terms of food preparation for consequent chemical
digestion, than grinding or crushing, allhough some work assumes that it is. More work is
needed to address these problems. Rensberger (1973) has made some important contributions
in this area.

When food is masticated by teeth, there are physical laws and processes which operate, both
on the teeth and the food. The shape and structure of the tooth will be influenced by these
laws. Analysis of the process of breaking food particles, and the effect on teeth, have recently
adopted a more detailed approach. Wainwright et al. (1976) and French (1988) are introducing
engineering and design concepts to the understanding of the properties of food materials and
teeth, while Lucas (1979, 1980, 1982) and Lucas & Luke (1983, 1984) in particular and others
(e.g. Sheine & Kay 1977, Kay & Sheine 1979, Rensberger 1973) have begun to apply this
knowledge to understanding tooth function. Fortelius (1985) concisely reviews this work.

The properties of biological materials and their reaction to loads has been treated by
Wainwright er al. (1976), French (1988), Gordon (1968, 1978) and Lucas (1979, 1980, 1982)
and Rensberger (1973) and are very briefly summarised here. Food, like any other material,
fractures, by the propagation of cracks, or divides, by the separation and flow of material, upon
the application of force. The amount of force required will vary, depending on the area over
which it acts, and the physical properties of the food. When material is compressed by a tooth,
a load is applied which causes a stress, measured as force per unit area. The result is a
deformation or strain, which is measured as the change in length of the material divided by the
original length. Hookean materials are those where the stress is proportional to the strain,
which is initially elastic when the strain is fully recoverable. The ratio of stress to strain over
this clastic range is Young's Modulus of Elasticity, E. If the applied stress is maintained, the
matcrial enters a phase of plastic deformation, which is not recoverable and eventually the
material will fail. Many biological materials, particularly softer tissues, are non-Hookean in
that there is initial plastic deformation followed by an elastic phase. Eventually, however, the
material will still fail. The area under the stress/strain curve is a measure of the work that
must be done in order to break the material. In other words, it represents the amount of energy
that the material can absorb before it fails.

Hard, stiff, brittle materials have an E approaching 1. They can absorb an enormous
amount of stress with very little strain. However, they can only absorb a little stress once the
material enters the plastic phase of deformation, and so they tend to shatter. Crystals are a
good example. Ductile materials, with a very low E, tend only to have a small range of elastic
deformation but can absorb much more energy in the plastic phase before failure than a brittle
material. This means that brittle materials, such as tooth enamel, fail at a higher stress than
ductile materials, but, depending on the material, it may take less work. Since stress is force
per unit area, sharp cusps, which have a small surface area of contact, generate very high
stresses at their tips and in the material they are penetrating. All materials obey these laws,
and masticatory systems must provide sufficient stress over such an area that the material fails
before the teeth do. Consequently, the shape of the tooth is likely to be controlled to some
extent by selection for a particular configuration that matches a particular food for an amount of
available occlusal force.

By concentrating force over a very small area there is the risk of exceeding the sustainable
stress of tooth enamel, resulting in fracture of the tip. If the tip does fracture, it becomes
rounder, the force may be distributed over a larger area and, if the stress is temporarily reduced
below the failure threshold, it will not wear further. This explains why sharp cusps wear faster
than blunt ones. Wear is accelerated when inclusions such as silica are caught between the

PREDICTING FOSSIL MAMMAL DIETS - 207

teeth. Silica crystals can absorb higher stresses before failure than enamel, and so they wear
out teeth. However, with a worn tooth it obviously takes more force to break the ductile
foodstuff, because the available force is being distributed over a broader area. Tooth
morphology is a compromise between economically distributing sufficient force over the
occluding facets in order to fracture the food and reducing the wear on the tooth. Different foods
have different physical properties, and so tooth morphology is likely to be adaptive and
specific. There are very good reasons for expecting that tooth morphology adapts to the
physical properties of the food and that the evolution of tooth form is driven by selection, as a
clade adapts to a new food resource. The kinds of adaptations involved in resisting wear are
discussed in detail by Janis & Fortelius (1988).

THE INTERACTION OF MECHANICAL AND CHEMICAL
DIGESTION

Chemical digestion takes place in the gut. It occurs by the action of endogenous or
bacterially derived exogenous enzymes on the substrate. The greater the surface area, the more
enzymes can be employed and the faster the digestion. Access to cell contents requires the
rupture of the ccll wall, while digestion of plant cell wall requires the attachment of
microorganisms and the penetration of their enzymes. Arnold (1985) suggested that the
availability of substrate for these organisms is likely to be the rate-limiting step in digestion.
Thus, factors such as food particle size and mastication, by influencing substrate availability,
are important determinants of cell wall digestion rate. Since smaller objects have relatively
larger surface area to volume ratios, compared to larger objects, there is an enormous advantage
in mechanically reducing the particles to smaller and smaller pieces, at least in non-ruminants.
This latter process is dependent on many factors (sce Fortelius 1985).

Herbivores which digest plant cell walls, mainly composed of cellulose, hemicellulose,
pectin and lignin, have a symbiotic relationship with bacteria, fungi and/or Protozoa. The
position of the fermentation chamber, is either anterior to the truc stomach, and such animals
are referred to as foregut fermenters, or posterior to the small intestine, which are hindgut
fermenters (Janis 1976, Fortelius 1985). The relative position of the site of fermentation and
the main absorptive areas of the gut has important consequences for the kind of dentition
usually associated with the two kinds of "digestive strategies" (Janis 1976, Fortelius 1985,
Janis & Fortelius 1988). Hindgut fermenters and most foregut fermenting nonruminants,
typically chew the food only once. Ruminants may regurgitate and chew the food several
times, the properties of the food becoming more uniform.

Symbionts can breakdown and metabolise cell wall and, if the cell contents are still present,
the cytoplasm will be metabolised as well, providing necessary substrates for bacterial growth.
Exposed cell contents are presumably absorbed before reaching the hindgut fermentation site,
which has energetic advantages for the host. However, digestion of cell wall may be less
efficient in the hind gut fermenter than the ruminant foregut fermenter (Bel! 1971). It should
also be noted that ruminants, mainly as a result of a rate limiting step at the reticulo-omasal
orifice, maximise mechanical and enzymatic breakdown of the cell wall to small, uniform
particles. Particles must be reduced below a critical size before they can pass through to the
hind stomach and another meal can be taken. It is significant that digestibilities of tropical
grasses in vitro were increased by a reduction in the size of the particles (McLeod & Minson
1969) and chewing, in ruminants, facilitates the bacterial disintegration of structural cellulose
(Baker & Harriss 1947).

Cell contents are exposed during this process but are probably metabolized by the bacteria
before the host can digest and absorb them, Hindgut fermenters absorb the bulk of the cell
contents in the small intestine and tend to maximise the digestion of that component of plant

208 - SANSON

tissue. This process must occur before the fermentative action of the symbionts erodes the cell
wall, which would aid in the exposure of cell contents. Koalas derive about 90% of their daily
energy requirements from the cell contents (Cork & Hume 1983), which must be exposed by
the teeth (Lanyon & Sanson 1986, Cork & Sanson in press). Other hindgut fermenters may
derive considerably more of their daily energy requirements from fermentation of cell wall.

Rensberger (1973) and Fortelius (1985) suggest that the mechanical requirements of
"pulping" and primarily exposing cell contents and those of reduction of particle size, are
different. Fortelius (1985) notes that if the cell walls and other protective structures are broken,
digestion of the cytoplasm can proceed independently of the breaking down of the wall. While
this may be true it is not known if masticatory systems can optimise both functions.
Fortelius (1985) has predicted that hindgut fermenters will have a dental morphology that
reflects their dietary adaptations more than is the case for foregut fermenters. He notes that
"molar morphology is more diverse among the nonruminant artiodactyls and the perissodactyls
ihan the ruminating (selenodont) artiodactyls, in which differences are mainly of crown height
but not of occlusal design" (p. 20). This point is interesting with regard to the discussion
about Vorontsov's conclusions above, because Hofmann (1968) showed that East African
ruminants have significant structural differences in the stomach which relates to diet, yet their
teeth are remarkably uniform.

Fortelius (1985) also argues that the macropod marsupials have a "singularly uniform dental
design" despite varied dietary specializations. Further, the macropods are foregut fermenters,
some of which do ruminate. I would contend that the macropods do not have a particularly
uniform dental design and that macropods that are omnivores, browsers or grazers, have
demonstrably different dentitions (Sanson 1980, 1989). While they are certainly foregut
fermenters, Hume (1982) and Barker et al. (1963) argued that the regurgitation, sometimes
observed is not analogous to rumination and the term should not be used in connection with
kangaroos. The macropod forestomach does not have a structure equivalent to the rate limiting
reticulo-omasal orifice of ruminants (Hume 1982, Langer 1988). Following Fortelius'
argument, it might be expected that the teeth would reflect the diet more than is the case in the
ruminants. The macropods do support Fortelius' contention that animals that do not regularly
regurgitate and rechew their food, have a dentition that reflects their diet.

Fortelius (1985) also argues that hindgut fermenters should have a relatively greater
masticatory performance capacity than ruminants with the same dict. This should be reflected
in relatively larger occlusal area and/or larger masticatory muscles. There is evidence to
support this,

BODY SIZE, METABOLISM AND ENERGY REQUIREMENTS

Just as teeth react to changing properties of the dict, so does the gut. Van Hoven &
Boomker (1985) point out that new vegetation types, grasslands and shrublands, developed
from the original, widespread tropical forests, and are correlated with cooler, drier, and more
seasonal climates. Throughout evolution, animal tissues presumably retained similar nutrient
demands while plants became very diverse in composition and structure. With the development
of grasslands, readily available nutrients became progressively more diluted with fibre. Fibrous
material tends to be bulky, and digestion is slow. To meet their nutritional demands, ungulates
and other herbivores, evolved enlarged and complicated compartments in the digestive tract.
This serves to increase the amount of feed digested by microbial action at any point in time,
and increases the retention time, exposing digesta to microbial action for longer periods.
There are many good reviews of digestion that indicate the different patterns that exist (e.g. Van
Hoven & Boomker 1985).

Within vertebrates, mass-specific basal metabolic rate, measured as oxygen consumption
per unit body mass, decreases as body size increases (Kleiber 1961). The consequences of body-

PREDICTING FOSSIL MAMMAL DIETS - 209

weight related energy requirements on dietary preference and intake have been pointed out (e.g.
Eisenberg 1981, Parra 1978, Ripley 1984, Kay & Covert 1984). Janis (1976) and Bjornhag
(1987) relate these factors to gut structure. Briefly, small mammals have relatively higher
energy demands than large ones and must consume more nutritious and readily digestible foods.
Feeding on animal tissue as well as concentrated energy sources such as nectar, seeds, fruit, and
plant storage tissue, predominates in small mammals. The very smallest mammals are
probably restricted to such diets. Plant material rich in structural carbohydrates can only be
utilised by larger mammals. Digestibility can be increased by more efficient preparation and
this is often reflected in the dentition. Herbivores with the same kinds of teeth, but of different
body size, may then be eating different things in order to compensate for the rate of energy
extraction from relatively refractory dicts. There are body size limits below which mammalian
herbivores cannot be sustained. The size depends on the digestive strategy, hindgut fermenters
being able to exist at a smaller body size than foregut fermenters, particularly the true
ruminants (Janis 1976).

Carnivores are less likely to be constrained by these considerations. However, problems of
scale do affect them as well. The very smallest carnivores, the insectivores, are eating animals
close or equal to themselves in body size. The resistance of the exoskeleton may be
prohibitive without very precise and subtle dental modifications (e.g. Freeman 1979). Insects
are patchy in distribution, but once obtained will be a large resource to a small predator. Larger
carnivores do not need special dental adaptations for chewing insects because they have excess
power in their masticatory systems. Indeed, the largest insectivores are often edentulous.
However, insects are too small and patchy in their distribution to economically sustain large
carnivores, and vertebrates or other resources are often included in their diet. The exception is
the predation of social insects such as termites, and the scale of the animals involved suggests
that the prey can be crushed in large numbers rather than individually chewed. Kay & Covert
(1984) give an interesting illustration of the necessary foraging time larger insectivores require
and the constraints this places on body size.

BEHAVIOUR

It is possible for certain behavioural traits, particularly in the exertion of diet selection, to
compensate for the lack of morphological adaptations. For example, compared with other
baboons, the Gelada, Theropithecus, is exceptionally specialised with grass contributing about
90% of the diet (Dunbar 1977). Dunbar has noted that although Theropithecus seems to be
extremely efficient at handling the high cellulose content of the grass, it is not clear how it is
achieved. Handling efficiency was not defined. Dunbar suggests that this species may have
acquired a specialized internal flora. However, the gut is little different to that of other Papio
baboons (Hill 1970). The stomach is of the usual cercopithecid pattern while the colon and
caecum closely resembles that of Papio, the caecum being only 7.6 cm long. It does not
appear to show the kinds of gut specializations associated with other grazers of this size.

Dunbar (1977) considered that the large, hypsodont cheek teeth perhaps could have
"thoroughly pulverized" the grass blades prior to ingestion. The assertion that Theropithecus
has hypsodont molars is questionable. Swindler (1976) indicates that the molars are
bilophodont and conform to the molar morphotype of the cercopithecids. He cites Jolly (1970)
who compares the low, rounded cusps of Mandrillus with the the large, high crowned molars of
Theropithecus. Szalay & Delson (1979) describe the cheek teeth as having especially high
crowns with greatly increased relief. They do not describe the teeth as bilophodont, rather
noting that the cusps are somewhat columnar in form, as a result of their separation from one
another by the deep basins. They do argue that the delayed eruption pattern, high relief and
molar complexity, produce a distinctive wear pattern which prolongs the functional life of the
tooth row and is eminently adapted to the mastication of grass blades, seeds and rhizomes that

210 - SANSON

make up the diet. James (1960) illustrates the dentition of Mandrillus and Theropithecus.
Theropithecus molars do appear to be slightly more robust and slightly higher crowned.
However, when compared to grazers of the same body size from other mammalian groups, it is
hard to accept that they can reasonably be considered as having high crowned, lophodont teeth.
There is simply no comparison to macropod marsupials, for example, in the development of
bilophodonty, and, while they may be robust, they do not compare with other classic
hypsodont teeth.

Theropithecus is apparently a highly skilled and selective feeder, plucking the grasses with
its dextrous opposable thumb and forefinger. Coarse, reedy grasses are never eaten, and even
during the dry season, when much of the grass cover is dry and brown, only green leaves are
used (Dunbar 1977). This takes time, and geladas spend about 40-45% of the day feeding,
which is higher than the 25-30% reported for Papio baboons. It is possible that this behaviour
and mode of feeding, or “hand grazing" (Szalay & Delson 1979), allows the selection of young,
nutritious shoots while avoiding the highly fibrous, abrasive components of grasses. Other
grazers of similar body size have much more pronounced morphological adaptations, reflected
in the dentition and gut, enabling them to successfully cope with a grass diet without being so
selective. It is extremely doubtful that the diet of Theropithecus would have been correctly
predicted from a knowledge of its tooth morphology.

TOOTH WEAR AND STRIATION PATTERN AND STRUCTURE

Tooth wear produces scratches or striations on the surface of the tooth. This has been used
to analyse occlusal direction and contact. It has been argued that the kind of striation (depth,
degree and frequency of pitting) reflects the dict since the properties of the diet cause the wear.
The study of tooth microwear is, therefore, a potentially useful tool for determining diet (e.g.
Ryan 1979, Grine 1981, Rose et al. 1981, Walker 1981). There has been some debate as to
the limits of the information generated. Covert & Kay (1981) argued that the teeth of
opossums fed grit in their diet were distinguishable from opossums fed chitin and other plant
material. They considered that while grazers, which normally consume siliceous or high grit
foods, might be distinguished from browsers, the possibility of discriminating finer differences
was doubtful (Kay & Covert 1984).

Recent work, using more refined techniques, has indicated that a finer level of resolution can
be achieved (e.g. Teaford & Walker 1984, Teaford 1985, Kay 1987). In spite of these improved
techniques, which are invaluable and informative, tooth microwear must be interpreted
cautiously. Walker et al. (1978), in an important study, distinguished between grazing and
browsing hyrax on the basis of tooth microwear. However, they concluded that "examination
of microwear on a fossil tooth would give information about the diet of an individual only for
the period just before it died" (p. 910). This problem can be avoided by taking large samples,
but that is not always possible, and it is time consuming. In addition, seasonal differences in
diet, which can be quite marked (viz Sanson et al. 1985), may not be appreciated. It is equally
true, of course, that dietary preferences determined from other data have the same drawbacks. A
combination of techniques used to reconstruct dictary preference and using microwear to test
those hypotheses, as Walker et al. (1978) do, may be the best solution.

THYLACOLEO: PREDICTING THE DIET OF A UNIQUE FOSSIL
WITHOUT LIVING RELATIVES

Thylacoleo, the marsupial "lion", has long interested palaeontologists because of its unique
dentition. Consequently, there has been considerable debate conceming its diet (see Finch 1982

PREDICTING FOSSIL MAMMAL DIETS - 211

and Wells et al. 1982). The problem is that the thylacoleonids are of herbivorous phalangeroid
stock yet have no equivalent dental adaptations. The first two upper and lower premolars are
vestigial. The third premolars are remarkable, hypertrophied, sectorial blades (Fig. 2). No
other mammal has carnassials, if that is what they are, developed to such an extent. The first
lower molar contributes to the sectorial blade of the P3 while M! and M, are very reduced and

have many similarities, in form and possibly function, to those of felids. The lower incisors
are often described as caniniform and may have been used for stabbing prey.

Wells et al, (1982) have made the most detailed analysis of the possible diet. Because there
are no analogues, living or extinct, the problem cannot be resolved fully. Nonetheless, Wells
et al.'s analysis of microwear of thylacoleonid teeth and their comparisons with other known
carnivores, in terms of overall tooth morphology, precise diet, including the capacity or
tendency to consume bones, and killing behaviour is comprehensive and much of it is
convincing. They conclude that Thylacoleo was a flesh eater and, like sabretooth felids and
cheetahs, probably did not consume much, if any, bone. This can be attributed to the lack of
blunt cusps or well developed anterior premolars. The sectorial premolars certainly appear
marvellously adapted for shearing muscle, tendons and skin, and as such there would be an
advantage in protecting the edge of the blade from being chipped on bone.

The principles of material breakdown discussed above are pertinent here. It is clear that
brittle material, like bone, can be cracked with the same force by blunt or sharp cusps (Lucas
1979, 1982; Hill 1985). The principle of breakage requires the propagation of a crack through
the material (Gordon 1968). The energy required at the propagating crack tip is provided by the
applied load. The sharpness of the cusp or blade providing the load is not relevant, except that
sharp cusps will chip and wear faster than blunt cusps. It is interesting then that carnivores
that crack bones have blunt, well developed premolars. Domestic dogs use the rounded anterior
face of the carnassial tooth, but not the sharp blade (personal observation) for cracking bones.
This is entirely consistent with Wells et al.'s (1982) discussion of specialist meat eating
carnivore habits which avoid bone, compared to generalist meat eaters and bone crushers.
Thylacoleo, without apparent bone crushing adaptations, is presumed to be a specialist meat
eater, avoiding bone, like modern cheetahs. The predominant lack of Thylacoleo remains
occurring with incised bones of other species seems to support this (Wells e¢ al. 1982).

Wells et al. (1982) provide evidence from microwear to support their contentions. Grazers,
consuming grasses, loaded with grit and silica, produce fine striations; browsers they examined
had even finer striations, while the carnivores had much coarser ones than either browsers or
grazers. There was an "order of magnitude difference" between the mean size of striae of
herbivores and carnivores. Interestingly, Thylacoleo had striae on average five times larger than
the carnivores. Unfortunately, neither the number of animals examined, nor their dietary
history before death, is recorded. This may be important, as the larger striae in Thylacoleo
when compared to the herbivores, and presumably the other carnivores, is attributed to
fragments of bone or grit in the fur. Facsimile production of striae by Wells et al. (1982) on
model teeth, fed different materials, supported this.

There appears to be a contradiction between the observed gross morphology and the striation
results reported by Wells et al. (1982). The gross morphology indicates flesh eating and no
bone crushing, The striae suggest more bone crushing than in other carnivores and scavengers
(e.g. Canis familiaris, Felis catus, Thylacinus and Sarcophilus). The difficulty may be
resolved by wider sampling of Thylacoleo and animals with known dietary histories just before
death. A problem still remains concerning the selective pressures that led to such an extreme
specialization when all other highly successful carnivores have retained a notched carnassial.
The importance of such a notch is discussed by Mellett (1977). The notch separates the
anterior from the posterior cusps, which form the ends of the carnassial cutting edge. This
configuration means that the anterior cusps penetrate and hold material being separated by the
carnassial blades. Without this, the material would tend to be squeezed out in

212 - SANSON

Figure 2. Thylacoleo carnifex. A, ventral view of the cranium showing upper incisors and well developed
sectorial third premolars; B, occlusal, and C, lateral views of left mandible; x 0.6. Note the hypertrophied

sectorial blade. (After Woods 1956).

PREDICTING FOSSIL MAMMAL DIETS - 213

advance of the occluding cutting edge. An analogy would be a pair of scissors cutting a match.
The match is pushed down the blades. Pruning shears have a "cusp" to hold the material and
prevent it from being pushed out. Thylacoleo has a blade without a notch (Fig. 2). Thylacoleo
is an interesting example of the problems encountered in analysing unique fossils without any
good analogues. The problem is exacerbated when the available evidence is conflicting.

PROPLEOPUS: PREDICTING THE DIET OF A UNIQUE FOSSIL
WITH LIVING RELATIVES

Propleopus (Fig. 3) was a gigantic Pleistocene rat-kangaroo of the family Potoroidae. It
approached the megafaunal sthenurine kangaroos in size, yet other potoroids are small, having a
mass of only a few kilograms. The giant forms have been recently revised by Archer &
Flannery (1985). It is generally agreed that Propleopus is closest to Hypsiprymnodon, the
Musky Rat-kangaroo, the smallest of the living macropodoids. Hypsiprymnodon has a large
plagiaulacoid, sectorial premolar and the molars are tubercular and tunodont. It is unique

Figure 3. Propleopus oscillans. A, lateral and B, occlusal views of left mandible.; x 1. Note the well
developed plagiaulacoid premolar and bunodont molars. (After Woods 1960).

among the kangaroo group in having a simple non-sacculated stomach, suggesting that it does
not ferment plant fibre. At 360-680 g (Strahan 1983), fermentation of plant fibre is an
unlikely method of supplying its energy requirements. Its diet consists of insects and other
invertebrates, seeds and tubers (Johnson & Strahan 1982).

Propleopus has similar dental features to Hypsiprymnodon, only the teeth are much larger
(Fig. 3), and this led Pledge (1981) to conclude that the bunodont molars, like humans, pigs
and bears, suggest an omnivorous or browsing vegetarian diet. The large premolars were

214 - SANSON

probably associated with omnivory, possibly being used to cut flesh as well as vegetable
matter.

Archer & Flannery (1985) went further, suggesting that Propleopus may have been
carnivorous. They noted the omnivorous diet of Hypsiprymnodon and Burramys parvus, both
species having large plagiaulacoid premolars and bunodont molars, and observed that
Propleopus was 200 times heavier than Hypsiprymnodon. The significance of this size
difference was not commented on, but is important for the reasons discussed in previous
sections. It is unlikely that an animal the size of Propleopus could sustain itself on a diet of
insects, seeds and fruits, like /7ypsiprymnodon, because such items are patchily distributed in
space and time (Norbury et al. 1989), and large amounts would be required. On the other hand
vertebrate prey or carrion, possibly supplemented with fruits and other high quality, but rare,
foods, would suffice. This argument is supported by the rarity of Propleopus in deposits,
which might be expected if the species is near the top of the food chain (Archer & Flannery
1985). Archer & Flannery do, however, caution that other, presumably herbivorous, species
are equally rare.

Archer & Flannery (1985) argue that low crowned molars would be rapidly destroyed if the
animals fed predominantly on siliceous grasses. This is almost certainly true. The molars are
not high crowned, a common adaptation for the extreme wear imposed by grasses, nor do they
have the usual development of fine cutting edges to deal with the parallel and dense fibre
bundles. However, grasses are not the only plant material available. Low abrasive, low fibre
dicotyledonous leaves must also be considered. With the exception of the selenodont
artiodactyls, browsers of this size usually have molars with broad grinding surfaces.
Propleopus has a steep molar gradient with reduced lophs suggesting that the molars are not
adapted to grinding large amounts of plant matter (Archer & Flannery 1985). This might be
true but really requires an analysis of the functional molar occlusion of the genus.

There are physiological reasons why some animals may need less food than might
otherwise be expected. Marsupials have a lower basal metabolism than eutherians (Dawson &
Hulbert 1970), and this has an effect on energy requirements (Hume 1982). Kangaroos have
lower maintenance energy requirements than sheep of the same body mass (Hume 1974).
Kangaroos, as a result of a curved tooth row and molar progression, only occlude about two
molars in each quadrant at a time (Sanson 1980). Sheep, on the other hand, occlude all their
molars at once. Browsing macropods occlude all their molars at once, but that may be as a
result of their heritage. They may not need to - grazers do not. This aspect requires further
work before a browsing condition can be excluded on the basis that the molars are not adapted
for grinding large amounts of plant matter.

Archer & Flannery (1985) report that the premolars have very coarse wear striae. The larger
striae in Propleopus are approached only by those seen in Thylacoleo, These striae are more
similar to those seen on the cheek teeth of the carnivores, than on those of herbivores,
examined by Wells et al. (1982). While this is interesting, it must be remembered that the diet
of Thylacoleo is a matter of some contention, and the difference between the microwear on
Thylacoleo teeth and other carnivores was significant. A detailed microwear analysis of
Propleopus molars may help resolve this dilemma.

Archer & Flannery (1985) report that other living potoroids occasionally take meat, but do
have a specialized forestomach. Since Hypsiprymnodon lacks a specialised stomach, they
conclude that it is "probable, in view of our present phyletic understanding, that the even more
primitive species of Propleopus would have lacked this specialised feature also" (p. 1346).
However, if Propleopus had independently evolved a fermentation capacity, its diet could have
been fundamentally different to the living potoroids, including, of course, Hypsiprymnodon.

As previously pointed out some kind of fermentation chamber, either foregut or hindgut,
could compensate for apparently inferior teeth. Pledge (1981) mentioned that Propleopus has
molars with features typical of bears, and pandas. Moreover, giant pandas seem to cope with a

PREDICTING FOSSIL MAMMAL DIETS - 215

simple stomach, intestines remarkable for their shortness, as short as any known carnivore, no
caecum, and a short colon (Davis 1964). Pandas are very inefficient digesters of bamboo, and
do not rely on the microbial degradation of plant material (Dierenfeld et al. 1982). Giant
pandas seem to meet their nutritional requirements from high levels of intake combined with
fast gut passage rates (Dierenfeld et al. 1982). The molars are fundamentally similar to Ursus,
and, though the teeth are bigger and have secondary tubercles, Davis (1964) explains the
differences on allometric grounds. He states "Any relation between the "bunodont" character of
the molars of Ailuropoda and its diet is fortuitous" (p. 130). On the other hand Deirenfeld et
al. State that the large, flat cheek teeth with elaborate crown patterns are dentition characteristics
typical of a herbivore. This assertion is difficult to accept when the array of herbivore dental
adaptations are reviewed. The low crowned, but very broad, molars are teeth that would seem
to be poorly adapted to chewing plant matter, particularly bamboo. Davis (1964) reports the
findings of field observers who have emphasized the poorly chewed and undigested condition of
pieces of bamboo in the droppings of giant pandas. Davis considers that the exclusively
herbivorous diet is an extension of a non-carnivorous dietary trend already present in the group
= la it is derived. Would the diet of the panda have been correctly predicted from its
teeth!

The analysis of the relationships of the propleopines, hypsiprymnodontines, potoroines and
macropodines is important, because if the potoroines and macropodines are monophyletic and a
sister group to the hypsiprymnodontines, it suggests that foregut fermentation evolved once. If
so, the propleopines and hypsiprymnodontines need to have their rank elevated. However, if
the propleopines are placed within the Potoroidae (Archer & Flannery 1985, based on a
synapomorphic feature of the masseteric canal; Flannery & Rich 1986), it means that foregut
fermentation evolved twice - in the potoroine and macropodine lineages. Alternatively, it
evolved once at the base of the macropodoids, and Hypsiprymnodon has secondarily lost it. If
foregut fermentation evolved twice, it is hard to argue that it could not have evolved three
times. Langer (1980) suggests that the potoroine and macropodine stomachs evolved
independently, while Hume (1978) considered that the macropodine condition may have
developed from the potoroine type. It is not certain that Propleopus was not a foregut
fermenter with all the dietary consequences that that would entail. In addition, the diet of the
giant panda, with few herbivorous adaptations, encourages caution. Although Archer and
Flannery's arguments are interesting and persuasive, more detailed analysis of microwear on
molars and premolars and functional occlusion of the dentition is warranted. Even then the
answer may be equivocal.

FOSSIL STHENURINAE AND MACROPODINAE: PREDICTING
THE DIET OF FOSSILS WITH LIVING RELATIVES

The tooth morphology and associated dietary preferences in the living Macropodinae are
fairly well understood (Sanson 1989). The Sthenurinae are a sister group of the Macropodinae
and mainly comprise extinct megafaunal elements. There are close similarities in tooth
structure between the two groups, and our knowledge of tooth function and relationship of
many of the species allows some detailed predictions to be made. While Ride (1959) attempted
to infer diet from a knowledge of morphology of extant species, based largely on his analysis
of the incisors and premolars, other authors have given little reason for their assertions about
the diet of Sthenurus, Procoptodon and Troposodon.

Sthenurus (Sthenurus) species are considered to be grassland or savanna grazers (Tedford
1966), while the short faced Sthenurus (Simosthenurus) and Procoptodon species are considered
to be browsers (Ride 1959; Bartholomai 1963; Tedford 1966, 1967). Bartholomai (1967)
considered that the tooth structure of Troposodon minor indicated a secondary reversal from

216 - SANSON

grazing macropodines to a semi-browsing habit, but did not elaborate. Campbell (1973)
suggested that Troposodon kenti occupied a grazing or grazing-browsing "niche".

Troughton (1947) generally considered the Macropodinae as a whole are adapted for grazing.
This probably stemmed from observations of where macropods were found and a lack of
knowledge as to what they actually ate. The notion of a secondary reversal from grazing
predecessors follows from Ride (1959) who, following Troughton, also considered that the
Macropodinae as a whole are adapted to grazing. Later evidence, reviewed in Sanson (1980),
challenges this broad generalisation. Ride (1964), however, did recognise problems with this

ANT.

A B LAB. LING.

mlk
ml

hid
mlk

Figure 4. Occlusion in a representative browsing macropod. A and C are lateral views of upper (thick
outline) and lower (thin outline) molars in occlusion; B and D are plan views of upper (thick outline) and
lower (thin outline) molars in occlusion at the same stage as A and C respectively. Horizontal hatching
represents area of lower tooth in contact with upper tooth; vertical hatching in stage of occlusion represented
by respective lateral views. Therefore, total area of tooth which comes in contact with occlusal counterpart
represents total horizontal or vertical hatched areas in B and D. Arrow indicates movement of lower tooth.
Abbreviations: ANT, anterior; flk, forelink; hld, hypolophid; LAB, labial; LING, lingual; ml, metaloph; mlk,
midlink; pl, protoloph; pld, protolophid.

concept and linked Wallabia bicolor, Dorcopsis, Dorcopsulus and Dendrolagus as forest- and
thicket-dwelling wallabies and tree kangaroos and noted that they possess simpler molars than

PREDICTING FOSSIL MAMMAL DIETS - 217

their "grazing adapted relatives". For a fuller discussion of the development of these arguments
and the evolution of mastication in the macropodines see Sanson (1989).

The functional incisor and premolar morphology is not as well understood as the molar
system, which does not support the dietary predictions referred to above.

Living browsing macropodines have a suite of dental characteristics associated with the
physical properties of the diet (Sanson 1989). The main characteristics are that the molars have
very weak longitudinal ridges, the links, connecting the transverse lophs (Fig. 4), which means
that the molars occlude over a broad surface area. There is a large sectorial permanent premolar,
never naturally lost, which, with the four molars, occludes along a plane line of contact (Fig.
5). This maintains a large contact area between the occluding check teeth.

plane of occlusal
contact

M4 M3 M2 Mi P3

Figure 5. Lateral view of the mandible of Wallabia bicolor, a browsing macropod, drawn from a radiograph.

Living grazing macropodines have elaborated the longitudinal link, increasing the cutting
edge length of each molar and reducing the surface area contact. Unlike the condition seen in
the browsing macropodines, the initial forward movement of the lower jaw is followed by a
transverse lingual movement (Fig. 6). The high links prevent close interdigitation of the lophs
at right angles, the transverse movement maintaining this contact. There is a reduction in the
size of the premolar, and it is often lost by the forward progression of the molars. The lower
tooth row is curved, so that it meets the upper tooth row at a tangent on the anterior portion of
the row (Fig. 7). This further reduces the surface area contact. Posterior molars are brought
forward into occlusion by molar progression. Browsing macropodines have a procumbent first
upper incisor and a small third upper incisor. The lower incisors meet the first incisors, often
lying just inside the upper incisor arcade. It is likely that this pattern is associated with the
finer manipulation required in browsing. Grazing macropodines have increased the length of
the third incisor, and the lower incisors meet the uppers over a larger contact area. These teeth
are involved in gathering grass material, not processing it, so the significance of the increased
contact area of the grazing incisors should not be confused with the opposite effect in the
grinding molars. Flannery (1983) has suggested that there is a taxonomic distinction in the
incisor structure between the Macropodinae and the Sthenurinae. It is not yet clear what
functional significance these differences have.

218 - SANSON

The weak link structure of the bilophodont molars, wear facet formation and occlusal
mechanics of Sthenurus (Sthenurus) (Figs 8, 9) and Troposodon are all very similar to the
morphology of W. bicolor. The molars of this species have been shown to be primarily adapted
to low fibre vegetation and not high fibre plant tissue in the form of grass (Sanson 1980).
Equally the molar patterns in Procoptodon are fundamentally the same as those in Macropus
giganteus, a species primarily adapted to a diet of grass.

Sthenurus (Simosthenurus) have ornamented enamel lophs (Figs 10, 11), which mainly
occur in an area involved in grinding in the similar kind of tooth of W. bicolor. This may
serve to increase the efficiency of this type of occlusion, but it is very doubtful if it is capable
of comminuting fibre.

The simplest Procoptodon morphology is found in P. pusio (Fig. 12), which shows a link
and loph structure very similar to the large Macropus species. The wear facet formation is very
similar in the two genera, and there is every reason to believe that the teeth operate in the same
way, that is bringing opposing cutting edges together. When related to the inferred functional
morphology of the tooth, the crenulation patterns in S. (Simosthenurus) are not the same as

t ANT.

flk milk
pl ml

pid hid
flk milk

ANT.
LAB. + LING.

a

ml

Figure 6. Occlusion in a representative grazing macropod. For explanation see legend to Fig. 4.

PREDICTING FOSSIL MAMMAL DIETS - 219

those in the large Procoptodon species. The molar crenulations in P. rapha and P. goliah
(Fig. 12) are in positions which increase the number of cutting edges in occlusion, a common
feature of grazers, but at the same time preventing the broad surface area contact occurring in
Sthenurus, a browsing feature. In many respects the functional molar morphology of
Procoptodon is a slightly more elaborate version of that in M. giganteus, and it is hard to see
why this indicates a browsing habit. The increased development of cutting edges in

line along upper tooth row
region of occlusal ie
contact M2 M1

=
Sees

M3 == : P3
-M4 ¥
i an
yd I J) ft line along lower tooth row

Figure 7. Lateral view of the mandible of Macropus giganteus, a grazing macropod, drawn from a
radiograph.

Procoptodon may be a function of size, the larger Procoptodon having to ingest absolutely
more food. It would be interesting if the degree of development of extra links is allometrically
related to body size. Within Macropus, increased body size is associated with more complex
links. At Lake George, in New South Wales, Procoptodon cf. goliah has been found in what
was predominantly a cold, dry grassland, supporting the suggestion that Procoptodon may have
been more of a grazer than previously thought (Sanson et al. 1980).

One problem with the interpretation of Procoptodon as a grazer is that it has a flat tooth
row (Fig. 13). This is not a feature of advanced grazing macropodines. However, it is
suggested here that the development of links in the sthenurine Procoptodon is a grazing
adaptation convergent on the condition found in the grazing macropodines. There is no reason
to expect that all of the other grazing adaptations of the macropodines, such as the curved tooth
row, should also be found in Procoptodon. Powerful jaw muscles, indicated by the massive
skull, may have compensated for the large surface area contact consequent on retaining the
ancestral flat tooth row. Following this argument, it is notable that Sthenurus
(Simosthenurus), which has crenulated molars, also has a brachycephalic skull. Brachycephaly
may have been influenced by the selection for larger and more powerful jaw muscles. The
more anteriorly directed coronoid process in these forms should provide a greater mechanical
advantage for the masseteric muscles.

220 - SANSON

Figure 8. Sthenurus (Sthenurus) tindalei, right upper incisors and cheek tooth dentition, Lake Menindee,
New South Wales. A, labial and B, occlusal views (after Tedford 1966).; x 0.8. Note relatively simple
molars with poorly developed links.

ae ee See MOR

Figure 9. Sthenurus (Sthenurus) atlas. A, labial ; B, occlusal view of right ramus with part of lower incisor,
P2, DP3 and M1-2; C, labial; D, occlusal; E, lingual, views of unerupted left lower P3, xl, Lake Menindee
(after Tedford 1967). Note the flat tooth row, sectorial premolar and molars with poorly developed links.

PREDICTING FOSSIL MAMMAL DIETS - 221

CONCLUSION

___ There are basic difficulties in interpreting the dict of fossils from the teeth and jaws. This
is because there is abundant evidence that the gut and behaviour of an animal can compensate
for the dietary adaptations of the teeth, which are most likely to be the parts fossilized. Only
in very rare cases like Chaeropus (Wright et al., Chap. 8, this volume), where there is evidence
from the gut to test the dictary hypotheses, can one be really confident. Tooth microwear can
also be used to test hypotheses, but a broad data base is required because of the problems
alluded to above.

; Janis (1984) presents examples of how cumulative patterns of molar wear in living
primates are related to diet, enabling the prediction of the dict of extant and extinct species.
Janis does show correlations between the tooth form and the diet. The critical question is

Figure 10. Sthenurus (Simosthenurus) occidentalis restored right upper incisors and cheek teeth. A, lateral,
and B, occlusal views (after Tedford 19660; x 0.8. Note the massive premolar and the molar with crenulations

on the faces of the lophs.

222 - SANSON

Figure 11. Sthenurus (Simosthenurus) occidentalis restored left mandible. A, lingual; B, occlusal; and C,
labial views; x 0.75; Western Australia (after Tedford 1966). Note the large premolar, the flat tooth row, the
molars with crenulations on the faces of the lophs and the vertical anterior margin of the coronoid process

associated with brachycephaly.

whether the diet would always be accurately predicted from the form of the tooth. Kay &
Covert (1984) note that Old World colobine monkeys have a bilophodont molar condition
which aids in cutting up plant fibre. However, the bilophodont condition is also seen among
fruit eating species consuming low fibre diets. They argue that an inference about dietary
adaptation is not warranted simply because an animal has bilophodont molars. They state with
some justification that for whatever reason bilophodonty evolved, it has served as a basis for
many different sorts of dietary variations. This may be because such a pattern, evolved early on
in a lineage and acting as a heritage factor, is adequate for many diets. It may not be adequate
for the harsher demands of say, grazing, but it still precludes dietary predictions. In addition,

PREDICTING FOSSIL MAMMAL DIETS - 223

en ‘ore zi
Ss

SQ
SX aul
NC : gu

a

\s

Ze

Figure 12. Occlusal view of left mandible . A, Procoptodon pusio; B, Procoptodon rapha; and C,
Procoptodon goliah,; x1; Bingara, New South Wales (after Stirton & Marcus 1966). Note the reduced premolar
and well developed links.

Maier (1984) argues that the absolute body size of primates at least, has been shown to
influence tooth structure independently of dietary adaptations. Finally, there are many species
with peculiar dietary adaptations, for example Theropithecus and the giant panda, that are
compensated for by other adaptations which could not be predicted.

This paper seeks to demonstrate that the prediction of diet can, and should, take into account
many interacting factors. Our current knowledge of digestion and nutrition allows a more
detailed and sophisticated approach than was available in the past. However, there are clearly
considerable problems involved in determining the diet of extant animals when the dentition,

224 - SANSON

gut structure and behaviour, acting as compensatory mechanisms, are known. There is no
evidence to suggest that these compensations are predictable. With fossils, only the dentition
and masticatory system are known, and this must confound any conclusions concerning diet to
the extent that the diet will never be known with certainty. The more we know about the
complex interactions of parts of the digestive system the more we must be aware of the
complications that exist in predicting diet from just part of that system.

ACKNOWLEDGEMENTS

I am indebted to Drs. Clare McArthur, Grant Norbury and Bill Foley, and Suzanne Moore and
Janet Lanyon for much valued discussion and comments. I do not presume that they share all
of the views expressed here. In particular, Clare and Suzanne's reservations about dietary
inference from the dentition, and Suzanne and Bill's criticism of the manuscript, have been
influential.

A angular iL.
process SJ \mas me
we 9 anterior mental

/ \ fossa cingulum foramen

masseteric
crest

mandibular .”
foramen

postalveolar
B inferior Process
dental
angular
angular fossa _ Pek fede dhe Om

process _—

protolophid

midlink

hypolophid
masseteric angular
foramen process
A / / digastric

C “_-_ sulcus
__- ~~ digastric
process

Figure 13. Procoptodon goliah, right mandible. A, lateral; B, occlusal; C, medial, views (after Tedford
1967). Note the flat tooth row and the massive coronoid process with a vertical coronoid process.

PREDICTING FOSSIL MAMMAL DIETS - 225

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Mechanical Design in

Diet and teeth: dietary hypotheses and human evolution. Phil. Trans. R. Soc. Lond.

CHAPTER 8

THE DIET OF THE EXTINCT
BANDICOOT CHAEROPUS
ECAUDATUS

Wendy Wright!, Gordon D. Sanson? and C. McArthur2

INtrOductiOn ............cecececececeeccecncececececeeeeeeees 230
Analytical Methods ..............cccscesescececeeeeeeceees 230
Molar Morphology and Occlusion............... 230
Examination of Gut Contents...................+ 231
DISSECTION .......... cece cece scence cecec cece ee eeees 231
Fixation of Gut Contents....................- 231
Sectioning and Microscopy.............:.20+ 231
RESUS Fc sos eda cess Te oveesecatilednoonsenste etvede scene sls 232
Molar Morphology and Occlusion..............- 232
Morphology ...........0ceceeccecsceeececesececees 232
OCCIUSION............cccecsecccneceecececcnceeeeees 234
Examination of Gut Contents.................... 234
DiSSECtION .........2.2 0c ec eccecececeececeneeeeeeees 234
Light Microscopy............seceseeeeeeeereeees 234
Transmission Electron Microscopy
CTEM) 20 oe cetenectbeewsnestneoecsieresenton 235
DISCUSSION Loc orcndoestec eden stncenesecbensescuetal 236
Summary and Conclusions ..............2cseseeeeeeeees 238
Acknowledgemenls...............cecececceeceecceecenseees 239
REPETENCES SY, Mav cid sae) cles asada Was'edecl. dcTiaasecsecd 239
Plates <2, Rn AR eras eda vel taenddtceesecedelpcectess 240

9 IIE EEE ———————————————————————————E—————EE——————————————
1 Department of Zoology, University of Reading, Whiteknights, P.O. Box 228, Reading, R6 62 AJ, England.
2 Department of Botany and Zoology, Monash University, Clayton, Victoria 3168, Australia.

230 - WRIGHT ET AL.

INTRODUCTION

The Pigfooted Bandicoot, Chaeropus ecaudatus, is a recently extinct member of the
Peramcelidae, not having been reported since 1926 (Ride 1970). Literature on its diet is scarce
and conflicting, the earlicr reports mainly consisting of anecdotal observations. Krefft (1865)
considered Chaeropus 10 be herbivorous, while Sturt (1848) described it as a flesh-eater (in
Dixon, 1988). Gould listed insects and their larvae as well as plant material as dictary items
(in Dixon, 1988), while a gut content analysis by Dixon (1988) suggested that Chaeropus was
herbivorous.

Taking some important considerations into account, a great deal about the dict of a mammal
can be inferred from its teeth (Sanson this volume, Chap. 7). Chaeropus molars, while
showing some similarities to those of other bandicoots, are the most highly specialised
(Bensley 1903) within this group. Their devclopment of shearing crests is unique.
Consequently, there are no appropriate analogues to compare them with. However, it is
possible to obtain information about occlusion and the action of the tecth from a study of the
wear on their surfaces (e.g. Butler 1952, Mills 1967, Greaves 1972, Rensberger 1978).
Coupled with information about the physical properties of dictary items and the shape of tooth
morphology best suited to break them up (e.g. Lucas & Luke 1984, Frazetta 1988), a
knowledge of functional dental morphology can allow predictions regarding the sorts of food
that particular teeth would be most effective at reducing in size. For example, Hill (1985)
found that ductile matcrials are best cut by sharp blades or points which concentrate force over a
small surface area. A blunt surface tends to deform matcrials without penetration, unless a
larger force is applicd. Britthe matcrials can be shattered duc to a force applied by a blunt or a
sharp surface and eventual failure is due to cracks which form and grow within the material
(Gordon 1976). In extant animals, predictions regarding dict can be confirmed by dictary
studies. Sanson (1978, 1980), from a study of macropodid molar morphology, concluded that
Macropus rufogriseus and Thylogale billardieri were predominantly grazers and browsers
respectively, although detailed dictary studics for the two specics were unavailable.
Subsequently, Fletcher (1980) confirmed these predictions, giving us some confidence that the
techniques have predictive power.

Some spirit preserved specimens of Chaeropus survive, and one of these was kindly made
available by Ms J. Dixon of the Muscum of Victoria. This allowed examination of the
morphology of the digestive system and of the gut contents. It is not known whether the
animal was kept alive and fed in captivity before it died. Thus, the gut contents may not reflect
the natural dict. However, in this case there is much corroborative evidence to suggest that this
species was herbivorous.

ANALYTICAL METHODS

MOLAR MORPHOLOGY AND OCCLUSION

Upper and lower tooth rows of one relatively worn (Muscum of Victoria, MV C468) and
one relatively unworn (MV C2900) specimen of Chaeropus ecaudatus were photographed using
Kodak Panatomic X film on a Wildphotomacroskop M400 camera. Specimens were coated
with sublimated ammonium chloride to reduce reflection and illuminated using a Volpi
fibreoptic lamp. The negatives were placed on the stage of a Nikon V20 Profile Projector, and
the image, produced by transmitted light, was traced onto transparent sheets. Tooth wear facets
were examined under reflected light and marked on the tracings. The specimens were also

DIET OF CHAEROPUS ECAUDATUS - 231

examined under a Hitachi $570 scanning electron microscope. Because of their rarity, the
specimens were not sputter coated, but the teeth had sufficient conductance to produce images
without too much flaring. Micrographs were taken of occlusal surfaces using Ilford XP1 400
film. Facet development and striation patterns were recorded on the occlusal maps. The
tracings of upper and lower molar tooth rows were superimposed and manipulated to reproduce
occlusion ensuring that facets and striae direction matched.

EXAMINATION OF GUT CONTENTS

Dissection

A spirit-preserved specimen (MV C5858) was collected during the Spencer-Gillen
Expedition (1901-1902) from Barrow Creek, Northern Territory and had since been stored in
70% ethanol. The abdominal cavity had been opened previously and the gut was, therefore,
accessible without further damage to the specimen. Minimal cuts in the mesenteries were made
to allow the digestive tract to be displayed. Both the anterior mesenteric and the posterior
mesenteric circulatory systems were severed. Photographs of the displayed gut were taken
using Kodachrome 25.

Gut contents were removed from various regions of the digestive tract after short
longitudinal cuts were made in the gut wall. Ligatures prevented the loss of digesta from
unsampled areas and were left in place following the dissection. Samples were taken from the
stomach, the proximal colon, proximal caecum, and distal caecum. Very little material was
present in the stomach, and the contents of this organ were flushed out with a syringe.
Samples were placed in separate vials in 70% alcohol.

Fixation of Gut Contents

The samples were rehydrated by repeatedly changing the ethanol solution, reducing the
concentration of ethanol by 10% at each change. Samples were centrifuged between each step,
allowing liquid to be drawn off with a pipette without losing any of the sample. The digesta
remained in each solution for 5 minutes. After reaching a solution of 10% ethanol, the
samples were rinsed twice in distilled water and centrifuged. Excess water was removed with a
pipette. 2.5% gluteraldchyde in PIPE's buffer was added to each sample and left for 1 hour.
Samples were then centrifuged and the gluteraldehyde/buffer solution removed with a pipette.
After two changes of PIPE's buffer, 1% OsO, in PIPE's buffer was added to each sample. An
hour later, the samples were rinsed twice with distilled water and centrifuged. Excess liquid was
removed.

Dehydration of the samples involved repeatedly changing the ethanol solution, increasing
the concentration of ethanol by 10% at each change until a concentration of 90% alcohol was
reached. After two changes of 95% ethanol, three changes of dry ethanol were made.

Sectioning and Microscopy

Spurr's resin was added to cach sample until the solution consisted of 10% resin and 90%
ethanol. This was left for 1 hour. The concentration of resin was then increased dropwise to
approximately 80%. The suspensions were stirred regularly during this process. Samples were
centrifuged and the resin-ethanol solution removed. 100% resin was added to each sample. The
samples were then left for 36 hours. Fresh resin replaced the first 100% change, and the
samples were pipetted into plastic moulds and placed in an oven at 70°C for 24 hours.

232 - WRIGHT ET AL.

_The hardened blocks were trimmed and sections for light microscopy were cut with glass
knives on a Reichert OMu2 microtome. These were placed on glass sides and stained with
toluidine blue at pH 9 (O'Brien & McCully 1981). Paraffin oil and a coverslip were added and
the sections were viewed under a light microscope at X100 and X400. Photographs were taken
of sections from each area of the gut using Kodak Ektachrome 160 for a tungsten light source.

Sections for transmission electron microscopy (TEM) were cut with glass knives on a
Reichert-3 microtome, stained with methanolic uranyl acetate followed by lead citrate (O'Brien
& McCully 1981) and examined in a JEOL 200 CX transmission electron microscope.

RESULTS

MOLAR MORPHOLOGY AND OCCLUSION
Morphology

The occlusal surface of the upper molar is rectangular in shape (Fig. 1) and the teeth are
separated by gaps or embrasures which extend to the buccal edges of the teeth. In no other
peramelid does the embrasure between the upper molars extend this far. The major features of
the upper molar are similar to those of other bandicoots, such as Jsoodon (Wright 1988),
although certain structures have been emphasised while others have been reduced. The upper
molar is 'tiered' and can be divided into three areas: the stylar shelf, the buccal tier and the
lingual tier (Plate 1a). In Chaeropus the lingual tier is much reduced and is the smallest and

oe a Stylar shelf
St.B St.D y re e
pr.pac po.mec t
po.pac pr.mec
Buccal tier
pa me B
A—+—P
pr.prpr saptees L
po.prcr pr. hy ,
pr hy Hnguat tier

Figure 1. Occlusal view of an upper molar of Chaeropus. Abbreviations: A, anterior; B, buccal; hy,
hypocone; L, lingual; me, metacone; P, posterior; pa, paracone, po.hyc, posthypocrista; po.mec,
postmetacrista; po.pac, postparacrista; po.prcr, postprotocrista; pr.hyc, prehypocrista; pr.mec, premetacrista,
pl.pac, preparacrista; pr, protocone; pr.prcr, preprotocrista; St.A, stylar cusp A; St.B, stylar cusp B; St.D,
stylar cusp D; St.E, stylar cusp E.

lowest of any peramelid, with a very small hypocone and protocone. Concomitantly, the
buccal tier has expanded lingually compared to that of the other bandicoots. The cusps of the
buccal tier (metacone and paracone), and the crests associated with these, are the major features
of the tooth. The post-paracrista is well developed in Chaeropus, extending to stylar cusp B
and emphasizing the groove between the post-paracrista and pre-metacrista. The crests of the

DIET OF CHAEROPUS ECAUDATUS - 233

buccal tier are almost parallel and form two adjacent open triangles. The shearing edges show
evidence of tooth-tooth contact; that is they have planar attrition facets (Pl. 1B). With wear,
the crests become rounded and show evidence of tooth-food-tooth abrasion (Pl. 1C). Attrition
facets are planar with even, generally parallel striations. Abrasion facets, caused by the tooth-
food-tooth contact, are rounded with uneven multidirectional striae (Sanson 1980).

The small protocone and hypocone are linked by the post-paracrista and the pre-hypocrista.
These two crests and the pre-protocrista and post-hypocrista enclose the lingual tier and define
the protocone and hypocone. The lingual sides of these cusps are polished, and the flat area of
the lingual tier, between these, does not appear to occlude (PI. 1D).

Like other peramelids, the occlusal surface of the lower molars of Chaeropus is composed
of two adjacent open triangles, one in the talonid and another in the trigonid regions of the
tooth (Fig. 2, Pl. 2A). The triangles are better defined in Chaeropus, and the pre-hypocristid is
well developed, leading from the hypoconid to a point anterior to the base of the entoconid.
The height difference between the tiers of the upper tooth effectively prevents the protocone and
hypocone from occluding with the lingual cusps of the lower molars and allows effective
contact of the pre-hypocristid against the post-paracrista.

=— Trigonid —-~~—— Talonid ——~

hyd
prd
pacd
po.hycd
mecd —— pr.hycd
a.c
pad — hyld
med end
B
A—1—P
L

Figure 2. Occlusal view of a lower molar of Chaeropus. Abbreviations: A, anterior; a.c., anterior cingulum;
B, buccal; end, entoconid; hyd, hypoconid; hyld, hypoconulid; L, lingual; mecd, metacristid; med, metaconid;
P, posterior; pacd, paracristid; pad, paraconid,; po.cyd, posthypocristid; prd, protoconid; prhycd,
prehypocristid.

With wear, the buccal cusps become lower than the lingual cusps and, of the buccal cusps,
the hypoconid wears faster than the protoconid. The shearing crests (paracristid, metacristid,
pre-hypocristid and post-hypocristid) show attrition striations, becoming rounded and showing
evidence of tooth-food-tooth abrasion with wear (Pl. 2B). Only the buccal sides of the lingual
cusps are worn.

234 - WRIGHT ET AL.

Occlusion

The occlusal cycle in Chaeropus emphasises fine shear, as in grey kangaroos, rather than
the shear and grind seen in Perameles and Isoodon. The lingual tier, used as a grinding
platform in other peramelids, is greatly reduced, and the shearing crests are emphasized in both
height and length. Shear occurs when two edges pass each other, and material caught between
them is loaded with two compression forces not in the same plane. Brittle materials fail due to
forced crack propagation. Ductile materials break due to laminar flow of material away from
the points where force is applied. Eventually, the material tears, or fails in tension, as half
flows one way and half flows the other way. Crushing occurs between two surfaces brought
together with no translational motion along the plane perpendicular to the axis of occlusion.
Grinding is essentially crushing with an added translational component so that material is
subjected to both compression and torsion. These definitions are modified after Rensberger
(1973), Rosenberger & Kinzey (1976), Frazetta (1988) and Sanson (1989).

Molar occlusion in Chaeropus consists of the lower molars moving antero-lingually across
the uppers as the jaw closes (Fig. 3A,B). The apices of the lower open triangles, the
protoconids and hypoconids, pass between the paracones and metacones. The protoconid passes
between the paracone and metacone of adjacent upper teeth and along the embrasure bounded by
the postmetacrista and preparacrista. The embrasure between the uppers allows this to occur.
Similarly the hypoconid passes between the metacone and paracone of the opposing upper
molar. Wear facets and altrition striations along the cristae and cristids indicates tooth-tooth
contact, and it is interpreted that shearing occurs between these edges and their occluding
counterparts. Abrasion striations on the flat areas of the buccal tier enclosed by these crests
indicates tooth-food-tooth contact. Food caught in this arca would be stretched across the
triangles, cut by the shearing edges as the teeth move through one another and deposited in the
intervening valleys. Presumably, the paracone and metacone grind such material caught in the
valleys of the lowers, and the protoconid and hypoconid would do the same in the upper
valleys.

EXAMINATION OF GUT CONTENTS

Dissection

The specimen, drained of alcohol, had a body mass of 300 gm and a body length of
approximately 20 cm. The caecum was found to be a simple, unsacculated organ
approximately 5 cm in length, and was relatively empty in this particular specimen. The colon
was relatively long for such a small animal, about 210 cm. The tissucs were hardened by the
preservative but appeared to have been considerably expanded in life (Pl. 3). Material from the
colon contained pieces of fibrous plant material. Digesta from the caecum was made up of fine
particles. All samples contained substantial quantities of grit.

Light Microscopy

Sections of digesta from the distal caecum contained very fine particles of plant material,
the largest of which were only a few cells across. Most of the cells at the edges of these
particles were lysed and empty. Most of the inner cells still contained cell contents. Some
plant fragments contained fibre bundles. Groups of bacteria were apparent (Pl. 4A), and their
presence confirmed by TEM.

DIET OF CHAEROPUS ECAUDATUS - 235

B
upper molars
A—+— P
——— lower molars L

Figure 3. Stages in the occlusal cycle of Chaeropus. A, the beginning of occlusion with the buccal cusps
of the upper and lower molars aligned: _B, towards the end of the occlusal cycle with the lower molars having
moved lingually and slightly anteriorly across the upper molars.

Particles of plant material found in the proximal caecum were somewhat larger than those
of the distal caecum. Often the cells remained attached to a midrib. Again, lysed cells appeared
to have lost their cell contents, although this could have occurred during preparation. These
empty cells were most often at the edges of the fragment, or surrounding a crack in the
fragment. Large clumps of bacteria were again present (PI. 4B). Histologically, plant material
in samples from the caecum appeared to be from a grass (T.P. O'Brien, pers. comm.). Particles
in sections from the proximal colon were much larger than those in the caecum. Sections of
grass nodes were visible in some preparations (PI. 4C), but the presence of small quantities of
very dark brown-staining material, probably tannins, suggests that dicotyledonous plant
material made up part of the diet, since grasses do not normally contain tannins,

Very little digesta was found in the stomach. Fine particles of plant material were present,
along with small rafts of bacteria (Pl. 4D). There was no evidence of the animal having
ingested anything other than plant material and grit.

Transmission Electron Microscopy (TEM)
TEM confirmed the results given above. Bacteria were found to be present in sections

prepared from the proximal caecum (PI, 5A), the distal caecum (PI. 5B), the proximal colon
(Pl. 5C) and from the stomach (PI. 5D). Sections of plant material were found only in the

236 - WRIGHT ET AL.

distal caecum, but the presence of plant fragments in light microscope sections from other parts
of the gut indicates that this was merely a sampling error.

DISCUSSION

The molars of Chaeropus show a suite of dental characters which suggest that it is a
specialist herbivore. The post-paracrista and the pre-metacrista are prominent in the upper
molars, and they are of similar importance to the pre-paracrista and post-metacrista. This is not
the case in any of the other peramelids, where the two central crests (post-paracrista and pre-
metacrista) are lower than the pre-paracrista and post-metacrista. The other bandicoots are
insectivorous or omnivorous taking fungus, seeds, root nodules and vascular plant material in
their diet (Quin 1985, Harrison 1963, Opie 1980, Lobert 1985). None are recorded as taking
grass. In Chaeropus the lower tooth has lost its talonid basin and is composed of two sets of
crests, which form open triangles and oppose those of the upper molar in occlusion. The
lingual tier is much reduced, the high shearing crests extending almost to the lingual edge of
the tooth, so that the teeth are essentially one-tiered. Gaps between the upper molars allow the
protoconid to fit between adjacent upper molars. If these gaps did not exist, the crests of the
protoconid could not shear past those of the upper tooth, As the upper molars of Chaeropus
are square in outline, separation of the upper teeth in particular allows the crests to pass cach
other.

How the gaps are maintained between the upper molars is unknown. It is possible that the
presence of the protoconid in the embrasure resists the tendency of transseptal fibres to pull the
teeth together. Such a system would theoretically keep the anterior and posterior crests of
adjacent upper molars in close occlusion with the crests of the protoconid.

The shear exhibited by the dentition of Chaeropus can be described as ‘fine shear’ (Sanson
1989). By contrast, ‘coarse shear' occurs in systems like the occlusion of the large sectorial
premolars of potoroids and allows thick stems and midribs of plants to be cut, due to a high or
deep shearing crest (Sanson 1989). The crests of Chaeropus, while high compared with those
of the other bandicoots, are low compared with the thickness of twigs, stems and midribs of
dicotyledonous browse and would be relatively inefficient at cutting these materials. The
thickness of the material would prevent the curved edges contacting each other and generating
shear. Thin, laminar leaves of grasses are effectively reduced by the fine shearing teeth of
grazing kangaroos (Sanson 1989), which have crests on a similar scale to those of Chaeropus.
It is interesting to note that Dasyurus, with molars of similar size to Chaeropus, and with an
important shearing component in its occlusal cycle, is also able to shear grasses, reducing them
to small particles (Wright 1988). Chaeropus, the peramelid which Bensley (1903) considered
highly specialised with regard to the dentition, and furthest modified from the tribosphenic
dentition, appears to have been able to shear grasses even more finely than Dasyurus (Wright
1988).

Few studies are available which deal with the diet of Chaeropus. Gerard Krefft, during an
expedition to the Murray-Darling plains, was informed by local Aborigines that these animals
fed on coarse barley grass. He kept captive animals on lettuce, barley grass, bread and bulbous
roots and noted that Chaeropus, unlike other peramelids, did not eat mice or meat which were
presented. Krefft described the dung of wild Chaeropus to be entirely composed of grass, and he
considered them herbivores (Krefft 1865).

Dixon (1988) cites Sturt (1848) as reporting Chaeropus to be partial to flesh, and Gould
(1863) noted that the diet consisted of insects and their larvae and some vegetation (in Dixon,
1988). Dixon (1988) examined two preserved specimens from the Museum of Victoria,
concluding that the faecal pellets consisted almost entirely of grasses. This study, and that of
Dixon (1988), supports the hypothesis that Chaeropus was primarily herbivorous, with grass
perhaps being an important part of the diet. However, the specimens used in both of these

DIET OF CHAEROPUS ECAUDATUS - 237

studies may have been held in captivity for some time before preservation. If this is the case,
results based on gut content analyses will reflect the food items fed to the captive animals,
rather than the natural diet. However, it is perhaps unlikely that an insectivore or omnivore
would readily eat grass even if that was all that was offered.

If Chaeropus was dealing with grasses, tooth wear and the capacity to extract enough
nutrients from a nutritionally poor diet would have been major problems. The shearing crests
of Chaeropus wear vertically, becoming lower. The height of the lingual part of the buccal tier
of the upper molars and the lingual cusps of the lower molars wear first (Pls 6A,B). This is to
be expected since these are the important shearing crests. Wear results in the crests becoming
low and rounded, but they may be kept in tight occlusion by the tendency for the upper molars
to drift together and the presence of the protoconid keeping them apart (Fig 4). It is
noteworthy that the hypoconid, which occludes onto the surface of the upper molar, wears more
than the protoconid, which occludes in the embrasure between successive upper molars.

The problem of such a small animal gaining enough nutrients from a diet of plant material
may have been avoided or iessened by processes such as selective grazing, fermentation and
coprophagy, which appear possible from aspects of this study. Light and transmission electron
microscopy provided reasonable evidence for a large population of hindgut flora, suggesting
that Chaeropus fermented plant material in the caecum and the large colon. Parallels with other
small hindgut fermenters with separation of contents in the colon (Bjornhag 1972), some of
which are caecotrophic, are interesting.

Lee & Cockburn (1985) considered herbivory in marsupials to be confined to animals which
have an adult body weight above 600 g. This is due to the necessity for the digesta to spend a
certain time in the gut to permit bacterial fermentation of the cell wall components.
Chaeropus has a body weight of approximately 300 g. Small herbivores tend to be highly
selective, feeding on relatively easily digested food items rich in energy and protein. Chaeropus
may have been a selective grazer, taking only the nutritious parts of plants and avoiding fibrous
regions of the leaf and the stem, low in readily digestible matter, The presence of grass nodes
in some of the sections supports this argument, as the nodes of grass plants are high in
concentrations of amino acids and proteins (T.P. O'Brien, pers. comm.). Demment (1980) also
noted that small herbivores often select high quality foods, stating that they often chew food
more completely than larger herbivores, to increase its digestibility, since there is a shorter gut
retention time. He also suggested coprophagy as a mechanism whereby small herbivores can
extend the period that the food spends in the digestive tract.

The presence of bacterial rafts in the stomach of Chaeropus suggests that these animals may
have been coprophagous, reingesting faecal or faccal pellets which contain nutrients provided
by the hindgut flora. This is similar to the situation observed in ringtail possums (O'Brien et
al. 1986).

Fine shear is emphasised in the dentition of Chaeropus at the expense of other functions,
and it appears that this bandicoot was able to reduce grasses to fine particles. Shearing in
Chaeropus is extremely fine, and it is unlikely that such fine shear has developed for the
communition of insects, which can be effectively reduced by relatively coarse-shearing crests
such as those of Dasyurus (Wright 1988) and Dasyuroides (Moore, 1986). Grasses are rich in
silica and cause excessive wear in mammalian teeth. Mature plants and older leaves have a
higher silica content than do younger plants and leaves (Lewin & Reimann 1969). An animal
as small as Chaeropus has an advantage over large herbivores in terms of the scale of the
substrate to its absolute energy requirements. It has been suggested that some small herbivores
are able to survive in this way (Norbury e¢ al. 1989). For example, the small Hare Wallaby
Lagorchestes hirsutus has been shown to select the very tips of grasses (Bolton & Latz 1978).

Chaeropus is described here as a specialist herbivore, probably taking a mixture of herbs and
grasses. Selective herbivory, fermentation and coprophagy are suggested traits which may have
enabled this small animal to meet and sustain its metabolic requirements. Behavioural and

238 - WRIGHT ET AL.

A

Figure 4. Wear facets on the shearing crests of Chaeropus molars. A, occlusion of the crests of the upper
and lower molars along wear facets (shaded); B, edges of the two opposing shearing crests along the line "b"
in (A). Dotted lines indicate successive states of wear, changing from planar facets with relatively sharp

edges to facets with rounded edges.

nutritional studies to test these proposals are prevented because of the recent extinction of this
animal, although investigation into the digestive strategies of the extant omnivorous
bandicoots may provide some insight as to how Chaeropus maintained itself on a diet of
relatively lower nutritional quality plant material.

SUMMARY AND CONCLUSIONS

A study of the occlusal morphology and wear of the molars of the extinct bandicoot
Chaeropus ecaudatus, suggests that this animal may have been a herbivore, and more precisely,
a grazer. Examination of a spirit-preserved specimen and light and transmission electron
microscopy of the gut contents supported this hypothesis.

Fine shear was emphasised in the occlusal cycle of Chaeropus, reducing all other functions.
The high crests provided enough shearing amplitude so that Chaeropus was able to reduce plant
material, including grasses, to fine particles. The shearing molars probably released cell

DIET OF CHAEROPUS ECAUDATUS - 239

contents for digestion in the small intestine and prepared plant material for fermentation in the
hindgut. If so, Chaeropus was unique among the Perameloidea, with the capacity for
fermentation of plant material in the hindgut. Coprophagy is suggested as a mechanism
whereby Chaeropus may have prolonged the period that plant material spent in the digestive
tract, enabling more efficient digestion.

Recently Burbidge e¢ al. (1988) have reported dietary information about various extinct or
rare mammals, obtained from interviewing Aborigines from the central deserts and surrounding
areas. They did not present data unless it was corroborated by two or more groups of people.
They state that the reported food of Chaeropus was “termites and ants, including honey-pot
ants" (p. 20). We are unable to assess the information in that report, and can only indicate that
it is not supported by this study.

This study emphasises the value of preserved specimens in the determination of possible
dicts of extinct species, but it also shows that observation of molar wear patterns can lead to
useful speculations about diets of such animals.

ACKNOWLEDGEMENTS

We wish to acknowledge the support and generosity of Ms Joan Dixon, Curator of
Mammals, Museum of Victoria. We also would like to thank Dr T.P. O'Brien, Dr W.J. Foley
and Ms S. Moore, for their advice, comments and discussion. The work was supported by a
grant from the Museum of Victoria.

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(Marsupialia: Macropodidae). Aust. J. Zool. 28: 341-365.

SANSON, G.D., 1989. Morphological adaptations of teeth to diets and feeding in the Macropodoidea In
Kangaroos, Wallabies and Rat-kangaroos. G. Grigg, P. Jarman & I. Hume, eds., Surrey Beatty.
Sydney: 151-168.

WRIGHT, W., 1988. Functional Molar Morphology in Four Genera of Marsupial Bandicoots. Unpublished
Honours Thesis, Zoology Department, Monash University, Clayton.

PLATES

Plate 1. Scanning electron micrographs of the upper molars of Chaeropus. A, Upper tooth row looking
posteriorly from M2 to M4; BB, postmetacrista of M? on a relatively unwom specimen; note the planar facet
with attrition striae; C, preparacrista of M? on a relatively worn specimen; note the rounded abrasion facet.
D, occlusal view of the narrow lingual tier between the paracone and metacone. Abbreviations as for Fig.1.

Plate 2. A, scanning electron micrograph of the lower molars Mz and M3 of Chaeropus; B, metacristid of
the worn lower molar of Chaeropus; note the rounded nature of the crest and the well developed attrition striae.
A = anterior.

Plate 3. Gut of Chaeropus ecaudatus (MV C5858) indicating caecum and expanded colon and showing sites
from which samples of gut contents were taken: 1, distal caecum; 2, proximal caecum, 3, proximal colon;
4, stomach.

Plate 4. Light micrographs of sections prepared from the gut contents of Chaeropus ecaudatus (C5858)
(X480). A, plant material and bacteria prepared from the contents of the distal caecum; B, plant material
prepared from the contents of the proximal caecum; C, section of a grass node from the contents of the
proximal colon; D, plant material and bacteria from the contents of the stomach.

DIET OF CHAEROPUS ECAUDATUS - 241

Plate 5. Transmission electron micrographs of sections prepared from the gut contents of Chaeropus
ecaudatus (C5858). A, bacteria and digesta from the contents of the proximal caecum (X14,000); B, bacteria
and digesta from the contents of the distal caecum (X27,000); C, bacteria and digesta from the contents of the
proximal colon (X20,000); D, bacteria surrounded by digesta from the contents of the stomach (X27,000).

Plate 6. Scanning electron micrographs of wom molars of Chaeropus ecaudatus. A, lingual view of M2
showing reduced height of the tooth on the lingual edge; note the embrasure between adjacent molars; B,

posterior view of the left upper tooth row showing the reduced height of the lingual part of the buccal tier.

PLATE 1

242 - WRIGHT ET AL. PLATE 2

PLATE 3

PLATE 4 DIET OF CHAEROPUS ECAUDATUS - 243

244 - WRIGHT ET AL. PLATE §

PLATE 6 DIET OF CHAEROPUS ECAUDATUS - 245

A

1.2mm 1.2mm

246 - WRIGHT ET AL,

Ektopodon was an enigmatic animal. In this restoration it is shown about to catch an insect,
but its diet is far from understood (see Pledge, this volume). The genus is known to range from
the Miocene of central Australia into the Early Pleistocene of coastal Victoria. (From Rich &
van Tets 1985, with permission of The Museum of Victoria).

CHAPTER 9

RECONSTRUCTING THE
NATURAL HISTORY OF
EXTINCT ANIMALS:
EKTOPODON AS A CASE
HISTORY

Neville Pledge!
IntrOdUCHION....eFetsseidie eos seosee Fags tee Poo See See oie tie 248
Reconstruction and Restoration...............:eeeeeee 248
Methods and Examples............ccccccsseeceeeeees 248
Case Study - Ektopodon..........cccccce ce eeeee eee ences 251
History of Discovery ........... cc ece cece eee ee teen ees 251
Diversity and Range of Ektopodontidae ........ 253
Morphology ....... cece cece eseeeeeceeeeeeseeseeseeenes 254
Relationships ...........eceeceseecee ee neeceneeseeeeees 256
Interpretation Of EkLOPOdON..........ccceeeeeeee eens 256
Habitat... oss bs nde stead bea hiee ese de bs See 256
DDT Co aes Ri Ret okes oateeine da Deady nesdaceet egg e'es obede 256
Teat-Batings esol sichweceiiss elbseenleee ee’ 257
Insect-Eating..... eee ceeeeseeeeceeeeeeeeee ees 257
Aquatic-Invertebrate Eating...............242+ 258
Sed Bane in. ce sentedee. Saindveeecasdeenacedd eee 258
Modern Counterparts ...........ceeeeceeeeeeeeeee eee 259
SUMMMALY. feo 5 book ceteet AAR tebe eeins cence sind sae'e' 259
IRELETENCES 27s. heer PE eee ieos eh e dor eouetss 259
Plates Meee oe tl AAs Mi. came dhdestweasl tse cdeet 261

1 Department of Palaeontology, South Australian Museum, North Terrace, Adelaide, South Australia 5000,
Australia.

248 - PLEDGE
INTRODUCTION

Many people regard the restoration of extinct animals as the acme or culmination of a
palaeontologist's work, in that it summarizes graphically the total knowledge of the animal's
body form, habits, habitat, diet and even social life.

Frequently, today, we see artists' renditions of dinosaurs, and other prehistoric animals,
often fanciful or exaggerated to fill some market requirement (in the case of children’s toys and
models). Yet, for the most accurate reconstructions, complete skeletons (or at least
representatives of essential bones) are needed, and, for most tetrapod species, this kind of
material is rarely available. Most vertebrate fossil species are represented by incomplete
skeletons, or even by only isolated teeth.

For well over a century, it has commonly, but mistakenly, been believed that
palaeontologists can reconstruct an animal from as little as a single bone or tooth. This myth
arose from the undeniable skill of Baron Georges Cuvier (Simpson 1953), who demonstrated
that it was possible to identify a species from such scanty material. However, it is not
possible to reconstruct extinct forms accurately until most of the skeleton is known. How,
therefore, can palaeontologists make such statements as they do, about these extinct species?

The answer lies in detailed morphological comparisons of the fossils with perceived related
species (living, if possible) or ecological analogues.

RECONSTRUCTION AND RESTORATION

Strictly speaking, the term "reconstruction" should be reserved for the process of
reassembling the skeleton of a fossil animal, while "restoration" is the more subjective matter
of depicting the soft tissue - muscles, tendons, skin and pelage - and its habits and habitat,
based on interpretation of skeletal anatomy, sedimentology, other fossils and, indeed,

palaeogeography (Harris 1987).

METHODS AND EXAMPLES

In the best of circumstances, with a complete skeleton to work with, the bones are first
reassembled in their correct anatomical positions. The palaeontologist, or his anatomist/artist,
then depicts this graphically and after carefully studying the muscle attachment areas (scars) on
the bones, draws in the muscles in the relative bulk indicated by those scars (e.g. Murray
1984). Allowance is made for fat and other soft tissue based on experience with modern
animals, and the "skin" is added graphically. This gives an idea of the "bald" animal. For
mammals, fur/hair is added at the artist's discretion. Climatic indicators from the geological
setting may dictate how much hair is needed, and sometimes also the size of the ears - organs
that otherwise are purely speculative.

When only a small portion of the skeleton is preserved, the problem is much more difficult
and the result much more speculative, involving, as it does, so many assumptions. The main
assumption made is that the unknown species must be similar, in greater or lesser degree, to its
closest known relatives. For mammals, such relationships are best exemplified by their teeth,
which besides indicating dietary preferences, bear many subtle characters that can link or
distinguish species.

Palaeontologists are helped by the fact that the teeth in a mammal's jaws are differentiated
into serving various functions: nipping incisors, stabbing canines, and cheek teeth (premolars
and molars), which may be adapted for cutting, as in carnivores, or grinding as in horses and
cattle. Carnivores do not need to grind up their meat, as flesh is readily digested, but plant
material can vary considerably and may need extensive preparation before digestion is efficient.

NATURAL HISTORY OF EXTINCT ANIMALS - 249

Insectivores generally have teeth similar to carnivores in morphology and function, as they
need to penetrate the tough chitinous skeleton of the insects. Their molars, however, may be
modified somewhat for crushing their prey to extract the nutrient, and may also be adapted to
specific types of prey (Sanson 1985).

® Riversleigh

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L

SOUTH AUSTRALIA|
L. Eyre f° L. Ngapakaldi
¢ L. Palankarinna_

L. Biome @ u Tarkarooloo
L. Pinpa

t

ADELAINE
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Figure 1. Locality map, showing distribution of ektopodontid fossil finds.

Animals with omnivorous tastes, i.e. those able to eat a wide variety of foods - such as
bears, pigs, humans and brushtail possums - generally have bunodont molar teeth, with low
crowns and relatively simple cusps. Those that are primarily herbivorous have molars with
more complex structures of ridges, which act against each other like shears to cut or rasps to
grind the vegetation into small particles thus efficiently breaking down the cell walls. The
teeth of browsers, living on leaves and other soft vegetation, remain relatively low crowned and
simple, but those of grazers such as horses and grey kangaroos, living primarily on grass,
which contains microscopic particles of organic silica (phytoliths) and surface dust, have high-
crowned teeth, often growing continuously throughout life (e.g. horses, rabbits, wombats). It
is noted that amongst Australian marsupials at least, obligate arboreal leaf eaters, such as
koalas and ringtail possums, have selenodont molars, in which the cusps have developed into

250 - PLEDGE

longitudinally oriented crescentic crests (selenes) instead of the transverse lophs of kangaroos
and diprotodontids.

In addition, most specialized herbivorous mammals (such as horses, cattle, rabbits, koalas,
kangaroos and diprotodontoids) also possess a gap or diastema between the canines and the
cheek teeth (premolars or molars). This is generally believed to allow the tongue to
manipulate the wad of partially chewed vegetable matter efficiently without being bitten. The
diastema is even more pronounced in taxa that have lost (by reduction) their canines.

It is often difficult to determine with any confidence what the dictary preferences of an
extinct herbivore are. Sanson (1978, 1980, 1982) has shown how interpretations of some
fossil kangaroos may have been wrong. Most of the large living kangaroos are primarily
grazers, preferring grass to leaves, but amongst the large extinct species only Procoptodon
seems to have been primarily a grazer, the others (Sthenurus, Simosthenurus, Protemnodon,
Prionotemnus and Troposodon) having relatively low-crowned, simple-crested molars appear to
be adapted to lower-fibre vegetation. Conclusions about diet based on zoo observations of
captive animals can be erroneous, as preferences may be for artificial or not-naturally available
foods (e.g. Clemens & Kielan-Jaworowska 1979).

For totally extinct groups, such as the multituberculates with which Ektopodon was once
compared, we must rely heavily on analogy. Indeed, the ecological nature of multituberculates
has long been controversial, with agreement only on one genus, Taeniolabis, which is
considered for various dental reasons to have been a rodent-like herbivore (Clemens & Kielan-

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Figure 2A, B. Extracts from R.A. Stirton's field note book, 28 July 1962 and 3 August 1962. (With
permission, University of California).

NATURAL HISTORY OF EXTINCT ANIMALS - 251

Jaworowska 1979). The others, with their sectorial premolars, are now believed to have been
generally herbivorous, or omnivorous, based in part on comparisons with the living rat
kangaroo, Bettongia.

With Ektopodon, we are near the opposite end of the spectrum, so far as restoration goes,
as the species are known from such limited material, the family has no living representatives,
and only distant relatives are perceived.

CASE STUDY - EKTOPODON

HISTORY OF DISCOVERY

The existence of the mammal family Ektopodontidae was first realised in 1962 when five
very odd teeth were recovered from a fossiliferous conglomerate in the Miocene Wipajiri

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Formation, a fossil stream deposit exposed at the Leaf Locality, Lake Ngapakaldi, east of Lake
Eyre, South Australia (Fig. 1).

252 - PLEDGE

Initial interpretation by the late Professor R.A. Stirton, whose party had found the
specimens, was that the teeth possibly represented a monotreme or even the long extinct
Multituberculata (R.A. Stirton, Field notes, 28 July 1962; Fig. 2A), whose molar teeth are
characterised by rows of numerous similar tubercular cusps (Fig. 3). However, as more
specimens were found, it was realised that, whereas in multituberculates those rows lie
longitudinally on the tooth crowns, the wear facets and roots of the newly discovered teeth
indicated that the rows of cusps had a transverse orientation (ibid., 3 August 1962; Fig. 2B).

Figure 3. Lower teeth of the multituberculate Meniscoessus robustus, showing longitudinal rows of cusps on the
molars, in occlusal view. (From Clemens 1963).

Consequently, it was several years before the teeth were described and named (Stirton,
Tedford & Woodburne 1967) - Ektopodon means odd tooth - during which time they had been
compared with many groups. Finally, with some doubt, they were allied with the
monotremes, primarily because the platypus (Ornithorhynchus anatinus) in its juvenile stages
has some unusual teeth with rows of tubercles. Several workers, however, were still
dissatisfied with this interpretation, belicving that the tecth must represent some unusual type
of marsupial. Discoveries in Oligo-Miocene sediments of the Namba Formation of Lake Pinpa
in 1971 and of the Etadunna Formation at the Tedford Locality, Lake Palankarinna in 1972,
strengthened this belief. A handful of teeth of a new genus (Chunia) similar to, but less
advanced than Ektopodon, were found (Woodburne & Tedford 1975, Woodburne & Clemens
1986a). They showed features akin to those of the phalangeroids, particularly Phalanger, the
cuscus (Archer 1976, Woodburne & Clemens 1986a).

In 1976, proof of its marsupial nature came when I discovered a fragmented jaw of
Ektopodon at yet another locality (Mammalon Hill) at Lake Palankarinna in younger beds of
the Oligo-Miocene Etadunna Formation. This jaw contained the last premolar and first three of
four molars, in a typically diprotodontan marsupial dentition, and further demonstrated just how
odd the teeth were. Within weeks another discovery of this new species was made in slightly
older beds (upper Namba Formation), 330 km away at Lake Tarkarooloo near Lake Frome
(Pledge 1982, 1985, 1986). Eventually nearly 40 isolated or fragmentary teeth were found,
together with three toothless mandibles, and a possible incisor - the largest sample of the taxon
yet obtained. Continued work at Lake Palankarinna has yielded a maxilla with a partial molar,
a first upper molar and a mandible fragment with a molar, of Chunia (Woodburne & Clemens
1986a), and several more teeth and another jaw of Ektopodon (Fig. 4, Pl. 1).

The geographic and temporal ranges of the family have been greatly extended, first, by the
discovery of several associated molars of a second new genus (Darcius, Rich 1986) near
Hamilton, Victoria. These specimens were found by T.H. Rich and Party in a palaeosol
developed on the Grange Burn Formation that was overlain by a basalt flow dated at 4.46
myBP. Secondly, in 1986, a fragment of molar was found in the rich mid Miocene deposits at

NATURAL HISTORY OF EXTINCT ANIMALS - 253

Figure 4. A, dentary of Ektopodon stirtoni, SAM P19509, from Mammalon Hill, Lake Palankarinna Fossil
Reserve, South Australia, showing twisted occlusal plane; B, E. sp. cf. E. stirtoni, P19963, first and
second upper molars, showing imbricated bifurcating plate-like cusps arranged in transverse lophs, and
referred premolar (NMV P48768).

Riversleigh, northwest Queensland. (Archer, pers. comm.). The youngest ektopodontid appears
to be an undescribed molar from the Early Pleistocene of Portland, Victoria (T. Rich & M.

Whitelaw, pers. comm.).

DIVERSITY AND RANGE OF EKTOPODONTIDAE

The diversity and stratigraphic range of the Family Ektopodontidae can be seen in Table 1.
Fossils of the ektopodontid species are relatively rare and almost entirely restricted to isolated
teeth. There is a maxilla (upper jaw) fragment of Chunia illuminata, with half of a molar
preserved, and a fragment of dentary, also with one molar. Ektopodon sp. cf. E. stirtoni is
represented by several toothless dentaries, while Ektopodon stirtoni itself is known from two
dentaries, preserving between them the full lower cheek dentition, and there are numerous loose

254 - PLEDGE

teeth of these species. Ektopodon serratus is known only from isolated teeth and a toothless
jaw, and Chunia omega from a single tooth. Darcius duggani is also known from isolated
teeth, but some are known to be associated as they were found in correct anatomical position
with only a fragment of the rotted jaw bone remaining.

NN eee eee

Table 1 Stratigraphic Range of Ektopodontidae

RIVERSLEIGH LAKEEYREBASIN TARKAROOLOO BASIN HAMILTON

- 1100 KM - - 330 KM - - 700 KM -

Pliocene Darcius
duggani
(>4.46
myBP)

Middle Ektopodon Ektopodon

Miocene _ sp. indet. serratus

and

Ektopodon n.sp.

Ektopodon
stirtoni
Early Ektopodon sp. cf.
Miocene to E. stirtoni
Late Oligocene and
Chunia omega
Chunia illuminata Chunia sp. cf.

C. illuminata

es SSS:2795°0—00O“aO09aoaaoooa_—a—=

MORPHOLOGY

To judge by the jaws found at Mammalon Hill and elsewhere, Ektopodon was a marsupial
the size of acat (Felis catus).

The molar teeth of the Ektopodontidae all have a basic bilophodont pattern (i.e. have two
transverse ridges or loph(id)s), with the loph(id)s composed of numerous similar overlapping
cusps. Another characteristic of the family is that the first upper molar (M2) also has an
additional short anterior loph with a few cusps. Blade-like crests extend radially (in Chunia) or
longitudinally (Ektopodon and Darcius) from the cusps, and may bifurcate. In posterior molars
these crests become lower and reticulate. Species differ in the number and complexity of the
cusps. The number varies from 5 or 6 in Chunia to 8 in Ektopodon stirtoni, 8 or 9 in
Ektopodon serratus and even more in an undescribed species of Ektopodon, while there is a
concomitant simplification of crests. The youngest described species, Darcius duggani, is
anomalous in having only 3 or 4 cusps on the lophids, but relatively simple crests, indicating
a different line of descent.

The genus Ektopodon is notable in that its anterior lower molars are very wide - the width
equal to or greater than their length - and almost as wide as the occluding uppers: they actually
overhang the inner face of the dentary! This is unlike most other marsupials. There is a

NATURAL HISTORY OF EXTINCT ANIMALS - 255

marked molar size gradient (also present in upper molars of Chunia), and the overall occlusal
surface of the cheek teeth in the dentary shows a distinct helicoidal twist (Fig. 4a).

The upper and lower molars of Ektopodon species are remarkably similar, but may be
distinguished by the lophids of lower molars being slightly oblique to the midline of the tooth
row, the upper teeth having three or four roots, and the lowers two roots. The teeth are
relatively short and broad, upper teeth being broader than long, and even the lower molars of E.
serratus are wider than long. In both upper and lower molars, size decreases markedly from M2
to MS, the loph(id)s becoming lower and obtuse posteriorly, so that MS has a very low
protoloph(id), and the other loph is almost non-existent. The loph(id)s each have six to eight
cusps. The largest cusps are lingual on the lophs and labial on the lophids, and grade to the
smallest and most appressed at the opposite ends. From the apex of each cusp there is at least
one anterior and one posterior longitudinal crest (pre- and postcrista), which sometimes divide
basally. Adjacent cristae may be joined by fine transverse ribs or struts, and this tendency is
increased away from the main cusps, and also in more posterior teeth, where it culminates in
MS having an irregular network of anastomosing cristae and struts.

The first upper molar, M2, is exceptional in having a pentagonal outline with a short
additional loph, anterior to the main ones and supported by its own root (Fig. 4B). This loph
has three cusps, and with the contiguous upper premolar (P3), may form a "carnassial" blade to
occlude with the short, bladed lower premolar. However, in the Mammalon Hill jaw, the P3 is
tilted forward out of the plane of molar occlusion, so it is difficult to imagine how it would
have functioned as a cutting tooth.

The referred lower incisor is phalangeroid in basic form, rather short and laterally
compressed, and of the right order of size and cross-section to fit the alveolus of the jaw.

The dentary itself is solidly built, thickest and deepest under the first two molars, and short.
There is a very short diastema of only a few millimetres, and the alveolus of the large lower
incisor suggests it curved up fairly sharply. There was also, apparently, a tiny I2 or canine
immediately adjacent to the incisor. The premolar (P3) in front of M2 is relatively small but
solidly built. The mandibles do not appear to have been firmly ankylosed, but probably were
not as flexibly joined as those of Macropus.

Nothing is known of the palate of Ektopodon, but we may assume, because of the dental
similarity, that it was like that of Chunia illuminata. Assuming the cheek-tooth rows are
parallel, we may mirror-image the known maxilla of Chunia to make a composite palate (PI.

1A). This is very wide and short. The premolars align with the oblique outer face of M2 to
produce an extended cutting edge, and it can be seen that the face is very broad and blunt
subtending an angle of 120°, with only a tiny muzzle protruding from it. We have no idea of
the size of the premaxillae, and hence of the nasal region, Presumably the upper incisors were
small.

The maxilla preserves the root of the zygomatic arch (cheek bone) and, therefore, the lower
part of the orbital cavity (eye socket). This is quite large, and because of the wide facial angle,
is directed forward in such a manner as to give almost, if not entirely, full binocular,
stereoscopic vision. Because the dentaries of Ektopodon appear to complement the dental
arrangement of the maxilla of Chunia, we can reasonably assume that Ektopodon also was very
short-faced, blunt-snouted and large-orbited (Pl. 1B).

Many teeth show occlusal wear facets. On the molars, these are situated symmetrically on
each face of the loph(id)s, but generally do not affect the apical crest which tends to be rounded,
and do not extend to the bottom of the transverse valley. Relatively coarse, irregular, more or
less vertical gouges may be seen on these facets, more commonly towards the apex, while very
fine, close, parallel, transverse striae show best on the lower parts. Sanson (1980) has
summarized recent work on mastication and occlusal wear. The coarser gouges would seem to

256 - PLEDGE

be produced by the initial "puncture-crushing" phase, while the fine striae are the result of
tooth-tooth contact during chewing.

RELATIONSHIPS

The discovery of a dentary of Ektopodon stirtoni (Pledge 1986) proved the marsupial nature
of the Ektopodontidae. The dental formula of one incisor, a premolar and four molars is
characteristically diprotodontan. The family was assigned to the Phalangeroidea on the basis of
remnants of the cristid obliqua and stylid cusps in lower molars of Chunia illuminata
(Woodburme & Clemens 1986a, Woodbume 1987), these features being seen in some Phalanger
species (Archer 1987).

Within the family, the cristid obliqua is most strongly developed in Chunia sp. cf. C.
illuminata, which is considered to be the oldest species. This feature is not seen in Ektopodon
spp. or Darcius duggani (Woodbure 1987). The cristid obliqua and stylid cusps are, therefore,
seen to be primitive character states - the latter occurring in Chunia species and Darcius, and as
very reduced structures in M2 and M3 of E. stirtoni. Consequently, the number and
complexity of loph(id) cusps can be seen to be definitive character states, with fewer and more
complex cusps in Chunia species, evolving in two directions to give Ektopodon species with
more and simpler cusps, and Darcius duggani with fewer cusps of less complexity than those of
Chunia. There is also a general, though not consistent, size increase apparent, and in
Ektopodon species a widening of lower molars.

Thus, Ektopodontidae are considered to be a sister group to the Phalangeridae, and derived
from a phalangerid-like ancestor. Two branches developed, one leading to Chunia species and
the other to Ektopodon species. These relationships are summarized by Woodburne &
Clemens (1986b). They conclude that Darcius probably arose near the base of the Ektopodon
line, but I believe it is more closely related to Chunia, since it seems to show a simplification
of Chunia characters and a conservatism of lower molar proportions not seen in Ektopodon
species.

INTERPRETATION OF EKTOPODON

HABITAT

Stereoscopic vision is a characteristic feature of most arboreal mammals, since the ability
to judge distances accurately is essential for moving about in trees. It is also a feature of many
hunting mammals. Primates (including Man) and cats are the best examples of vertebrates
with stereoscopic vision, although this feature may also be seen to a lesser extent in the
Australian possums (and perhaps best developed in the ektopodontids). The stereoscopic
arrangement of the eyes of the ektopodontids, therefore, argues strongly for such a habitat or an
ancestry with such habits. Arboreal animals tend to have considerable manual dexterity. The
large size of the orbits suggests a nocturnal habit.

DIET

The unusual teeth of Ektopodon beg the question: What did it eat? A number of
suggestions for the diet of Ektopodon are posed below, together with reasons for and against
them, but no definite conclusions can be drawn. These possibilities are restricted by the
morphology and wear surfaces of the teeth and also by features of the maxilla.

NATURAL HISTORY OF EXTINCT ANIMALS - 257
Leaf-Eating (also flowers and fruit).

Archer (1981) suggested an arboreal, browsing niche for Ektopodon, based on its
phalangeroid relationships, and this is supported by the molar tooth morphology. Amongst
marsupials, obligate arboreal folivores, such as koalas and pseudocheirids, have selenodont
molars so that a shearing mode in food preparation is maximized (see Kay & Hylander 1978).
The ektopodont dentition could be interpreted as an extreme form of this condition, with
multiple en chevron selenes formed by the slightly curved pre- and postcristae.

The closest living relatives of Ektopodon are considered to be phalangerids (Woodburne &
Clemens 1986), particularly species of Phalanger s.l., the Cuscus. Some cuscuses have
crenulations in the molar tooth enamel that seem to parallel the much stronger development of
cristae in the ektopodontid molars (Archer 1976) (Pl. 2). Phalanger spp. subsist on a diet
mainly of leaves and fruit, which are crushed between their bunolophodont molars. They are
also opportunistic carnivores. The rasp-like surface of the molars of Ektopodon, moving
transversely as indicated by wear striae, could presumably deal with much coarser leaves.
Cuscuses also have relatively blunt faces with large eyes, and are crepuscular or nocturnal in
habits.

It should be noted here that there is more than a slight resemblance between the molars of
ektopodontids and the Giant Panda Ailuropoda, particularly M! and M2 of the latter (see
Gregory 1936) where two transverse rows of cusps are developed (PI. 3). The Giant Panda
feeds bamboo stalks into the side of the mouth and thoroughly masticates them a section at a
time (Chorn & Hoffmann 1978).

Insect-Eating

Insectivores, such as the small dasyurids, typically have teeth with sharply pointed cusps
for gripping and piercing their prey, and a rather sectorial occlusion for cutting the tough
cuticle. Differences in dentitions may reflect dietary selection (Sanson 1985). Although
generally herbivorous and having teeth unspecialized for the food, many possums relish any
insects that come their way (Strahan 1983). Ektopodon could have done likewise, its heavy
teeth crushing the cuticle efficiently to extract the nutrients.

Most insectivores have long, sharply-pointed snouts with which they can probe nooks and
crannies for their prey. Ektopodon obviously could not do this, although it may have been
able to compensate by using its claws (which are still unknown), in the same way that the
specialised primate Daubentonia (the Aye-Aye, Pl. 4; Owen 1863) and the Striped Possum,
Dactylopsila trivirgata, do (Hildebrand 1974: 636-7; Strahan 1983: 144-5).

However, this modification may not have been necessary for Ektopodon, as there was
probably an abundant and easily caught prey available in the forests that covered the interior of
Australia during the Miocene. A large proportion of animal biomass in modern tropical and
subtropical forests consists of insects (Norris 1970: 110), much of it in larval form, i.e.
caterpillars and grubs, that might be more active at night when bird predators were absent.
This situation probably characterised the treetops of the Oligo-Miocene forests where
Ektopodon lived. Caterpillars are a slow prey, often abundant, and easily eaten. Ektopodon's
tasping molars would despatch them efficiently.

A variant of this hypothesis is that Ektopodon foraged on the forest floor for grubs and
insects. Its teeth, as indicated above, could handle such a dict, but the foreshortened face would
seem to argue against the idea. It would seem also to be a “waste” of stereoscopic vision,
unless this were a left-over feature from some ancestral form.

Insectivory is considered here because of the unusual, almost unparalleled structure of the
teeth of ektopodontids. However, apart from the morphological differences, the large body size

258 - PLEDGE

argues against a purely (or largely) insectivorous diet. The largest modern insectivorous
phalangeroid (Cercartetus caudatus) is much smaller than Ektopodon would have been.
Arborcal insectivores are limited in size by the abundance of food (Kay & Hylander 1978).

Aquatic Invertebrate-Eating

A variation on the preceding hypothesis is supported by the stereoscopic vision that is
essential in hunting, and by the crushing/grinding molars. We may compare Ektopodon with
Enhydra, the sea otter (PI. 5; Estes 1980), and Ornithorhynchus, the platypus (Woodburne &
Tedford 1975), which have broad crushing teeth.

The large orbits (and hence large eyes) of ektopodontids may seem at variance with this
habitat, since beavers and platypus have small eyes, but seals and otters do have rather large
eyes. Otters also have rather blunt faces and stereoscopic vision.

No known aquatic marsupials exist in Australia, the role being filled by the platypus and
the lately-arrived water rat, /7ydromys. During Oligo-Miocene times, however, there was an
abundance of permanent streams and lakes in the interior of Australia, inhabited by the
primitive platypus Obdurodon insignis (Woodburne & Tedford 1975, Archer, Plane & Pledge
1978), crocodiles, turtles and fish. While fish remains are abundant in the Miocene sediments,
aquatic invertebrates are also recorded (Stirton, Tedford & Miller 1961), and must have been
plentiful to support the rich aquatic bird faunas known to have existed there. There is no
reason why a marsupial should not have exploited this niche also, as does the otter-like water
opossum or Yapok, Chironectes minimus, in central and South America (Walker 1964:25).
However, it is probably only a coincidence that most ektopodontid remains have come from
stream channcl or lake deposits.

Seed-Eating

Until the arrival in Australia of rodents, in Pliocene times (Archer & Bartholomai 1978),
the "gnawing" mammal niche seems to have been vacant. Yet it is likely that some mammal
group was engaged in seedeating, and Sanson (1985) notes that Burramys parvus cats seeds of
Hovea sp., using its plagiaulacoid premolars to break them open. Burramys wakefieldi (Pledge
1987) is known from the same beds as Ektopodon at Mammalon Hill. Elsewhere I have
suggested (Pledge 1986) that Ektopodon occupied the carpophagous niche, although the early
radiations of seedeating parrots may have precluded major exploitation of this food source (Rich
1975, Archer 1981: 1480).

Rodents typically have rootless incisors for grinding through the tough pericarp of seeds,
while the molars in most groups are rooted and have a complicated rasping pattern of enamel
(Romer 1966).

In Ektopodon, the unknown upper incisors probably were small and relatively weak and
unfit for the task of breaking away hard seed coats. The referred lower incisor also seems
inappropriate for this, although its rather short spatulate or spoon-shape may have been good
for nipping seed pods or scraping grains from seed heads.

Breaking open seed cases could have been accomplished by the combination of the small
but sturdy premolars and the buccal crest of M2 and possibly also by direct pressure of the
M2's. The more posterior molars are lower crowned and seem relatively weak, but gouges seen
on the wear facets may support such a puncture and crushing process. Fine transverse striae
also seen indicate that in the chewing process the jaws moved from side to side. Sanson (1980)
notes that rats "rarely, if ever, puncture-crush their food, possibly due to the efficiency of their
incisors" in preparing it for chewing. Ektopodon, therefore, differs functionally from rodents,
but this does not rule out seed/grain eating.

NATURAL HISTORY OF EXTINCT ANIMALS - 259

As a source of food, grasslands were still a minor part of the vegetation (Truswell & Harris
1982), but other plant groups such as the Acacias, Cassias and other Leguminosae, Casuarina
and conifers probably produced an abundance of seed. Such seed gathering could have been an
arboreal, nocturnal task.

MODERN COUNTERPARTS.

Dentally, there is no modern mammal like Ektopodon, which, thereby, earns its scientific
name. Ecologically, however, it may be matched by some primates and by some cuscuses,
being a binocular-visioned arboreal herbivore, living primarily on leaves and fruit, but also on
seeds and nuts and occasional insects and small vertebrates.

SUMMARY

Ektopodon was a marsupial with a particularly unusual dentition: bilophodont molars
having numerous, transversely rasp-like longitudinal cristae. A maxilla of the related genus
Chunia indicates that these animals had large eyes, placed stereoscopically in a very broad face,
suggesting both nocturnal habits and an arboreal habitat. An aquatic lifestyle is also possible.
These few data allow several dietary preferences to be proposed but, in the light of such limited
information all are speculative.

1) leaves and fruit,

2) insects (particularly caterpillars and grubs),
3) aquatic invertebrates,

4) seeds and grains.

It is likely, however, that, as with its nearest living relative, the Cuscus, Ektopodon was
fairly omnivorous and opportunistic in its feeding habits.

The interpretation of the sparse remains of Ektopodon, therefore, is an example of the
frustration, and indeed futility, of attempting detailed reconstructions and restorations with only
limited data. This is not to say that the task should not be attempted - perhaps it should be
done more often - but rather that the limits of confidence should also be spelled out.

REFERENCES

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ARCHER, M., 1981. A review of the origins and radiation of Australian mammals. In Ecological
Biogeography of Australia. A Keast, ed., Junk, the Hague, Monogr. Biol. 41: 1437-1488.

ARCHER, M., 1987. A possum of a very different kind. In The Antipodean Ark. S. Hand and M. Archer,
eds., Angus & Robertson, Sydney: 67-69.

ARCHER, M. & BARTHOLOMAI, A., 1978. Tertiary mammals of Australia: a synoptic review. Alcheringa
2(1): 1-19.

ARCHER, M., PLANE, M.D. & PLEDGE, N.S., 1978. Additional evidence for interpreting the Miocene
Obdurodon insignis Woodbume and Tedford, 1973, to be a fossil platypus (Omithorhynchidae :
Monotremata) and a reconsideration of the status of Ornithorhynchus agilis De Vis, 1985. Aust. Zool.
20(1): 9-27.

CHORN, J. & HOFFMANN, R.S., 1978. Ailuropoda melanoleuca. Mammalian Species 110: 1-6.

CLEMENS, W.A., 1963. Fossil Mammals of the Type Lance Formation. Part 1. Introduction and
Multituberculata. Univ. Calif. Publs. geol. Sci. 48: 1-105.

CLEMENS, W.A. & KIELAN-JAWOROWSKA, Z., 1979. Multituberculata. In Mesozoic Mammals. J.A.
Lillegraven, Z. Kielan-Jaworowska & W.A. Clemens, eds., Univ. Calif. Press, Berkeley. 99-149.

ESTES, J.A.,1980. Enhydra lutris. Mammalian Species 133: 1-8.

260 - PLEDGE

FLANNERY, T.F., ARCHER, M. & MAYNES, G., 1987. The phylogenetic relationships of living
phalangerids (Phalangeridae: Marsupialia) with a suggested new taxonomy. In Possums and Opossums,
eat in Evolution, M. Archer, ed., Surrey Beatty & Sons Pty Ltd. and R. Zool. Soc. N.S.W.: 477-

GREGORY, W.K., 1936. On the phylogenetic relationships of the Giant Panda (Ailuropoda) to other Arctoid
Camivora. Am. Mus. Novit. 878: 1-29.

HARRIS, J.M., 1987. Introduction. In Dinosaurs Past and Present, Vol. 1, S.J. Czerkas and E.C. Olson, eds.,
Nat. Hist. Mus. Los Angeles Co. and Univ. Washington Press, Seattle and London: 1-6.

HILDEBRAND, M., 1974. Analysis of Vertebrate Structure. John Wiley & Sons, New York.

KAY, R.F. & HYLANDER, W.L., 1978. The dental structure of Mammalian Folivores with special reference
to Primates and Phalangeroidea (Marsupialia). In Arboreal Folivores. G.G. Montgomerey, ed.,
Smithsonian Inst. Press, Washington, D.C.: 173-191.

MURRAY, P., 1984. Australia’s Prehistoric Animals. Methuen Australia, Sydney.

NORRIS, K.R., 1970. General Biology. In The Insects of Australia, Melb. Univ. Press, Melboume: 107-
140.

OWEN, R., 1863. On the Aye-aye (Chiromys, Cuvier; Chiromys madagascariensis, Desm.; Sciurus
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PLEDGE, N.S., 1982. Enigmatic Ektopodon: a case history of palaeontological interpretation. In The Fossil
Vertebrate Record of Australasia. P.V. Rich and E.M. Thompson, eds., Monash University, Clayton:
480-488.

PLEDGE, N.S., 1985. Ektopodon serratus Stirton, Tedford and Woodbume, 1967. The 'Strange-toothed’
Marsupial. In Kadimakara. Extinct Vertebrates of Australia. P.V. Rich & G.F. van Tets, eds., Pioneer
Design Studio, Lilydale: 214--218.

PLEDGE, N.S., 1986. A new species of Ektopodon from the Miocene of South Australia. Univ. Calif. Publs.
geol. Sci. 131: 43-67.

PLEDGE, N.S., 1987. A new species of Burramys Broom (Marsupialia : Burramyidae) from the Middle
Miocene of South Australia. In Possums and Opossums: Studies in Evolution. M. Archer, ed., Surrey
Beatty & Sons Pty Ltd. and R. Zool. Soc. N.S.W., Sydney: 725-728.

RICH, P.V., 1975. Antarctic dispersal routes, wandering continents, and the origin of Australia's non-
passeriform avifauna. Mem. natn. Mus. Vict. 36: 63-126.

RICH, T.H.V., 1986. Darcius duggani, a new Ektopodontid (Marsupialia: Phalangeroidea) from the early
Pliocene Hamilton Local Fauna, Victoria. Univ. Calif. Publs. geol. Sci. 131: 68-74.

ROMER, A.S., 1966. Vertebrate Paleontology. Univ. Chicago Press, Chicago.

SANSON, G.D., 1978. The evolution and significance of mastication in the Macropodidae. Aust. Mammal.
2: 23-28.

SANSON, G.D., 1980. The morphology and occlusion of the molariform cheek teeth in some Macropodidae
(Marsupialia : Macropodidae). Aust. J. Zool. 28: 341-365.

SANSON, G.D., 1982. Evolution of feeding adaptations in fossil and Recent macropodids. In The Fossil
Vertebrate Record of Australasia. P.V. Rich & E.M. Thompson, eds., Monash University, Clayton:
490-506.

SANSON, G.D., 1985. Functional dental anatomy and diet selection in dasyurids (Marsupialia : Dasyuridae).
Aust. Mammal. 8: 239-247.

SIMPSON, G.G., 1953. Life of the Past. An Introduction to Paleontology. Yale Univ. Press, New Haven.

STIRTON, R.W., TEDFORD, R.H. & MILLER, A.H., 1961. Cenozoic stratigraphy and vertebrate paleontology
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STIRTON, R.A., TEDFORD, R.H. & WOODBURNE, M.O., 1967. A new Tertiary formation and fauna from the
Tirari Desert, South Australia. Rec. S. Aust. Mus. 15: 427-462.

STRAHAN, R., 1983. Complete Book of Australian Mammals. Aust. Mus. and Angus & Robertson, Sydney.

TRUSWELL, E.M. & HARRIS, W.K., 1982. The Cainozoic palaeobotanical record in arid Australia: fossil
evidence for the origin of an arid-adapted flora. In Evolution of the Flora and Fauna of Australia. W.R
Barker and P.J.M. Greenslade, eds., Peacock Press and Aust. Syst. Bot. Soc., and ANZAAS, Frewville:
67-76.

WALKER, E.P., 1964. Mammals of the World. 3 vols. Johns Hopkins Press, Baltimore.

WOODBURNE, M.O., 1987. The Ektopodontidae, an unusual family of Neogene phalangeroid marsupials. In
Possums and Opossums: Studies in Evolution. M. Archer, ed., Surrey Beatty & Sons Pty Ltd. and R.
Zool. Soc. N.S.W., Sydney: 603-606.

WOODBURNE, M.O. & CLEMENS, W.A., eds., 1986. Revision of the Ektopodontidae (Mammalia,
Marsupialia, Phalangeroidea) of the Australian Neogene. Univ. Calif. Publs. geol. Sci. 131.

WOODBURNE, M.O. & CLEMENS, W.A., 1986a. A new genus of Ektopodontidae and additional comments
on Ektopodon serratus. Univ. Calif. Publs. geol. Sci. 131: 10-42.

WOODBURNE, M.O. & CLEMENS, W.A., 1986b. Phyletic analysis and conclusions. Univ. Calif. Publs.
geol. Sci. 131: 94-102.

NATURAL HISTORY OF EXTINCT ANIMALS - 261

WOODBURNE, M.O. & TEDFORD, R.H., 1975. The first Tertiary monotreme from Australia. Am. Mus.
Novit. 2588: 1-11.

PLATES

Plate 1. A, photographically reconstructed palate of Chunia illuminata using right maxilla fragment with half of

M?, and left M“, indicating a very broad face and short snout; B, restoration of Ektopodon skull in ventral view;
and C, skull and jaws in lateral view, based on Chunia sp.

Plate 2. Palate of Phalanger orientalis, a cuscus, indicating relatively short snout and wide face. The molar
teeth bear several crenulations on the weak lophs. (From Flannery, Archer & Maynes 1987).

Plate 3. Palate of Ailuropoda melanoleuca (Giant Panda) showing upper molars with multiple cusps arranged in
two transverse rows. (From Gregory 1936).

Plate 4. Daubentonia madagascariensis, the Aye-Aye. Note the short face, forward facing eyes, and elongate
third finger used in winkling beetle larvae from their burrows in tree limbs. Molar teeth are typically bunodont in

the primate fashion, but the incisors are rodent-like and ever-growing, used to gnaw holes in branches in search of
grubs. (From Owen 1863).

Plate 5. Palate of Enhydra lutris, Californian Sea-Otter, showing the short snout, wide face, and bunodont
molars enlarged for crushing shellfish. (From Estes 1980).

262 - PLEDGE PLATE 1

PLATE 1 (CONT.) NATURAL HISTORY OF EXTINCT ANIMALS - 263

~< “ \
Sreartinonieatte %
wis \

PLATE 2

264 - PLEDGE PLATE 3

NATURAL HISTORY OF EXTINCT ANIMALS - 265

PLATE 4

266 - PLEDGE PLATE 5

CHAPTER 10

THE TAPHONOMY OF LATE
QUATERNARY CAVE
LOCALITIES YIELDING
VERTEBRATE REMAINS IN
AUSTRALIA.

Robert F. Baird!

The Study of Bias in the Vertebrate Fossil

Record: an Introduction.............. 268

Caves as Favourable Environments for the

Accumulation and Preservation of
Vertebrate Bone.............. cece eeeee 275
How is Material Accumulated? ......... 275
Formation and Classification of Caves:
Why They Preserve Vertebrate

BONES... 0.chedetbewelfesvant sat eteecy dite. 276
Distribution of Caves in Australia.....277
Natural’ Trap iisi...c ced oee.eeeasraeosase 279
Habitual Cave-dwelling Species........ 281
Water-transported Material ............... 282
Material Accumulated by Animals..... 283

Material Accumulated by
Invertebrates ............ceceeee eens 283

Material Accumulated by
Vertlebrates..........cceeccsseeeeeeee
Non-carnivore Accumulators of
Vertebrate Bone ...............666+
Primary Carnivore/Scavenger
Accumulators of Vertebrate

BONG? ciliate ce Sass vensaed soaetees

Avian CarnivOres .............2eeeeeeee
DISCUSSIONS igo. hc eebagelerele dete asedetleeesivaee’
RefCrenceS.......cccccceceececeececceeceececeseeuees

Appendix I: Mammalian Carnivores
and SCAVENBETS .........e ce ceeee eee eee
Appendix II: Avian Accumulators...........
ACCIPIUTi dae ..... eee eee cence eee eeeeees
TP YtOm1d aes fo cgteede cca. aisotecenpangetenes tect

1 Department of Earth Sciences, Monash University, Clayton, Victoria 3168, Australia.

nn nn eee yaa EEE EEIEE EEE EES

268 - BAIRD

THE STUDY OF BIAS IN THE VERTEBRATE FOSSIL
RECORD: AND INTRODUCTION

To date no single reference has endeavored to summarize the taphonomic studies concerning
birds and cave deposits, although there are several which cover various subdisciplines within
the field (Brain 1981). Behrensmeyer & Kidwell (1985) have summarized the history of
workers in this discipline. It is my intention here to provide a survey of the state of
knowledge of taphonomy in relation to terrestrial avian communities and caves in Australia.
This synopsis provides an introduction to the concepts involved in the study of taphonomy and
outlines the range of environments capable of concentrating vertebrate bone (see Fig. 1). It
should set the stage for the discussion of caves as environments favourable to the concentration
and preservation of vertebrate bone. Included is a brief summary of the range of taphonomic
studies available, followed by a detailed consideration of caves and a discussion of the range of
Australian biota which are considered likely to accumulate vertebrate material in caves. The
introduction and most of the remaining chapter focuses on microvertebrate assemblages with
the occasional reference to macrovertebrate studies where pertinent. Although I concentrate on
skeletal material, it should be recognized that items such as eggs can also provide a wealth of
information regarding palaeoecology (Hayward MS). Most of the unpublished data come from
my own studies on avian assemblages from caves across southern Australia (Baird 1986), and
form the basis of many of the generalities.

Taphonomy has been succinctly defined by Behrensmeyer & Kidwell (1985) as, "...the study
of processes of preservation and how they affect information in the fossil record." The
understanding of the processes involved in fossilization allows interpretation of the usefulness
of certain assemblages in both palaeocenvironmental reconstructions and palaeocommunity
structure (see Patterson 1981). The prima facie assumption is that all accumulators of
vertebrate bone are biased, with some exercising a more extreme bias than others. One should
recognise that the biases are against fossilization and should never be considered to favour
preservation.

The study of taphonomy usually begins with the living community or group of
communities (biota), because some terrestrial communities will be in situations more
favourable for preservation than others. An appreciation of primary biases impinging on
fossilization of vertebrate material can be had by understanding these modern analogues. For
example, any community living near water may be preserved preferentially, because, if an
individual falls into the water, it is possible that it will be rapidly deposited within sediments
after death, therefore decreasing the likelihood that its elements are destroyed. In fact, areas near
water bodies are more favourable for attritional concentration of bone than surrounding bushland
(Behrensmeyer 1983). Those communities least likely to be preserved include forest and
woodland inhabitants, away from fluvial environments. This is associated with increased
availability to predator and scavenger destruction, insect destruction and chemical weathering
through exposure to the sun, soil acids and the general lack of rapid sedimentation (Bickart
1984). Animals in such habitats would find few places where preservation occurs, and,
therefore, members of certain communities will have a lower probability of being preserved and
fossilized.

Taphonomy also involves the determination of the proximity of the species and/or
assemblages to their original place of residence, relative to their final resting place. Prior to
their final burial, individuals may be subject to a number of different forces which can influence
the location of their corpses relative to their home ranges when living (e.g. deposition within
the area which they lived (autochthonous species: see Fig. 2) or transported away from the area
within which they originally lived, either when they are living or subsequent to their death

AUSTRALIAN CAVE TAPHONOMY - 269

(allochthonous species: Fig. 3). If a number of individuals are affected by the same taphonomic
influence, and are deposited together then, this assemblage may either remain in the area of
original deposition (i.e. autochthonous assemblage: see Fig. 4) or be shifted and re-deposited
elsewhere (i.e. allochthonous assemblage: see Fig. 5). Determining which of these situations
occurred can be difficult. Shotwell (1955, 1958) considered that the percentage composition
within an assemblage, and degree of sorting and fragmentation of bony elements could reflect
the proximity of the original habitat of each species represented. This hypothesis has been

discussed and largely discounted by a number of authors (e.g. Wolff 1973, Korth 1979),
particularly for small mammal assemblages.

Lost: life LIVING Lost: communities
processes COMMUNITIES infrequently sampled
pate ALLOCHTHONOUS AUTOCHTHONOUS Betsior
usually DEATH DEATH usually
preserved ASSEMBLAGE ASSEMBLAGE preserved
HARD
PRESERVABLE
PARTS
Lost: hard
parts not HARD Lost: destroyed
preserved PARTS by geological
PRESERVED processes

Lost: bones BONES

not yet NOT
available DESTROYED
BONES Lost: bones
AVAILABLE nat
iy collected
Lost: bones ess
discarded ess

Figure 1. Diagrammatic representation of the processes involved during the fossilization of vertebrate
bone. A, bias by accumulating force (incomplete sampling of the biocenose); B, preferential preservation
(physical and chemical action); C, reworking; D, bias by collection techniques.

270 - BAIRD

Also important to the study of taphonomy is knowing the possible causes of death. These
may include predation, disease, physical accident, poisoning, starvation, dehydration,
intraspecific conflict and natural death, amongst others (Clark et al. 1967).

In theory both of the two death assemblages (i.e. autochthonous and allochthonous) could
provide information about the palaeoecology of the community, that is if one could determine
the degree of allochthony. In practice, however, only the autochthonous assemblages may
provide this information, due to the attritional nature of such deposits (e.g. animals of the
floodplain; Voorhies 1969). A possible course of events yielding an autochthonous assemblage
might allow for a Genyornis (a large extinct ground bird), for arguments’ sake, to become
bogged and subsequently covered by sediments whilst crossing a floodplain. This animal, if
preserved, would, therefore, be in situ. An assemblage of these types of animals would be
more likely to reflect community structure, because all age classes may be represented, because
of the attritional nature of deposition, and all those species frequenting the area are also capable
of being represented. Therefore, attritional autochthonous assemblages are suitable for the
analysis of palaeocommunity structure (Damuth 1982).

AUTOCHTHONOUS SPECIES

HABITUAL CAVE DWELLING SPECIES
(e.g. swallows, falcons, owls)

GROUND DWELLING SPECIES

(e.g. emus, rails, quail, etc.) an accumulation of which would be an
assemblage of autochthonous species

4 Limestone Sediment @@ Bone

Figure 2. Those species (and their habits) which are most likely to make up the autochthonous species
component in cave deposits.

Individuals included in the other group (i.e. allochthonous assemblages), by definition, have
been transported after initial burial to their final place of burial. The same force which has
carried one individual, or bone of an individual, would presumably carry others. The result

AUSTRALIAN CAVE TAPHONOMY - 271

would be an accumulation of elements from different animals. These animals would not
necessarily be from the same habitat, for the collecting agency would be indiscriminant about
the habitat. Sorting would occur based on the elements’ sizes and shapes, which depend on
hydrodynamic parameters (see Voorhies 1969, Table 5 in Dodson 1973) or may be biased by
the accumulating animal, be they invertebrate or vertebrate, primary-predators or scavengers

(Dodson 1973, Wolff 1973, Mellett 1974, Horton 1978, Fisher 1981). A scenario for such a
case might be that the Genyornis, mentioned previously, having died in the river, would have
been carried downstream, either slowly decaying and dropping bits to the bottom or deposited in
toto at one locality (Dodson 1973). If the river had a large catchment, then its course might
have crossed a number of different habitats, and, therefore, could have sampled animals from a
number of different communities. The bones of the individuals, assuming similar size and
shape, would be deposited together (hydrodynamic sorting), therefore, making up an assemblage
of animals that would not necessarily, in life, have been interacting.

ALLOCHTHONOUS SPECIES

Species brought in by taphonomic
accumulating agents
(e.g. water and animals)

an accumulation of which would be an
assemblage of allochthonous species

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Figure 3. The process by which allochthonous species are incorporated into a cave deposit.

Lost to both the autochthonous and allochthonous assemblages would be those soft parts
that decay during the natural course of decomposition, therefore, leaving only hard parts which
are not as easily broken down (Voorhies 1969, Wolff 1973). Bias is not only due to inanimate
accumulating forces, for animate forces may also bias both the size and the individual
abundances of elements. An example of the latter is what is termed the "schlepp effect." This
concept was originally used to explain the abundance of certain skeletal elements and the
absence of others (Perkins & Daly 1968). It suggests that animals which were too large to be

272 - BAIRD

carried were dismembered at the place of capture or death, and during this process certain body
parts were favoured over others. These were subsequently carried back to the locality where soft
parts were eaten and hard parts were eventually discarded. Although this idea was put forward to
explain the bias in deposits of human origin, it may also be relevant to the behaviour of other
animals that bring food to some habitual place away from the place of capture.

AUTOCHTHONOUS DEPOSIT

ES aes OER a) Oe A [ JT J J)
“JT JI J J [J ] [ J JT JT JT 4
a CAE EE ES Ce Ce es a =) 2 ee
2 ee ee [ J] J-T J JT 7)
i ee eS Gee [TJ J JT J
i i a [ J Jy JT JT JT
I J J] 4 ee ae OS
[J / 2 ae eS eee [ J Jy J J] J J
[JT TT TT tT JT JT TT JT JT JT
[ J) a SE GS GS Ce Soe Pe oe a
a ay, \L IJ JT JT Jy Jy TT JT JY Ty J
[ w7; LT I Tf J] J] yT Jy Jy JT JT J 4
[ T] c oe tN ~~. T JT T J J JT J [ J J J
[ ] =. <x] _J.-}. ~[._}-_[ Jf J} 4
[1 —=L_] I tI tr Td
— “J JT JT TT 4
[J eS a
[J a a
a Ss a
= 2
a \ I JT 4)
[J i
[J i yf
LZ as oe
ke =]
L =
ry — 7)
fo Limestone Seo Ae So a
vt TT
LC

2] Sediment an insitu accumulation of
@m@ Bone vertebrate material

Figure 4. Any deposit of vertebrate material which has remained in situ since the time of deposition is an

autochthonous deposit.

Of those hard parts capable of preservation a certain number will be eroded both physically
and chemically. Many would not survive beyond the carcass stage. Predators, scavengers,
acidic environments, weathering, all will destroy the potentially preservable parts before they
can be preserved or enter an environment favourable for preservation (Haynes 1980). Preferential
preservation of certain elements, presumably due to their superior structural competence, has
been noted for both bird elements (Napawongse 1981) and mammal elements (Voorhies 1969)
within fluvial environments. The avian elements frequently represented in these types of
environments include humeri, femora, tibiotarsi and tarsometatarsi (Fig. 6). Preferential
preservation has also been noted even within a cave deposit proposed to be pitfall in origin.
Murray & Goede (1977) found that the frequencies of large mammal elements varied, with
tibiae, femora, pelvic elements and mandibles most abundant and scapulae, neurocrania, fibulae
and radii exhibiting very low frequencies. The authors suggested that differential preservation
occurred, because the lighter, thinner elements were less likely to be preserved due to their large
surface area and small bulk, therefore, decomposing very rapidly (Murray & Goede 1977).

Assuming an element is not destroyed or removed during deposition, fossilization may
occur. Fossilization includes “all processes involving the burial of a plant or animal in
sediment and the eventual preservation of all, part, or a trace of it" (Bates & Jackson 1980).
Preservation of hard parts usually occurs by the replacement of calcite and apatite through

AUSTRALIAN CAVE TAPHONOMY - 273

carbonization, silicification, calcification, etc., although the intake of elements into the calcite
eae matrix is not uniform, even within a single fragment of bone (Henderson et al.

Once preserved, the bone may still be destroyed (through reworking) or lost to the fossil
record through processes such as sorting, abrasion, weathering and diagenesis. If the specimen
is deposited in a non-destructive environment, it can still be reworked and liable to destruction
through the same processes at a later time.

ALLOCHTHONOUS DEPOSIT

AS
al

Figure 5. Any deposit of vertebrate material which has been reworked since the time of deposition is an
allochthonous deposit.

The time encompassed in any one accumulation of fossils is frequently large. This results
from the amount of reworking and redeposition that can take place along with the amount of
time attritional deposits take to form. Behrensmeyer (1982) has studied the resolution and
accuracy in dating changes in the fossil record and has determined that hypothetically both fossil
soils and attritional channel deposits are capable of yielding information of a 100 to 10,000
year scale. In reality, however, the lowest limit is approximately 1000 years (Fig. 7).

At the collection stage, there will be only a certain number of fossils available at any one
time. There are others which may be too deep in the earth or not yet weathered out that may be
available at a later date. This class of fossils will always be unattainable. Another bias may be
introduced by the collector for frequently he will be most interested in a specific group, such as
mammals, birds, or fish, and other preserved elements of unwanted taxa may be passed by due
to lack of space or interest. These fossils may be lost by the time another person has an
opportunity to collect them.

274 - BAIRD

The sample may also be biased through the particular collecting technique used. An
example of this is sieving boxes with one quarter inch (0.6 cm) mesh, a standard method of
searching fine sediment for bone (Hibbard 1949, McKenna 1962). This size mesh allows many
elements, small enough to fit through, to be missed (Olson 1982). If one determines the size
of the mesh used by the size of the elements excavated then the only elements collected will be
those of the size looked or collected for. In many cases the only certain way of retaining all
vertebrate bone is to sieve the matrix in the field with fly screen (1.5 mm or less) and then

Carpometacarpus

—j SAN

aw,

Sternum

Tibiotarsus

|

Tarsometatarsus

\

f)
f
ff

LS
LY — Fe
mea Se a C= i ay
“Ly
SSS

Figure 6. Bird skeleton illustrating those elements most resistant to destruction (in black) in simulated
fluviatile environments (tumbler experiments of Napawongse 1981: from Rich & van Tets 1982).

AUSTRALIAN CAVE TAPHONOMY - 275

transport the concentrate back to the laboratory for wet screening and sorting under a dissecting
microscope (Hope 1973, Olson 1982). Often this is not possible, and in early times elements
were, in fact, sieved from their surrounding sediment through the hands of the excavator.
Because of the constraints of the excavation, a distinctive distribution of bone related to the size
and length of the bones available is produced with this method (see Rich & Baird 1986).

To fully comprehend the prehistory of a deposit, one must have a full appreciation of the
range of different forces that accumulates fossil material. In discussing these I will concentrate
on vertebrate material in the Australian context and specifically on birds, but I will refer to
foreign studies where pertinent. In general, there has been a host of papers written on the
taphonomy of mammal rich deposits, but little has been written on birds. To provide a starting
point, I have used the taphonomic classification system set up by Bishop (1980: Table 1).
Although the taphonomic history of many deposits is rarely as clearcut as his system indicates,
it is still important to understand the total range of possible accumulating forces.

Using this classification, it is easy to place the cave deposits I will discuss. Given that
caves, especially those formed in limestone, makeup 68% of all Quaternary deposits studied to
date in Australia (Baird in press), I have concentrated on the taphonomy of these deposits.

TAPHONOMIC MODEL FOR
BONE INPUT TO FLUVIAL DEPOSITS

BONE BURIAL
CARCASS ORIGINALLY IN FLOODPLAIN
IN CHANNEL -
OVERLAND BONE }
TRANSPORT va .
~

DESTRUCTION !
raga INSOIL | 4

1ae
la
EROSION

Figure 7. "Model for taphonomic input of bones to deposits of fluvial systems with channels which
periodically or continuously erode portions of the contemporaneous floodplain sediments. Letters indicate
three distinct taphonomic pathways’ through which bones are incorporated into the fluvial deposits: A,
autochthonous burial on the floodplain (followed in some cases by A':alteration by soil processes and A”,
erosion from channel banks into active channel; B, overland transport into the channel, C, origination in the
channel as part of a carcass." (From Behrensmeyer 1982).

CAVES AS FAVOURABLE ENVIRONMENTS FOR THE
ACCUMULATION AND PRESERVATION OF VERTEBRATE
BONE

HOW IS THE MATERIAL ACCUMULATED?

Vertebrate material accumulates in caves in many of the same ways as in other areas
suitable for preservation of vertebrate bone, via both autochthonous (e.g. natural trap) and
allochthonous mechanisms (e.g. water transported material, invertebrate and vertebrate
accumulated material). For the most part, it is the position of the cave, the process of the
cave's formation or its microclimate that encourages the material to be accumulated. Therefore,
it is important to understand all these aspects of modern day caves.

276 - BAIRD

Table 1: Classification of fossiliferous palaeoenvironments in continental regimes (adapted from
Bishop 1980).

A. Tectonic situations inducing rapid and D. Inland basins and traps, temporary burial
frequently cyclic sedimentation sites
1. Molasse and fan deposits in mountain girt 1. Lacustrine, fluvio-lacustrine, swamp, and
basins and on the flanks of growing spring-eye margins
mountain chains a. deltas
2. Rift valley slumps and grabens b. channel lag ("dropout" situations)
B. Volcanic fields yielding tephra of particular c. lakes, peat bogs, and tar pits
petrological composition and lava flows d. springs (petrification)
1. Primary carbonate fallout onto land 2. Caves
surfaces a. karstic settings : favorable chemical
2. Secondary deposits derived from volcanic burial environments
sources b. lava caves
a. calcareous tuffs/steppe limestones c. sandstone caves
b. trachytic pumices d. granite caves
Cc. Continental margins 3. Impedance and sedimentation on
1. Distal ends of rivers, especially those planation surfaces yielding
flowing from semi-arid hinterlands concentrations of surface sweepings
2. Shallow water estuarine, deltaic, and 4. Deserts
lagoonal deposits, especially those a. hot desert dune fields
associated with marine transgression b. fringing deserts; coastal dunes
a. bay bars c. cold deserts, permafrost
b. bone beds

c. fissure fills
d. sea caves

FORMATION AND CLASSIFICATION OF LIMESTONE CAVES: WHY

THEY PRESERVE VERTEBRATE BONE

Although the term cave indicates to most people some sort of subterranean cavern, the
definition of what a cave is varies. I regard the following as appropriate: "cave [speleo] (a) A
natural underground open space, generally with a connection to the surface and large enough for
a person to enter" (Bates & Jackson 1980).

The formation of caves and the matrix within which caves form varies immensely and
seems only to require that the surrounding rock be structurally competent enough to hold itself
up once the cavern is formed within. The formation of caves varies with matrix-type, and the
agents of formation include "...lava flow, wave attack, landslides, and movement and melting of
glaciers..." (Jennings 1971). But as Jennings (1971) goes on to say "... caves formed by karst
processes are the most numerous, largest and most complex" and, in fact, all the caves
discussed in this section were formed through karst processes. Therefore, emphasis in this
chapter has been placed on the understanding of these processes.

The process of cave formation within limestone depends upon a varied number of factors.
These have been listed succinctly in Sweeting (1973:133):

"{1] The form of the primary capillary. [2] The petrological and chemical
character of the limestones. [3] The structure of the limestones, such as the dip,
joints and faults, etc. [4] The type and amount of waterflow through the
passages, i.e. whether the flow is forced or free (phreatic or vadose). [5] The

AUSTRALIAN CAVE TAPHONOMY - 277

regional physiography of the area. [6] The influence of preceding developments
in the caves, i.e. the history of the cave. [7] The climate and past climatic
variations. [8] The influence of cave deposits" [information within the brackets
added] (Fig. 8).

The genesis of all karst features seems to depend upon the percolation of acidic ground
waters through the limestone. The acidity necessary for the initiation of solution is usually
produced by meteoric waters percolating through soils, resulting in higher carbon dioxide
content in these waters. Subsequent solution, although enhanced by the acidity of the ground
waters, is largely due to differential solubilities of carbon dioxide in the water (Sweeting 1973:
see Fig. 8).

The above covers primary cave formation, where the initial cave formation and ensueing

passages are largely determined by the flow of water through, or, the water table in, the rock.
More important to caves acting as natural traps is secondary cave formation or enlargement.
The driving force behind this process is spalling from the roofs of caves, particularly block
breakdown, rather than other forms of breakdown (i.e. slab, plate and chip) (see Davies 1952).
This allows those caverns which were once subterranean to migrate topographically upwards
and eventually breach the surface. The entrance of these types of caverns is typically apical and
allows for the capture and entrapment of animals which are incapable of either scaling the walls
of the cave or flying out. Cave entrances can also be indicators of the history of formation, in
that those which are small and smooth are considered to be entirely erosional and those which
are large and irregular result from rockfall (Jennings 1971). Solution of caves, and their
surrounding limestone, particularly in areas of high rainfall, will generally continue until the
caves have lost their surrounding matrix and are fully exposed to the surface. Then they no
longer form a 'cave' and can simply be regarded as the remnants of one (e.g. Hope 1982).
Cave formation in limestone is often dependent upon the flow of water through the rock. The
different water flow types, therefore, present the most satisfactory way of classifying these
caves (Sweeting 1973:158). The two most common forms of caves are phreatic and vadose.
As defined by Sweeting (1973), phreatic caves are those formed within the zone of limestone
which lies at or just beneath the level of permanently saturated rock, and vadose caves are those
caves formed by water circulating under gravity above the level of permanently saturated rock.
The bulk of cave genesis typically occurs at or above the saturation level in the karst
limestone. The two most common surface features pertaining to karst in Australia are collapse
and solution dolines. Dolines are simply closed depressions. They form either through the
failure of the limestone, usually the ceiling of an underground cavern (i.e. collapse dolines) or
through "...the pronounced surface solution of the karst bedrock around some favourable point
such as a joint intersection" (Jennings 1971), which subsequently enlarges, typically forming a
conical shape (i.e. solution doline). Most other caves are exposed to the surface through
riverine erosion of karstic limestone exposing the caverns within, especially at meanders.

Preservation of bone is encouraged by the decreased amount of weathering within caves and
their alkaline environment (Frank 1971, BMNH 1974).

DISTRIBUTION OF CAVES IN AUSTRALIA

The distribution of caves around Australia parallel that of the rock substrate which favours
their formation. Volcanic caves are restricted to the east coast of Australia, particularly western
Victoria and northeastern Queensland (see Fig. 9). Sandstone caves and rockshelters are present
in western Victoria, northern Western Australia, the Northern Territory and in select localities
around the coast. These are too scattered for inclusion in Fig. 9. Caves in limestone occur
right across Australia and are particularly abundant near the coast, where a higher than average
rainfall provides the water necessary for solution of the limestone (Fig. 9).

278 - BAIRD

E

eee (pad =
Eee

4 ;

Figure 8. Hypothetical sequence of events leading to the development of a cave system of vadose origin.

: ypo
(From Waltham 1974).

AUSTRALIAN CAVE TAPHONOMY - 279

Stalagmite etc.

Clay sediment

NATURAL TRAPS

In Australia, the idea of natural traps typically conjures up images of apical entrance dolines
where animals fall into a cave from which escape is impossible. But natural trap mechanisms
include any natural situation that repeatedly causes animals to be trapped and to subsequently
die at the point of capture. These include subaqucous deposits, unctious sediments and pitfall
and deathtrap caves, to name a few. For example, subaqueous deposits could be formed by the
following scenario: flocking birds could come down to drink, soak their feathers and
subsequently drown (e.g. Baird 1985). An assemblage produced in such a way should exhibit

280 - BAIRD

several features, including: a general lack of damage (except through solution of the bone
calcium), a preponderance of flocking and terrestrial species and proportional representation of
body parts, both cranial and post-cranial (unless sorting in a water column or down a cone of
talus occurs). What will be noticeable by their absence are flying animals, excepting those
which naturally occur in caves. Unctious sediment along lake and wetland margins can cause
both mammals and birds to become trapped (e.g. Gillespie et al. 1978). This type of natural
trap should not be ruled out for caves, for many act as temporary or permanent catchments of
water, and similar conditions could be found within the cave as are found around wetlands
outside. Characteristics of these types of assemblages depend upon whether they are found in
situ or not. Those in situ reflect the positions of the animals at the time of death; most
elements should be present in the relative positions as in life, little to no damage should occur

to any of the elements (except for weathering for those elements exposed to the surface) and
mainly terrestrial forms should be represented.

Y limestone caves e g

volcanic caves

Figure 9. Distribution of the major regions exhibiting cave development in Australia.

AUSTRALIAN CAVE TAPHONOMY - 281

By far the most common form of natural trap involving caves is the gravity induced pitfall
or deathfall situation. As discussed in the section on cave formation, many of the doline caves
act as natural traps during some part of their evolution. These would naturally form pitfalls or
deathtraps and would sample the passing traffic of animals. Once the collection of animals had
begun, scavengers would be preferentially attracted to the cave, thereby introducing a bias for
these within the assemblage.

An assemblage accumulated in such a way is considered attritional, but in some rare cases it
may be due to some catastrophic event. One way of distinguishing catastrophic from attritional
mortality can be through the study of age classes, which, although of little use in birds, can be
helpful with mammals (Van Valen 1963, 1964; Voorhies 1969; Klein & Cruz-Uribe 1984).
The catastrophic assemblage should exhibit the proportions of juvenile, adults and senile
individuals as they occur in life, whereas the attritional assemblage would exhibit greater
proportions of those age classes not capable of surviving the rigours of life (Z.e. juveniles and
senile individuals), Characteristic damage to individuals within a deathtrap/pitfall situation can
be incurred through falling. Therefore, broken limbs, ribs and skulls can be present in the case
of entrance to the cave via a fall. These breaks will show little, if any, healing and are
characteristic of breaks in fresh bone. Abundances of elements should be largely equal or
slightly subequal unless some other mechanism for redistributing the elements is present (e.g.
talus cone slumping). Animals represented should be largely terrestrial forms or, in the case of
birds, those volant species which are poor or weak fliers, incapable of vertical flight. For birds,
weight distribution would be inconclusive, for it is not the weight of the animal that causes it
to be trapped but its inability to escape the trap.

The assemblages provided by these accumulating environments are regarded as
autochthonous in that the animals accumulated live in the area of deposition.

HABITUAL CAVE-DWELLING SPECIES

Habitual cave-dwelling species are those members of the animal community represented in
deposits in caves due to their preference for using caves as nesting sites, roosts, foraging areas
(Hamilton-Smith 1965) and heat escape (Attwood 1982). These are regarded as autochthonous
species to the assemblage, for they are living in the area of deposition.

Characteristics of elements that form this portion of any deposit include: juveniles present
(and not on the upper end of the body weight range for the assemblageas with some predator
accumulated assemblages), associated skeletons present, most elements undamaged and in
subequal abundances and species of known speleophiles.

The list of bird families and species recorded as using caves in Australia include: Falconidae,
Falco peregrinus (Peregrine Falcon: Richards 1971), F. cenchroides (Nankeen Kestrel: LeSouef
1928, McGilp 1932, Collins 1934, P.R. Penney in litt.), F. berigora (Brown Falcon: Richards
1971); Aegothelidae, Aegotheles cristatus (Owlet nightjar: Richards 1971, P.R. Penney in
litt.); Platycercidae, Platycercus elegans (Crimson Rosella: P.R. Penney in litt.); Tytonidae,
Tyto alba (Barn Owl: Parker 1977), T. novaehollandiae (Masked Owl: McGilp 1932, Collins
1934, McKean 1963, Parker 1977); Strigidae, Ninox novaeseelandiae (Boobook Owl: Dwyer
1966); Apodidae, Aerodramus spodiopygius (Grey Swiftlet: Pecotich 1974); Alcedinidae,
Dacelo novaeguineae (Laughing Kookaburra: P.R. Penney in Jitt.); Hirundinidae, Hirundo
neoxena (Welcome Swallow: Richards 1971, P.R. Penney in litt.), Cecropsis ariel (Fairy
Martin: Richards 1971), C. nigricans (Tree Martin: Wall 1984); Acanthizidae, Origma solitaria
(Rock warbler: Gannon 1966); Climacteridae (tree creepers: Hamilton-Smith 1965); Corvidae
(crows: Richards 1971).

282 - BAIRD

Table 2: Characteristic transport susceptibility groups of avian skeletal elements from
experiments completed with the use of a flume, including Group I = immediately removed,
transported by saltation or floatation, Group II = removed gradually, transported by traction and
Group III = lag deposit (from Napawongse 1981).

Group I Intermediate Group II Intermediate Group III

Ribs Phalanges Furcula Sterna Skulls
Vertebrae Pelves
Mandibles
Coracoids
Scapuli
Humeri
Ulnae
Radii
Carpometacarpi
Femora
Tibiotarsi
Tarsometatarsi

WATER-TRANSPORTED MATERIAL

Water transported material may occur in any type of cave but is especially pertinent to
limestone caves, where the method of formation is largely caused by the dissolution of
limestone by water flowing through joints in the rock. Vertebrate material may be accumulated
once the joints are sufficiently enlarged and sediment carried by the water begins to become
deposited, within the then primordial cave. The variety of environments of deposition that
occur in fluvial systems outside caves can largely be reproduced within caves, but usually on a
smaller scale. The two major factors determining the type of deposit include whether the source
of the water is permanent (e.g. a formation stream) or ephemeral (e.g. a swallow hole at the
center of a local basin). A deposit from a permanent stream will exhibit characteristics of other
fluvial deposits (see Voorhies 1969, Wolff 1973, Napawongse 1981, Behrensmeyer 1982). On
the other hand, a deposit from a swallow hole would not have as much influence from the
moving water, and, therefore, may not have a sufficient number of diagnostic characters to
interpret with confidence.

The bone accumulated through these forces may be characterized by hydrodynamic sorting
(Table 2) and/or damage through abrasion (Napawongse 1981). Because water-transported
material, when it is associated usually represents attritional assemblages, the weight
distribution of the individual species would exhibit a wide range, with all forms having an equal
probability of representation depending on abundance within the community although
hydrodynamic sorting can skew this in actual deposits. According to Korth (1979), the
following characteristics are considered diagnostic for a fluvial accumulation of microvertebrate
bone:

"1) the sediment enclosing the fossils should generally show primary
sedimentary structures of alluvial origin, 2) the modal size of sand grains in the
deposit should be predictable from the size and hydraulic characteristics of the
skeletal elements present, 3) the bones should show varying degrees of stream

AUSTRALIAN CAVE TAPHONOMY - 283

abrasion, and 4) the percentage representation of bones in the entire fauna
should be low".

Although fluvially accumulated species could be autochthonous, I regard them as
allochthonous, because these animals have been transported to their place of deposition and did
not necessarily live there.

It should be kept in mind that fluvial activity can redistribute and sort previously deposited
material of any of the other accumulating agents. Archer (1974a) investigated this possibility
with respect to the redistribution and mixing of charcoal and bone of different ages, he found
that "experiments using a controlled laboratory situation ... demonstrate that small skeletal
elements differ in their ability to be transported by water, and charcoal is more mobile than all
bone when transported by water." A good example of the mixing of non-contemporancous
material in a cave situation is discussed in Wells et al. (1984). In this example, the cave acted
as a natural trap (i.e. pitfall, for the macrovertebrate assemblage, based on the age classes of the
fauna), but the bony material was later distributed down the cone of talus by slumping and
sheet wash.

MATERIAL ACCUMULATED BY ANIMALS

All species included in animal accumulated assemblages should be regarded as
allochthonous, because all have been transported from the place of capture and are not
necessarily indigenous to the area immediately around the site of deposition.

The biases introduced by animal accumulators of vertebrate bone are many and highly
complex. They depend on the natural histories of both the accumulating animal and its prey.
Much caution should be used in the interpretation of these types of assemblages of bone (see
below).

Material Accumulated by Invertebrates

Although not particularly pertinent to caves, the accumulation of vertebrate material by
animals other than vertebrates is a well known phenomenon. For example, some harvester ants
collect vertebrate teeth and bone, which are used as cover for the ant colony's mound to stop
wind and water erosion (Shipman & Walker 1980). Any grain of a particular size is used,
therefore, some teeth and bone are of the right size to be collected. Fossil accumulating habits
are associated with the excavation of the colony as well, and, similarly, the fossils are found
adorning the top of the mound (Clark e¢ al. 1967). The habit of adorning the mound surface
with debris is known for harvesting ants of the Australian arid zone (Morton 1982) and may be
useful in locating fossil localities.

Material Accumulated by Vertebrates

Bias from accumulators of vertebrate bone affects the amount and condition of material
capable of being preserved and the types of prey captured or scavenged. Because vertebrate bone
accumulators are biased, several things should be evident in studying any vertebrate accumulated
assemblage. Of particular importance is that the wear or damage to the elements might be
sufficiently diagnostic to determine the primary accumulator (Dodson & Wexlar 1979). The
effects diagnostic of each species of accumulator, as reflected on prey bone, are basically three-
fold. First is the action brought to bear on the bone whilst a prey species is being captured,
manipulated for swallowing, or masticated. This includes the predigestion/masticatory stage
where the prey may be dismembered, chewed up, or otherwise manipulated to aid swallowing
and digestion. For mammals this is a particularly important phase because, of the material

284 - BAIRD

swallowed, much will be well chewed. Some parts may be left behind uneaten, but the bulk
may be masticated to varying degrees. The second stage involves digestion, where the amount
of degradation to the bone depends on the pH of the stomach acid. The third stage is the
ejection of the bone from the body, post-digestion.

Herbivores, carnivores and scavengers are known to chew on bone. Of the herbivores,
camels Johnson & Haynes 1985), deer (Sutcliffe 1973), and kangaroos (Abbie 1952) chew
bones regularly, for either calcium or phosphate (Sutcliffe 1973). Similar behaviour is recorded
for rodents, the evidence of which is common in many cave deposits (Archer et al. 1980, Brain
1980). It is largely carnivore and scavenger activity, however, that characterizes bone
assemblages in caves, and, therefore, I will concentrate on their signatures .

Important to the understanding of the differences between the remains left by mammal and
bird accumulators is the process by which these animals capture, ingest and eject remains of
their prey. Depending on the size of the prey captured, mammalian predators or scavengers
either ingest the whole prey item or dismember the prey into ingestable pieces. Subsequently
this food is masticated and then swallowed. The food ingested usually includes bone. In fact,
one Australian mammal (i.e. Sarcophilus harrisii, the Tasmanian Devil) has a dentition adapted
for bone crushing (Werdelin 1986). Some damage due to teeth of carnivores can be
distinguished from man induced damage, via stone tools, by the microscopic features (Shipman
1981), Also, carnivore tooth-marks typically follow the contour of the bone instead of being
deeper on convexities and shallower on concavities, as with stone tools (Binford 1981:
Appendix I, Homo sapiens, for more information).

Fragmentation of the bone increases the surface area prone to chemical attack, and,
therefore, bone ingested by mammalian predators is typified by extensive corrosion from
stomach acid. Two characteristics of bone ingested by mammalian carnivores have been
recognized, including "[1] surface rounding of the bone, where the whole surface is smoothed
over and broken edges and articular surfaces are rounded; and [2] corrosion, where the surface
rial is removed and the underlying bone is partly dissolved away" (Andrews & Nesbitt-Evans
1983).

For mammals, digestion occurs in the stomach, with a pH that is variable, but generally,
low. Upon passage through the digestive system, the undigested material is defecated, and,
therefore, any bone surviving the digestive process occurs in the faeces. Occasionally
mammals may regurgitate bones, but apparently this occurs only when sharp points form
during the process of dissolution (Behrensmeyer et al. in press).

The method of ejecting bone after digestion of the muscle and soft tissue differs between
birds and mammals. Depending on the size of the prey item, avian predators or scavengers (at
the point of capture or discovery) cither ingest the whole prey (if small) or dismember it into
ingestable pieces (if large). In the case of prey items consisting of birds, the prey may be first
plucked of its primaries, secondaries and rectrices. Prior to swallowing, the mandibles close on
the pieces ingested but usually little mastication occurs. Therefore, damage to the bone of
prey species is largely incurred through the process of dismembering. After the prey is
swallowed, initial digestion occurs in the proventriculus, The pH in the stomach of a bird is
variable but is generally high. There is a sphincter between the proventriculus and the
ventriculus, which in some species only allows the transfer of liquids due to its small size and
high placement on the wall of the proventriculus. Upon the completion of digestion, the hair,
feathers and bone are formed into a bolus, and then regurgitated (Reed & Reed 1928). No bone
is found in faecal material (Farner 1960) but instead occurs in the regurgitated pellets.

Because of their different modes of ingesting and digesting their prey, mammalian- and
avian-accumulated deposits differ in the degree of fragmentation and the amount of corrosion
exhibited on the bone, which is usually greatest in mammals. The timing of fragmentation and

AUSTRALIAN CAVE TAPHONOMY - 285

sh aan corrosion exhibited by bone is also important to the interpretation of its taphonomic
istory.

Interpretation of the approximate time at which the damage was done to the bone, relative to
the death of the individual, is largely based upon certain attributes, which together suggest that
the bone was either fresh, dry or fossilised at the time the fracture or break occurred (see Table
3, from Morlan 1984; Kuhne referred in Mellett 1974). Confusing the issue is damage to
elements incurred by a reworking of material already deposited, producing secondary or perhaps
tertiary signatures.

Table 3: "Attributes of limb bone fractures to bone condition at the time of fracture."
Morlan 1984, modified after Bonnichsen 1979, Morlan 1980 and Stanford et al. 1981).

(From

Attribute Fresh bone Dry bone Fossilised bone

Negative impact scars Present or absent Present or absent Absent

(loading points)

Texture of fracture
surface

Angle of fracture

Smooth Smooth or rough Rough ("pebbly")

Acute, obtuse Acute, obtuse Right

with outer surface or right or right
of bone
Termination of At or prior to May cross-cut May cross-cut
fracture at epiphyses epiphyses epiphyses
epiphyses
Colour of fracture Same as Little or no contrast Often contrasts sharply

outer surface
Straight, diagonal,

curved, spiral;
generally smooth

Outline form of
fracture

with outer surface
All outlines seen in
fresh and fossil bones,
often perturbed by
curved, rarely spiral

with outer surface
Usually straight,
transverse, Or

longitudinal; can be

curved, rarely spiral

often perturbed by
split lines

split lines
———

Hypothesized secondary signatures are pertinent to any deposit that was reworked (e.g. by a
scavenger like Sarcophilus (Dortch & Merrilees 1973)), where characteristic damage of the
secondary accumulator would be superimposed on the damage done by the primary accumulator.
It might be possible to determine whether the deposit exhibits overprinting by referring to the
other characteristics of the assemblage.

Biases can be inferred through knowledge of the natural history of the predator species . No
predator or scavenger is wholly indiscriminate when it comes to the capture of its prey or in the
pattern by which it discovers its food. Such biases include the size of prey captured, particular
species captured (due to their lifestyle or captureability: see Hope 1976), and the habitat covered
by the predator (or vegetation structure type within that habitat; e.g. Tyto tenebricosa, Sooty
Owl, and arboreal mammals). Of the types and size of prey being eaten, primary carnivores in
Africa (and presumably in Australia) seem to prefer prey within a restricted weight bracket
(Vrba 1976, Brain 1980). Scavengers, on the other hand, seem to be more opportunistic and
sample a wider range of weights. Avian predators may also select prey from a certain weight
range not only because they are primary predators but because of limitations imposed by wing
loading. Possible elements in the natural histories of the prey species that would increase the
likelihood of capture include gregariousness, irruptive nature and nocturnality or diurnality.

286 - BAIRD

The range of prey species represented is also restricted either by the biases of the predator
species (palatability), individual preference, the type of habitat covered during hunting and the
pattern of hunting (e.g. in open vs. closed vegetation formations).

The final information relevant to determining the signatures of each predator concerns the
lifestyle exhibited by the living predators, which may, for example, suggest that a certain
distribution of bone is characteristic of a particular predator (Hill & Behrensmeyer 1984).

Signatures of the accumulating agents should reflect all of the above criteria, including; i)
damage to the elements, ii) type and size of prey, iii) lifestyle of the prey, and ultimately, iv)
the lifestyle of the predator. The presence of the accumulator in the deposit and coprolites
attributed to the accumulator may be evidence for the identity of the accumulator, although the
presence of a suitable predator is not a priori evidence for its accumulation of the deposit.
Secondary signatures are particularly important, because secondary damage may mask the
primary signature (i.e. that imposed by the primary accumulator). In some cases this can be
clearly seen, as with rodent damage to bone (Archer et al. 1980), but in other cases it may be
impossible to separate primary and secondary damage (see Dortch & Merrilees 1973). Roots
and rootlets can also impose secondary signatures, and according to Morlan (1984) "...mark the
bone with a distinctive network...the opposite edges of a rootlet mark are often scalloped, and
the cross section is usually U-shaped". These always follow the contour of the bone surface.

Although many animals may mark bones with a characteristic signature, many of these
animals only do so opportunistically (e.g. Thylacoleo, Marsupial Lion), and few actually
accumulate the bone in places favourable to fossilization.

Non-Carnivore Accumulators of Vertebrate Bone

Material accumulated by vertebrates is not restricted to carnivores. Hystrix (Old World
Porcupine: Brain 1980) and Atherurus (New World Porcupine: Dixon 1984), or Neotoma
(Packrat: Harris 1985) all collect and chew both wood and bone. This habit may serve to
supplement phosphorus in their diet and/or to curb the development of their ever-growing
incisors. There are no known non-carnivore accumulators of vertebrate material extant in
Australia today, although their former presence must always remain a possibility.

Primary Carnivore/Scavenger Accumulators of Vertebrate Bone

Included in this section is data for all of the carnivorous accumulators of vertebrate bone
occurring in Australia within the late Quaternary. For some of the fossil species, however, the
taphonomy is either unknown or hypothetical (see Appendix I). A number of possible
accumulators of vertebrate bone were suggested by Balme et al. (1978) including man, dingos,
thylacines, Tasmanian devils, native cats, smaller carnivorous marsupials, water rats, hawks,
falcons, owls, herons, cormorants, crows, kingfishers, butcherbirds, currawongs, magpies,
[bowerbirds], monitor lizards, large scincid lizards, fresh-water turtles, pythons, large venomous
snakes and large frogs. Balme et al. (1978) commented, "some of these possibilities seem very
remote in view of the habits of the species concerned" and a number of members can be deleted
from this list based on their natural histories. The reasons for these deletions include any one,
or a combination, of the following factors: 1) except under extremely unusual circumstances,
the species does not occur in caves, 2) vertebrates form an insignificant part of their diet, 3)
they do not ingest bone, 4) ingested bone is usually completely digested, as is the case in all of
the lower vertebrates and some higher vertebrates (Fish: Barrington 1942, Phillips 1969,
Bellairs 1970; Reptiles: Benedict 1932, Pope 1961, Dmi'el & Zilber 1971, Dandrifosse 1974,
Skolcezylas 1978; Birds: (herons: Ardeidae) Glue 1970). These criteria would exclude all but
the following group of vertebrates from consideration (in alphabetical order): Dasyurus spp.
(native cats), Falco berigora (Brown Falcon), F. cenchroides (Nankeen Kestrel), Homo sapiens
(humans), Macroderma gigas (Ghost Bat), Sarcophilus spp. (devils), Thylacinus cynocephalus
(Marsupial Wolf), Thylacoleo carnifex (Marsupial Lion), Tyto alba (Barn Owl), T.

AUSTRALIAN CAVE TAPHONOMY - 287

novaehollandiae (Masked Owl) and T. tenebricosa (Sooty Owl). All have occurred within fossil
deposits of the late Quaternary (except Tyto tenebricosa) and all may have been important in
concentrating material. Canis familiaris dingo (Dingo) is not included, because it is not
considered to have dispersed to Australia until the late Holocene (Milham & Thompson 1976)
and the use of caves as dens and lairs by canids is very ephemeral (i.e. only during breeding up
to the time the young are capable of going out on kills, Binford 1981:203). Although not
pertinent for most cave localities, crocodiles should certainly be considered in fluvial and
lacustrine situations (see Fisher 1981).

In Appendix I, I provide the known information on the predators and scavengers important
in Australian cave deposits. Each taxon's section includes the characteristic taphonomy or
damage to bone, natural history (i.e. time of forage, type of accumulator, and any pertinent
information), habitats frequented, prey, dates of first and last occurrence in the fossil record,
foraging range and fossil sites attributed. Although both mammals and birds are included in
Appendix I, in the text I emphasize the avian accumulators, especially the tytonids, because
they play by far the largest role in the accumulation of vertebrate material.

Avian Carnivores

A vast array of bird species are known to eject pellets, including, most frequently, those in
the orders Accipitriformes (i.e. hawks and falcons of the families Accipitridae and Falconidae)
and Strigiformes (i.e. owls of the families Tytonidae and Strigidae: Mayhew 1977); with
apparently more than 60 families and 330 species producing pellets regularly (Glue 1970).
Regurgitation pellets are the "...accumulations of the undigested portions of bird food items
which, instead of being excreted with other waste materials are regurgitated and ejected through
the mouth in compact units" (Glue 1970). Pellets are largely composed of "...bones of
mammals, reptiles, amphibians and birds; claws, beaks and teeth; insect head parts, wing cases,
legs; seed husks and the like. Such hard parts are usually enclosed by softer substances such as
mammal fur, bird feathers and vegetable fibre..." (Glue 1970).

For the sake of this discussion we need only concentrate on birds that take vertebrate prey.
Within Europe the avian families known to take mammals include Tytonidae, Strigidae,
Accipitridae, Falconidae, as their main source of food, and Laridae, Corvidae, Ardeidae and
Laniidae eat mammals regularly (Glue 1970). While Australia lacks any laniids, only those
species which use caves regularly are considered here, therefore restricting the list to the
Falconidae, Strigidae and Tytonidae.

There seems to be a large variation with regard to the degree of digestion of vertebrate prey
items between the Accipitriformes and Strigiformes. Duke et al. (1975) demonstrated that on
average, owl pellets yielded 46% bone and that from accipitrids only 6.5% bone. Dodson &
Wexlar (1979) correlated this to the differences in stomach pH where the average basal pH of
accipitrids is 1.6 and that for strigids is 2.35. Because of this, the bone in accipitrid rejection
pellets is characterized by being well digested. Although mammal teeth are frequently
regurgitated without being severely altered, much of the long bone is largely unrecognizable due
to the chemical corrosion, where "...crosion of limb bones takes place chiefly at the ends
producing sharp pointed diaphysial fragments..." (sce Mayhew 1977). Variation also occurs in
the digestion within Strigiformes, and this is discussed in the following section.

The faecal material of birds has been found to be completely devoid of bone (Farner 1960),
and so accumulations referred to this class of vertebrates can only have resulted from
regurgitated pellets.

The avian predators currently living in southern Australia are assumed to have occurred there
some time within the late Quaternary. At least one extinct, undescribed accipitrid from Mair's
Cave and one from Green Waterhole Cave were also present. Although many avian
accumulators have been accounted for in cave deposits, several (i.e. Milvus migrans (Black
Kite), Aquila audax (Wedge-tailed Eagle)) may have been attracted there by the remains of dead,

288 - BAIRD

dying or semidevoured carcasses. Therefore, their input into the actual assemblage was
probably minimal, This is reinforced when actual cave dwelling animals are taken into
account. Very few species live or roost in caves, and it is only these species that will input
bone through attrition or actively accumulating bone as part of their lifestyle.

Reversed sexual size dimorphism (i.e. females larger than males) is important to the
consideration of avian predators as accumulators of vertebrate material. The most widely
accepted hypothesis for the presence of this phenomenon is that it allows the predators to
sample more than one type of prey, whether it be size or habitat (niche) specific (Earhart &
Johnson 1970). In Australia the range of sexual size dimorphism for diurnal raptors has been
documented by Baker-Gabb (1984). For some diurnal raptors the dimorphism can be quite
large, but for those members considered below (e.g. Falco berigora and F.. cenchroides) the
amount of dimorphism in not quite as marked and, therefore, is not considered to be a major
determinant of prey size. Although similar studics have not been completed on Australian
nocturnal raptors, sexual size dimorphism is known to be quite marked (Schodde & Mason
1980). Unfortunately, the data presented in Appendix I are only of a preliminary nature and do
not take into account the effects of sexual size dimorphism on discrimination of prey size,
although this is discussed.

Species within the Accipitriformes differ from those in the Strigiformes in that the former
arc, in general, diurnal (except for Elanus scriptus (Letter-winged Kite) in Australia), and the
latter are largely nocturnal or crepuscular. There are two major points of contention to the idea
of members of the Accipitriformes accumulating bones in caves. The first and probably most
minor of the two is that only two species are regular occupicrs of caves, and then only within
the twilight sections of the cave. The most common is Falco cenchroides, which nests in
many of the sinkholes of the Bunda Plateau. The least common is Falco berigora, which is
only recorded from a few of the caves of the Bunda Plateau. The second obstacle to
Accipitriformes accumulating bones in caves is that their stomach acid pH is so low that very
little of the bone ingested actually comes out again (Duke et al. 1975, Cummings et al. 1976,
Mayhew 1977, Dodson & Wexlar 1979). Were it not for their habit of snipping off the wings
and legs of their prey and occasionally discarding partial carcasses, their accumulations would
largely be barren of any identifiable elements (Sharland 1931).

Suiviformes

By far the most frequent accumulators of vertebrate bone in caves are members of the avian
order Strigiformes (owls). Owls are implicated as the main accumulators of vertebrate material
because i) they use caves regularly, ii) they prey on vertebrates and it) they regurgitate pellets
containing vertebrate bone. Some strigiform species have been distinguished by the
characteristic damage they impose on avian postcranial clements (Mourer-Chauvire 1983) in
combination with the mean size of the prey species involved (Brain 1981). Dodson & Wexlar
(1979) listed eight characters, which, if combined, would characterize "...accumulations of bone
resulting from food processing activities of several species of owls", including (descriptions of
damage to individual elements refers to mammalian clements and may not be similar to that in
birds):

"1.) abundance and high quality of bone; 2.) possible extreme inequitability of
species distribution; 3.) good representation of all skeletal parts including both
mandibles and vertebrae that contrast strongly with their transport behavior
(Dodson 1973); 4.) high representation of mandibles (possibly with damage to
the articular region) and complete femora relative to other skeletal parts; 5.)
highly fragmented skulls with isolated maxillae and premaxillae well
represented, or intact skulls with occiput and cranial vault removed; 6.) heavily

AUSTRALIAN CAVE TAPHONOMY - 289

damaged scapulae and pelves; 7.) breakage of the proximal humeri, distal radii,
ulnae and femora; and 8.) heavy damage to either end of the tibiae."

Additional studies by Hoffman (1988) stress the use of quantitative analyses.in order to
determine the exact taphonomic accumulator,

Two families within the Strigiformes can be considered likely candidates for accumulating
bone in Australia, the Strigidae and the Tytonidae. Although the Tytonidae are the main
speleophiles, at least one species of strigid owl has been recorded in Australian caves, Ninox
novaeseelandiae (Dwyer 1966). Ninox novaeseelandiae is not considered to be a major
accumulator of vertebrate material due to the paucity of records of this species using caves, the
small amount of fossil material referable to N. novaeseelandiae in cave deposits (see Baird, this
volume), and the fact that its diet consists largely of invertebrates. Deposits accumulated by N.
novaeseelandiae might be distinguished from those of the tytonids by an abundance of insect
remains (if preserved), which are preferred during the non-breeding season, and by the size of the
vertebrates that it selects (e.g. Planigale sp., Antechinus sp., Sminthopsis sp., Mus musculus
(House Mouse), and small birds: Fleay 1925, Vestjens 1973). An additional characteristic
allowing separation of material accumulated by N. novaeseelandiae from that taken by tytonids
is that the condition of bone in pellets of strigid and tytonid owls is different. This has been
studied by Raczynski & Ruprecht (1974) and Dodson & Wexlar (1979), who found that
strigids, on average, digest bone to a greater degree and cause more damage to skeletal elements
than tytonids.

The Tytonidae are, in fact, the group of avian predators most likely to have their
accumulations of pellets preserved because i) the amount of manipulation of prey is minimal
prior to ingestion, ii) their stomach pH is high and iii) several members inhabit caves in
Australia (i.e. T. alba and T. novaehollandiae). Although T. tenebricosa has not been recorded
as using caves, it docs range over karst regions in Australia (see Fig. 5) and, therefore, should
not be totally disregarded. In addition, Schodde & Mason (1980) point out that members of
this species "...roost in sheltered enclosures..." and, therefore, may use caves. Tyto
longimembris, the Grass Owl, with its restricted habitats of large expanses of tall tussock grass
or sedges excludes it from being a speleophile and, therefore, from consideration here.

Characteristics of mammalian assemblages accumulated by tytonid owls, include: 1) the
largest animals are represented by juveniles, 2) most animals tend to be small, with the largest
the size of a small rabbit, 3) there is no evidence of acid digestion, 4) most bones are whole, 5)
jaws and skulls retain teeth, although the teeth may be loose in their sockets, 6) skulls are
either complete or have a crushed bases (i.e. basicranium, occiput and bullae) or, the occiput
and bullae are simply separate from skulls, with no breakage, and, 7) assemblages are made up
largely of nocturnal or crepuscular mammals (especially rodents) and birds (compiled from
Lundelius 1966, Korth 1979).

The territorial nature and foraging patterning of most owls may be a significant factor to the
range of prey captured and the palaeoenvironment interpreted using prey items included in the
assemblage. This has been explained by Martin (1986) through the following:

"Many features of the natural history of the Tawny Owl become explicable in
the light of the sensory limitations and the behavioral and cognitive aspects of
nocturnality discussed above. Thus, the high degree of territoriality may be
interpreted as essential to permit prey capture and general mobility when light
levels become limiting for the immediate visual guidance of flight and other
behaviour. To stray out of the territory (for example in response to a shortage
of optimal size prey) is of no advantage since it is specific knowledge of
landmarks and regularly used perches that is essential for prey capture and

290 - BAIRD

movement under restricted sensory input. For similar reasons invasion of an
adjacent territory is of little value. The use of the "perch-and-pounce" hunting
technique, employing a limited number of perches, is also comprehensible,
since the use of these regular perches will facilitate the accumulation of the
topographical knowledge required to mediate prey capture using audition alone."

Martin (1986) puts forward the idea that detailed knowledge of the topography within an
owl's territory is vital to the success of that individual due to the inability of owls to see any
more than gross objects on nights of low ambient light. Those species of owls inhabiting
vegetation formations with closed canopies should, therefore, exhibit a greater degree of
territoriality than those inhabiting vegetation formations with no canopy. This is indeed the
case in Australia for T. tenebricosa which is a highly territorial species that inhabits mainly
closed forests and Eucalyptus tall open forests, while T. alba is an irruptive nomad, which
inhabits mainly chenopod low shrubland, Acacia tall scrub and Eucalyptus woodland.

DISCUSSION

This chapter is meant to provide a reference point from which one can now compare the
fossil avian assemblages available. Of the three broad categories (e.g. material from natural
traps, fluvially transported material and animal accumulated material) each has its own set of
characteristics, which are in Table 4.

Table 4: Gross characteristics for assemblages of birds collected by the four major categories
of taphonomic accumulator for caves.

Sub-assemblage Bone
Natural Trap Terrestrial forms Little or no predator
or flocking species induced damage
Habitual Cave-dwellers Habit of frequenting Little or no predator
caves in modern forms induced damage
Kluvially Transported No obvious bias, Damage due to abrasion;
depends upon local sorted
attrition
Animal Accumulated Strongly biased by Characteristic damage
the accumulating through capture,
agent (varies) ingestion and digestion

ESS

Given these characteristics useful in the identification of the accumulator of the fossil
vertebrate assemblages (i.e. wear, size of individual and habits of the prey species), I can now
summarize the accumulators of the cave deposits known to date as interpreted from the avian
assemblages.

Deposits interpreted as natural traps include the avian assemblages in both Amphitheatre
Cave in Victoria (Baird in press) and Green Waterhole Cave in South Australia (Baird 1985).
These assemblages are considered to have been accumulated by gravity-based collecting
agencies. This determination is founded on the following characters i) general lack of any

AUSTRALIAN CAVE TAPHONOMY - 291

damage to the elements, other than in situ decomposition of the bone, ii) wide range of body
weights exhibited by the species represented (Table 9 and Fig. 10) and iii) species composition
of the assemblages. The two deposits are different in the type of trap involved. Amphitheatre
Cave is interpreted as a pitfall, based on the large percentage of terrestrial species in the
assemblage (86%), and Green Waterhole is interpreted as a water accumulated assemblage, based
upon the large percentage of flocking species (70%: see Baird 1985).

Collecting bias is most prominent in these two deposits (Amphitheatre Cave and Green
Waterhole Cave), presumably because the specimens were retrieved using the collectors’ fingers
to sieve the elements from the sediment, and, therefore, the longest elements were preferentially
collected (Fig. 11).

There are no assemblages included in this chapter which are interpreted as being accumulated
by fluvial activity, although the material in Koonalda Cave on the Nullarbor Plain exhibits
minor sorting of the long elements (Fig. 12).

Of the assemblages interpreted as being accumulated by animals, only a section of one
deposit is referred to a mammalian carnivore or scavenger accumulator. The assemblage from
Layer 3 of the Mabel Cave deposit in Victoria is interpreted as accumulated by cf. Dasyurus
maculatus (Tiger Cat) based upon, i) the amount of fracturing exhibited by elements
represented, ii) the lack of chemical corrosion to these elements and iii) the wide range of prey
body sizes represented (Table 5; Figs 6, 12).

Table 5: Statistics of the mean weights for prey items identified from cave deposits across
southern Australia compared with a modern sample in the case of Tyto alba (data from Baird
1986). AU-8 = Skull Cave, EB-2 = Clogg's Cave, EB-1 = Mabel Cave, G-4 = Curran's Creek
Cave, G-2 = Amphitheatre Cave, L-81, Green Waterhole Cave, M-89 = Pyramids Cave, N-4 =
Koonalda Cave, N-62 = Madura Cave and WI-6le = Devil's Lair. MNI = minimum number of
individuals.

Taphonomic Agent Cave oi Weight Statistics
D4 Range (MNT)
Tyto alba
Modern Sample 70 12 - 156 694
M-89 74 23 - 220 28
EB-2 81 6 - 181 696
EB-1 69 8 - 138 69
G4 57 8 - 112 21
N-62 49 10 - 212 43
N4 34 19 - 219 64
WI-6le 73 8 - 219 153
AU-8 49 8 - 219 67
cf. Dasyurus maculatus
EB-1 216 43 - 700 89
Natural Trap
G-2 545 43 - 1143 67
L-81 658 78 - 3653 47

The bulk of the deposits interpreted as being accumulated by animals are referred to the
avian primary carnivore, Tyto alba. Assemblages referred to this species are characterized by
their, i) relatively good preservation with complete and incomplete elements outnumbering
those of the terminal ends (Fig. 12 for an example), ii) body size distribution concentrated
below 100 g, and body weight ranges between 5 and 220 g (Table 9 and Fig. 13), and iii) prey

292 - BAIRD

assemblages attributed to T. alba include, i) the specific abundances of elements, which are very
uniform across a range of prey species (e.g. Coturnix sp. (Quail sp.), Turnix varia (Painted

cf. Dasyurus maculatus

90 EB-1
(3rd layer)

Prey Body Weight (g)

Natural Traps

100 300 500 700 900 1100 1300
Prey Body Weight (g)

L-81

100 300 500 700 900 1100 1300 3500 3700

Prey Body Weight (g)

Figure 10. Histograms of weight distributions for prey species in three avian assemblages from caves,
including that from EB-1, Mabel Cave attributed to cf. Dasyurus maculatus (unpublished data from Baird 1986) and
those from G-2, Amphitheatre Cave and L-81, Green Waterhole Cave attributed to Natural Traps.

AUSTRALIAN CAVE TAPHONOMY - 293

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Figure 11. Graphic distribution of postcranial elements of Gallinula mortierit (Top) and Dasyornis broadbenti
(Bottom) from Amphitheatre Cave demonstrating the bias for long bones by the collecting method of sieving with
hands [A , element abundances and B , element total lengths]. H , humeri; U, ulnae; C, carpometacarpi; F,
femora; T, tibiotarsi; Ta, tarsometatarsi and Co , coracoids.

Button-quail), T. cf. velox (Little Button-quail), Glossopsitta porphyrocephala (Purple-crowned
Lorikeet): see Baird 1986) and a number of deposits (i.e. Clogg’s Cave, Mabel Cave, Madura
Cave, Devil's Lair, and Skull Cave), and ii) the relative percentages of certain avian behavioural
groups. Of the avian behavioural groups identified the terrestrial/crepuscular animals that are
irruptive, most are commonly represented (>50%), with terrestrial, diurnal species and non-
terrestrial, diurnal species that are gregarious being represented subequally next most abundant.

294 - BAIRD

50 -
B ao Turnix cf. velox (N-4)
Q
5 30 - a _—+@
E 2) ze ;
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H U Cc F T Ta Co
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Total number

complete elements

proximal ends

Figure 12. Abundances of seven commonly represented post-cranial elements of Turnix sp. cf. T. velox and
Phaps chalcoptera from the excavations of two caves: N-4, Koonalda Cave and EB-1, Mabel Cave, respectively.
That from Koonalda Cave is an owl deposit, which has been altered by fluvial activity and that from Mabel Cave is
attributed to cf. Dasyurus maculatus. Included in each are the sums of complete elements and most common
terminal ends (top: data points are connected to facilitate visual cognition of changes) and the proportions of
fragmentary specimens against complete (and incomplete) specimens (bottom). H, humerus; U, ulna; C,
carpometacarpus; F, femur; T, tibiotarsus; Ta, tarsometatrsus and Co coracoids. (Data from Baird 1986).

AUSTRALIAN CAVE TAPHONOMY - 295

90 Tyto alba 90 T. novaehollandiae
70
ge 50
30
10
20 40 60 80 100 120 140160 180 200 100 300 500 700 900 1100 1300 1500
Prey Body Weight (g) Prey Body Weight (g)
EB-2
20 40 60 80 100 120 140 160 180 200 20 40 60 80 100 120 140 160 180 200
90 90
EB-1 G-4
A9 (1st layer) 70
50
30
10
20 40 60 80 100120 140 160 180 200 20 40 60 80 100 120 140 160 180 200
90 90
N-62 N-4

20 40 60 80 100 120 140 160 180 200

WI-61E Au-8

20 40 60 80 100 120 140 160 180 200 20 40 60 80 100 120 140 160 180 200

Figure 13: Histograms of weight distributions for prey species in eight avian assemblages from caves as

compared with those from modem prey items of Tyto alba and T. novaehollandiae, demonstrating their
similarity to the former. Including M-89, Pyramids Cave, EB-2, Clogg's Cave, EB-1, Mabel Cave, G-4,
Curran'’s Creek Cave, N-64, Madura Cave, N-4, Koonalda Cave, WI-6le, Devil's Lair and AU-8, Skull Cave.
(Data from Baird 1986).

296 - BAIRD

Bias towards certain types of prey by tytonid owls is noted from studies of living owls (see
Appendix I), and additional information may be provided from fossil assemblages themselves.
It appears that animals most accessible to the crepuscular and nocturnal predators depends on i)
the habitat of the prey species, ii) the social structure of the prey species (i.e. whether
gregarious or not), iii) the amount of noise made during roosting, iv) whether the animal is
diurnal, nocturnal or crepuscular, and v) the regularity of population irruptions. Based on the
lists of fossil forms from the caves studied so far, it is apparent that a very small number of
species actually make up the bulk of the individuals preyed upon.

Species of birds most likely to appear in owl accumulated deposits include those species
exhibiting a tendency to population irruptions (¢.e. Coturnix sp., Glossopsitta
porphyrocephala, Turnix varia, T. sp. cf. T. velox), which are most abundant in fossil deposits,
whether the species are terrestrial or arboreal. Also of importance are terrestrial species,
particularly those associated with wetlands and heaths (e.g. rallids (rails), Pezoporus wallicus
(Swamp Parrot), Alrichornis spp. (scrub-bird spp.), Dasyornis spp. (bristle-bird spp.), etc.
These may be important because 7. alba is known to demonstrate regular patterning during
foraging over streams and rivers (Taberlet 1983). A third group of prey species important to T.
alba are diurnal, non-terrestrial species which are gregarious (e.g. psittaciform species (parrot
sp.), Artamus spp. (woodswallow spp.), etc.). They may be particularly accessible because of
noise made during roosting or due to their gregarious nature, making their relative abundance
greater than that of other non-gregarious species.

What should be most apparent from this chapter is that not only are we capable of
determining the accumulator of most cave deposits but also to some degree of interpreting the
biases involved in the collection of prey items by those accumulators. This knowledge is
critical in further studies involving the vertebrate assemblages (e.g. determination of
palacoenvironments, extinction sequences, palaeobiology, etc.) in order to understand the biases
in our data set.

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APPENDIX I: MAMMALIAN CARNIVORES AND
SCAVENGERS.

Dasyuridae
Sarcophilus species

Taphonomy. The characteristics of Sarcophilus accumulated material are severe crushing, fracturing and corrosion, presence
of circular punctures, frayed fractures of element ends and a wide range of body sizes for the animals represented (see below).
The surface corrosion occurs through dissolution by stomach acid, resulting in the removal of the surface cortex and the
exposure of the trabecular tissue (Douglas et al. 1966, Lundelius 1966).

In the fossil record there are at least two separate species of Sarcophilus (e.g. S. harrisii and S. laniarius), with the
possibility of a third, smaller, species (see Dawson 1982). The taphonomic characters of all three cannot be separated. It is
assumed here that they had similar feeding behaviours and foraging habits to the living species, S. harrisii.

Natural History. Sarcophilus harrisii is a nocturnal scavenger unable to efficiently capture and kill living prey (Buchmann
& Guiler 1977). It is considered to be non-territorial but to have regular scavenging routes (Guiler 1970). Only the young are
capable of climbing trees (Guiler 1970).

Buchmann & Guiler (1977) discussed the feeding behaviour of S. Aarrisii and mentioned that small prey (e.g. rats and small
birds) were consumed in a single five to ten minute session whereas larger carcasses were fed upon over a longer period of
time. Pieces of these large carcasses were often detached and consumed completely, including all large bones except the most
robust (e.g. pelves). The authors described the large bones as being crushed prior to swallowing. Werdelin (1986) stated that
S. harrisii is similar to the hyaenids in its cranial adaptations to cracking bones.

Caged S. harrisii individuals have a habit of removing food to secluded parts of their pens before consuming it (Buchmann
& Guiler 1977). In the wild similar behaviour may have occurred with the food items being eaten in the caves. Bone deposits
resulting from these species, therefore, are largely the accumulation of uneaten portions of carcasses and of bone in faecal
material.

Habitats. Sarcophilus harrisii currently ranges throughout Tasmania. They will occur anywhere with sufficient cover, but
favour sclerophyll forest and coastal scrub (Guiler 1970). Fossil specimens have been found over the whole of Australia
(Calaby & White 1967, Archer & Baynes 1972, Dawson 1982).

Prey. "It has been shown that Sarcophilus does not gain most of its food by predation (Buchman & Guiler 1977) and most of
its food is obtained from carrion” (Guiler 1978). Mooney (1983a) mentioned that Sarcophilus will eat the chicks of both
Falco berigora and Falco peregrinus when the nests are located on accessible cliff faces in Tasmania. In general, avian remains,
except for terrestrially adapted birds, are only a small proportion of the Sarcophilus diet and then accessible only to juvenile
Sarcophilus individuals due to their ability to climb trees (Guiler 1970). Because of its scavenging habit, the wide range of
body sizes of the items consumed is not unusual (see Table IA), and the accumulated food items of this species may represent
an attritional assemblage.

First and Last Occurrences. The latest dated remains of Sarcophilus on the Australian mainland are from Cave 1, Turner
Brook, south west Westem Australia at 4304160 yBP (Archer & Baynes 1972). The earliest dated remains include those from
Lancefield, Victoria at 26,000 + 500 yBP (Gillespie et al. 1978) and those from Dempsey's Lake, South Australia at >30,000
yBP (Wells 1978).

Foraging Range. Given an abundant supply of food Sarcophilus harrisii travels, on average, 3.2 km [2 miles] during the
course of an evening, but during times of scarce food they can travel up to 16.1 km [10 miles] (Guiler 1970). Therefore, I will
assume that the foraging radius ranges from approximately 3 km to 16 km depending upon the abundance of food, with a mode
of 3 km.

Fossil Sites Attributed. The fossil sites attributed to Sarcophilus include: Seton Rock Shelter (K-30), Kangaroo Island
(Hope et al. 1977); cave in the Dunstan's Limestone Kiln Quarries north of Wanneroo, Western Australia (Douglas ef al. 1966);
Devil's Lair (WI-6le), Wester Australia (Lundelius 1960, Dortch & Merrilees 1971); and Wedge's (SH-14) and Webb's (N-132)
Caves, Western Australia (Lundelius 1960, 1963).

Dasyuridae

Dasyurus maculatus
Taphonomy. Lundelius (1966) characterizes the faeces of D. maculatus as being 4.5 x 1.5 cm and tapered at both ends. The
bone that passed through the digestive system is highly fragmented.
Natural History. This species is both noctumal and diumal, and both primary predator and scavenger. It is capable of
climbing, but spends most of its time on the ground (Edgar in Strahan 1985).
Habitats. Sclerophyll forests and rainforests are considered the optimal habitats for D. maculatus (Edgar in Strahan 1985).
Prey. Dasyurus maculatus feeds upon both terrestrial and arboreal animals, including arthropods, reptiles, birds, rats, gliding
possums, small macropods (Edgar in Strahan 1985). No studies on the dict of D. maculatus were found in the primary
literature.
First and Last Occurrences. The earliest record for D. maculatus is from Seton Rock Shelter, Kangaroo Island at 16,110
+ 100 yBP (Hope et ai. 1977), and this species is still recorded on continental Australia (Edgar in Strahan 1985).
Foraging Range. Unknown.
Fossil Sites Attributed. None to date.

AUSTRALIAN CAVE TAPHONOMY - 303

Dasyuridae

Dasyurus viverrinus (also D. geoffroyii and D. hallucatus).
Taphonomy. Lundelius (1966) characterizes the faeces of D. geoffroyii as 8 x 1.5 cm, tapered at both ends, and the bone
that passed through the digestive tract as highly fragmented. Apparently this species skins its vertebrate prey before eating
(Buchmann & Guiler 1977).
Natural History. Dasyurus viverrinus is both nocturnal and diumal, and both an opportunistic scavenger and a primary

camivore. In a study of the diet of D. viverrinus in southern Tasmania, Blackhall (1980) found that that population "...fed
almost exclusively on the ground."

Table IA: List of avian and mammalian prey items recorded for Sarcophilus harrisii (in alphabetical order: as
compiled from Guiler 1970). * indicates dependance on these prey items (Guiler 1970).

Weight (gms)

Bos taurus approx. 200000
Canis familiaris approx. 11000
Cercartetus nanus 24
Corvus coronoides 600
Dasyurus viverrinus approx. 6000
Equus caballus approx. 200000
Felis catus approx. 3000
Gallus gallus 1000
Tsoodon obesulus 800
*Oryctolagus cuniculus 1600
*Ovis aries approx. 50000
Perameles gunnii 650
Potorous tridactylus 1100
*Pseudocheirus peregrinus 900
Rattus lutreolus 122
Sarcophilus harrisii 7000
Tachyglossus aculeatus 5000
*Thylogale billardieri 4500
Trichosurus vulpecula 3000
*Vombatus ursinus 26000
*Macropus rufogriseus 17000

Habitats. The range of habitats of D. viverrinus are dry sclerophyll forest (Ride 1970), woodland, scrub, heathland and
cultivated land (Green 1973).

Prey. Food consists largely of plants and invertebrates (e.g. insects: Blackhall 1980) but also includes small herpetofauna,
(e.g. frogs and lizards: Sharland 1962), and small mammals and birds (e.g. Malurus cyaneus (average weight 9 gms), Mus
musculus (average weight 15 gms): Blackhall 1980, Godsell 1982). Although Godsell (in Strahan 1985) mentioned that
"ground nesting birds and small mammals such as bandicoots, rabbits and rats are frequently eaten and the carcasses of larger
animals such as wallabies, possums and sheep are scavenged..." direct evidence from the primary literature indicates that the
average size of vertebrate prey items tends to be much smaller, approximating the size of Mus musculus. Anecdotal reports
also suggest that the species is capable of capturing animals up to the size of a domestic chicken (Buckland 1954), although
this is considered an unnatural situation (J. Nelson, pers. comm.).

First and Last Occurrences. The oldest record of this species group occurs in Madura Cave, Western Australia, dated at
37,880 + 3,880 yBP (Lundelius & Tumbull 1978). Dasyurus geoffroyii still occurs in south west Wester Australia. D.

viverrinus has been recorded in various parts of Victoria since European settlement.

Foraging Range. The foraging range for D. viverrinus males extends up to 1 km and females up to 100+ m from the den
(Godsell in Strahan 1985).

Fossil Sites Attributed. Fossil sites attributed to this species group are restricted to three caves in the Buchan area of
Victoria, including: M-27, M-28 and layer 3 of Mabel Cave (Wakefield 1960a, 1960b).

Hominidae

Homo sapiens

Taphonomy. The criteria defining the signature for humans in Australia are produced by both tools and manual breakage. I
am not aware of any study completed on bone from faecal material or, in fact, whether faeces are deposited near the living site.
It has been suggested, however, that both human- and Sarcophilus-accumulated deposits may be indistinguishable, as pointed
out in the following passage from Balme etal. (1978): "Field observations made by J.E. Stanton (pers. comm.) support a
suggestion made by Baynes et al. (1976 p.102) that bone might be extensively chewed and fragmented by human beings as

304 - BAIRD

beings as well as devils:- "Westem Desert Aborigines at the present day commonly crack up and chew quite large bone such as
those from kangaroos, emus and goats, and seem to swallow quite large pieces (perhaps up to 3 cm long and 1.5 cm wide).
Bone of small animals such as rabbits or goannas are commonly chewed into small pieces and swallowed with the meat.”
Morlan (1984) defined criteria by which human caused (via stone tools) and animal caused marks on bone can be
distinguished, including: "(1) anatomical element that is marked; (2) the position on the element, (3) the gross morphology
of ae mark; (4) microscopic features of the mark; and (5) comparison between the contour of the mark and that of the bone
surface.”

For each of these criteria the human-caused mark is decidedly different from animal-caused marks (see Morlan 1984).
Although this has not been tested in the Australian situation, it may provide a framework around which future tests may be
run.

If tools were used in the butchering of the larger animals (this probably is not realistic in the case of most
microvertebrates) then diagnostic microscopic patterns of wear may occur which are discernible through use of scanning
electron microscopy as has been found on bone from other continents (Shipman 1981). Slicing marks using a stone tool, for
example, may be discemed by their V-shaped cross-section, and multiple fine striations may occur within the main groove that
lie parallel to its long axis. This differs from tooth scratches, which are variably V-shaped and U-shaped and which always
have a smooth bottom. Unfortunately, this may still be confused with some trampling marks and caution should be
maintained at all times (Behrensmeyer et al. in press).

Other criteria that may suggest that a particular deposit is attributable to humans include, vertebrate bones with broken at
marrow rich areas (Noe-Nygaard 1977, Archer et al. 1980); completely bumt bone, instead of just one side (Merbs 1967, Coy 1975);
and bone considered allochthonous mixed with shells (Dortch & Merrilees 1973) or in association with a ”,..discrete scatter of mussel
shells" (Hope 1972, 1978).

For those societies known to split bone for marrow a complicating factor in determining presence or absence of human

accumulators would be rockfall from roof spalling (Dixon 1984).
Natural History. Humans are considered to be diurnal, primary carnivores. It is unknown whether caves formed their home bases
or were only considered transit sites. Caves appear to have been litle used by aboriginals during the Pleistocene on continental
ee (Bowdler 1977) but probably had increased usage in those areas where severe weather occurred (e.g. Tasmania, Kiernan et
al. 1983).

Of particular pertinence to microfauna is that the natives of New Guinea eat small birds whole, crunching up the bones and flesh
all together (T. Flannery, pers. comm.) and, unlike the butchering practices hypothesized for a "megafauna” (Hope 1984), small to
medium sized birds would probably be carried back to home base whole. ;

The apparent lack of small to medium sized birds in non-coastal aboriginal middens, particularly caves, may result from these
having been eaten whole and voided in areas away from the living area, or the bones may be so digested that they are unrecognizable
as avian (Dortch & Merrilees 1973).

Prey (avian). Avian prey items consumed by Aboriginals is largely unknown, except for scattered anecdotal reports.

First and Last Occurrences. The oldest record of humans in Australia is from Upper Swan, Western Australia and is thought to
be older than its associated radiocarbon date of 39,500 +2300/-1800 yBP (Pearce & Barbetti 1981).

Foraging Range. Aboriginals typically have two areas of forage, site exploitation territory and site catchment; therefore, a strict
definition of foraging range is tenuous. If occupancy of the cave lasted more than a day, then the site exploitation territory would be
appropriate where "a ‘site exploitation territory’ is the territory surrounding the site which is exploited habitually by the inhabitants
of the site” (Vita-Finzi & Higgs 1970). In this case then a 2 hour or 10 km foraging radius can be assumed to be appropriate (Baynes
et al. 1976, Bailey 1983). On the other hand cave use may have only been transitory, occurring as stop overs along travels which
averaged twelve to fourteen kilometers a day (Gaughwin & Sullivan 1984); therefore, foraging range must be regarded as an average
of these distances.

Cave Sites Attributed. There are several cave deposits attributed at least in part to Homo sapiens, some of which include: Burkes
Cave, New South Wales (Allen 1972), Cave Bay Cave, Hunter Island, Tasmania (Bowdler 1977), Fraser Cave, Tasmania (Kiernan et
al. 1983), Devil's Lair, Westen Australia (Balme ef al. 1978) Mammoth Cave, Western Australia (Archer et al. 1980) and
Putjamarpa Rock Shelter, Western Australia (Gould 1968).

Thylacoleonidae
Thylacoleo carnifex

Taphonomy. Cuts in the bone which were contemporancous with or prior to fossilization and which exhibit the
following combination of characters are considered diagnostic for T. carnifex (from Horton & Wright 1981): "1. The marks are
in opposition. 2. They are of a regular appearance, that is they repeat the same shapes. 3. Where deep enough they are Y-
shaped in cross section. 4. Where long enough they may show a gap between two ends of the cut, and the two ends may be
angled to each other.”

The cuts can exceed 2 cm in length (Runnegar 1983, Flannery & Gott 1984). The first two characters are appropriate for
any carnivorous animal (vs. human), and the rest are considered specific to Thylacoleo,.

Natural History. That Thylacoleo was a primary carnivore has not always been taken for granted. In fact, in the early
20th century this species was regarded as a herbivore. Wells et al. (1982) argue convincingly that Thylacoleo was a camivore
through the study of reconstructed jaw mechanics, microstriations on the teeth and review the past literature on the subject,

It has been hypothesized by Wells et al. (1982) that Thylacoleo, like the Leopard (Panthera pardus), preyed upon ground
dwelling animals, and subsequently dragged the carcasses into trees to avoid competition from scavenging forms like
Sarcophilus. This would be good reason for the lack of its influence on cave deposits where Thylacoleo can be abundant.
Wells et al. (1982) further suggest that the marsupial lion was often trapped in cave sites (pitfalls) and did not use them as
dens. This hypothetical behavior would not exclude them as being significant accumulators in caves, for the kills of leopards
are very important to any discussion of carnivore accumulators in Africa (Brain 1981).

Horton & Wright (1981) suggest that in feeding habits Thylacoleo may have been more like the Cheetah (Acinonyx
jubatus) causing litle damage to skeletal elements. ‘This is based on the small number of bones exhibiting the diagnostic

AUSTRALIAN CAVE TAPHONOMY - 305

damage of Thylacoleo, the small number of elements broken or cut straight through (unlike deposits associated with habitual
bone crushers like Sarcophilus), the lack of bone-laden coprolites associated with Thylacoleo, and a tooth morphology that
can be best described, through functional morphology, to be adapted to meat shearing.

Habitats. Unknown.

Prey. Unknown.

First and Last Occurrences. The earliest dated occurrence of Thylacoleo from Henschke's Bone Dig, South Australia at
33,800 + 2,400/-1,800 yBP and the youngest occurrence is from Spring Creck, Victoria at 19,800 + 390 yBP Archer et al.
(1984) summarize the distribution of localities, yielding fossils of this genus, throughout southern Australia.

Foraging Range. Unknown.

Fossil Sites Attributed. Localities with vertebrate elements demonstrating the diagnostic damage associated with
Thylacoleo include: Darling Downs, Queensland (DeVis 1883, 1900); Reddestone Creek, New South Wales (Runnegar 1983);

Lancefield, Victoria (Gillespie et al. 1978, Horton et al. 1979); Spring Creek, Victoria (Flannery & Gott 1984); and various
unspecified localities in Victoria (Spencer & Walcott 1911).

Thylacinidae
Thylacinus cynocephalus

Taphonomy. Unknown.

Natural History. Thylacinus cynocephalus is a largely nocturnal primary camivore (Rounsevell in Strahan 1985).

Keast (1982) presented the idea that Thylacinus was a pursuit camivore similar to Canis lupus, but this has been since
challenged by Werdelin (1986), who noted that if skull shape, and not size, were taken into account then T. cynocephalus is
most similar to Vulpes vulpes, not Canis lupus. Werdelin (1986) subsequently stated, “this result is especially interesting in
view of the very different modes of predation of V. vulpes, which is an ambush predator, and C. lupus which is a pursuit
predator. It is evident that other analogues than C. Jupus must be considered in any discussion of the behavioral ecology of T-
cynocephalus."

Smith (1982) also suggests that the limb proportions of T. cynocephalus would be better adapted to stalking in dense
cover with short bursts of speed rather than to prolonged chasing.

Conceming its ability to accumulate vertebrate material Thylacinus cynocephalus may not have been a consumer of bone,
which is suggested by the the lack of bone-laden coprolites associated with that species (Horton & Wright 1981).

Habitat. Using the fossil record, Smith (1982) stated that "the widespread fossil distribution proves that, even allowing for
climatic change, its range of habitats was broad, a fact confirmed by the fossils with which it is associated" and that “its
alleged preference for dense forest is quite likely a result of persecution".

Prey. The prey items of T. cynocephalus are largely unknown, except for anecdotal reports of them eating Tachyglossus and
preying upon domestic stock, as reviewed in Smith (1982).

First and Last Occurrences. This species was last recorded from the mainland from 3090 + 90 yBP (Archer 1974b) and
3,120 + 100 yBP (Calaby & White 1967). The date of 0 + 180 yBP has been questioned by Archer (1974b), who suggests that
the Thylacinus material is not necessarily associated with the dated material. 7. cynocephalus was last recorded from Victoria in
Clogg's Cave between 17,000-23,000 yBP (Flood 1973). The carliest record for the genus is in the Miocene (Woodbume
1967). A listing of the fossil localities from which T. cynocephalus is recorded was presented in Smith (1982).

Foraging Range. Unknown.

Sites Attributed. None to date.

Chiroptera
Macroderma gigas

Taphonomy. Unknown.

Natural History. Macroderma gigas is a noctumal, primary camivore. According to Richards (in Strahan 1985), it is the
only carnivorous bat in Australia. The prey is usually captured on the ground and subsequently taken back to an established
feeding locality, often a rock shelter or small cave (Richards in Strahan 1985).

Habitat. Habitats frequented by Macroderma gigas include those associated with the "...arid Pilbara region... [to] ...the lush
north Queensland rainforests..." (Richards in Strahan 1985).

Prey. Richards (in Strahan 1985) lists large insects, frogs, lizards, birds and small mammals (including other bats), as the
main food items of this species.

First and Last Occurrences. Macroderma gigas currently lives on continental Australia.

Foraging Range. Unknown.

Sites Attributed. None.

APPENDIX II: AVIAN ACCUMULATORS

Accipitridae

Falco cenchroides

Taphonomy. Unknown for this particular species although studies have been completed on a falcon of similar size from North
America (i.e. F. sparvarius) where the researchers describe the following behaviour (Dodson & Wexlar 1979): "This small falcon,
holding the mouse in its talon, consumed the prey mouthful by mouthful, commencing at the snout and systematically working its way
caudally. The damage resulting from this process is ... only 20% of bone escapes breakage.” ;

Although unstudied it is suspected that the avian portions of deposits accumulated by F_ cenchroides will be characterized by the
high abundances of distal limb elements due to its habit of snipping off the wings and legs of its prey (Sharland 1931). If their habuis
are similar to that of the North American F. sparvarius then a variety of other elements may be represented because "depending upon

306 - BAIRD

ot of hunger kestrels [F. sparvarius] may cease feeding at any time, abandoning a variably-sized carcass” (Dodson & Wexlar

Natural History. This species is considered a diurnal, primary camivore and hunts from both perches and on the wing.

Habitats. Falco cenchroides is distributed throughout Australia, with a marked preference to open formations (Hollands 1984).

Prey. Through most of the non-breeding season this species is mainly an insectivore, but during the breeding season it is known to

take more vertebrates, including mammals, birds and reptiles (Lea & Gray 1935). Similar to F. berigora, F. cenchroides is

considered a generalist with regards to food preference, where a generalist is defined as "...take[ing] both invertebrate and vertebrate

prey. Apart from exploiting prey species which may be locally common, litde if any dietary specialization occurs” (Czechura 1979).
Hollands (1984) recorded Anthus novaeseelandiae (Richard's Pipit), Cinclorhamphus cruralis (Brown Songlark) and Sturnus

vulgaris (Starling) as prey items brought to the nest and Czechura (1979) included Coturnix australis (Brown Quail) and C, chinensis

(King Quail) to the list of avian prey items collected by F. cenchroides.

. Apparently this species will take advantage of plagues in any of a wide variety of prey (Blakers et al. 1984) and in the arid

interior move into areas of recent rain to breed (Brooker et al. 1979).

Oldest Known Occurrence. This species is recorded from Koonalda Cave, South Australia at a level dated between 19,300 + 300

and 23,700 + 850 yBP (Baird 1986).

Foraging Range. Unknown.

Sites Attributed. None to date.

Accipltridae
Falco berigora

Taphonomy. Unknown.

Natural History. Falco berigora is considered to be a diumal, primary predator and scavenger. Its movements are largely
unknown, where in southem Australia Blakers ef al. (1984) regard it as sedentary and Frith (1969) regards it as migratory. This
species feeds from exposed perches or by walking through grass.

Prey. Falco berigora is considered to be a food generalist and "takes a wide variety of prey: small mammals, rats, mice,
small rabbits, birds, particularly pipits, and other small ground birds but also capable of catching such birds as Starlings and
woodswallows in flight; lizards, small snakes and large insects which are mostly taken on ground” (Hollands 1984).

Hollands (1984) also mentioned that F. berigora "will come down to carrion.”

Changes in the diet according to seasons have been recorded (Blakers ef al. 1984), where the diet consisted of 50%
mammals in winter (with the other 50% composed of birds, reptiles, insects and carrion) to 50% insects in summer-autumn.
This species is also known to follow plagues of both insects and mammals (Appleby 1976).

Oldest Known Occurrence. Falco berigora is recorded from Koonalda Cave, South Australia at a level dated at between
13,700 + 270 and 19,300 + 300 yBP (Baird 1986).

Foraging Range. Unknown.

Sites Attributed. None to date.

Tytonidae
Tyto tenebricosa

Taphonomy. The taphonomy of deposits accumulated by this species is unknown, except for the general
characteristics associated with tytonids. Schodde & Mason (1980) mentioned that "Sooty Owls have the same feeding habits
as other masked owls. The heads of warm-blooded prey are first nipped off and swallowed whole, followed by the viscera and
forequarters.”
ont this species seems to have a preference for arboreal mammals and samples a broad range of prey body weights (see
below).

Natural History. Tyto tenebricosa is considered a sedentary and territorial, primary predator (Blakers ef al. 1984). Schodde
& Mason (1980) state that it is “...restricted in Australia to the south-east coast and adjacent slopes of the Great Dividing
Range...”

avordiiig to Martin's (1986) hypothesis this owl should be the most highly sedentary of all the tytonids in that the
canopy cover in its territories is very dense, therefore allowing little ambient light. Because of this, these owls must rely
upon a detailed local knowledge of their surroundings for the capture of prey. This may possibly explain the reason for the
preponderance of arboreal prey in their diet, as capture at or just below canopy level would allow for a greater amount of
ambient light to be available.

Habitat. Schodde & Mason (1980) describes the habitat of T. tenebricosa thus "...they favour galleries and pockets of tall,
wet, gully forests along creeks, particularly those overtopped by great eucalypts... [which] have [a] substorey of rainforest
trees and treeferns.”

Prey. Blakers et al. (1984) reported the diet of T. tenebricosa as consisting of “...Mammals : mainly terrestrial (rodents and
bandicoots) and, to a lesser extent, arboreal (possums and gliders) in a wide range of sizes (see Table IIA). They also take
birds..."._ I can find no published evidence to support this definition of diet. In fact, based upon other sources this species
seems to take largely arboreal mammals (Seebeck, pers. comm., "mainly take arboreal mammals and perching birds, as well

as canopy insects.").

AUSTRALIAN CAVE TAPHONOMY - 307

Table ITA: Percentage contributions of animals of different weights to the range captured by Tyto tenebricosa. N = total
MNI (52).

Weight Classes (g) N %
0 - 20 1 2
20 - 40 8 15
40 - 60 1 2
60 - 80

80 - 100
100 - 120
120 - 140 2A 46
NYY
280 - 300 8 15
Wn
560 - 580 1 2
ANY

700 - 720 6 12
Wy

900 - 920 3 6

Oldest Known Occurrence. No fossil record.
Foraging Range. Unknown.
Sites Attributed. None to date.

Tytonidae
Tyto alba

Taphonomy. The taphonomic characteristics defining tytonid deposits are based upon studies of T. alba (see above). Fossil
vertebrate assemblages accumulated by T. alba may be differentiated from those of T. novaehollandiae and T. tenebricosa by the
narrow range of body weights of prey species and the small mean body weight for the whole assemblage (see below and
Discussion).

Similar to other tytonids, T. alba typically feeds in the following manner (description from Dodson & Wexlar 1979): "A

barn owl held the mouse in its talon, carefully severed the head from the neck (opening the cranium in the process?) and
swallowed it whole. It then opened the thoracic cavity and fastidiously removed through the enlarged thoracic inlet the
thoracic and abdominal viscera, which it neatly consumed: finally it swallowed the hollowed out carcass, neck first. Observed
destruction of bone during ingestion was thus minimal.”
Natural History. Tyto alba is considered a primary camivore, which is crepuscular and nocturnal, but rarely diurnal.
Schodde & Mason (1980) considered this species "...essentially irruptive nomads, gathering opportunistically at plagues or
abundances of native and introduced rodents..." It is a solitary hunter which uses both gliding, and perch and pounce methods
(Blakers et al. 1984).

Tyto alba demonstrates regular patterning during foraging and frequently uses flight paths over streams, river, etc.
(Taberlet 1983). This species has been reported as using caves (see Parker 1977 for a listing).

Habitats. Tyto alba is cosmopolitan and seems to favour open habitats over closed habitats throughout the world (Fast &
Ambrose 1976). This trend is also seen in Australia where the range of habitats occupied by T. alba includes "...woodland,
forest or rainforest" (Blakers ef al. 1984). Its preferred habitat is "...light woodlands, the edges of timbered watercourses, and
savannahs..." (Schodde & Mason 1980) and heavily wooded areas seem to be avoided (Dickison 1941, Mees 1963, Schodde &
Mason 1980).

Prey. Tyto alba in southem Africa "...typically feeds on prey weighing approximately 60 g per item” (Brain 1981:125). A
North American study found that the mean weight for bird species collected by T. alba was 64.6 gm (Fritzell & Thome 1984).
A similar prey size range is exhibited by Australian 7. alba individuals (Table I[B), although the figures may be heavily
influenced by the introduced Mus musculus, which makes up 1408 MNI in the 0-20 gm weight class in Table IIB. What is
important is that most of the prey captured by this species are below 100 gm, and weight range from 6 to 320 gm.

Brain (1981 : 126) stated "it is abundantly clear that, although bam owls may have some dietary preferences, they will
feed on the most readily available food source as long as it is palatable and of manageable size”. Australian 7. alba
individuals also take advantage of irruptive prey species and this is exemplified by the following: “During the summer and
autumn of 1970 (before the survey was started) a plague of house-mice Mus musculus occurred...During that period the contents
of a number of stomachs of Bam Owl Tyto alba, Boobook Owl Ninox novaeseelandiae, Tawny Frogmouth Podargus strigoides,
Laughing Kookaburra Dacelo gigas and Nankeen Kestrel Falco cenchroides were examined and revealed that all these birds had
fed on mice only" (Vestjens 1973).

308 - BAIRD

In both North America and Africa Tyto alba are known to feed on birds in winter and rodents and small mammals in
summer (de Graaf 1960, Fritzell & Thome 1984). In North America there is also a shift from mammalian prey to avian prey
during rodent population declines (Hawbecker 1945, Otteni et al. 1972).

Oldest Known Occurrence. Oldest dated occurrence of T. alba in Australia occurs in Koonalda Cave, South Australia
between the dates of 13,700 + 270 and 19,300 + 300 yBP (Baird 1986).

Foraging Range. The most recent study involved in determining the foraging radius for T. alba is Taberlet (1983). In his
study Taberlet found that the foraging radius for 7. alba was at least 2.5 km, which would agree well with the 1.5 km radius of
Schmidt et al. (1971) and the 2 km radius of Geroudet (1978). Therefore, 2.5 km will be regarded as the mode. As has been
pointed out by Brain (1981, below), although the foraging range of the Bam Owl may vary with the abundance of prey it is
usually restricted to that area directly surrounding the roosting site. "The range is likely to be greatly affected by the
availability of food; when prey is abundant, the owls range will be smaller than in times of food scarcity” (Brain 1981:127).
"Thus, from the point of view of environmental reconstructions, owl prey should be regarded as coming from within a few
kilometers of the roost. When prey is normally abundant, some recent evidence suggests that owl hunting in the Transvaal
may be extremely localized” (Brain 1981: 127).

Table IIB: Percentage contribution of animals of different weights to the range captured by Tyto alba. N = total MNI (2152).

Weight Classes (g) N %
0 - 20 1608 75
20 - 40 221 10
40 - 60 50 2
60 - 80

80 - 100

100 - 120

120 - 140 269 13

140 - 160

Wn

280 - 300 3 <1

300 - 320 1 <1

Mean Body Weight (with Mus musculus) = 34 g
Mean Body Weight (without Mus musculus) = 70 g

Sites Attributed. These include the caves of Chambers Gorge, South Australia (Smith & Medlin 1982) and Cave 2 of
southwestern Westem Australia (Archer & Baynes 1972).

Tytonidae
Tyto novaehollandiae

Taphonomy. Characteristics defining tytonid species are based upon studies of T. alba (see above). Deposits accumulated by
T. novaehollandiae may be differentiated from those of T. alba and T. tenebricosa by the wide range of body weights for prey
species and the large mean weight for the whole assemblage (see section on prey, below, and Table IIC).

Natural History. This species is considered to be a sedentary, primary carnivore which is both crepuscular and nocturnal.

Tyto novaehollandiae has been recorded from a number of caves, including Marble Arch, New South Wales (Hall 1975);
Clogg's Cave, Victoria (McKean 1963); and several on the Bunda Plateau (summarized in Parker 1977).

Of the three species of tytonid owls considered, this species would probably be the most intermediate with respect to

Martin's (1986) hypothesis, in that it would be moderately territorial, based upon the amount of ambient light available. The
canopy cover of its preferred habitat would only limit ambient light during nights of little to no moonshine and/or heavy
cloud cover.
Habitats. The range of vegetation formations inhabited by T. novaehollandiae includes, ™...eucalypt forest and woodland,
requiring partial clearing or forest edges for hunting but roosting in the dense cover of gullies or caves." (Blakers ef al. 1984).
Tyto novaehollandiae seems to prefer "...heavy forests and woodlands of eucalypts...” and "...requires a greater diversity of
habitat than the others [tytonids] using closed eucalypt forest for roosting and nesting, and forest edge and open woodland for
henting. Because of this their home ranges, each the permanent property of the pair, are large and cover about 5000 - 1000
hectares in eastern Australia (inferred from Hill 1955 and Hyem 1979)" (Schodde & Mason 1980).

AUSTRALIAN CAVE TAPHONOMY - 309

Soon Percentage contributions of animals of different weights to the range captured by Tyto novaehollandiae. N, total

Weight Classes (g) N %
0; 30 32 16
50 - 100 1 <1

100 - 150 62 32
150 - 200
200 - 250
250 - 300
300 - 350 1 el
350 - 400
400 - 450
450 - 500 2 1
500 - 550
550 - 600
600 - 650 9 5
650 - 700
700 - 750
750 - 800 20 10
800 - 850
850 - 900
900 - 950 19 10
950 - 100

1000 - 1050

1050 - 1100 8 4

1100 - 1150

AY

1500 - 1550 4 2

1550 - 1600 37 19

Mean Body Weight = 633 g

Prey. The range of species and range of sizes of prey items for T. novaehollandiae is far greater than that for T. alba and T.
tenebricosa (although only little is known for the later species) (see Table IIC). From available data about half of the prey
items for this species appear to fall below the 100 g weight class and the other half are evenly distributed from this point to
1600 g.

Mooney (1983b) mentioned that in Tasmania T. novaehollandiae is known to eat a variety of bird species (i.e. Falco
berigora Brown Falcon, Platycercus caledonicus Green Rosella, P. eximius Eastern Rosella, Gymnorhina tibicen Australian
Magpie, and Sturnus vulgaris Starling), as discovered from both the pellets and remains left beneath roosts of this species.

Sexual size dimorphism is important in this species. Schodde & Mason (1980) mention that, "the great difference in
talons between male and females implies that, like the Sooty Owl, each takes prey of different average size." This may
explain the large size range in the prey associated with this species.

Foraging Range. The foraging range is unknown for T. novaehollandiae, so that for T. alba will be used (e.g. 2.5 km).
Oldest Known Occurrence. The oldest dated occurrence of T. novaehollandiae is from Koonalda Cave, South Australia
between 13,700 + 270 and 19,300 + 300 yBP (Baird 1986). The species is also recorded from both Pyramids Cave, Victoria,
which is considered to be greater than 30,000 years old, and Devil's Lair, Westem Australia, from the reworked deposit, which
would be older than the oldest radiocarbon date of 37,750 + 2,500 yBP.

Sites Attributed. Included in the sites attributed to T. novaehollandiae are Mabel Cave (Layer 1), and Pyramids Cave, in
Victoria (Wakefield 1960a, 1972) and Cave 3 (AU-25), southwestem Western Australia (Archer & Baynes 1972). Mees (1964)
suggested that “it is almost certain that the deposits of mammal bones in caves on the Nullarbor Plain described by Lundelius
(1963) and others are the work of Tyto novaehollanidae..." although results presented in Parker (1977) may suggest otherwise.

310 - BAIRD

Thylacoleo carnifex, one of several marsupial lions known from the Australian continent. This species
is restricted to the Pleistocene, and in this illustration is preparing to drag the carcass of a newly killed
Grey Kangaroo into a tree out reach of other carnivores and scavengers. (From Rich & van Tets 1985,
with permission of The Museum of Victoria).

CHAPTER 11

PRESERVATION OF
BIOMOLECULAR
INFORMATION IN FOSSILS
FROM AUSTRALASIA

Merrill Rowley!
ATPOGUCHOMY 52s cs hove Bese eens ne vege lessened 312 Extraction and Characterization of
Sources of Biomolecular Information ColaGens AP scan eden caesssey es 320
INMEOSSIIS., coke vetase Ss ecveedceseeslceseer ess 313 Preparation of Antisera............... 320
Preservation of Organic Material....... 313 Extraction of Collagen from the
Biomolecular Information in Soft Tissues: Mummified Skin of a Moa......... 321
Collagen and Albumin............... 313 Immunoreactive Collagen in Bone.....321
DOIN AN ela ata coca ceaassfesteceh gs ec eseie cise 314 Immunoreactive Osteocalcin in Bone..323
Biomolecular Information in Bone: Effect of Environment on Collagen
(Gro) F:17-) 1 eee oe 316 Survival in Bome.............cc:eeeeees 324
OStEOCAICIN: .csicenaeFerzeeesencceaueoss 317 Loss of Collagen in Fresh Bone
Biomolecular Information in Immediately after Death.............. 326
Australasian Fossils...............0+ 318 Extraction of Collagen from Miocene
Analytic Techniques ............2:sceseeeee 318 BONG yeh oo aldckescedec anes ceeds dione oe 326
Radioimmunoassay .............600+0 318 Applications..........cceseccceeeceeeeneeeeeen ees 330
Immunoblotting...................066+ 319 Conclusions ...............cceceeeeececeeeeeeeeeee 332
Preparation of Fossils.............0. 320 Acknowledgements...........:.sececeeeceenee ees 332

REfELENCES: ccrcecsceedseces vocdesessosistoaesegeoses 333

nnn EEE

1 Centre for Molecular Biology and Medicine, Monash University, Clayton 3168, Australia.

312 - ROWLEY

INTRODUCTION

Phylogenetic relationships among living and extinct species have traditionally been
established morphologically. However, such an approach has been hampered by the problem of
convergent evolution, meaning that groups of organisms, not closely related genetically, have
developed morphological similarities as a result of environmental pressures. Moreover, the rate
of morphological divergence may vary greatly between groups of animals. In an attempt to
overcome the problems associated with a classification based entirely on morphological
grounds, taxonomists have turned increasingly to the examination of genetic similarity, by
comparing DNA or RNA or proteins.

Genetic comparisons are based on the recognition that DNA sequence changes
(substitutions, insertions, deletions and rearrangements) are a major source of phenotypic
variation in evolution, since by affecting genes or the regulation of genes, such changes
influence biochemistry, development, morphology and behaviour. Evidence has accumulated
that the probability of base substitutions in the DNA, and hence of amino acid substitutions in
a particular protein, is a function of time, so that evolving proteins serve as "molecular
clocks”, and may provide a measure of how closely two organisms are related, and possibly
how recently they evolved from a common ancestor (reviewed in Wilson et al. 1977).
Although it is likely that the rate of DNA sequence evolution does differ between taxonomic
groups (Wu & Li 1985, Britten 1986, Li & Tanimura 1987) molecular comparisons are
making increasing contributions to taxonomy.

A variety of techniques has been used to measure molecular differences. Proteins have been
sequenced directly (Dayhoff 1972, Ibrahimi et al, 1979, Jolles et al. 1979), or they have been
compared by physical means, such as electrophoresis (Sibley & Ahlquist 1972, Sibley &
Frelin 1972), in which changes in amino acid sequences are recognised by altered mobility in
an electric field. Proteins also have been compared by immunological techniques, in which the
degree of reactivity of a protein with a specific antibody is measured, and compared with that of
the same protein from a different organism (Maxson et al. 1975, Goodman 1976, Kirsch 1977,
Ibrahimi et al 1979).

Comparisons of DNA from different sources have been carried out by directly sequencing
isolated genes and computerised data banks of gene sequences are available to facilitate such
comparisons. Alternatively, a much larger part of the genome may be compared by DNA
hybridization (Sibley & Ahlquist 1983, 1986) in which single-stranded DNA from one
organism is allowed to reassociate with the DNA from a second organism. "Hybrid" double-
stranded molecules form between homologous sequences. These hybrid duplexes contain
mismatched as well as matched base pairs, because of base sequence differences that have
evolved since the two species diverged from their most recent common ancestor. Hence, the
hybrid molecule will dissociate at a lower temperature than would either of the parent
molecules. The greater the degree of genetic divergence between the two organisms, the more
the temperature of dissociation will be lowered.

These techniques are being used to reevaluate the phylogenetic relationships of living
animals, e.g. Man and primates (Lowenstein & Zihlman 1984) or the Giant Panda, bears and
raccoons (O'Brien et al. 1985). However, the lack of suitable genetic material has limited their
use when dealing with phylogenies with extinct members. Nevertheless, under favourable
circumstances, many of the components of living tissues may be preserved long after the death
of the animal.

The first part of this article will draw together much of the evidence for the persistence of
proteins and DNA in ancient materials. In the second half, I review data on the persistence of
collagen in Australasian fossil material, and the interpretations derived from it.

BIOMOLECULAR INFORMATION IN FOSSILS - 313

SOURCES OF BIOMOLECULAR INFORMATION IN FOSSILS

PRESERVATION OF ORGANIC MATERIAL

It is well recognised that, under certain conditions, organic material can survive unchanged
for long periods. For example, the germination of seeds up to 2000 years old (Keilin 1959)
indicates that the genetic code stored in the DNA must have survived intact. Blood group
antigens have been determined from ancient mummy material (Boyd & Boyd 1937, 1939;
Candela 1939; Otten & Florey 1964; Berg et al. 1975), with the necessary reservation that
bacteria are capable of synthesizing structures with the same antigenicity as human blood group
antigens (Hakomori 1974). Moreover, the stability of dried protein is such that Loy (1983)
could demonstrate the presence of blood on prehistoric stone tools 1000 to 6000 years old, and
he has been able to identify the animal origin of the haemoglobin detected.

The two fossil proteins which have been most widely studied are the structural protein
collagen, and the serum protein albumin. Collagen is the most abundant protein in the body
and is very resistant to degradation. Hence, it has been most readily demonstrated in preserved
tissues. However, collagen is a highly conserved molecule and is relatively non-immunogenic,
making it difficult to prepare a range of antisera with which to test samples. In addition,
collagen is made up of three polypeptide chains, with a defined three-dimensional structure, and
in rabbits this "native" molecule stimulates a much stronger antibody response than the
molecule denatured by mild heating, at 43°C. For these reasons, collagen has not been the
protein of choice for molecular comparisons. Instead, albumin has proven the useful
alternative where soft tissues are preserved, and proteins other than collagen may be present.

Albumin occurs naturally in high concentrations in all tissues. It is a single polypeptide
chain of 580 amino acid residues. Although it has a well-defined three dimensional structure its
amino acid sequence is not highly conserved, and it is highly immunogenic across species.
Furthermore, Wilson and colleagues (1977) have carried out thousands of genetic comparisons
on living species, and this large body of comparative information provides an invaluable
framework for analyzing data obtained from the albumins of extinct species.

BIOMOLECULAR INFORMATION IN SOFT TISSUES: COLLAGEN AND
ALBUMIN.

Much of the genetic information which has been obtained so far has come from preserved
soft tissues, such as skin, either naturally preserved, or from museum specimens, mummified
bodies, or animals preserved in the permafrost. These specimens, which are relatively young
geologically, are likely to contain a range of protcins, and are the most likely source of intact
DNA.

Much preservation of soft tissues has taken place by desiccation and mummification,
whether naturally occurring, or induced artificially as a part of funeral rites. Histopathological
examinations have been performed on both Egyptian mummies and on dried bodies from other
ancient cultures (Williams 1927, Reyman et al. 1976). Histochemical (Sandison 1963,
Montes ef al. 1985) or immunofluorescence staining techniques (Wick et al. 1980) have
demonstrated the presence of recognisable proteins, particularly elastin and collagen. Similarly,
collagen has been demonstrated by immunofluorescence in skin from a moa (Megalapteryx
didinus) mummified naturally (Rowley et al. in prep.) (Fig. 1). Using scanning electron
microscopy, the characteristic striations of collagen also have been detected in specimens of

314 - ROWLEY

muscle from the Magadan mammoth (Barnhart ef al. 1980), after being entombed in the
Siberian permafrost for approximatcly 40,000 years.

Figure 1. Collagen demonstrated by immunofluorescence in acetone-fixed frozen sections of Moa skin. A,
reaction with rabbit antiserum to chicken collagen; B, reaction with normal rabbit serum.

Although there are no techniques for the histological or electronmicroscopic demonstration
of albumin in tissues, albumin has been detected in various tissues. This has been
accomplished either by direct assay using an antibody to albumin (Lowenstein et al. 1981) or
by indirect methods in which the tissue in question is injected into rabbits, and the resultant
antiserum shown to react with purified albumins (Prager et al. 1980, Shoshani et al. 1985).
By such means, the presence of albumin has been demonstrated in mammoth muscle (Prager et
al. 1980, Lowenstein et al. 1981), dried muscle or skin from Thylacines (Lowenstein et al.
1981), and Quagga skin (Lowenstein & Ryder 1985).

There have been few attempts to extract and purify these proteins, and the proteins extracted
have been variably degraded. Biochemical analysis of the protein of the Magadan mammoth
showed that the most abundant protein material in the tissue studied was degraded collagen,
although some undegraded, possibly native type I collagen chains were also present (Goodman
et al. 1980). The albumin, also, had undergone post-mortem change (Prager et al. 1980), and
only about 20% of it appeared to be similar in size to native monomeric albumin: much of the
remaining albumin was aggregated. Similarly, the collagen in the moa skin (Rowley et al. in
prep.) was partially degraded, with random intra-chain breakages, producing peptides of a wide
range of molecular weights. Yet in each case, although the protein was partially degraded, it
retained its immunological identity, and reacted specifically with the appropriate antiserum,
anti-albumin, or anti-collagen.

BIOMOLECULAR INFORMATION IN SOFT TISSUES: DNA

As with collagen and albumin, DNA has been detected in ancient tissues. Higuchi e¢ al.
(1984) obtained mitochondrial DNA sequences from dried muscle of the Quagga, a zebra-like

BIOMOLECULAR INFORMATION IN FOSSILS - 315

species that became extinct in 1883. They were able to show that the sequences differed by 12
base substitutions from the corresponding sequences of mitochondrial DNA from a Mountain
Zebra, an extant species. The number, nature and location of the substitutions implied that
there had been little or no postmortem modification of the Quagga DNA sequences and that the
two species had a common ancestor 3-4 myBP. The DNA was extracted from dried muscle and
connective tissue attached to the salt-preserved skin of an animal which died 140 years
previously, and had been stored in the Museum of Natural History at Mainz, West Germany.
The recent origin of the specimen, and the special conditions of preservation, may have
provided particularly favourable conditions for the preservation of DNA.

DNA has been cloned and sequenced from a 2,400-year-old mummy of an Egyptian child
(Padiibo 1985). In this study tissues from 23 different mummies and mummy fragments,
ranging in age from the Sixth Dynasty (2370-2160 BC) to late Roman times, were examined,
but only samples of the epidermis and several subcutaneous structures from one individual, a
less than l-year-old boy, contained DNA. The mummification process in ancient Egypt
consisted of dehydration of the eviscerated body by embedding it in crystalline salts. It may be
relevant that both this mummy and the Quagga skin were salt preserved, and hence rapidly
dehydrated and maintained in dried form: freshly prepared, purified DNA is stable indefinitely
after drying. The general histological preservation of the various mummy tissues was much
better in superficial tissues and peripheral parts of the body than in the more deeply situated
tissues and no DNA could be extracted from any of the deeper tissues.

Attempts to clone DNA from tissues preserved under less favourable conditions have been
less successful. Thus, although DNA was cloned from the Magadan mammoth, after 40,000
years in the Siberian permafrost (Higuchi & Wilson 1984, Jeffreys 1984), most of the DNA
isolated appeared to be of recent microbial origin, and probably introduced after excavation.
Elephant-like DNA sequences were present in very small quantities, and much of the DNA was
severely degraded. Any sequence information recovered from such material is likely to be
seriously distorted by post-mortem modification. I also have been unable to extract DNA from
naturally preserved skin from a moa (Megalapteryx didinus) found in a cave near Cromwell in
the South Island of New Zealand (unpublished observations).

The apparent lack of DNA in these samples is in contrast with the presence of extractable
proteins from soft tissues of similar age e.g. the mammoth and the moa skin. However, DNA
was extracted from human brain tissue obtained from the Windover archaeological site a
swampy pond in central Florida (Doran et al. 1986) dated at about 8,000 years. The yield of
DNA from the brain tissue from the Windover site was only about 1% of that from fresh
tissue, and the quality of the DNA was not good. The DNA digested poorly with restriction
enzymes, probably due to base modifications or other damage. In contrast, the DNA obtained
from both the Quagga and the human mummy was of good quality and had apparently
undergone little post-mortem modification, although the yield was only 1-5% of that expected
from fresh tissue. Swampy conditions do not give uniformly good preservation, however,
since no DNA could be extracted from muscle obtained from the much younger British bog
body, Lindow Man (Hughes et al. 1986).

Although these studies of DNA in preserved tissues have aroused considerable interest, they
are of minimal relevance to palaeontologists, for two reasons. First, the oldest samples tested
which have given unequivocal results are only 8000 years old, a mere instant on the geological
time scale. Studies on older material do not indicate that DNA will be preserved, and the
chemistry of DNA degradation over geological time periods is unknown. Second, the
overwhelming majority of vertebrate fossils consist only of skeletal material and other tissues
are very rarely preserved. Bone is a relatively acellular tissue, and even fresh bone is not a good
source of DNA. For example, we were unable to extract DNA from bone of a King Island
Emu (Dromaius ater), possibly no more than 150 years old, even though the bone contained
levels of collagen similar to those of modern bone (Rowley et al. 1986).

316 - ROWLEY

BIOMOLECULAR INFORMATION IN BONE: COLLAGEN

A much larger potential source of biomolecular information is the wealth of fossil bones
and teeth, ranging in age to hundreds of millions of years. The most abundant proteins in bone
would be the structural proteins of bone, collagen and osteocalcin (Hauschka 1980), which are
embedded in the inorganic matrix of the bone and may be expected to be lost only very slowly.
The retention of serum albumin in bone is much less likely.

The survival of collagen in bones has been implicit in the radiometric dating of fossils
using !14C (Berger et al. 1964), and in the use of amino acid racemization as an alternative
method of dating (reviewed by Williams & Smith 1977). The unique amino-acid composition
of collagen (in particular the 33% glycine, high proline and hydroxyproline) enables it to be
easily identified. Many studies have demonstrated that amino acids can be extracted from fossil
bone (Ho 1965, Cantaluppi 1975, Dungworth et al. 1975, Carmichael et al. 1975, Dungworth
et al. 1976, Stafford et al. 1982, Armstrong et al. 1983).

The most extensive studies of amino acids in bone would be those of Wyckoff (1972), who
examined fossils from many sites, ranging from Pleistocene mammal bones from the Rancho
la Brea tar pits to Jurassic dinosaur bones from the Morrison Formation in Wyoming, U.S.A.
Several important points have come from Wyckoff's work. First, there may be wide differences
in the preservation of amino acids in fossils, even from the same environment, and of similar
age. Wyckoff compared the amino acid composition of numerous bones from Rancho la Brea,
and found that although each had a similar distribution of amino acids to that found in modern
collagens, the total quantity extracted varied widely. In some cases the amount of collagen was
comparable with that of modern bone, but some bones contained 5% or less of that found in
modem material. Second, although the residue from each of the bones from Rancho la Brea had
the composition of collagen, this has not been true for fossils from all other sites. For
example, in a Pliocene site (Matter et al. 1970) in which many animals had been collected from
a sand and silt matrix of lacustrine origin, many of the bones tested had amino acids
characteristic of collagen. However, others had a composition approximating that of bacteria.
Wyckoff's (1972) interpretation was that the residues of most of the bones were the result of an
incomplete rotting of the original carcass. As microscopic study of the bones gave no evidence
of recent microbial attack, and there was no detectable protein in the matrix, this was thought
to have occurred at the time of fossilization. Third, Wyckoff's studies (1972) do not suggest
that the age of the fossil is a good indicator of the amount of amino acids remaining in the
fossil, although generally there was a decrease in amino acids with increasing age, and non-
collagenous residues were more frequent in fossils older than the Pleistocene. However, the
decline in the amino acid content of bones appeared to be slow, and the amino acids
characteristic of collagen have been detected in some dinosaur bones from the Cretaceous and
the Jurassic. Furthermore, the amino acid content of many of the Jurassic bones was as high
as that in the younger Cretaceous samples. Fourth, the environment of fossilization was
important for the preservation of protein, Amino acids were more likely to be found in fossils
preserved in alkaline than in acidic rocks, and the most successful extractions were from bones
embedded in calcitic matrices (Wyckoff 1972, 1980). Preservation in a dry environment may
assist the preservation of collagen, since, when boiled with water, collagen is solubilised into
gelatin, but, when dried, the hydrolytic decomposition which occurs with heat and even traces
of moisture does not occur. Although most of these experimental studies have involved
temperatures far above those likely to occur in the natural environment, such changes may be
expected to take place, more slowly, at much lower temperatures over geological time scales.

Although Wyckoff's studies (1972) provided the first systematic observations on the
preservation of proteins in fossil bone, measurements of amino acids cannot be used for
molecular comparisons, and these studies did not show whether the amino acids remained as
intact proteins, as substantial peptides or as amino acids per se. However, collagen in fossil

BIOMOLECULAR INFORMATION IN FOSSILS - 317

bones has been detected by electronmicroscopy, as fossil bone samples have shown the 640 A
striations characteristic of collagen (Little et al. 1962, Isaacs et al. 1963, Wyckoff & Doberenz
1965, Doberenz & Wyckoff 1967, Tuross ef al. 1980). Although preserved ultra-structure may
be an unreliable indicator of the survival of protein in the tissue (Towe & Urbanck 1972, Towe
1980), proteins containing the characteristic amino acid composition of collagen have been
extracted from a number of the bones showing such striations (Tuross et al. 1980).

Various methods have been used to extract the collagen from bones. The bone samples
have been cut into small pieces, or ground to a powder, and demineralised by extraction with
ethylenediaminetetraacetic acid (EDTA) (Hedges & Wallace 1978, Tuross et al. 1980) or by
dilute acid (Gurtler et al. 1981), and residual protein has been solubilised in saline, dilute acetic
acid, or with pepsin digestion in acetic acid. With modern material, only a small amount of
recently formed, non-crosslinked collagen would be soluble in either saline or acetic acid, but
pepsin digestion is a standard method of collagen preparation (Miller 1971, Chung & Miller
1974). Samples have been further characterised by electrophoresis using sodium dodecyl
sulphate polyacrylamide gels (SDS-PAGE) (Tuross et al. 1980, Gurtler et al. 1981) or
Sephadex gel filtration (Hedges & Wallace 1978), to determine molecular weights of proteins.
Peptides also have been examined after digestion with cyanogen bromide, which cleaves
polypeptide chains at the amino acid methionine (Hedges & Wallace 1978, Tuross et al. 1980).
Furthermore, the collagen remaining may retain sensitivity to enzymatic cleavage with
collagenase or other proteases, which may liberate characteristic peptides for further study
(Armstrong et al. 1983, Rowley et al. 1986),

Using these methods, bones ranging in age from 200 years to >53,000 years have been
shown to contain collagen, although much of the collagen was degraded. Degraded collagen
was reflected both in the changes in the pattern of peptides seen after electrophoresis (Tuross et
al. 1980, Gurtler et al. 1981), and also in the increasing amount of soluble material found in
the older fossils (Hedges & Wallace 1978, Tuross ef al. 1980). It is likely that extensive
peptide cleavages occurred along the collagen molecules in the fossil bone samples even though
the collagen has apparently remained in situ in the bone. In these studies, the age of the bone
did not provide a good indicator of the amount or quality of preservation of the collagen, since
protein bands corresponding to alpha chains could be seen in the extract from the oldest bone
tested, a whale bone from Baffin Island, aged >53,000 years, but not in other much younger
bones (Tuross ef al. 1980). There may be an increased tendency for the remaining protein to
aggregate, reflected in the greater amount of protein which did not enter the gel during
electrophoresis on SDS-PAGE (Gurtler e¢ a/. 1981, Tuross et al. 1980).

These studies clearly show that biochemical investigation of fossil proteins by directly
isolating and sequencing individual peptides would be a formidable task. However, the use of
immunochemical techniques may overcome both the quantitative and qualitative problems
associated with collagen extraction from fossils. With suitable immunoassays, even minute
amounts (nanograms or picograms) of proteins can be detected, and even degraded proteins may
retain some of their original species-specific sequences and Lowenstein (1980, 1981) has
exploited such methods. Using a solid-phase radioimmunoassay (Fig. 2) and antibodies to
collagen raised in rabbits, he has shown that immunologically reactive collagen could be
detected in extracts of human fossil bones ranging in age up to 1.8 million years. The reaction
with these fossils was species specific, in that reactions with antibody to monkey, bovine or
rat collagens were persistently weaker than with antibody to human collagen.

BIOMOLECULAR INFORMATION IN BONE: OSTEOCALCIN

Although studies of the preservation of proteins in bone fossils have overwhelmingly been
studies of the preservation of collagen, about 1% of the total bone protein is osteocalcin.
Osteocalcin is a small molecule with a molecular weight of about 6000 daltons, containing a

318 - ROWLEY

high content of the amino acid gamma-carboxyglutamate. In several studies (King 1978a, b;
King & Bada, 1979) this amino acid has been shown to persist in fossils up to 50,000 years,
in the absence of leaching or weathering. More recently, using specific antibodies and a
sensitive radioimmunoassay, osteocalcin itself has been demonstrated (Huq et al. 1985,
Hauschka unpublished observations), and immunoreactive osteocalcin has been chemically
extracted from specimens of bone from the moa Pachyornis elephantopus, 4000-7000 years old
(Huq et al. 1985).

wks

Collagen binds Antibody binds Radioactive probe
to plastic. to collagen. binds to antibody.

Figure 2. Solid phase radioimmunoassay for collagen. Collagen binds firmly to the polyvinyl wells, and
antibodies bound to the collagen can be detected using a radioactive tag. The amount of radioactivity is
proportional to the amount of collagen present.

BIOMOLECULAR INFORMATION IN AUSTRALASIAN
FOSSILS.

The previous section has reviewed the published evidence for the preservation of proteins
and DNA in fossils. However, these studies have, almost without exception, originated from
the Northern Hemisphere, and nothing is known about the effect of the Australian
environment on the preservation of biomolecules. This section describes the results of studies
on the preservation of collagen in fossils from Australasia. These studies have been carried out
in the past 5 years within the Department of Earth Sciences at Monash University in
Melbourne, Australia. The methods used have been standard throughout, and are described in
detail, although they are similar to many of those used in the studies reviewed above.

ANALYTIC TECHNIQUES

Radioimmunoassay.

A solid-phase radioimmunoassay (RIA) for collagen, based on that of Lowenstein (1980)
was carried out on flexible polyvinyl microtitre plates (Fig. 2). Wells were coated with 50
microlitres (ul) of the sample to be tested and held overnight at 4° C in a moist chamber to
allow any protein present in the sample to bind to the plastic. The plates were then washed 3
times with phosphate buffered saline pH 7.3 (PBS) containing 1% skimmed milk powder and
0.05% Tween 20, and 6 times with distilled water, exposed to 200 ul of the wash solution for

BIOMOLECULAR INFORMATION IN FOSSILS - 319

2 hours at room temperature, to coat residual sites on the plastic, and again washed as above.
Fifty 11 of antibody to collagen was added to each well, and the plates were kept overnight at
4° C, then washed as before. Antibody bound to the plate was detected using a radioactively
labelled probe, which bound specifically to the antibody. Rabbit anti-collagen antibodies were
detected by adding 50 wl of protein A from Staphylococcus aureus, labelled with !251
containing 50,000 counts per minute (specific activity 40 microcuries/microgram). Sheep
antibodies, which do not bind protein A, were detected using 50,000 counts per minute (cpm)
of donkey anti-sheep antibodies (specific activity 10 pici/ugram). After the plates were kept
overnight, unbound radioactivity was washed away, and the radioactivity bound to the wells
was counted on a gamma counter. Under the conditions of the assay, the amount of
radioactivity bound to the plate was proportional to the amount of collagen bound to the plate.
Each sample was tested in quadruplicate, using anticollagen antibodies, and a corresponding
antiserum from an unimmunised animal to measure non-specific binding. In addition, each
serum was tested on uncoated wells of the plate, to determine the "background" binding
observed in the absence of fossils. Each sample was counted for ten minutes. The sensitivity
of this technique as measured using a standard curve of purified collagen was high, detecting
less than 0.1 g/ml, corresponding to | ng of collagen in 1 mg of bone powder.

Immunoblotting.

Similar in principle to the radioimmunoassay, immunoblotting provided a rapid and
convenient method of screening valuable fossils, since it required only a few milligrams of
bone. Many samples could be tested at once , handling was simple, and fossils which deserved
further investigations were easily recognised. Five microlitre samples were spotted onto
nitrocellulose, which provided the solid-phase support. After blocking, using PBS pH 7.3
containing 5% skimmed milk powder, the nitrocellulose was incubated with either rabbit anti-
collagen antibodies, or normal rabbit serum, and then !%°I-labelled protein A and
autoradiographed with x-ray film in the dark at -70° C. The presence of radioactivity, and hence
collagen in samples was recognised by the dark spot on the film (Fig. 3). Background binding
with rabbit serum was low, and after 2 weeks of autoradiography, the sensitivity was such that
0.4 ng of collagen applied to the filter could be detected.

X - ray film

Antibody binds Radioactive probe
to collagen. binds to antibody.

Collagen binds
to nitrocellulose.

Figure 3. Immunoblotting for collagen. Similar in principle, to the radioimmunoassay, the collagen is
bound to nitrocellulose membrane. After reaction with antibodies, and with the radioactive tag, the
radioactivity bound is detected on X-ray film.

320 - ROWLEY
Preparation of Fossils.

Fossils were ground to a fine powder, decalcified with 10 volumes of 0.5 M EDTA, pH
7,5, then re-extracted with 10 volumes of 0.5 M acetic acid. The remaining bone powder was
resuspended in 10 volumes of phosphate buffered saline (PBS) pH 7.3, for testing. Extractions
a mee out in siliconized glassware throughout, to minimize loss of protein on the sides
of the tubes,

Extraction and Characterization of Collagens.

The major collagen component of bone is type I collagen, which is also a major component
of skin. For this reason, modern collagens for comparison with fossil samples were prepared
from skin samples by pepsin digestion and purified by differential salt precipitation (Chung &
Miller 1974). Proteins present in samples were examined by SDS-polyacrylamide gel
electrophoresis under reducing conditions, according to the method of Laemmli (1970). After
electrophoresis, gels were stained using 0.1% Coomassie blue R-250, or by the silver/
Coomassie bluc double staining technique of Dzandu et al. (1984, 1985). After separation by
SDS-PAGE, collagens were electrotransferred to nitrocellulose for immunoblotting, which was
carried out as described above.

The physical form of the fossil collagens, whether retaining the native triple-helical
conformation, or denatured, was assessed in several ways. First, selected samples were reacted
with purified bacterial collagenase, which specifically degrades native collagen. Samples were
exposed to | mg/ml collagenase in 0.05 M Tris pH 7.5, 0.2 M NaCl, 0.002 M CaCl,
overnight at 37° C. Second, samples were tested in the radioimmunoassay by coating the
plates at 50° C, which denatured the collagen. As the rabbit antisera used reacted far more
strongly with native collagen, heating greatly reduced the binding of antibody to control
collagen. Third, selected samples were also tested by immunoblotting as described by
Ramshaw & Werkmeister (1988), under conditions in which collagen will retain the native
triple-helical conformation, Electrophoresis was carried out in a 3% gel, in the absence of
SDS, and both electrophoresis and clectrotransfer to nitrocellulose were carried out in 1% lactic
acid at pH 3.

Preparation of Antisera.

Rabbits were immunized sub-cutaneously with 5 mg of collagen in complete Freund's
adjuvant initially, and in PBS 4 weeks later, Booster injections of collagen in PBS were
repeated as necessary, to maintain high levels of antibody in the sera, Antibodies prepared in
rabbits reacted predominantly with native collagen, and minimally with denatured collagen.
Antibodies to denatured collagen appeared only after months of antigen exposure, and never to
high titre (Fig. 4A), Denatured collagen itself did not stimulate antibody production, even after
4 injections of heat-denatured collagen in adjuvant. Although a series of antisera were
produced, to collagens from various animals, one rabbit antiserum was used for the following
studies. This was chosen, because it contained the greatest reactivity to denatured collagen of
any antiserum tested. Although it reacted most strongly with chicken collagen, it gave 70-80%
of that reactivity with emu and other avian collagens, and less than 50% of that reactivity with
various mammalian collagens.

Additionally, as sheep produce antibodies to collagen which react similarly to both native
and denatured collagen, a widely reactive antiserum to avian collagens was prepared in a single
sheep immunised with | mg of a mixture of avian collagens in complete Freund's adjuvant,

BIOMOLECULAR INFORMATION IN FOSSILS - 321

followed by 3 subsequent injections of 1 mg of the collagen mixture in incomplete Freund's
adjuvant at fortnightly intervals (Fig. 4B).

EXTRACTION OF COLLAGEN FROM THE MUMMIFIED SKIN OF
A MOA.

Although moas were found throughout New Zealand at the time when the Maoris
arrived, about 1000 years ago, most, if not all, were extinct by the arrival of the first
Europeans. Much remains unknown about the living moas and their phylogenetic
relationships with other flightless birds. Recently, it has been possible to examine a piece of
naturally mummified moa skin for the presence of collagen. Collagen, detectable in this
specimen by immunofluorescence (Fig. 1), was extracted with acetic acid and pepsin for further
examination.

After separation by SDS-PAGE on a 10% gel, the bands observed did not correspond with
those of modern collagen. Much of the protein was present as a smear throughout the gel,
indicating considerable degradation (Fig. 5A). After transfer to nitrocellulose and
immunoblotting under denaturing conditions, some reactivity occurred with high molecular
weight material, but the staining was weak, and none of the protein bands stained specifically.
Using conditions of electroblotting and electrophoresis to retain native collagen, a single spot
of protein was observed, of slightly greater mobility than that of the collagen control (Fig.
5B). By radioimmunoassay, the pepsin digest of moa skin contained significant collagen
reactivity, which was destroyed by heat treatment at 45° C. These results indicated that the
collagen remaining in the moa skin retained considerable helical structure. Although
obviously severely degraded, as evidenced from the SDS-PAGE protein profile, the moa
collagen reacted specifically with antibodies, and the reactivity was inhibitible to varying
extents with collagens from other birds and animals (Fig. 6).

IMMUNOREACTIVE COLLAGEN IN BONE.

In an initial study (Rowley et al. 1986), avian fossils, usually Emu (Dromaius
novaehollandiae), from 9 separate Pleistocene sites, and ranging in age from at least 150 years
to about 2.5 million years were examined, using a rabbit antiserum to chicken collagen.
Kangaroo or wallaby fossils from 5 sites were tested for comparison. Both emu and wailaby
bones were obtained from 2 sites. Unidentified bone from Tom O's Quarry, a mid-Miocene site
in central Australia, thought to be approximately 10 million years old, was also tested. Details
of fossils and sites are shown in Table I. By dot immunoblotting, reactivity was detectable in
many of the fossils after treatment with antibodies to collagen, but not after treatment with
normal rabbit serum (Fig. 7). This reactivity was strongest in the most recent samples, but
the oldest sample tested also gave a weak, but definite signal. Reactivity was apparently
specific for collagen, as the reactivity was abolished by overnight incubation with collagenase.
Results obtained in the RIA were similar to those obtained by immunoblotting, and heat
treatment significantly reduced antibody binding to several of the fossils (Table 2). As the
rabbit antiserum reacted strongly with native collagen, but only weakly with denatured
collagen, if the collagen detected in the fossils retained its tertiary structure, heat treatment
should cause a drop in reactivity. However, if the collagen was already denatured, heat
treatment should cause very little change in the amount of radioactivity bound. In the fossils

322 - ROWLEY

>

RABBIT ANTISERUM

100,000

10,000

1000

100

Radioactivity Bound (CPM)

100 1000 10,000
Serum Dilution

38

SHEEP ANTISERUM

100,000
=
Oo
O 10,000 OO 0 a
E one
1000 Sen
ps ry
=
iS 100 @® Native Collagen
2 © Denatured Collagen
©
oO

100 1000 10,000 100,000
Serum Dilution

Figure 4. Levels of antibodies to native collagen, and heat denatured collagen in serum tested at varying
dilutions. In this assay, the amount of radioactivity is proportional to the amount of antibody present. A,
antibodies in the serum of a rabbit immunized with native collagen react much more strongly with native
collagen than with denatured collagen; B, antibodies in the serum of a sheep immunized with native collagen
react similarly with both native and denatured collagen.

BIOMOLECULAR INFORMATION IN FOSSILS - 323

Figure 5. Polyacrylamide gel electrophoresis and Wester blotting of the Moa collagen extract. A,
electrophoresis on 10% SDS-PAGE, stained with Coomassie blue. 1. Moa, pepsin extract. 2. Emu collagen
control, 10 micrograms; B, Western blot of proteins separated on a 3% gel for native collagen. After
autoradiography for 24 hours there was strong reaction with both Moa collagen (1) and the collagen control
(2) and the background radioactivity was very low.

treated there was not only a decrease in total antibody bound with increasing age, but a relative
increase with age in the amount of denatured collagen in the fossil.

IMMUNOREACTIVE OSTEOCALCIN IN BONE.

Several of the fossils tested for collagen, were also tested for osteocalcin (Hauschka &
Rowley, unpublished data). These included emu bone samples, and the unidentified mid-
Miocene bone from Tom O's Quarry. A sample of modern Emu bone was included for
comparison. Each sample was tested in a competitive radioimmunoassay, using an antibody to
chicken osteocalcin, raised in a rabbit. Although there was not good cross reactivity between
chicken and emu osteocalcins, and modern Emu gave only 1.3 % of the reactivity expected with
fresh chicken bone, osteocalcin was detected also in those bones which contained substantial

324 - ROWLEY

collagen (Table 3). As with collagen, osteocalcin was clearly demonstrable in bones aged up to
10,000 years.

Table 1: Fossils examined for the presence of collagen. Bone from emu (e), unidentified bird
(b), kangaroo (k), or wallaby (w), and unidentified bone (b) were tested. Sites represented a
variety of environments: dune (D), ignimbrite (I), fluviatile (F), lacustrine (L), cave (C), or
swamp (S). Sites were dated by l4c (*) or by relative data with associated fauna (**).

Sample Site Environment Age
le King Island D >150
2b Tower Hill I 6-10,000
3e Kings Creek 2F?L 23-41,000*
4e Wombeyan Quarry Cave Cc >18,000*
5e Callabonna L *>36,000**
6e Lancefield South S ~26,000**
Tu Tom O's Quarry F 10,000,000
Se Strathdownie Cave Cc Pleistocene**
De McEachern's Cave Cc 2-28,000*
10e Lancefield S 26,000*
11k Lancefield S 26,000*
12k Foul Air Cave Cc Pleistocene**
13k Lake Colongulac L Pleistocene**
14w King Island D >150
15k Wyandotte Creek F 2,000,000

EFFECT OF ENVIRONMENT ON COLLAGEN SURVIVAL IN BONE

Factors which affect the preservation of collagen are not understood, but the ability to
predict which samples are most likely to contain collagen would reduce sample destruction and
preparation time during phylogenctic studies of fossil species using collagen. For this reason,
a study was carried out to test the effect of environment on the preservation of collagen in
fossils from a range of Australian sites. Specimens were obtained from museum collections,
and to avoid destruction of valuable specimens, "scrap" bone was obtained from each site, as
our previous study (Rowley et al. 1986) had shown that bone scrap may contain significant
amounts of collagen. Wherever possible, multiple samples were tested, to test variability
within a single environment, Sites tested ranged in age from a few thousand years back to the
mid-Miocene, and from a range of environments, including caves, alluvial deposits, central
Australian salt lakes, swamps, etc. (Table 4).

Of the 39 sites tested, one or more samples from 15 of them tested positively for collagen.
As noted previously by Wyckoff, (1972), not all samples from any one site contained collagen.
Thus, some samples were completely negative, while another from the same site reacted quite
strongly with the antibody. There was poor correlation between the age of the fossils and the
amount of collagen detected. Bones which apparently contained collagen were obtained from
several Miocene sites around Lake Palankarinna, but bones {rom the Wellington Caves, carbon
dated at about 2,500 years did not contain detectable collagen.

BIOMOLECULAR INFORMATION IN FOSSILS - 325

Table 2: Heat treatment of fossils tested for collagen in the radioimmunoassay. The rabbit
antiserum used in this study reacted most strongly with conformational epitopes. Thus, heat
treatment significantly reduced binding to purified Emu collagen. If the collagen detected in the
fossils was already denatured, heat treatment should cause very little change in the amount of
radioactivity bound. In the five fossils tested, there was not only a decrease in total antibody
bound with increasing age, but a relative increase with age in the amount of denatured collagen in
the fossils. Sample numbers in this Table refer to those in Table 1.

Sample Age (years) Counts Bound
Before heating After heating Ratio
Enu 0.5 pg/ml 0 133,000 5100 26
Collagen
Fossil le >150 104,000 4700 23
Collagen 2b 6-10,000 9100 2100 4.3
10e 26,000 2100 1500 1.4
12k Pleistocene 890 850 1.1
14w >150 7000 3600 1.9

SS SBhBhnDhnDnDDDS=E_—_—_—_————

In this study, as in the first study (Rowley et al. 1986), caves generally seemed to provide a
poor environment for the preservation of collagen, as bones were tested from 10 cave sites, and
of these, 8 did not contain detectable collagen. However, most of the material examined for
collagen had been from caves with abundant moisture, and many of the bones examined were
powdery and easily ground. A second study (Baird & Rowley 1990) was, therefore, undertaken
to see whether, when well-preserved bones were selected, the cave environment could be shown
to influence the survival of collagen. The caves from which material was studied included
Clogg's Cave and McEachern's Cave, Victoria, Koonalda Cave, South Australia, and Madura
Cave and Devil's Lair, Western Australia. Clogg's Cave, McEachern's Cave and Devil's Lair
represented caves with moist environments, and Madura and Koonalda caves occur in dry
environments. The bones studied were mostly well-preserved humeri from Turnix species
(button-quail), and the primary accumulator was the Barn Owl (Tyto alba) for all except

—==—_=<=_ =_—hhmi"_—“<—— OO ESESESaEaE=EE_—_— eae

Table 3: Osteocalcin retention in fossil avian bone. Bone powder was extracted in 0.5M
EDTA overnight at 4° C. and the supernatant was analysed for osteocalcin in a
radioimmunoassay with a chicken antiserum. Sample numbers refer to specimens listed in Table
1. Results are expressed as a percentage of the osteocalcin detected in a sample of modern Emu
bone.

Sample Age (years) % Osteocalcin
Relative to Modern
Emu 0 100
le >150 1.38
2b 6-10,000 2.38
Emu >150 1.81
4e >18,000 0
Se >36,000 0
8e Pleistocene 0
10e 26,000 0

ee SS0000$0$@mS _—=

326 - ROWLEY

McEachern's Cave, which appeared to be of fluvial/pitfall origin. Bones ranged in age from
8,720+230 to 37,880+3520 years, as determined by dating using 14C. In this study, however,
samples from both "wet" and "dry" caves contained significant residual collagen: one sample
aged 18,990+220 years contained 92% of the collagen detected in a modern bone sample.
Thus, the humidity of the caves did not appear to be the critical factor in the preservation of
collagen. In contrast to our previous studies, however, these avian bones had been extremely
well-preserved, with an undamaged surface and minimal breakage. Such good survival of small
bones, each weighing less than 1 g, suggests that the microenvironment may have been
particularly favourable. It should be noted that good bone quality is not always a reliable
indicator of collagen preservation. The only sample tested which did not contain significant
collagen was also rated as very well preserved.

LOSS OF COLLAGEN IN FRESH BONE IMMEDIATELY AFTER DEATH.

The previous study of the preservation of collagen in fossils of various ages, and in varying
environments, had indicated that neither the age of the fossil, nor its appearance, was a good
guide to the preservation of collagen. Wyckoff (1972) suggested that the amino acids
remaining in fossils may reflect the changes which took place immediately after the death of
the animal. For this reason, the collagen remaining in a series of modern bones which have
undergone decomposition under natural conditions, have been examined. The bones selected
were from Puffinus tenuirostris (Short-tailed Shearwater), and were collected from among the
sand dunes behind the beach at Forrest caves, Phillip Island, in Victoria. The sand dunes along
this stretch of coast line are a favourite nesting site, and as a result of predation from feral cats
and foxes, skeletal remains are common. Ulnae from seven different birds were examined,
selected to represent a range of stages of preservation. The best preserved specimen, from an
articulated skeleton buried in the sand at high tide level, was taken to represent the collagen
found in fresh bone. Other samples had been subject to varying degrees of weathering, as
indicated by bleaching, cracking and growth of algae on the surface. Details are shown in Table
5. In these modern bones, as in the fossil bones, the amount of collagen remaining varied,
ranging from 11% to 94% of that in the best specimen. Here also, the appearance of the bone
was a poor predictor of collagen preservation, in that the bone containing 94% of the collagen
of fresh bone was bleached, cracked and algae covered. Several of these bones contained less
collagen than those of the fossils of Turnix varia from caves up to 32,000 years old.

EXTRACTION OF COLLAGEN FROM MIOCENE BONE.

Although immunoreactive "collagen" has been detected in avian and mammalian fossils, up
to 10 million years old, the reactivity detected has not been directly demonstrated to be due to
the presence of intact collagenous protein. Reactivity with these very old fossils has been
weak, and has not been convincingly removed by collagenase, or altered by heat treatment. It
is likely that such old collagen would have undergone considerable changes and react quite
differently from modern material. However, it is also possible that such weak reactivity may
be no more than artefactual, non-specific antibody binding. To demonstrate that collagen may
indeed be preserved for millions of years in fossils, proteins in fossil bone from two Miocene
sites were extracted (Rowley et al., in prep.) ,

The samples tested were selected solely on the availability of sufficient bone for analysis.
Sample 1 was 60 g of turtle bone from Lake Namba in South Australia, believed to be about
15 million years old. These bones were collected from the dry bed of the lake, and washed free
of surrounding matrix. Sample 2 was 120 g of long bone, probably dromornithid, from
Bullock Creek in the Northern Territory. This bone was encased in limestone, and had to be

BIOMOLECULAR INFORMATION IN FOSSILS - 327

extracted manually before testing. The bones were ground to a powder, and extensively dialysed
against 0.5 M EDTA pH 7.5 and 0.5 M acctic acid (Sample 1) or against 0.5 M acetic acid
(Sample 2) to solubilise the collagen and remove the limestone. The supernatants were
concentrated, and examined for reactivity to collagen in the RIA, and directly by polyacrylamide
gel electrophoresis.

Before extraction, neither of the samples contained significant collagen, measured as a
three-fold excess of reactivity with antibody to collagen over reactivity measured using a
normal rabbit serum control. After concentration, only Sample 2, from Bullock Creek showed
reactivity with the antibody (Table 6 ). By polyacrylamide gel electrophoresis under reducing
conditions, both samples contained high molecular weight material which did not enter the
stacking gel, and low molecular weight material which migrated with the dye front. However,
in sample 2, faint bands were identified with similar mobility to the bands of molecular
weights 300,000 and 200,000 observed witha modern rhea collagen control (Fig. 8). In

100
80

60

40

20

PERCENT INHIBITION

1 10 100 1000
INHIBITOR (ng)

Figure 6. Inhibition of the reaction between moa collagen and sheep antiserum to avian collagen, using
purified collagens. A solid phase radioimmunoassay was carried out in which plates coated with moa
collagen, 20 pg/ml overnight at 4° C were reacted with a sheep antiserum to avian collagen, at a standard
dilution of 1:50,000, and with varying amounts of collagen as inhibitor. Percent inhibition was obtained by
comparison of the amount of radioactivity bound in the presence of inhibitor and the amount of radioactivity
bound without inhibitor. Linear regression lines are shown for each inhibitor, calculated using the computer
programme Sigmaplot (Jandel Scientific, Sausalito, California). Inhibitors used were collagens from moa (M),
ostrich (OQ), rhea (R), emu (E), cassowary (CA), and chicken (CH). As expected, the reaction was most
strongly inhibited by increasing concentrations of moa collagen, but it was also inhibited by ostrich, rhea,
and emu collagens. In this assay, chicken and cassowary collagens were not significantly inhibitory.

328 - ROWLEY

Figure 7. Fossils tested for collagen by immunoblotting on nitrocellulose membrane, after 2 weeks
autoradiography. Fifteen fossils (F) and Emu collagen control solutions (C) were tested on each filter.
Control solutions were tested in doubling dilutions from 10 pg/ml. Filter 1 (A) was reacted with rabbit anti-
chicken collagen. Filter 2 (B) was reacted with normal rabbit serum. Reactivity with normal rabbit serum may
reflect the presence of bacterial contaminants in the fossil sample. With the exception of fossil 10, the
reaction with normal rabbit serum was much less than with immune serum, and developed later during
autoradiography.

BIOMOLECULAR INFORMATION IN FOSSILS - 329
i _______...____ eee

Table 4: Fossils from 49 sites were tested for the presence of collagen. Several samples of
bone were tested from each site. Mammalian bones were tested unless otherwise noted.
Environments included dune (D), fluviatile (F), lacustrine (L), cave (C), swamp (S) or marine (M).
The presence of collagen was assessed by RIA using a rabbit anti-collagen antiserum or normal
rabbit serum as a control. Samples were tested in quadruplicate, and were considered positive if
they showed both a statistically significant difference (P<0.05) and at least a three-fold increase
in counts with anti-collagen antibodies over control counts.

Number Site Environment Age Samples Samples
tested positive
1. King Island D 2 2
2. Goulden's Hole Cave Cc Holocene 3 PA
3. Wellington Caves Cc 2,550 2 0
Cathedral Cave
4. King Island 10940+160 2 0
Seton Site K6/5
2: Wellington Caves Cc 14,300+730 3 0
Cathedral Cave
6. King Island C 2 0
Emu Caves
Dempsey's Lake L Late Pleist 1 0
7. Hookina Creek Late Pleist. 2 0
8. Baldina Creek Late Pleist. 2 0
9. Henschke's Cave Cc ~30,000 5 0
10. Henschke's Cave Cc >40,000 5 0
11. Couman Forest Cave LS Pleistocene 7 3
12. Goulden's Hole Cave Cc Pleistocene 4 0
13. Curramulka Town Cave Cc Pleistocene 3 10)
14. Morgan A Pleistocene 3 0
15. Lake Callabonna L >40,000 4 4
16. Port Pirie Pleistocene 3 1
17. Oakley Creek, Weetalibah A Pleistocene 3 1
18. Salt Creek, Normanville F Pleistocene 3 0
19. Tambar Springs A Pleistocene 3 1
20. Lake Kanunka L Plio-Pleist 5 3
Katipiri Sands
21. Brothers Island, Coffin Bay M Pleistocene 1 0
22 Calca Pleistocene 1 0
23. Curramulka Quarry Pleistocene 1 0
24. Wellington Caves Cc Early-mid 3 0
"The bone cave" Pleistocene
25. Nelson Bay M 0.94-1.74 Myr 6 1
26. Chinchilla A Plio.-Pleist. 4 1
ZF. Curramulka, Cc Pliocene ?older 4 0
Cora Lynn Cave
28. Sunlands, Waikerie M Early Pliocene 4 3
29. Dog Rocks, Geelong Pliocene >2Myr 5 0
30. Lake Palankarinna L Pliocene 7 1
Woodard Quarry
31. Alcoota Mio./Pliocene 8 i)
3.2%. Lake Ngapakaldi L M-L Pliocene 5 0
Leaf Locality
33. Lake Palankarinna L Mid-Miocene* 11 3

Mammalon Hill

330 - ROWLEY

Table 4: (Continued)

Number __ Site Environment Age Samples Samples
tested positive

34. Lake Palankarinna 16 Mid-Miocene* 7 3
Tedford Locality

3D Lake Palankarinna ju, Mid-Miocene* 6 3
Croc Pot

36. Tom O's Quarry F Mid-Miocene* 1 0

37. Lake Namba L Mid-Miocene* 6 0

38. Riversleigh Mid-Miocene* 1 0
Pancake Site

39. Bullock Creek Mid-Miocene* 6 0

*May be Late Oligocene - Middle Miocene

Table 5: Loss of collagen in fresh bone subject to natural decomposition and weathering
immediately after death. Ulnae from 7 specimens of Puffinus tenuirostris were examined. Results
were expressed as a percentage of collagen found in the best preserved specimen, for which the
skeleton was still articulated.

No. Description Percent
Collagen
ie Excellent preservation, fully articulated. 100%
2. Bone with tendons attached.
Algae in marrow cavity 57%
3. Excellent preservation 53%
4. Bleached, cracked, algae in marrow cavity. 94%
Ds Bleached, algae covered. 29%
6. Bleached, cracked. 51%
Rs Very cracked, algae covered 11%

very faint bands were identified, of 100,000 molecular weight, corresponding to the 100,000
bands of the control, although these bands could not be reproduced photographically. No such
bands were visible with Sample 1. Acetic acid alone, concentrated similarly, did not contain
any measureable protein.

APPLICATIONS

Using albumin, it has been possible to measure immunological distances between the
mammoth, modern Indian and African elephants, and the Sea Cow (Prager et al. 1980), while
the relationships between the mastodon and the mammoth have been examined using both
collagen and albumin (Shoshani et al. 1985). Immunological comparisons have also been

BIOMOLECULAR INFORMATION IN FOSSILS - 331

Table 6: Samples were tested for collagen in the RIA. Results are expressed as mean + 1
standard deviation, of counts per minute of radioactivity bound. Before concentration, neither
sample reacted significantly with the anti-collagen antibody, as defined by three-fold greater
reactivity with anti-collagen antibody than with normal rabbit serum. After concentration, the
sample from Bullock Creek became positive (*). N.D. = Not done.

Site Sample Anti-collagen NRS
Lake Namba Before Concentration 1,166 + 43 623 +42
Concentrate 1,056 + 272 517 +18
Filtrate N.D. N.D.
Bullock Creek Before Concentration 1,124 + 382 642 +145
Concentrate 3,674 + 573* 1,180 +169
Filtrate 974 + 44 567 +24
__/ —— resend a)
aaa
Seer
94—
67—
43—
30—
20

Figure 8. Electrophoresis of collagen extracted from fossils, on 10% SDS-PAGE stained with both
Coomassie blue and silver. A, Lake Namba concentrate; B, Bullock Creek concentrate; C, Modem collagen
control, 10 pg. High molecular weight material, which did not enter the stacking gel, and low molecular

weight material, running with the dye front were found in both fossils. The sample from Bullock Creek also
showed bands with the mobility of the 200 KD and 300 KD collagen bands.

332 - ROWLEY

used to derive a phylogeny based on albumin for the Thylacine, or Tasmanian Wolf
(Lowenstein et al. 1981) and for the extinct Steller's Sea Cow (Rainey et al. 1984), while the
protein-based phylogeny of the Quagga (Lowenstein & Ryder 1985) agreed very closely with
that derived from DNA (Higuchi et al. 1984).

Techniques for identifying proteins have uses other than for creating molecular phylogenies.
They have also been used to identify museum specimens of doubtful authenticity. For
example, when the skull and lower jaw, presumed to be that of Piltdown man, was shown to
be a forgery, consisting of a human skull, and a suitably "aged" jaw from a Chimpanzee or an
Orangutan, the exact origin of the jaw was hotly contested. Using radioimmunoassay,
Lowenstein et al. 1982, was able to show that the jaw, and the canine tooth found nearby, were
those of an Orangutan. Similarly, radioimmunoassay using albumin and serum proteins, has
been used to confirm that two shrunken heads in the British Museum were, indeed of human
origin, and not forgeries made from horse skin (Lowenstein 1985).

CONCLUSIONS

This review has examined the extent of the biomolecular information available from fossils,
but has not considered the question of whether molecular or morphological data are more useful
for the determination of phylogenetic relationships. A recent review by Hillis (1987) has
examined the two approaches, setting out the advantages of each technique, and the regions of
conflict. No single technique is applicable to every problem, and morphological techniques
will continue to be applied to a huge range of museum and fossil material. Nevertheless,
appropriate molecular comparisons are very powerful tools, which may be tailored for
application to a wide range of problems. Thus, protein and DNA sequencing have been used to
compare enzymes from organisms as widely different as man and bacteria (Griffin et al. 1988),
and DNA analysis and hybridization has been used for "molecular fingerprinting" to identify
individuals and to determine paternity (Jeffreys et al. 1985a, b). At present the application of
techniques derived from molecular biology to palaeontology is hampered by the small amount
of protein or DNA remaining in most fossils, but the field is rapidly evolving, and newer,
more sensitive techniques are constantly becoming available. In particular, the development of
the polymerase chain reaction, in which specific short regions of a gene can be greatly
amplified in vitro from as little as a single molecule of DNA (Saiki et al. 1985, Mullis &
Faloona 1987) has provided a means whereby biological samples whose DNA content is too
low, or too degraded for direct analysis may be amenable to genetic analysis after
amplification. The sensitivity is such that, using the polymerase chain reaction, the DNA
from a single hair may be analysed (Higuchi et al. 1988). Paabo (1989) has demonstrated the
use of the polymerase chain reaction to amplify a specific fossil DNA sequence in the presence
of a vast excess of heavily modified DNA. Moreover, the ability to selectively amplify regions
of the DNA, will allow the selective examination of fossil DNA in the presence of large
amounts of contaminating bacterial DNA.

ACKNOWLEDGEMENTS

I would like to thank Dr. T. H. Rich, Mr. N. Pledge, Dr. K. McNamara and Dr. L. Dawson
for providing fossil samples for analysis; Drs. P. V. Rich and R. F. Baird for helpful
discussions, and Associate Professor P. Hauschka for permission to use unpublished data in
this review. The work was supported by grants from the Australian Research Grants Scheme,
and I was the recipient of a National Research Fellowship.

BIOMOLECULAR INFORMATION IN FOSSILS - 333

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336 - ROWLEY

Fossil fish from the Late Devonian upland streams and lakes of central Victoria. Bothriolepis
cullodenensis (above) and the phyllolepid (below) are two placoderms that are characteristic of
this time period in Australia, and are useful biostratigraphic tools. (From Rich & van Tets 1985,
with permission of The Museum of Victoria).

CHAPTER 12

THE LONG HISTORY OF
AUSTRALIAN FOSSIL FISHES

John Long!
Introductions nc eteetshcslesthy sesso eect 338
Major Collections of Australian
Bish®ehOssils es fecte nce ecceece ucts seeeet- 338
Systematic ReVICW............cccccccccesesceceeececeeeees 343
Agnathane = Une, eke tee 343
Gnathostomata - Jawed Vertebrates.............. 345
Chondrichthyes ................seeeeeeeeeeeeeeeee 345
ACAnthOdany s2i8. .0.0..28edoet neceetsibereceteneees 349
Placoderminss..M0 058 a 8 352
Arthrodires.............cccsccesceecesseecees 354
AMIUIATCHS fos. sds ds ee telc. coestescdee vee s 362
Petalichthyids................ceecseeeeeeees 364
Acanthothoracids................cceeeeeees 367
Ptyctodontids ...............ceeeeeceeeee eee 367
Osteichthyes .............cccceecececenceceeeeeeees 367
ACUNOPICTYBiL.......... cece cee eeeeeeeeeees 368
Devomian................2cececececeeeees 368
CarboniferouS.............2.sceceeneee 370
PEEMPAN See ii os kes leer dee nevcexs 371
“PRASSICME us feet BREA oh ote woe 374
UTASSIC eee OL ot eecle tetas 376
CretaCeOuS ..........ceceeeeeeceeeeceees 377
TST ary-see abee hoe FA icc ebdetc a tee Bee 382
Crossopterygians ............0ceeeeeeeeeee 386
Dipnot, he och BA ele 3 391
Australian Palaeozoic Fish Biostratigraphy
and Biogeography..............cccccsseeceeeceeeceeees 397
References Suh siiec. stb etlcs tulesieetiesel steseestebad 403
Appendix I: Classification ...............eseeeeeeee eee 414
Appendix II: Abbreviations Used
AEP US ULC Stage oe nc sitewide odine sete con ists nue deswseene rd 419
PlateSme 1 ake cle, Mate vies teccretetin Shea tositod 420

1 The Western Australian Museum, Francis St., Perth, Western Australia 6000, Australia.

338 - LONG

INTRODUCTION

In this paper the Australian fossil fish record (Fig. 1) is extensively reviewed and more
amply illustrated than my earlier review (Long 1982a), with both stratigraphic and systematic
cross referencing. In this work groups are treated systematically in order from Agnatha to
Dipnoi, as outlined in the section on classification below. A classification of fishes is given
in Appendix 1.

Depositional environments containing fossil fishes fall into two categories: undoubtedly
marine, and all others. Marine deposits in the Palaeozoic are easily identified by lithology
(usually limestones or limey shales) and accompanying marine invertebrate faunas. Redbed
deposits of Late Devonian-Carboniferous age contain fishes but lack marine invertebrates, and
because of associated terrestrial plant fossils, have been designated terrestrial deposits. The Mt
Howitt site in Victoria contains fish preserved in finely varved black shales, and occurs
stratigraphically in a sequence of sandstones and conglomerates. For this reason, and the lack
of invertebrate fauna, it has been identified as a lake deposit. European workers dealing with
black shales of marine origin, and red bed facies thought to be marine or littoral deposits have
suggested that some Australian Devonian fish deposits thought to be terrestrial could
alternatively be marine, and, for this reason, it is necessary for more detailed sedimentological
studies to be undertaken at some of these sites. References treating the depositional
environments of Australia Palaeozoic fish sites include Marsden (1976) and Long (1982b) for
Victoria (Late Devonian and Carboniferous), Connolly (1965), Connolly et al. (1969),
Fergusson et al.(1979) and Powell (1983) for New South Wales (Late Devonian), and Olgers
(1972) for Queensland (Late Devonian and Carboniferous). Cas (1983) summarizes the facies
distribution for the Palaeozoic of south-eastern Australia and includes reference to sites
containing fish. Most Mesozoic and Tertiary deposits are marine fossiliferous limestones or
shales (Cretaceous of Queensland, Tertiary of Victoria, South Australia and Western Australia),
lacustrine diatomites (Tertiary of New South Wales and Queensland), lacustrine shales
(Cretaceous of Victoria) or coaliferous shales in thick fluviatile or deltaic sediments (Permo-
Triassic of New South Wales and Queensland). Some references treating depositional
environments of Australian Mesozoic fish sites are Waldman (1971a), White (1981) and Banks
et al. (1978); and for Tertiary sites are Hills (1943a,b, Gill (1957) and Abele (1976).

A complete listing of Australian fossil fishes with comprehensive bibliography can be
found in Long & Turner (1984). Other reviews of Australian fish groups are Turner et al.
(1981) and Turner (1982d) for thelodonts, Turner (1982a) for Palaeozoic Chondrichthyes, Kemp
(1982) for Tertiary Chondrichthyes and Long (1984a) for placoderms. Turner (1982c) has listed
fossil fishes in Queensland, including specimens in the Queensland Museum, but,
unfortunately, this task has not been completed for any other major collection.

Fig. 1 shows a rough classification of the major fish groups and their current record from
Australia, and Fig. 2 details some of the main fossil fish localities in Australia. The
significance of Australia's major fish faunas is indicated with their distribution in time in Fig.
3. For students a glossary of some palacoichthyological terms can be found in Long (1982a).

MAJOR COLLECTIONS OF AUSTRALIAN FISH FOSSILS

A requirement necessary for understanding fossil fish are good collections. The major
collections of Australian fish fossils in this country are in the following institutions.

LONG HISTORY OF FOSSIL FISH - 339

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The Australian fossil fish record. Black portions of the columns represent the known occurrence

of the group in Australia.

Figure 1.

340 - LONG

The Australian Museum, Sydney. Staff includes Dr Alex Ritchie and Mr R. Jones.
Collections include material from New South Wales; the Antarctic Devonian; Gogo, Western
Australia; Mt Howitt, Victoria; Ordovician of the Northern Territory; Permian from
Queensland; diverse Triassic and Jurassic faunas of New South Wales; and many specimens
from foreign locales such as Scottish Devonian and Silurian fishes of Europe.

The Bureau of Mineral Resources, Canberra. Staff includes Dr Gavin Young.
Collections include material from the New South Wales south coast Devonian; the Antarctic
Devonian; the Devonian of central Australia, Queensland and Western Australia; and superb
specimens from the Taemas-Wee Jasper Early Devonian.

The Geology Department, Australian National University, Canberra. Staff
includes Prof. Ken Campbell and Dr R.E. Barwick (Zoology Department). Collections include
Early Devonian Taemas lungfishes, Late Devonian Gogo and New South Wales lungfishes;
some other Gogo material; some Queensland Carboniferous and Permian fishes; and a
comparative collection emphasizing lungfishes but including other foreign specimens.

The Museum of Victoria, Melbourne. Staff includes Dr Tom Rich. Victorian
collections include Late Devonian material from Mt Howitt; Early Devonian fishes from
Buchan; Carboniferous fishes from Mansfield; Cretaceous fishes from Koonwarra; and Tertiary
sharks' teeth and fishes from throughout the State. Some overseas material, especially
Devonian from Scotland, is also held in this collection.

The Queensland Museum. Brisbane. Staff includes Dr Mary Wade and Honourary
Research Fellows Dr Susan Turner and Dr Anne Kemp. Collections include Queensland
Devonian-Carboniferous faunas; Cretaceous and Palaeogene fishes from Queensland; Triassic-
Recent lungfish toothplates and Palaeozoic microvertebrate assemblages from all over
Australia.

The South Australian Museum, Adelaide. Staff includes Mr Neville Pledge.
Collections include Tertiary sharks’ teeth; and well preserved Morgan Limestone Miocene
fishes.

The Western Australian Museum, Perth. Staff includes Dr Ken McNamara and
Dr. John Long. Collections include the type specimens, and much additional material of Gogo
fishes; some Cretaceous and Tertiary Chondrichthyes from Western Australia.

Figure 2. Australian fossil fish localities: m, marine; f, freshwater. 1, Gascoyne River region (m, Pemn.);
2, Gneudna Fm. (m, Dev.); 3, 4, Gogo Fm. (m, Dev.); 5, Billiluna (f, Dev.); 6, Hargreaves Hills (m, Dev.); 7,
Spirit Hill No. 1 Well (m, Dey.); 8, Dulcie Range (f, Dev.); 9, Mt Charlotte (m, Ord.); 10, Mithaka
Waterhole (f, Dev.); 11, Toomba Range (f, Dev.); 12, Hughenden region (m, Cret.); 13, Gilberton (f, Dev.);
14, Broken River region (m, Dev.); 15, Blackwater (f, Perm.); 16, Springsure (f, Carb.); 17, Redbank Plains
(f, Tert.); 18, Knocklofty Fm, (f, Trias.); 19, Leigh Creek (f, Trias); 20, Lake Eyre (m, Cret.); 21, Gosses
Bluff (f, Dev.); 22, Dare Plain (f, Dev.); 23, Mootwingee (f, Dev.); 24, Mt Jack (f, Dev.); 25, Jack's
Lookout (f, Dev.); 26, Mt Grenfell Station (f, Dev.); 27, Mt Deerina (f, Dev.); 28, Wuttagoona Station (f,
Dev.); 29, Bulgoo Station (f, Dev.); 30, Walgett (F, Cret.); 31, Warrumbungles (F, Tert.); 32, Talbragar (f,
Jur.); 33, Redcliff Mt, Grenfell (f, Dev.); 34, Newcastle (f, Trias.); 35, Gosford (f, Trias.); 36, Brookvale
(f, Trias.); 37, St Peters (F, Trias.); 38, Canowindra (F, Dev.); 39, Jemalong Gap (F, Dev.); 40, Taemas-
Wee Jasper (m, Dev.); 41, Braidwood (f, Dev.); 43, Eden (f, Dev.); 44, Genoa River (f, Dev.); 45,
Combienbar River (f, Dev.); 46, Orbost (m, Tert.); 47, Buchan (m, Dev.); 48, Mt Tambo (f, Dev.); 49,
Jemmy Point (m, Tert.); 50, Freestone Creek (f, Dev.); 51, Mt Howitt, Bindaree Rd. (f, Dev.); 52, Tatong (,
Dev.); 53, Koonwarra (f, Cret.); 54, Mansfield Basin, including So. Blue Range (f, Dev.); 55, Taggerty (f,
Dey.); 56, Balcombe Bay (m, Tert.); 57, Beaumaris (m, Tert.); 58, Janjuc (m, Tert.); 59, Otway Region,
Lome (f, Cret.); 60, Batesford, Waurn Ponds (m, Tert.); 61, Grampians (m, Dev.); 62, Hamilton (m, Tert.);
63, Carapook (f, Cret.); 64, Upper Murray Cliffs (m, Tert.), Many Tertiary sharks' teeth and other fish
remains have also been found along the southern coast of Australia (Nullarbor Plain Cliffs) and in the
Carnarvon Basin, Wester Australia. Some localities in Queensland were discovered after the figure was
completed (referred to in the text and in Kemp's chapter (15) in this volume. These are not shown on this
map.

LONG HISTORY OF FOSSIL FISH - 341

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342 - LONG

LOCALITY

Grange Bum (62)
Beaumaris (57)
Balcombes Bay (56)
Wurrumbungles (31)
Janjuc (58)
Redbank Plains (17)
Walgett (30)
Richmond area (12)
Koonwarra (53)
Carapook (63)
Talbragar (12)
St Peters (37)
Brookvale (36)
Gosford (35)
Knocklofty Fm. (18)
Gascoyne River (1)
Blackwater (15)
Newcastle (34)
Springsure (16)
Mansfield (54)
Worange Point (43)
Grenfell (33)
Canowindra (38)
Jemalong (39)
Dare Plain (22)
Freestone Creek (50)
Eden (43)
Mt Howitt (51)
Braidwood (41)
Taggerty (55)
Gogo Fm. (3 , 4)
Gneudna Fm. (2)

\~ Grampians (61)
Tatong (52)
Bunga Beach (42)
Broken River (14)

Hatchery Creek (40)

Mulga Downs Gp (23-29)

A= Peak Beds (10,11)
Buchan (47)

Taemas (40)
Mt Chanrtotte (9)

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ENVIRONMENT

ZFFESFEMmmNrnVNFZFEAFZFFmmrAMAMAMAMAUMAAUMAAAAAEs AAAANt NDE mMmT7MFE NT ZZFS

REMARKS

Edaphodon - youngest occurrence
isolated bones, otoliths, sharks’ teeth
sharks’ teeth, otoliths
entire fish
shark's teeth, otoliths
entire fish
Ceratodus
articulated skulls
entire fish; last palaeoniscid
entire fish
entire fish; earliest teleost Leptolepis
holostean dominated
entire fish, well preserved

except for skull regions
cosmopolitan taxa only
bradyodont teeth
entire fish, including sharks
Urosthenes
fragmentary remains
fauna currently under revision >8spp.
Remigolepis, Groenlandaspis only
sinolepid present
entire fish
okest amphibian bony remains
fragmentary remains
several horizons yield fossils
fragmentary remains
entire, well preserved fish >18spp.
undescribed fauna
first Late Devonian fauna described
3-D preservation, diverse fauna >34spp.
youngest thelodonts in the world
elasmobranch spines, denticles
undescribed fauna
fragmentary remains
undescribed fauna
equally okdest Bothriolepis in the world
Wuttagoonaspis - Turinia

faunas
3-D preservation, fish rare
3-D preservation, diverse fauna
oldest Southem Hemisphere vertebrates

|very diverse faunas

Figure 3. Stratigraphic sequence and significance of major fish faunas of Australia. Locality numbers refer

to Fig. 2. F, freshwater; M, marine.

LONG HISTORY OF FOSSIL FISH - 343

. The Tasmanian Museum, Hobart. Staff includes Mr. Noel Kemp. Collections
anes many T ertiary sharks’ teeth from throughout Australia, and Triassic fish faunas from
asmania.

SYSTEMATIC REVIEW

AGNATHA

The Agnatha are classically divided into two major groups, the Cephalaspidomorphi and
Pteraspidomorphi (Moy-Thomas & Miles 1971), although recent workers are doubting the
monophyly of these groups, and discoveries of unusual agnathans from the Early Devonian of
China (Liu 1965, P'an Jiang 1984) indicate that this simple division cannot be maintained
(Janvier 1981, 1985, Forey 1984, Maisey 1986). The five monophyletic groups which are now
recognized are the Thelodonti, the Anaspida, the Osteostraci, the Heterostraci and the
Galeaspida.

Osteostracans (included within the "cephalaspidomorphs") are characterized by having a
single opening in the cranium for the nares, termed a nasohypophysial foramen. The head of
Osteostraci and Galeaspida (a closely related group from China) is covered by a broad, flat
shield made up of one piece of bone with holes for the eyes and nares. To date no osteostracans
have been discovered in Australian Palaeozoic rocks, but there are two new genera of agnathans
from the Middle Devonian of the Amadeus Basin which may be related to the Galeaspida of
China (Dr. G. Young, pers. comm). This absence is not thought to be due to the
incompleteness of the fossil record but to the restriction of osteostracans to the European and
the American landmass ("Euramerica", see Young 1981a for biogeographic provinces) and
galeaspidans to the South China Province (and possibly East Gondwana -Australia) during the
Early Palaeozoic (Young 1981a, 1987c, 19874).

The Heterostraci are characterized by separated and paired nasal openings and the shield is
composed of several plates, which fit around the head and trunk region. Australia has some of
the earliest complete heterostracans in the world, from the Middle Ordovician, but after this
occurrence there is no younger record of the group on this continent. These are the
Arandaspidiformes, a primitive order known only from the Middle Ordovician Stairway
Sandstone, south of Alice Springs in the Northern Territory. Ritchie & Gilbert-Tomlinson
(1977) described two genera: Arandaspis (Fig. 4), known from almost complete armour and
trunk scales, and the poorly known Porophoraspis with a different type of surface ornament.
Dr A. Ritchie (1985) has indicated that up to four taxa of Arandaspidiformes may be present in
the Stairway Sandstone fauna.

The thelodonts, small scale-covered agnathans (see Turner, Chap. 13, this volume), are
poorly known, represented mostly by scale assemblages. They are thought to be closely related
to the ancestral line of the jawed vertebrates or Gnathostomata (Janvier 1981) or, alternatively,
thought to be a sister group to another extinct Northern Hemisphere agnathan group, the
Anaspida (Forey 1984), or to the Chondrichthyes (Turner 1985).

Heterostracans are a major part of Silurian-Middle Devonian faunas in the Northern
Hemisphere, with great diversity of forms that are utilized in biostratigraphic correlation (¢.g.
Blieck 1984, Blieck et a/.1987). Until recently the Arandaspidiformes represented the earliest
near-complete remains of vertebrates, because older fishes were known only from incomplete
pieces of bone (Nitecki 1978, Repetski 1978). They are the oldest known vertebrates from the
Southern Hemisphere. Arandaspis (Fig. 4A-C) has two simple large plates, a dorsal shield and
a ventral shield, which are separated laterally by a series of rectangular branchial plates,
reflecting that the number of paired gill openings was up to 15. This arrangement is perhaps

344 - LONG

the simplest pattern of heterostracan armour known and presumably the most primitive
(although see Elliott 1987). As the material is studied from latex casts made from natural
moulds where the bone had weathered away, there is no information on the histology of the
tissues. More generalized discussion of its morphology can be found in Ritchie (1985). Recent
finds of complete Upper Ordovician vertebrates from South America by Drs Gagnier and
Janvier of Paris should shed light on the relationships of the Arandaspidiformes.

Thelodonts are rarely preserved intact (Ritchie 1968, Turner 1982f). Half of the known
genera are based on scales. Each thelodont scale has an ornamented dentinous crown around an
acellular bony base with a pulp cavity (see Turner this volume for more details). Taxa are
described from different scale types and scale associations as the individual scales vary from
different body positions. Thus body scales of one form might be long with high crowns,
whereas the head scales of the same species could be short with a flat crown. In addition, there
are various special scales (e.g. around the gill slits) and transitional scales between differing
body regions. Care must be exercised in thelodont scale taxonomy with emphasis placed not
only on morphology but also on histology (thin sectioning is needed). Despite this,
thelodonts are very useful in biostratigraphic correlations particularly across the Silurian-
Devonian boundary, and in Australia throughout the Devonian (Turner & Dring 1981, Turner
et al. 1981, Turner 1982d, 1982e, 1986, Long et al.1988).

trunk
A branchial scales

dorsal shield plates

mouth

ventral
shield

Figure 4. Agnatha (Middle Ordovician, Northem Territory). Arandaspidiformes. Arandaspis prionotolepis
Ritchie & Gilbert-Tomlinson 1977. A, attempted restoration showing main feature; B, detail of dermal
omament; C, trunk scale, x2. (After Ritchie & Gilbert-Tomlinson 1977).

Thelodonts were first reported from Australia by Peter Jones (in Johnstone et al. 1967) and
unrecognized scales were figured from Western Australia by Seddon (1969). Later a detailed
study of Western Australian scales of Early Devonian age was published by Gross (1971) in
which a new species, Turinia australiensis was named. Turner & Dring (1981) described the
first endemic thelodont genus, Australolepis, from the Late Devonian Gneudna Formation,
Western Australia, making it possibly the youngest known member of the group world-wide
(recent re-assessment of the Turinia hutkensis fauna from Iran also indicates a Frasnian age for
these thelodonts according to Dr P. Janvier).

Finally, an unusual body fossil belonging to an agnathan has been identified from the
Emsian-Eifelian Wuttagoonaspis fauna in the Georgina Basin, Queensland (G. Young, pers.
comm.). The specimen is a complete shield with a long rostrum and may be related to the

galeaspids.

LONG HISTORY OF FOSSIL FISH - 345

GNATHOSTOMATA - JAWED VERTEBRATES

The superclass Gnathostomata includes all jawed vertebrates, that is all fish (except
agnathans) and all amphibians, reptiles, mammals and birds. Within the piscine division of the
gnathostomes there are some early groups whose relationships are uncertain, For convenience I
will treat the major gnathostome groups in the following order: Chondrichthyes (sharks, rays
and chimaerids), Placodermi (extinct armour-covered fishes), Acanthodii (spiny shark-like
fishes) and Osteichthyes (true bony fishes, including ray-finned and lobe-finned fishes).
Interrelationships of these groups have been treated recently by Schaeffer (1975, 1981), Jarvik
(1980), Janvier (1981, 1984), Goujet (1982), Gardiner (1984a, 1984b) and Maisey (1986).
These represent strongly differing views of early gnathostome phylogeny.

The gnathostomes are usually divided into two major groups (Elasmobranchiomorphii -
shark like fishes, placoderms and sharks; Teleosteomi - bony fishes, osteichthyans and
acanthodians) based on the presence or absence of an internal swim-bladder. Most living fishes,
such as the salmon, trout and goldfish are osteichthyans. Although internal organs are rarely
preserved in fossils the shape of the body and fins reflect how the fish produced its lift whilst in
the water. Development of large, wing-like pectoral fins and strongly heterocercal tails in
sharks, results from the lack of a swim-bladder, because lift is provided principally from the
thrust of the tail and broad aerofoil-sectioned fin shapes, particularly the wide pectorals
(Alexander 1967, Aleev 1969). Similar body form in the extinct placoderms suggests that
they, too, lacked a swim-bladder, although some forms may have possessed air sacs as
accessory respiratory organs (e.g. Bothriolepis, Denison 1941).

The acanthodians, an extinct group, have characteristics of both sharks and osteichthyans,
and their phylogenetic position is still hotly debated (Nelson 1968, Miles 1973, Jarvik 1977,
1980, Maisey 1986). Advanced acanthodians, such as the acanthodiforms, appear to have
essentially osteichthyan-like fin disposition and tail shape, suggestive of the presence of an
internal swim-bladder (Miles 1973). Maisey (1986) has recently suggested that the respiratory
mechanism of acanthodians was essentially like that of osteichthyans, based on hemibranch
morphology. For these reasons, they are probably closer to the osteichthyans, perhaps the
sister group of the bony fishes, than they are to sharks or placoderms, as once thought (e.g.
Watson, 1937).

Chondrichthyes

Fossil sharks and rays have long been known from the Tertiary and Mesozoic of Australia
(McCoy 1876; Etheridge 1888; De Koninck 1898; Chapman & Pritchard 1904, 1907), but
extensive discoveries of early chondrichthyans from the Devonian and Carboniferous have been
made only within the last decade. Some early records of Palaeozoic sharks (e.g. De Koninck
1898) have to be treated with care, as we now know much more about their affinities and tooth
variations in single species. Chondrichthyans are characterized by having a cartilaginous
skeleton with the only "hard" parts being the teeth, scales, fin-spines and calcified cartilage. As
such, their fossil record is mostly known from these parts with very rare preservation of
cartilage parts being confined to certain types of quiet, anaerobic depositional environments.
Sediments of this nature are very rare in Australia, with only four sites yielding articulated and
semi-articulated shark cartilage fossils. These are the Bunga Beds (Middle Devonian, New
South Wales), the Utting Calcarenite (Carboniferous, northern Western Australia), The
Blackwater Shales (Permian, Queensland) and near Sydney and Gosford (Sydney Basin, Triassic,
New South Wales). Despite the paucity of articulated specimens from these localities, they are
of great importance world-wide, as the Bunga material includes the oldest shark braincase
fossils known (Young 1982), and the material from the Sydney Basin represents one of the

346 - LONG

youngest occurrences of pleuracanth sharks. Recently Dr Alex Ritchie, of the Australian
Museum, has made exciting new discoveries of Triassic articulated sharks from Gosford, north
of Sydney. This new material appears to be of unusual pleuracanth sharks which lack the neck-
spine found in regular pleuracanths (Fig. 7).

prc.f.

Figure 5. Chondrichthyes (Middle Devonian, New South Wales). A, C-F, Antarctilamna prisca Young 1982:
A, restoration of braincase; C, tooth; D, fin-spine; E, scale; F, denticle; B, Xenacanthus sp., tooth. After
Young (1982). See Appendix IL for abbreviations used with figures.

The Chondrichthyes can be conveniently subdivided into two main groups: Elasmobranchii
(sharks and rays) and Holocephali (chimaeras, rabbitfishes), although there is uncertainty about
the systematic position of many fossil forms, such as "bradyodonts". Sharks and rays are
essentially carnivorous (eating other fishes), although rays are mostly durophagous (crushing
hard-shelled invertebrates) or microphagous (filter feeding on plankton or krill). The
elasmobranchs have five or more pairs of open gill slits and amphistylic jaw suspension
(hyomandibular braces the upper and lower jaw articulation) with well developed teeth.
Chimaerids adopted a durophagous life-style early in the Palaeozoic. They have an opercular
cover to the gill chamber and, amongst other specializations, have upper jaw cartilages fused to
the braincase and possess crushing tooth plates. Both groups have small placoid scales set in
the skin, and the males possess clasping organs for reproduction.

Australia's oldest sharks are from the Early Devonian limestones of New South Wales,
being represented by scales as yet not fully investigated ("Skamolepis"; Giffin 1980, Turner
pers. comm., 1982). Teeth of Mcmurdodus whitei were recently described from Early-Middle
Devonian limestones of the Cravens Peak Beds, Toko Syncline (Queensland) by Turner &
Young (1987). These teeth may have belonged to an early shark of the extant hexanchid

LONG HISTORY OF FOSSIL FISH - 347

group, and could represent one of the oldest members of the advanced living shark groups
(neoselachians). This can only be confirmed, however, by future study of the enamcloid
structure (Reif 1977).

One species is present in the Middle Devonian Bunga Beds Fauna, Antarctilamna prisca
(Fig. 5), which is represented by jaws, scales (Fig. 5E) and teeth (Fig. 5C), in addition to
neurocrania (Fig. 5A). Antarctilamna is thought to be a primitive xenacanth by Young (1982).
It is biostratigraphically important that this species is also known from the Devonian of
Antarctica, where it occurs with Mcmurdodus? featherensis (White 1968, Young 1982). It is
unusual that the Bunga Beds do not contain placoderms, which are prevalent in nearly all types
of Devonian sediment, but abound in shark remains, along with other rarer fossils including
acanthodians and osteichthyan bones. An extensive range of sharks teeth, including
holocephalans, has been described by Turner (1982a, 1982c, 1983) from Upper Devonian-
Lower Carboniferous deposits of Queensland and Western Australia. The fauna from the
Broken River embayment includes the species Harpagodens (Thrinacodus) ferox (Fig. 6A, B),
which, as the Latin name suggests, has teeth like small grappling hooks. This genus has
recently turned up in the Uppermost Devonian beds of Ghuizou Province, China (Wang &
Turmer 1985) and in northern Thailand (Long 1990c), and is known from the Carboniferous of
Europe and North America. Other sharks from Queensland and New South Wales are a new
species of Phoebodus (Long 1990c, Long & Burrett 1989, Turner 1982a; Fig. 6C) and
Stethacanthus ("Cladodus") thomasi (Fig. 6D; Turner 1982a), although the most characteristic
form is undoubtedly Ageleodus pectinatus which has a long root and splayed crown with four
or five broad separate cusps. This genus is also known from the Carboniferous of Europe,
North America and the U.S.S.R. Carboniferous sharks were earlier recorded from the Laurel
Formation, near Fitzroy Crossing, Western Australia, by Thomas (1957, 1959). Recent
collecting from the Canning and Bonaparte basins, north Western Australia, by the author has
produced a number of new fossil sharks! teeth, including species of Helodus and Orodus and
several unidentified types.

The Permian of Western Australia has produced tooth whorls of the edestid shark
Helicoprion as well as isolated "bradyodont" type teeth (Teichert 1940, 1943). Edestid sharks
are well known in Permian deposits throughout the world and are useful biostratigraphically.
The unusual lower jaw tooth whorls probably folded into a pouch below the jaw symphysis as
new teeth grew out (Fig. 6F). Bradyodonts are early holocephalans whose relationships are
not clear, and as they are known largely from teeth which are of similar form (flat crushing
pavement teeth), they may represent a paraphyletic collection of taxa. Recently an edestid
shark was discovered in the Upper Permian Blackwater shales of Queensland. It is currently
being studied at Macquarie University (Mr Mike Leu, pers. comm.). The Blackwater shales
have also produced a complete, new ctenacanthoid shark, Surcaudalus (Leu 1989).

Mesozoic chondrichthyans are known from the Triassic of New South Wales and the
Cretaceous of Queensland, South Australia and Western Australia. Australia's most complete
chondrichthyan fossils are large, almost complete pleuracanth sharks from the Lower Triassic,
St Peters fauna, near Sydney (Woodward 1908). Pleuracanth or xenacanth sharks have bicuspid
teeth with a median prominence on the oral surface of the base, and advanced forms usually
possess a large spine protruding from behind the skull (Fig. 7). Pleuracanthus parvidens
(Woodward 1908) requires further study in the light of Dr Ritchie's new discoveries from
Gosford, New South Wales, The Cretaceous Rolling Downs Group of Queensland has produced
a number of isolated sharks’ teeth, including Scapanorhynchus and Lamna species (Etheridge
1888, Jack & Etheridge 1892, Hill er al. 1968). Australia has two Mesozoic chimaerids
represented by single specimens only recently described. Edaphodon eyrensis from the Aptian
Bulldog Shale, near Marree, South Australia, represents one of the earliest known occurrences
of this genus, which occurs most abundantly in the Late Cretaceous (Long 1985b).
Ptykoptychion is an endemic genus from the Early Cretaceous (Albian) of Queensland (Lees

348 - LONG

Figure 6. Chondrichthyes (Upper Devonian-Permian). A-B, teeth of Harpagodens (Thinacodus) ferox Turner
1982a, (A-E all Upper Devonian-Lower Carboniferous); C, Protacrodus tooth; D, Stethacanthus (Cladodus)
thomasi Turner 1982a tooth; E, Phoebodus sp. tooth; F, restored lower jaw tooth whorl of Helicoprion,
similar to H. davisii from the Permian of Western Australia. A-E after Tumer (1982a); F, after Bendix-

Almgreen (1966). B-E, approx. x 1 mm.

LONG HISTORY OF FOSSIL FISH - 349

1987). Another chimaerid lower jaw is known from the Cretaceous Molecap Greensand, near
Gin Gin, Western Australia, which has some resemblances to Edaphodon eyrensis .

Tertiary chondrichthyans are common fossils in marine deposits all over the southern half
of Australia. Kemp (1982) (and this volume (Chap. 15)) reviews the taxa present, and lists
synonymies. Early workers greatly confused identifications of shark teeth by not taking into
account the positions of teeth in the jaw. Some recent finds which are not included in Kemp's
(1982) chapter are Palaeocene teeth from the Boongerooda Greensand, Giralia Anticline,
Western Australia. These teeth, collected in 1986, include a possible new species of
Hexanchus as well as Odontaspis and Otodus teeth. Microscopic remains of hybodontid sharks
(scales and teeth) and actinopterygians have recently been recovered from drill cores in the off-
shore Carnarvon Basin, Western Australia. These are of Middle Triassic age, and represent
Australia's only marine Triassic fish fauna.

neck spine

pelvic fin

pectoral fin

Figure 7. Chondrichthyes (Triassic). Reconstruction of a typical pleuracanth shark., x0.2.. A similar form
ocurrs in the Triassic of New South Wales. (After Burian & Augusta 1965).

Holocephalans are known from three species in the middle Tertiary of Victoria and
Tasmania: Edaphodon sweeti (Miocene-Pliocene, Victoria), E. mirabilis (Miocene, Victoria)
and Ischyodus newtoni (Oligocene, Tasmania), all described by Chapman & Pritchard (1907)
and Chapman & Cudemore (1924). Long (1985b) gives a new restoration of the mandibular
tooth of E. sweeti. E. mirabilis may not be a separate species but just an unusually large
variation of E. sweeti (N. Kemp., pers. comm).

Fossil stingrays are represented by two species of the cosmopolitan genus Myliobatis from
the mid-Tertiary of Victoria and Tasmania (Chapman & Pritchard 1904, 1907, Chapman &
Cudemore 1924). Although both these species were described from toothplates (crushing
combs), remains of tail stingers are also commonly found, especially from Beaumaris,
Victoria.

Acanthodii

Acanthodians are a poorly known group as a whole. Most of our knowledge of acanthodian
anatomy is based on the structure of the youngest and most specialized genus, Acanthodes,
from the Permian of Germany. Acanthodians are characterized by having fin-spines preceding

350 - LONG

all the fins except for the tail. The body is covered by small scales, which vary in shape and
histology, and the head of some primitive forms may have some dermal bones. Three higher
groups are recognized: Climatiida, which contains specialized forms with bony armour on the
shoulder girdle and often on the head; Ischnacanthida, which have teeth ankylosed to gnathal
bones in the jaws, and Acanthodida, which have only one dorsal fin and lack teeth, apparently
being specialized for microphagy or filter feeding. Australia's record of acanthodians is very
poor. Only two complete species have been described, although partially articulated remains of
a few other genera are known. A great number of acanthodian scales are being recovered from
mid-Palaeozoic marine deposits around Australia, and these include the oldest gnathostome
fossils, of Silurian age, from Australia (Turner & Pickett 1982).

dorsal fins
(ds)

ventral f-

pectoral pelvic aa ’
armour / fieepiné
(vpa) pectoral fin Fa (pel)
spine ait
(pfs)
B

Figure 8. Acanthodii, Climatiida (middle Palaeozoic). A, attempted reconstruction of Gyracanthides murrayi
Woodward 1906 (Lower Carboniferous, Mansfield, Victoria; B, Culmacanthus stewarti Long 1983b (Late
Devonian, Mt Howitt, Victoria.).

Climatiids are known from isolated scales and fin-spines in the Devonian of south-eastern
Australia (Young & Gorter 1981, Turner 1982c, d). Nostolepis striata (Pl. 1C) is commonly

LONG HISTORY OF FOSSIL FISH - 351

found from Early Devonian deposits (e.g. Philip 1965), and recently cf. Cheiracanthoides (PI.
1B) has been turning up from Middle Devonian marine limestones (Young et al. 1987). Two
groups derived from the main stock of climatiids - diplacanthids and gyracanthids - are known
from more complete material. Culmacanthus stewarti (Pl. 1D-E, Fig. 8B) was described from
articulated fossils of Late Devonian age from Mt. Howitt, Victoria (Long 1983b; restoration in
life habitat in Long 1983d), It is a deep-bodied diplacanthoid with large cheek plates bearing
strongly ribbed and spinose dermal ornament (PI. 1E). The snout is well preserved, being the
second acanthodian to have this region described, the other being Triazeugacanthus of similar
age from Canada (Miles 1966). Culmacanthid check plates are also known from Freestone
Creck in Victoria, from the Boyd Volcanic Complex (Pambula River), near Eden, New South
Wales, and from the Devonian of Antarctica (Young 1989a), suggesting the genus could have
been widespread in the Late Devonian of East Gondwana. Gyracanthides murrayi from the
Lower Carboniferous Mansfield Group, Victoria (Woodward 1906a; Fig. 8A) is relatively
completely preserved, except for the head. Resemblances to the European Carboniferous form
Gyracanthus led Woodward to date the Mansfield fossils, along with other fish he thought were
congeneric with Northern Hemisphere forms. Gyracanthides and Gyracanthus have very large,
robust pectoral fin-spines with an elaborate chevron pattern of ridges. Isolated Gyracanthides
spines have also been described from the Late Devonian of Antarctica (White 1968) and South
Africa (Chaloner et al. 1980). Isolated spines of Gyracanthides are also known from the Lower
Carboniferous rocks of the Bogantungan region of Queensland (Hills 1958).

Ischnacanthids were until recently almost unknown from Australia. Discoveries of
ischnacanthid gnathal bones by E. White and H. Toombs (British Museum Nat. Hist.) in the
1950's from Taemas, New South Wales, were described recently by Long (1986a). These
specimens (PI, 1F), along with another of similar Early Devonian age, from Buchan, Victoria
(Pl. 1G), represent the first definite record of this group from Australia. Otherwise only scales
attributed to ischnacanthids were recorded (Turner e¢ a/. 1981, Turner & Pickett 1982). The
Early Devonian jaws from Tacmas and Buchan belong to two endemic genera, Taemasacanthus
and Rockycampacanthus, both of which have complex arrangement of secondary cusp rows, and
a well developed median tooth row. The major significance of Taemasacanthus is in the
preservation of the mandibular joint, which differs from that of acanthodids and climatids.
This morphologic character incited a review of acanthodian interrelationships, which suggested
that ischnacanthids were the plesiomorphic sister group to climatiids and acanthodids (Long
1986a). Another ischnacanthid, known from isolated jaw bones, comes from the Upper
Devonian Hunter Siltstone, near Grenfell, New South Wales, but this genus has not yet been
described. Articulated small acanthodians from the Middle Devonian Bunga Beds, New South
Wales, may be ischnacanthids, although details of the jaws are missing from the specimens.
They have two dorsal fins and scem to lack the heavy armour of climatiids (Fergusson ef al,
1979).

Acanthodids were first recorded from Australia by Woodward (1906a), who nominated two
new species from the Lower Carboniferous Mansfield Group, Victoria, as Acanthodes australis
and Eupleurogmus creswelli. Acanthodes australis is represented by an almost complete fish,
although, as in Gyracanthides from the same fauna, the head is missing. It is clearly an
acanthodid, but cannot be reliably assigned to any genus and_ should be referred to as ef.
Acanthodes sp. Eupleurogmus was established on an incomplete section of acanthodian trunk,
which has larger scales flanking the lateral line. A similar arrangement of body scales is seen
around the ventrolateral sensory-line of Acanthodes bridgei (Zidck 1976), and it is likely that
Eupleurogmus is a nomen nudum. The best preserved acanthodids in Australia come from the
Upper Devonian Mt Howitt site (Long 1986b). //owittacanthus kentoni (Pl. 1A, Fig. 9) is a
moderate to small-sized acanthodid which closely resembles Protogonacanthus from the Middle
Late Devonian of Germany, differing in slight details such as the ossification of the jaw
cartilages and ornamentation of fin-spines. Howittacanthus is probably more closely related to

352 - LONG

Acanthodes (Long 1986b). Recently a new fish fauna was recovered by acetic-acid preparation
from the basal Carboniferous Raymond Formation, near Clermont, Queensland. This fauna
includes three-dimensional bones of acanthodids including the jaws, basisphenoid,
scapulocoracoids and fin-spines. The basisphenoid differs slightly from that of Acanthodes, the
only other known acanthodian with the ventral part of the braincase preserved. No Permian
acanthodians have yet been found in Australia - perhaps because suitable deposits have not yet
been sampled.

palatoquadrate
dorsal fin

scapular

Meckel's
caudal fin carttage

anal fin pelvic fins pectoral fins
(fin-spines)

Figure 9. Acanthodii, Acanthodida (Late Devonian). Sketch of a complete specimen of Howittacanthus
kentoni (Long 1986b), from Mt Howitt, Victoria.

Placodermi

Placoderms were armoured fishes which flourished from the Late Silurian to the start of the
Carboniferous, being the dominant vertebrates of the Devonian seas, rivers and lakes. The head
and trunk were covered by bony shields of overlapping dermal plates, which in most forms
were hinged by a condyle and trochlear joint at the neck. Some workers have placed placoderms
phyletically close to the Chondrichthyes (@rvig 1961; Stensié 1963, 1969; Maisey 1986) but
alternatively, they may be the sister group to Osteichthyes (Forey 1981, Gardiner 1984b).
Goujet (1984) and Schaeffer (1975, 1981) regard placoderms as phyletically equidistant to both
these groups. Young (1986) has recently reviewed the problem of placoderm affinities and
concludes there is no clear answer given the available data. The heavily armoured shields of
placoderms were amenable to fossilization and our Australian record of the group is
extraordinary. Some of the best placoderm material in the world comes from the Early
Devonian deposits at Taemas (New South Wales) and Buchan (Victoria) and from the Upper
Devonian Gogo Formation, Western Australia. Long (1984a) has reviewed placoderm
morphology with emphasis on the Australian record of the group. Here, the main placoderm
faunas of Australia, with emphasis on their stratigraphic and anatomical significance, will be

reviewed.

LONG HISTORY OF FOSSIL FISH - 353

nasal capsules ————=—==_ >=. ENDOCRANIUM

antorbital
olfactory bulb process
pituitary vien supraorbital
process

jugular vien

anterior,

y . posterior

ay
: Os Th postorbital
S& process

sacculus

cerebral
cavity

spino-occipital

nerves cucullaris

fossa

craniospinal
process

Figure 10. Placodermi (Early Devonian). A, B. Buchanosteus confertituberculatus,x 1 (Chapman 1916);
A, headshield, dorsal view; B, endocranium and cavities for soft tissues. (After Young 1979).

354 - LONG

_ Placoderms can be divided into five or six main groups according to phylogenetic
interpretation. The most successful group, forming about 60% of the known record, are the
arthrodires ("euarthrodires"), characterized by having a full complement of trunkshield plates and
a well developed dermal neck joint. Long (1984c) included phyllolepids, an unusual depressed
group of placoderms within the arthrodires, although Young (1984b, 1984c) regards
phyllolepids as a separate group. Antiarchs were bizarre placoderms with long trunkshields and
peculiar oar-like pectoral appendages. Their headshields had a single opening for the eyes and
nostrils, and the trunkshield was long and box-like. They were widespread in Middle and Late
Devonian times and are amongst the most common fossils of this age in freshwater deposits.
Acanthothoracids, petalichthyids and ptyctodontids are comparatively rare as fossils and are
characterized by having short trunkshields with unique patterns of headshield bones and sensory-
line canals (see Moy-Thomas & Miles 1971, Denison 1978 or Long 1984a for details of their
structure and reference to family names used below). Recent discussion of interrelationships
can be found in Denison (1978, 1983), Miles & Young (1977), Young (1979, 1980, 1981a, b,
1984a, b, 1986, 1988b), Goujet (1982, 1984a), Janvier & Pan (1982), Dennis & Miles (1983),
Long (1983a, 1984c, 1987b) and Gardiner (1984a). With many new discoveries on the
horzou, this pattern of differing phylogenetic opinions amongst workers will probably
continue!

Arthrodires

Several Early Devonian arthrodires have been described from the Taemas/Wee Jasper region
of New South Wales (Murrumbidgee Group) with others known from contemporaneous strata
at Buchan, Victoria. Perhaps the best known local placoderm of this age is Buchanosteus
confertituberculatus (Pls 2A-B, Figs 10, 12) from both these localities (Chapman 1916, Hills
1936b, Stensio 1945, White 1952, White & Toombs 1972, Young 1979, 1986).
Buchanosteus possessed a separate dermal rostral capsule in the snout, a primitive character for
arthrodires, but was quite advanced in the trunkshield being shortened. The braincase and
cranial anatomy were described in great detail by Young (1979) with discussion of their
implications to arthrodire phylogeny. Buchanosteid placoderms have recently been discovered
in the Northern Hemisphere (Dr E. Mark-Kurik, pers. comm., Goujet & Janvier 1984).
Taemasosteus (Fig. 11C-D), a large genus in the same fauna, is known by two species, T.
novaustrocambricus from both New South Wales and Victoria (White 1952, 1978, Long
1984c) and T. maclartiensis from Buchan, Victoria (Long 1984c). Taemasosteus, known from
abundant material, was more advanced than Buchanosteus in having a reduced trunkshield with a
posteriorly open pectoral incision, a condition which arose independently several times within
the Arthrodira (Denison 1983). Close relatives of Taemasosteus have been described from the
Emsian of Germany (Gross 1960) and recently from similar age in Morocco (LeLievre 1984a,
1984b). Other arthrodires from these Australian localities which are less completely known
but at a similar grade of organization as Buchanosteus, are Arenipiscis westolli (Fig. 11A, H),
Errolosteus goodradigbeensis and Toombsosteus denisoni (White 1978, Young 1981b, Long
1984c). Goodradigbeeon australianum (Fig. 11F, I) and Burrinjucosteus asymmetricus (White
1978) (Fig. 11B) are also known from the fauna but are of uncertain affinities. Williamsaspis
bedfordi (White 1952) (Fig. 12A, B) is the only arthrodire from Taemas which could be an
arctolepid. There are still new forms to be described from this fauna, some of which could be
homosteids (Dr G. Young, pers. comm.).

Placoderms of latest Early to earliest Middle Devonian age (Emsian-Eifelian) are also
known from several localities in the Mulga Downs Group (western New South Wales) and the
Georgina Basin (western Queensland). This fauna is dominated by the unusual endemic
actinolepid-like form Wuttagoonaspis fletcheri (Fig. 12H-I, 13B; Ritchie 1969, 1973), which
has a long headshield with small orbits and an unusual pattern of bones in the cheek area. The

LONG HISTORY OF FOSSIL FISH - 355

trunkshield of this species has a high crested Median Dorsal and varies in the degree of
development of a dermal neck joint (Dr A. Ritchie, pers. comm.). The distinctive meandering
ridge ornament caused Rade (1964) to mistake it for a phyllolepid. Occurring with
Wuttagoonaspis are new forms of actinolepids, primitive groenlandaspids (Ritchie 1975) and
larger arthrodires, which could be homosteids.

Middle Devonian placoderms are scarce in the Australian fossil record. The only arthrodires
to be described are from the Eifelian Hatchery Creek Conglomerate, New South Wales,
although others under study by Dr Gavin Young from the Broken River Embayment,
Queensland, will extend the list. Denisonosteus (Fig. 13A) is a small phlyctaenioid from the
Hatchery Creek fauna which has a long, strongly waisted nuchal plate. It is not unlike other
phlyctaeniids of this age from the United States or Europe (Young & Gorter 1981).
Indeterminate euarthrodire plates occur with Denisonosteus in the same fauna. Amongst the
Broken River placoderms (Fish Hill, Givetian) are several large brachythoracids including
Ailantidosteus (Dr G. Young, pers. comm.), also known from the Emsian of Africa (LeLievre
1984b).

A diverse fauna of over 20 species of Late Devonian arthrodires is known from the Gogo
Formation, in the north of Western Australia (Gardiner & Miles 1975, Long 1984a, 19874,
1988b, 1988c, 1988d, 1988f). This material consists of three-dimensional armours of
articulated fish and probably represents the best material of placoderms from anywhere in the
world. Most of the Gogo arthrodires are eubrachythoracids at the coccosteomorph grade of
organization, although some pachyosteomorphs, which have a trunkshield incised for the
pectoral fin, occur with them. Typical coccosteomorphs such as I/arrytoombsia (Fig. 14 E;
Miles & Dennis 1979) and several new undescribed forms from Gogo differ from the typical
Scottish , North American and Russian coccosteids in lacking a marginal-postorbital contact
with the central plate. This feature they share with the Frasnian Plourdosteus. All the
coccosteid-like arthrodires from Gogo are endemic genera. Bullerichthys, Kendrickichthys and
Bruntonichthys (Dennis & Miles 1980) were pachyosteomorphs that have jaws adapted for
crushing shelled invertebrates which were plentiful in the warm reef seas they inhabited.
Kendrickichthys may belong to the Mylostomatidae, a group known also from North America
and Africa (Young 1987c). Camuropiscis, Fallacosteus, Rolfosteus (Figs 14A, 15C) and
Tubonasus (Dennis & Miles 1981, Long 1983c, 1988f, in press 2) were streamlined and had
well produced rostra. These last three genera belong to the endemic family Camuropiscidae
and share other specializations, such as a reduced and strongly interconnected posterior cheek
unit and Anterior Lateral plates, which contact the Anterior Ventrolaterals and durophagous
dentition. Recently new forms of camuropiscids have been discovered (Long 1987d, 1988b, c,
d). Other arthrodires from Gogo include a snub-nosed form, Simosteus (Dennis & Miles
1982), large plourdosteids such as Kimberleyichthys spp. and other undescribed forms (e.g.
Fig. 15C; Gardiner & Miles 1975, Dennis-Bryan & Miles 1983, Long 1987c, 1988c).
Pachyosteomorph arthrodires are represented by Incisoscutum ritchie (Fig. 14D; Dennis &
Miles 1982), a form closely allied to the camuropiscids (Long 1988b) and the primitive
dinichthyid Eastmanosteus calliaspis (Pls 3C, 4A, Fig. 15A; Gardiner & Miles 1975, Dennis-
Bryan 1987). Eastmanosteus is widespread in the Frasnian, being known from North America,
Iran and Russia (Denison 1978). The fauna from Gogo contains nearly all endemic forms, with
the only cosmopolitan genera of euarthrodires being Eastmanosteus and the holonematid
Holonema, represented by the local species //. westolli (Fig. 15B; Miles 1971). Holonema
possessed strange concave toothplates with strengthened parallel ridging, possibly for sifting
food from the sediment or crushing a particular type of invertebrate, No large forms have been
found from Gogo unlike contemporary faunas from the Late Devonian of Morocco and North
America, (e.g. Dunkleosteus), which included the largest of the placoderms, reaching up to
about 5 or 6 m in length (Denison 1978). Recently a large placoderm skull roof was
discovered in the collections of the University of Western Australia, collected many years ago

356 - LONG

in the Canning Basin by Curt Teichert. Hills (1958, p. 90) refers to this specimen as a large
dinichthyid. The estimated size of the fish, Westralichthys uwagadensis (Long 1987b) would
have been about 3 m, and it is regarded as a dinichthyid phyletically close to Dunkleosteus.
Conodonts found with the skull indicate it is of mid-Famennian age, considerably younger than
the Gogo fishes (Long 1987e). Other discoveries of Famennian vertebrates from the Canning

Figure 11. Placodermi (Early Devonian). Euarthrodires from the Taemas district, New South Wales A, H,
Arenipiscis westolli Young 1981 headshield restored in dorsal (A) and ventral (H) views; B, Burrinjucosteus
asymmetricus White 1978, headshield; C, D, Taemasosteus novaustrocambricus White 1952; C, nuchal plate,
ventral view; D, headshield; E, Errolosteus goodradigbeensis Young 1981, partial headshield; F, I, restoration
of headshield of Goodradigbeeon australianum White 1978, in dorsal (F) and lateral (1) views; G, parasphenoid
of Buchanosteus. (A, E, G, H after Young (1979, 1981a). B, C, D, F, I, after White 1978). Scales approx. G
(x 2); all others (x 0.5).

LONG HISTORY OF FOSSIL FISH - 357

process for
articulation of
headshield

C

pectoral fenestra

Figure 12. Placodermi (Early-Middle Devonian). Euarthrodires from Taemas (A-G) and near Cobar (H, I),
New South Wales. A, B, trunkshield of Williamsaspis bedfordi White 1952 in lateral (A) and ventral (B)
views; C, trunkshield of Taemasosteus novaustrocambricus White 1952 in lateral view; D, Anterior Lateral
plate of Toombsosteus denisoni White 1978 in lateral view; EE, anterior Lateral plate of Errolosteus
goodradigbeensis Young 1981 in visceral view; F, reconstruction of the trunkshield of Buchanosteus
confertituberculatus (Chapman 1916) in lateral view; G, Median Dorsal plate of Arenipiscis westolli Young
1981 in ventral view; H, I, trunkplates of Wuttagoonaspis fletcher Ritchie 1973 in lateral view. A (x0.5); B
(x0.75); C (x0.25); D (x0.75); F, H, I (1.0); G & 0.75). (A, B, after White 1952; C, after White 1978; D,
E, G after Young 1981b; H, I after Ritchie 1973).

358 - LONG

mC PN . Ritchie
SO. Miles &
Young

ifc

Figure 13. Placodermi (Early-Middle Devonian, New South Wales). Euarthrodires. A, reconstructed
headshield of Denisonosteus weejasperensis Young & Gorter 1981, approx. x 0.75; B, headshield of
Wutlagoonaspis fletcheri Ritchie 1973, approx. x 0.5. (After Young & Gorter 1981) and Ritchie 1973).

Basin include large arthrodires as yet undescribed from the Yellow Drum Sandstone of the
Fairticld Group (Druce & Radke 1979). An incomplete skull of a large eubrachythoracid
arthrodire with coarse tubercular ornamentation was recently found in the Famennian Ningbing
Limestone, Bonaparte Basin, northern Western Australia, but it is too incomplete to allow
confident identification.

Freshwater Late Devonian faunas from Australia contain the arthrodire Groenlandaspis (Fig.
17B), which is widespread in both Frasnian and Famennian deposits throughout southeastern
Australia (Ritchie 1974, 1975, Long 1982a, 1983a), and the flattened phyllolepids. Several
species of Groenlandaspis are currently under study by Dr A. Ritchie, who is revising the
world-wide occurrences of the genus, The best material of this fish comes from the Frasnian
Mt. Howitt deposit where the articulated armours and tails of Groenlandaspis occur in all stages
of growth. Groenlandaspis is characterized by its high, flat, triangular Median Dorsal plate,
which varies in shape between species (Ritchie 1975), and the Posterior Dorsolateral plate bears
an inverted V-shaped lateral line canal groove. It is perhaps more easily recognized in
Australian Late Devonian faunas by its distinctive tubercular ornament, as other placoderms
occurring with Groenlandaspis have either reticulate or meandering ornaments (antiarchs such as
Bothriolepis and Remigolepis, phyllolepids). Recently Groenlandaspis sp. has turned up from
marine limestones in the Camarvon Basin (Frasnian Gneudna Formation).

Phyllolepids were, until recently, all thought to belong to the one cosmopolitan genus,
Phyllolepis and this genus was recorded from Australia mainly by its ornament and isolated
plates, by Hills (1929, 1931, 1932, 1936a, 1959). Recent description of the well preserved
Mt. Howitt phyllolepids resulted in the recognition of a new genus of endemic phyllolepid,
Austrophyllolepis (Figs l6A, C, 17C) which closely resembles PhAyllolepis in its ornament

LONG HISTORY OF FOSSIL FISH - 359

Figure 14. Placodermi (Late Devonian). Arthrodires from Gogo, Western Australia: A, Rolfosteus
canningensis Dennis & Miles 1979b; B, C, Camuropiscis concinnus Dennis & Miles 1979a; D, /ncisoscutum
ritchiei Dennis & Miles 1981; E, Harrytoombsia elegans Miles & Dennis 1979. (After papers by Miles &
Dennis 1979), Dennis & Miles 1979a, b, 1981).

360 - LONG

Figure 15. Reconstructions of Gogo placoderms: A, Eastmanosteus calliaspis; B, Holonema westolli; Cc,
Rolfosteus canningensis; D, Latocamurus coulthardi; E, Bothriolepis sp. Scale approx. A (x 0.1), B (x
0.12), C, D (x 0.5), E (x0.3). All original.

LONG HISTORY OF FOSSIL FISH - 361

Figure 16. Placodermi (Late Devonian). Phyllolepids: A, C, restored armour of Austrophyllolepis ritchiet
Long 1984b from Mt Howitt, Victoria, in dorsal (A) and ventral (C) views; B, D, same views of Placolepis
budawangensis Ritchie 1984 from New South Wales. Scale approx. x 0.5. (After Long 1984b and Ritchie

1984).

and general bone pattern, but differs in having a Posterior Median Ventral plate and in minor
details of the headshield (Long 1984b). The tail, jaws, cheek and outline of the braincase of
Austrophyllolepis were described for the first time in the group enabling revision of the
phylogenetic status of the phyllolepids. I concluded that phyllolepids are probably highly
specialized arthrodires in which the armour has degenerated as a response to extreme
dorsoventral flattening, although alternative placement of phyllolepids as sister group to
arthrodires is acceptable (Goujet 1984a). A primitive phyllolepid contemporancous with
Austrophyllolepis was described from the Comerong Volcanics, near Braidwood, New South
Wales by Ritchie (1984) as Placolepis budawangensis. Placolepis (Fig. 16B) has a headshield
with large marginal plates and a strongly arched front on the nuchal plate. The only other
possible phyllolepid is Antarctaspis of Middle or Late Devonian age from Antarctica (White

362 - LONG

1968, Denison 1978), suggesting that the stem group of phyllolepids originated from East
Gondwana and later spread to the Northern Hemisphere where they are restricted to one
specialized genus, Phyllolepis occurring in a narrow time zone of the Famennian (Westoll
1952; Young 1974, 1981a, 1987c; Lyarskaya 1978). Young (1988a) and Long (1990) have
recently described isolated placoderm plates from central and eastern Australia. More primitive
phyllolepids are anticipated to occur in Australian or Antarctic Early-Middle Devonian deposits.

Figure 17. Placodermi (Late Devonian). Attempted restorations of three placoderms occurring at Mt Howitt,
Victoria. A, Bothriolepis gippslandiensis Hills 1929; B, Groenlandaspis sp., C, Austrophyllolepis ritchiei
(Long 1984b). Scale approx. x 0.5. All original.

Antiarchs

The antiarchs make their earliest appearance in the Late Silurian-Lower Devonian of China,
represented by endemic primitive forms known as yunnanolepidoids (Zhang 1978, 1984).
Sinolepidoids are slightly more advanced in having well developed pectoral appendages attached
to the trunkshield by a brachial process ("euantiarchs" Janvier & Pan 1982). Recently the
pectoral joint was described in yunnanolepidoids (Zhang 1984), but is not known in detail on
sinolepids. Young (1984c, 1988b) relates sinolepids to bothriolepids in that they share a well
developed bisegmented pectoral appendage. Australia is the only country other than China to
have sinolepids. A new genus of sinolepid comes from the Late Devonian Hunter Siltstone,
near Grenfell, New South Wales (Young 1981a, Long 1982a, 1983a). This form, as yet
undescribed, has rectangular Median Dorsal plates, a long Nuchal plate and enormous Ventral
Fenestra.

The two most widespread antiarch groups are the asterolepidoids and the bothriolepidoids.
Asterolepidoids have long trunkshields with short pectoral appendages, but the small

LONG HISTORY OF FOSSIL FISH - 363

orbital facets

median
ridge

Figure 18. Placodermi (Middle-Late Devonian), Antiarchs: A, F, Sherbonaspis hillsi Young & Gorter 1981 (A,
B, C, F from the Middle Devonian Hatchery Creek Conglomerate of New South Wales). B, restored partial armour
of, and C, proximal segment of the pectoral appendage of Monarolepis verrucosa (Young & Gorter 1981, Young
1988b). D, E, median dorsal trunk plates of Briagalepis warreni Long 1983a. G, proximal pectoral appendage of
Bothriolepis gippslandiensis Hills. Scale approx. A-B (x 1); C-E (x 2); F (x 3); G (x1). (A, C, F, after Young &
Gorter 1981).

headshields feature large orbital fenestrae and broad, short premedian plates. The earliest known
asterolepidoids are from the latest Early Devonian Georgina Basin, Queensland (Young 1984a,
although recently doubts have been cast on the age of this fauna (Long, Turner & Young
1988). Asterolepidoids are abundant in the Middle and Late Devonian world-wide, but have
only recently been recognized from Australia, which appears to have a high proportion of
endemic types. Sherbonaspis hillsi (Fig. 18A, F) is a small pterichthyodid from the Eifelian
Hatchery Creek Fauna, New South Wales, which has a slightly cristate Anterior Median Dorsal
plate (Young & Gorter 1981). Another pterichthyodid is known from the Middle Devonian
Broken River region, Queensland (Young & Gorter 1981, Young 1990). The "Asterolepis"
plate from Gilberton, Queensland, described by Hills (1936a; figured in Hill et al. 1967) is
probably a Bothriolepis plate (Turner 1982c). The most common asterolepidoid fish from
Australia is Remigolepis (Pl. 3A) found throughout the Famennian deposits of New South
Wales and particularly well preserved from Canowindra and Jemalong Gap (Hills 1958, Ritchie
1969, Campbell & Bell 1977). This material, which remains to be described, includes
complete fish with the tails preserved. Remigolepis is very similar to Asterolepis, a
widespread genus from Northern Hemisphere faunas, but is most easily recognized by having
only one segment to the pectoral appendage, incorporating the bones of both parts of the

364 - LONG

normal asterolepidoid fin. There are probably several species of Remigolepis from the
Famennian of New South Wales, although none have yet been formally described. An endemic
asterolepidoid, Pambulaspis cobahdrensis, from the Frasnian Pambula River site, southern New
South Wales, has a separate posterior lateral plate, and a slight crest on the Anterior Median
Dorsal plate (Young 1983). The pectoral fin is not known in Pambulaspis so affinities with
Remigolepis are uncertain. Another asterolepidoid, as yet undescribed, comes from the
Pambula River Fauna. It is unusual in having deeply and irregularly pitted dermal ornament
(Dr G. Young, pers. comm.).

Bothriolepidoids had well developed long pectoral appendages and a longer Premedian plate
than most other antiarchs, with a divided chamber in front of the orbit for the nasal capsule
(preorbital recess). Bothriolepis, a cosmopolitan genus, was recently thought to occur earlier
in Australia and China than elsewhere, being found in the early Middle Devonian Hatchery
Creek Conglomerate fauna, New South Wales (Young & Gorter 1981). The species, B.
verrucosa (Fig. 18B, C), has recently been placed in a new genus, Monarolepis distinguished
by having a very small axillary foramen on the Anterior Ventrolateral plate, and primitive plate
arrangement in the pectoral appendage (Young 1984c, Young 1988b). Bothriolepis (Figs 17A,
19) occurs commonly in Late Devonian deposits throughout Australia, and although is
represented by many species, few have been described. The Mt Howitt site has yielded
articulated specimens with the tail preserved, enabling restoration of the whole fish (Figs 17A,
19E). B. gippslandiensis and B. cullodenensis from the Avon River Group, Victoria, have well
developed median dorsal crests on their trunkshields (Pl. 3B) and have a primitive type of scale
covered tail with a dorsal fin-spine present (Long 1983a, 1985e, Long & Werdelin 1986).
Other bothriolepids from Victoria include B. fergusoni, a typical species with low vaulted
armour; B. warreni (=Briagalepis warreni Long et al., 1990), an unusual small species with
isolated tubercular ornament (Fig. 18D, E); and B. bindareei (Fig. 19B), a high-vaulted short
species (Long 1983a, 1984a). B. gippslandiensis, B. cullodenensis, and B. fergusoni form an
endemic monophyletic group of relatively primitive species which all share the feature of
having large lateral pits on the visceral surface of the headshield, but differ from advanced
species like B. canadensis (from Canada) in retaining a scale-covered tail (Long 1983a, Young
1988b). B. tatongensis from the Holland Creek Rhyodacite, Victoria, is the only Victorian
species to have coarsely reticulate ornament and a trifid-shaped preorbital recess, an advanced
feature in bothriolepids where the cavity for the nasal capsules has three distinct lobes (Long &
Werdelin 1986). Well preserved bothriolepids of Famennian age are also known from
Canowindra and Jemalong Gap, New South Wales. These are more similar to Northern
Hemisphere species with regular shaped armour and trifid preorbital recesses, and could have
entered Gondwana following faunal interchange between Euramerica and Gondwana at the start
of the Late Devonian, as proposed by Young (1981a). A marine species of Bothriolepis occurs
at Gogo, Western Australia (Figs 17E, 19A, D), which has produced important anatomical
information on the homologies of antiarch jaws and braincase structures (Young 1984a), but as
yet remains to be formally described. Bothriolepis plates occur in many sites throughout
south-eastern Australia (Hills 1929, 1931, 1932, 1936a, Ferguson et al. 1979, Long 1983a,
Young 1983), Queensland (Turner 1982c), and central Australia (Hills 1959, Gilbert-
Tomlinson 1968, Young 1985b), most of which currently await detailed study. Articulated
bothriolepids with tails preserved have been reported from the Amadeus Basin, Northern
Territory, by Young (1985a). Recently a new species, B. billilunensis, was described from
fragmentary trunk plates by Young (1987a) from the ?Famennian Knobby Sandstone, Canning
Basin, Western Australia, and bothriolepid plates were recovered from the Frasnian Munabia
Sandstone, Carnarvon Basin, Western Australia, by the author and friends in late 1987.

Petalichthyids
Petalichthyids have a distinctive dermal skull roof pattern with a long narrow Nuchal plate
and two pairs of Paranuchals, and are usually found in Early-Middle Devonian deposits.

LONG HISTORY OF FOSSIL FISH - 365

\
J;
Uf orbital
((:-/ fenestra
YA
csl ‘ee 7

ae Re

oa.PMD

Figure 19. Placodermi (Late Devonian). Bothriolepids. A, D, Bothriolepis sp. from Gogo, Westem
Australia. A, headshield; D, anterior median dorsal plate; B, attempted restoration of the trunkshield of
Bothriolepis bindareei Long 1983a from Victoria; C, headshield of, and E, reconstruction of Bothriolepis
cullodenensis Long 1983a from Victoria; F, headshield of Bothriolepis gippslandiensis Hills 1929 from Mt
Howitt, Victoria. Scale approx. A (x 0.75); B (x 1); C-D (x 0.5) ; E (x 0.25); F (x 0.5). (A, B after material
in the Bureau of Mineral Resources; C-F, after Long 1983a and Long & Werdelin 1986).

366 - LONG

ri ¥ ‘ (
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posterior process — quadrate

Figure 20, Placodenmi (Devonian), Ptyctodontids, Petalichthyids and Palacacanthaspidoids, A, headshield

of Wydeaspixs warroovensis Young 1978 (AD, I, G from the Hartly Devonian neat Taemas, New South Wales);
B,C, Brindabellaspis stensios Young 1980, B, headshield, ©, restored armour in lateral view; D,

Notopetalichthys hillsi Woodward 1941, Ey LL, Clenurella gardinert Miles & Young 1977, 1, neck joint and

dorsal fin basala; 1, redtoration of xkull roof; J, upper jaw apparatus, K, L, clasping elements; VF, G,
Weejasperaxpis gavini White 1978, Fy trunkshield restored, G, front of Median Dorsal plate; HI,
Campbelloduy decipiens Miles & Young 1977, dorsal spine, Seale approx. A (x 0.5), B(x 0.3); C (x 0,25); D
(x O05), E(w 2) Fw 0.5), G (x 0.5), HG 1), EO 1) EO 2); JK (x 3), (A, after Young 1978; B, C, after
Young 1980; D, after Woodward 1941, 1H, 1, J, Ky L, after Miles & Young 1977; Ff, G, after White 1978).

Australia has few of these fish, all of which are known from incomplete headshields and trunk
fragments, Allof the petalichthyids described from Australia so far have come from the Early
Devonian limestones near Taemas, New South Wales, and Buchan, Victoria, Notopetalichthys
hillsi (Fig. 20D) was described from a partial headshicld by Woodward (1941), and is an
endemic genus, Wijdeaspis (Fig, 20A) is known from New South Wales and Victoria,
representing a new species, W. warrovensiy (Young 1978, Long 1984c; PL 2C), Lunaspis has
also been recorded from ‘Taemas (Young 1985a), Both Lunaspis and Wijdeaspis are otherwise

LONG HISTORY OF FOSSIL FISH - 367

known from the Early-Middle Devonian of West Germany and the U.S.S.R. (Denison 1978).
Another endemic form, Shearsbyaspis, comes from the Taemas district (Young 1985a).

Acanthothoracids

The unusual depressiform ray-like rhenanid placoderms from the Early Devonian of
Germany (Gemuendina, Gross 1963) are unknown in Australia, but the record of
acanthothoracids is very good. Both groups share dorsally open nostrils. Acanthothoracids
have short trunkshields, long headshiclds with dorsal eyes and nares and a premedian plate, the
latter used as evidence by Goujet (1984) to propose that they are related to antiarchs. The
dermal ornament of these fishes is often very elaborate, facilitating identification from
fragments of bone. Australian acanthothoracids include the best preserved specimens of cranial
material in the world from the Early Devonian Taemas and Buchan regions. Weejasperaspis
(Fig. 20F, G ; White 1978) and Murrindalaspis (ornament PI. 2G; Long 1984d) have high
median dorsal crests on the trunkshield, but their headshiclds remain unknown. A detailed
study of placoderm cranial anatomy based on beautiful specimens of Brindabellaspis stensioi
(Fig. 20B, C) by Young (1980) described cranial nerves, vessels and brain morphology, and has
important implications to the homology of placoderm braincase topography and to placoderm
interrelationships and gnathostome relationships in general. Recently, further remains of
Murrindalaspis were found at Taemas, which include, amongst other bones, a beautifully
preserved complete sclerotic capsule (ossified "eyeball"), as well as the first pelvic girdles
known in acanthothoracids (Long & Young 1988). This specimen (PI. 2E, F) shows
remarkable anatomical detail of the eye muscle attachments and arterial and venous pathways.

Ptyctodontids

Ptyctodontids are short-shielded placoderms with little bone cover on the head. They are
remarkably like modern chimacrids in body form, and other features such as crushing tooth
plates and holostylic jaw suspension. Ptyctodontids are the only placoderms known to possess
pelvic copulatory structures resembling chondrichthyan claspers. These structures and others
led @rvig (1961, 1962) to believe that ptyctodontids were ancestral to chimaerids, thus
implying chimaerids to be living placoderms. Other workers put these resemblances down to
convergence (Patterson 1965, Schaeffer & Williams 1977) but Miles & Young (1977) regard
the pelvic claspers of ptyctodontids as a primitive feature for placoderms and chondrichthyans.
Australia has beautiful material of Late Devonian ptyctodontids from Gogo, Western Australia
where two genera are found. Ctenurella gardinert (Fig. 20K, I-L) is congeneric with species of
similar age in Germany (@rvig 1961, Miles & Young 1977) but Campbellodus (Fig. 20H) is
an endemic Australian genus. Ctenurella from Gogo provides important anatomical
information on the braincase and jaw suspension of ptyctodontids (see also Forey & Gardiner
1986). A new, almost complete specimen of Campbellodus from Gogo shows unusual
development of two median dorsal plates as well as a high median dorsal spine (Long 19874,
1988c). Cf. Campbellodus? is also known from a single large upper jaw toothplate from the
Upper Frasnian Napier Formation, Canning Basin, Western Australia, (Long 1988a).
Fragments of ptyctodontids have been recovered from Early Devonian limestones near Tacmas
(@rvig 1969) but have not been formally described.

Osteichthyes

The Osteichthyes or "true bony fishes" represent the largest proportion of living and extinct
fishes, with over 23,000 extant species. They are characterized by a well-ossified internal and
external skeleton and a swimbladder, which in some forms became modified into a lung.
Amongst the primitive osteichthyans are the ancestors of tetrapods, and this lineage ultimately

368 - LONG

led to the evolution of man. Mammals, birds, reptiles and amphibians are, therefore, all
subdivisions of the Osteichthyes (Gardiner 1980). Although early workers generally recognized
three major groups of osteichthyans, recent debate over this rigid classification resulting from
recognition of paraphyly of traditional groups (Rosen et al. 1981, Gardiner 1984b). The major
groups are the Actinopterygii (or ray-finned fishes), the Crossopterygii (or lobe-finned fishes)
and the Dipnoi (or lungfishes). The latter two are often grouped together as the Sarcopterygii,
as both these groups have lobed fins, enamel on the teeth or toothplates and primitively
possessed a pore-canal system in the dermal skeleton (the tissue "cosmine", and other
characters, Schultze 1987). Actinopterygians and dipnoans are undoubtedly monophyletic
(Gardiner 1984b, Schultze & Campbell 1987), and within the Crossopterygii (believed
monophyletic by Schultze 1987) the following sub-groups are recognized: Osteolepiformes,
Struniiformes, Porolepiformes and Actinistia (coelacanths, the only living group of
crossopterygians). Recently, it was suggested that the Rhizodontida be treated as a separate
group outside of the Osteolepiformes (Long 1985a), although now I believe that
rhizodontiforms fit in at the base of the clade containing Osteolepiformes and tetrapods (Long
1989). Fig. 21 presents a recent view of osteichthyan interrelationships. The basic structure
of these groups will be briefly outlined preceding review of their Australian record. Until
recently, Palaeozoic osteichthyans were very poorly known in Australia, despite the oldest
reference to an Australian fossil fish being of a Permian actinopterygian (Dana 1848). There
are still very few well-known Palaeozoic osteichthyans compared to the fossil record in the
Northern Hemisphere countries like Britain or the U.S.S.R., although the Australian material
is well preserved and provides important data on the early radiation of certain groups.

Actinopteryli

The actinopterygians, or ray-finned fishes, have dominated the seas and rivers since the
Carboniferous Period. The earliest and most primitive members of this group, the
palaconiscoids, had evolved by the beginning of the Devonian, although articulated specimens
are known only from the Middle-Late Devonian. Palaconiscoids had long cheeks with fixed
maxillae and preoperculars, and thick rhombic peg and socket scales. Higher groups of
actinopterygians, the holosteans and teleosteans, developed mobile cheeks with reduced
maxillae as a response to the development of a specialized buccal pump mechanism of feeding,
enabling the jaws to move outwards (Schaeffer & Rosen 1961, Lauder & Liem 1983). Scales
became thinner and cycloid in shape, and the internal skeleton of the caudal fin was highly
modified. Australia's fossil record of actinopterygians includes the best preserved Devonian
paleoniscoids in the world, and a great diversity of Mesozoic forms. The Tertiary record is
poor, however, with most taxa based only on ear-stoncs, or otoliths, from marine deposits.

Devonian:

Palaconiscoid scales were recorded from the Early Devonian of Australia by Schultze
(1968). Ligulalepis toombsi (Pl. SA, Fig. 22B), from the Murrumbidgee Group, New South
Wales, is one of the world's first actinopterygians, although scales of proto-osteichthyans such
as lophosteiforms are known from the Late Silurian of Gotland (Gross 1969). Ligulalepis
scales are now frequently recognized in Early Devonian bone-rich acid residues (S. Turner pers.
comm.). Rare isolated actinopterygian bones are also known from the Emsian-Eifelian Mulga
Downs Group, preserved as impressions. The well preserved palaeoniscoids from Gogo
described by Gardiner (1973, Gardiner & Bartram 1977, Gardiner 1984b) have much
significance in solving problems of osteichthyan interrelationships (see also Patterson 1982).
Two genera are found at Gogo, one of which, Moythomasia (Fig. 22A), is also known from
the Late Devonian of Germany (Gross 1953, Jessen 1968), and possibly elsewhere if scale
morphology is reliable (Gardiner 1967, Blieck et al. 1982). Moythomasia durgaringa and
Mimia toombsi from Gogo show that the braincase of early actinopterygians was ossified as a
single unit but retains several embryonic fissures and lacks posterior myodomes for eye muscle

Onychodontiformes

S
>
jam
uJ
Ee
oO.
Oo
=
=
O
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DIPNOI
Actinistia

LONG HISTORY OF FOSSIL FISH - 369

OSTEICHTHYES

SARCOPTERYGII
Crossopterygil

Choanata

Porolepiformes
Rhizodontida
Osteolepiformes
Panderichthyidae

1-9
SYNAPOMORPHIES

lepidotrichia

marginal upper and lower teeth

long dentary and coronoids

hypohyal

shoulder girdle bone pattern

rhombic scales with peg and socket articulation
anteriorly directed infrapharyngobranchials
suprapharyngobranchials on first two gill arches
mesomenc skull roof with rostral bone mosaic
10. anocleithrum

11. true enamel on teeth

12. pectoral and pelvic fin- archipterygium type

13. cosmine (not known for some groups)

14. submandibular series

15. double-headed hyomandibular

16. intracranial joint

17. three extrascapulars

18. extratemporal bone

OHNOGAwN A

narrow submandibulars

fenestra ventro-lateralis

four infradentaries below a long dentary
three coronoid bones

plicidentine in teeth

supraorbital sensory-line joins cephalic
division of main lateral line

7 bones in cheek: squamosal large
polyplocodont infolding of teeth

humerus with rounded caput humeri
scapulocoracoid with three equally-sized
buttresses defining extensive fossae for
fin muscle attachment

caput humeni fits into groove on scapulocoracoid
choana (sensu Schultze 1987)

three pairs of mid-line skull roof bones
external naris close to upper jaw margin
flat skull with high dorsally-placed orbits

Figure 21. Possible interrelationships of the major groups of Osteichthyes, based on Schultze (1987) and

Long (1989).

370 - LONG

attachments. The Gogo palaeoniscoids are completely preserved, showing for the first time in
a Devonian actinopterygian the full gill arch series, internal morphology of the braincase, and
endoskeleton of the pelvic girdle and fin, Another well preserved palaeoniscoid, from lacustrine
deposits of Frasnian age at Mt Howitt, Victoria (Pl. 5B, Fig. 22C) is placed in a new genus,
Howqualepis, (Long 1988e), which is primitive in cheek structure but has a specialized snout
in which the premaxillae are separated by the toothed postrostral bone (also in some specimens
of Mimia, Gardiner 1984b). In many other respects, however, it is more primitive than the
Gogo palaconiscoids, e.g. in having a long body shape with long-based pelvic fins, long
maxillary blade, long dermosphenotic and a relatively small orbit. Howqualepis probably fits
into the basal radiation of the palaeoniscoids, somewhere between Cheirolepis, the oldest
known complete actinopterygian (Middle Devonian, Scotland; Pearson & Westoll 1979), and
the Gogo forms (Long1988e).

Figure 22. Osteichthyes, Actinopterygii (Devonian). A, Moythomasia a different species to this one occurs
at Gogo, Western Australia; _B, scale of Ligulalepis toombsi show ng typical palaeoniscoid dorsal peg (dp);
C, Howqualepis rostridens Long (1988e) from Mt Howitt, Victoria. Scale appr. A (x1); B (4); C
(x0.5).

Carboniferous:

Palaeoniscoids were described from Mansfield, Victoria, by Woodward (1902, 1906a) as
belonging to the Northern Hemisphere genus Elonichthys. Recent revision of this fauna (Long
1988e) reveals that there are three species present in the fauna: two are contained in the new
genus Mansfieldiscus (Pl. 5D), and one belongs to a new gonatodid, Novogonatodus (Fig.
23A). Turner (1982c 1982e) records Elonichthys? sp. from the Carboniferous of Queensland,
although identification of this genus is currently uncertain, due to the need for a world-wide
revision. Palaeoniscus randsi (Etheridge 1892), from near Boguntungan, Queensland, is now
known to be of Carboniferous age, although because the head is not preserved, positive
identification cannot be made and the species is now regarded as indeterminate (Turner & Long

LONG HISTORY OF FOSSIL FISH - 371

1987). Isolated palaeoniscoid bones have been found from the basal Carboniferous Raymond
Formation, Queensland, but identification of these specimens awaits more complete material.

Permian:

Actinopterygians occur infrequently in the coaliferous deposits of New South Wales and
Queensland. Ebenaqua ritchiei (Fig. 25) is a deep-bodied bobasatranid from the Blackwater
Shales, southern Queensland (Campbell & Le Duy Phuoc 1983). The original description
identified the large bone behind the maxilla as a suborbital, although re-examination of the
material has shown that on Ebenaqua and Bobasatrania the sensory-line canal on this bone is
not as shown by the authors (Campbell & Le Duy Phuoc 1983) but is a pit-line on the
quadratojugal (QJ, Fig. 25C). The course of the infraorbital sensory-line runs more
posteriorly. This implies that the bone in question is in fact a quadratojugal rather than a
suborbital.

Figure 23. Osteichthyes, Actinopterygii (Carboniferous). Two genera from the Lower Carboniferous
Mansfield fauna. A, Mansfieldiscus sweeti (Woodward 1906); B, the gonatodid Novogonatodus kazantsavae
Long (1988e). Scale approx. A (x 0.5); B (x 0.75).

Urosthenes is a poorly known Permian palaconiscoid from the Newcastle Coal measures,
New South Wales, which is known by two species U. australis (Dana 1848) and U. latus

372 - LONG

(Woodward 1931). Currently, these species cannot be diagnosed as distinct taxa, although a
new genus of palaeoniscoid attributable to the family Urosthenidae is now known from the
Upper Permian Blackwater Shale, Queensland (Fig. 24A; Le Duy Phuoc 1980, Mike Leu pers.
comm.), linked to the genus Urosthenes by features of the body shape and fins. The
Blackwater Shale, Queensland, has produced several new forms of actinopterygians, most of
which are currently being studied for a Ph.D dissertation by Mr M. Leu (Macquarie University,
Sydney) In the fauna are new redfieldiids (Fig. 24B), regular palaeonisciforms (Fig. 24C) and
the new urosthenid genus (Fig. 24A). The figures used here are all taken from Le Duy Phuoc
(1980, unpublished Masters dissertation, Australian National University, Canberra).

Fiure 24. Osteichthyes, Actinopterygii (Permian). Fishes from the Blackwater Shale, Queensland. A,
urosthenid, new genus; B, C, palaeoniscoids. Scales approx. A (x 1); B-C (x 2). (After Le Duy Phuoc, 1980
M.Sc thesis).

Acrolepid scales and bones are known from Permian marine sandstones near Aberfoyle,
Tasmania (M. Banks, pers. comm.).

- 373

LONG HISTORY OF FOSSIL FISH

Figure 25. Osteichthyes, Actinopterygii (Permian). Ebenaqua ritchiei Campbell & Le Duy Phuoc 1983. A,

restoration; B, ventral view of head; C, details of head, note new interpretation of cheek. Scales approx. A (x

0.75); B (& 3); C (x2). (After Campbell & Le Duy Phuoc 1983).

374 - LONG

Triassic:

A great diversity of actinopterygians have been described from the Triassic continental
sediments around Sydney and Gosford. Three main faunas are known: the youngest (Late
Triassic) is from the Wianamatta Group, which has produced the St Peter's fauna from the
Liverpool subgroup (Ashfield Shale). The Middle Triassic Brookvale fauna comes from the
Hawkesbury Sandstone, near Sydney. The oldest fauna is the Early Triassic Gosford fauna
from the Narabeen Group (Gosford Formation). Early work on these fishes by Woodward
(1890, 1902a, 1908, 1931) and Wade (1930, 1935, 1940, 1941, 1942b) established the diverse
nature and endemic character of the assemblage, but recent revision by Hutchinson (1973) has
clarified details of the systematic position of many of the taxa. As the taxonomy of
actinopterygians at the subholostean to teleostean level is very complex, readers should refer to
Hutchinson (1973) and Patterson (1973) for details. Taxonomic listing of these faunas can be
found in Long & Turner (1984). The palaconiscoids, the most primitive group of
actinopterygians, flourished during the Triassic, but were gradually phased out by subholostean
groups by the Jurassic, and such groups were already well established by the Late Triassic (St
Peters Fauna). Many of the Triassic fishes from Gosford, St. Peters and Brookvale are of
uncertain taxonomic status due to poor preservation of the cranial features. Because the fish are
well preserved as a whole unit, they were originally described mostly from body form, details
of the fin rays and scale ornament, and whatever could be deciphered of the head region.

Figure 26. Ostcichthyes, Actinopterygii (Triassic). Saurichthys, which is found in New South Wales,
Queensland and Western Australia freshwater Triassic deposits. (After Tumer 1982b). Scale approx. x 0.3.

Saurichthys (Fig. 26) is a slender, long snouted predator which has been found in the
Gosford (S. gracilis; Woodward 1890) and Brookvale (S. parvidens; Wade 1935) faunas of New
South Wales, as well as from the Triassic Rewan Group (Arcadia Formation) of Queensland (S.
sp. cf. S. gigas; Turner 1982b), from the Knocklofty Formation, near Hobart, Tasmania
(Dziewa 1980), and from the Blina Shale, northern Western Australia (Warren 1980).

Palaeoniscoids belonging to the Tegeolepidae (Apateolepis Woodward 1890) and
Acrolepidae (Leptogenichthys Wade 1935) are perhaps the most primitive osteichthyans from
the New South Wales Triassic faunas. Acrolepids have also been described from the Triassic
Knocklofty Formation, Tasmania (Acrolepis hamiltoni, A. tasmanicus; Johnston & Morton
1890, 1891, Dziewa 1980), with scales of acrolepids recently reported from the Permian Byro
Group of the Carnarvon Basin, Western Australia (Archbold 1980). Palaeoniscoids belonging

LONG HISTORY OF FOSSIL FISH - 375

Myers
oes ae =.
=

WSs .

Figure 27. Osteichthyes, Actinopterygii (Triassic, New South Wales). A, Schizurichthys pulcher Wade
1935; B, Molybdichthys junior Wade 1935; C, Manlietta crassa Wade 1935; D, Procheirichthys ferox Wade
1935. All approx. natural size. (All after Hutchinson 1973).

376 - LONG

to the family Palaconiscidae are represented by the endemic genus Agecephalichthys from
Brookvale (Wade 1935) and the cosmopolitan genus Palaeoniscus from the Wianamatta Series
(P. antipodeus, Egerton 1864; P. crassus, P. feistmanteli Woodward 1908), although this latter
genus requires revision. Palaconiscoids of indeterminate affinity were described from
Harrington, near Sydney by Woodward (1902b, Atherstonia) and Wade (1935, Belichthys spp.,
Megapteriscus, Mesembroniscus). Other palaconiscoids of currently uncertain taxonomic
status from New South Wales are the following: Myriolepis known by three species from
Campbelltown and Cockatoo Island (Egerton 1864, M. clarkei), Gosford (Woodward 1890, M.
latis) and St Peters (Woodward 1908, M. pectinata); and Elpisopholis from St Peters
(Woodward 1908). A body of a palaeoniscoid from the Triassic coal deposits near Leigh Creek,
South Australia was described by Wade (1953) as a new form, Leighiscus, but as the head is
missing, its taxonomic status is uncertain .

Redfieldiiformes are represented by the family Brookvaliidae (Fig. 28A, B), from which two
endemic genera have been described from the Hawkesbury Sandstone. Brookvalia (Figs 28, 29;
Wade 1935) is represented by four species, which differ by body shape and scale ornamentation,
and the genera Beaconia and Dictopleurichthys (Wade 1935) are now incorporated into the genus
(Hutchinson 1973). Phlyctaenichthys (Figs 28C, 29F; Wade 1935) is closely related to
Brookvalia. Woodward (1890) described three species of Dictopyge from this fauna, and
suggested that D. illustrans was closcly allied to Brookvalia. Other redfieldiiforms belonging
to the family Redfieldiidae, occur in the Brookvale fauna. Three of these (Geitonichthys,
Molybdichthys, Schizurichthys; Figs 27, 29) were redescribed by Hutchinson (1973).
Perleidiiforms in the fauna are Procheirichthys and Manlietta (Fig. 27C, D). A common taxon
in all Triassic faunas of New South Wales is the deep-bodied cleithrolepid Cleithrolepis
granulata (Fig. 29E; Egerton 1864, Wade 1930, Hutchinson 1973) with the similarly deep-
bodied Platysomus occurring only at St Peters (Woodward 1908).

More advanced actinopterygians belonging to the Order Semionotiformes are known from
the Brookvale and Gosford faunas (Enigmatichthys, Wade 1941; Semionotus; Woodward,
1890). The most specialized of the Triassic actinopterygians from Australia are probably the
parasemionotids, represented by two species of the genus Promecosomina: P. beaconensis
(Wade 1935) from Brookvale, and P. formosa (Woodward 1908). Overall the diverse Triassic
fish faunas of New South Wales require revision of many taxa before their true affinities and
biogeographic significance can be gauged.

Jurassic:

Australia's only fish faunas from this period are from the Upper Jurassic Talbragar site,
northern New South Wales, whose fauna was originally described by Woodward (1895) and
Wade (1930, 1942a) and from subsurface coal material in Queensland (Turner & Rozenfelds
1987). The Talbragar fishes are preserved in hard shale, which contains a diverse fossil flora
(White 1981). The fauna contains one palaconiscoid, Coccolepis australis, 3 genera of
holosteans and the first appearance of the teleosteans in the Australian fossil record. The
holosteans are the macrosemionotid Uabryichthys latus (Fig. 30D; Wade 1942a), which was
redescribed by Bartram (1977), and two forms belonging to an endemic family Archaeomaenidae
(Pholidophoriformes). These are Archaeomene and Madariscus (Woodward 1895). The family
Archaeomaenidae is discussed by Waldman (1971a). The teleostean Leptolepis (Figs 30B,
31D) is the most common element in the fauna, being represented by three species, L.
gregarious, L. lowei and L. talbragarensis (Woodward 1895, Waldman 1971a). These species
are considered in the review of leptolepids by Nybelin (1974). Fishes thought to be of Jurassic
age from Victoria by Woodward (1906b) and Chapman (1912) are now known to be Early
Cretaceous and are considered below. The Queensland Jurassic fishes consist of partial bodies
from unidentifiable genera.

LONG HISTORY OF FOSSIL FISH - 377

AA \WWwss a

SS

Figure 28. Osteichthyes, Actinopterygii (Triassic, New South Wales). A, Brookvalia gracilis Wade 1935;
B, B. spinosa Wade 1935; C, Phlyctaenichthys pectinatus Wade 1935. All approx. natural size. (All after
Hutchinson 1973).

Cretaceous:

Two areas have yielded good material of Cretaceous fishes in Australia: the Early
Cretaceous Koonwarra site (Strzelecki Group) in Victoria and localities in north-western
Queensland of similar age (Aptian-Albian Rolling Downs Group). These two assemblages
differ greatly due to their environments of deposition. Before describing the faunas, some
isolated specimens warrant comment. Leptolepis crassicaudata and Psilichthys selwyni were
described by Hill (1900) from singular, imperfect specimens from the Otway Group,
southwestern Victoria. Psilichthys was redescribed as a valid genus by Waldman (1971b).

The Koonwarra fauna is well preserved in finely laminated, varved shales representing a lake
which was thought to have regularly froze over killing the fish populations (Waldman 1971a,
1973). The most abundant taxon is the teleost Leptolepis koonwarri (Figs 30B, 31D), but

378 - LONG

Figure 29. Osteichthyes, Actinopterygii (Triassic, New South Wales). A, Molybdichthys junior Wade
1935; B, Brookvalia gracilis Wade 1935; C, Geitonichthys ornatus Wade 1935; D, Manlietta crassa Wade
1935; E, Cleithrolepis granulata Egerton 1864; F, Phlyctaenichthys pectinatus Wade 1935. Scale approx.
x2. (All after Hutchinson 1973).

primitive palaeoniscoids were still around with this species, as shown by Coccolepis
woodwardi (Figs 30A, 31A). One holostean belonging to the Archaeomaenidae is present,

LONG HISTORY OF FOSSIL FISH - 379

Figure 30. For caption, see next page.

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Figure 30. Osteichthyes, Actinopterygii (Jurassic-Cretaceous). A-C, fishes from the Early Cretaceous |
Koonwarra Fauna, Victoria; A, Coccolepis woodwardi Waldman 1971a; B, Leptolepis koonwarri Waldman

1971a; C, Koonwarria manifrons Waldman 1971a; D, Uarbryichthys latus Wade 1941, from the Jurassic
Talbragar Fauna, New South Wales. Scale approx. a (x 0.75); B (x 1); C (x 0.8); D (x 0.5). (A-C, after
Waldman 1971a: D, after Hutchinson 1973).

Wadeichthys oxyops (Fig. 31B; Waldman 1971a). The fauna thus closely resembles the Late
Jurassic Talbragar assemblage in containing Leptolepis, Coccolepis and an archacomaenid.
Recent excavations at the Koonwarra site have turned up a few interesting new forms, yet to be
described. Recently, excavations in the Aptian-Albian Otway Group of central southern
Victoria have yielded articulated cranial material of fish (Rich & Rich 1989).

Dpt Par Ex

=

Figure 31. Osteichthyes, Actinopterygii (Cretaceous). Koonwarra fishes, heads in detail: A, Coccolepis
woodwardi Waldman 1971la; B, Wadeichthys oxyops Waldman 1971la; C, Koonwarria manifrons Waldman
1971a; D, Leptolepis koonwarri Waldman 1971a. Scale approx. x 2. (AIl after Waldman 1971a).

The fishes from the Rolling Downs Group are more spectacular in their large size, but are
generally only preserved as isolated heads and pieces of body (e.g. vertebrae; Hill et al. 1968).
One or two rare specimens, such as one on display in the Queensland Museum, show the head
with a row of vertebrae still articulated. All of these forms are from marine deposits, and they

LONG HISTORY OF FOSSIL FISH - 381

\
lop Qd

Figure 32. Osteichthyes, Actinopterygii (Cretaceous, Queensland). A, head of Pachyrhizodus marathonensis
Etheridge 1905; B, C, Xiphactinus a different species to this one is found in Queensland. Scales approx. A (x
0.14); B « 0.05); C &@ 0.1). (A after Bartholomai 1969; B from Romer 1966; C, after Bardack 1965).

382 - LONG

all belong, except for one holostean, Belonostomus, to the Teleostei. The most primitive
genus is the clopiform (clopid) Flindersichthys from the Tambo Series near Richmond,
described by Longman (1932) from a complete head. Most of the fishes from this fauna are
more advanced teleosts belonging to the Clupeomorpha, represented by two families,
Ichthyodectidae and Pachyrhizodontidae. These groups are mainly very large predatory fishes.
Ichthyodectids present are Cooyoo (Xiphactinus) australis (Woodward 1894), commented on by
Bardak (1965), and redescribed as a new genus by Lees & Bartholomai (1987); and, of doubtful
identification, is Cladocyclus sweeti (Woodward 1894, sce also Patterson & Rosen 1977). A
typical xiphacyinid fish is shown in Fig. 32B, C. The best known fish from the Rolling
Downs Group is Pachyrhizodus marathonensis (Fig. 32A; Etheridge 1905, Bardack 1962),
which was redescribed by Bartholomai (1969) and commented on by Forey (1977) as being akin
to salmonids. Indeterminate ichthyodectid remains are also known from the Rolling Downs
Group (Turner 1982c). Recently new collections of the Rolling Downs fishes have been made
by the Queensland Museum, and these are currently being acetic acid prepared. The material
shows excellent details of braincase and all surfaces of the dermal bones. Some of the largest
specimens include skulls about 0.5 m long.

The Koonwarra fauna, having close ties to the Talbragar fauna of Jurassic age, is the link
with primitive freshwater actinopterygian assemblages such as prevail in the Triassic of New
South Wales. The marine faunas of Queensland on the other hand contain advanced teleosteans,
which were to dominate seas and rivers from that time onwards. The Cretaceous teleostean
faunas appear to have a cosmopolitan aspect to them from earlier descriptions, although the
new work under progress in the Queensland Museum will probably demonstrate that there is
actually a higher number of endemic genera present than previously believed (T. Lees, pers.
comm.).

Tertiary:

Actinopterygians are poorly known from the Tertiary of Australia. Complete fishes have
been described from freshwater deposits in southern Queensland and northern New South Wales
(Hills 1934, 1943b, 1946, Taverne 1973, 1976). Marine deposits which extend around the
southern margin of Australia have yielded a diverse actinopterygian fauna known only from ear-
stones (otoliths), with some isolated bones and rare articulated material also recorded (Chapman
& Pritchard 1904, 1907, Chapman & Cudemore 1924, Corbett 1980). A great deal of work
awails anyone atlempting to sort out the many isolated pieces of jaws and skull bones from
Beaumaris and other Miocene sites in Victoria, because a large osteological collection of
modern teleosteans is required as a basis for comparisons.

The Tertiary marine deposits of Victoria, South Australia and Western Australia have
yiclded few articulated fishes but a great number of otoliths, from which an overall picture of
the fauna emerges (Frost 1928, Stinton 1952, 1958, 1963). Articulated fishes of Early
Miocene age are known from the Longford Limestone that crops out along the Murray Cliffs,
near Morgan, South Australia, although none of the material has been described (e.g. Figs.
33L, M). One superb specimen belongs to a snapper very similar to the living genus
Chrysophrys. The head of a flathead, Neoplatycephalus cf. N. bassensis was described from the
Upper Oligocene limestones near Wynyard, Tasmania (Fig. 33A; Corbett 1980). Otoliths
from the Oligocene-Pliocene deposits around Victoria (Fig. 34) show that marine fish
populations then were not significantly different from those around today. Common forms are
the Whiting (Sillago), flathead (Platycephalus), flounder (Pleuronectes), red snapper
(Trachichthyedes), tropical marine eels (Uroconger, Astroconger, Muraenesox), ox-eye herring
(Megalops) and hake (Merluccius). Of interest is the whiptail (Coelorhynchus), which today
lives in deep waters (250-900 m; C. innovatis Coleman 1980). The cod (Gadus), lives in cold
to subtropical waters of the Northern Hemisphere today, although it used to occur in the
Oligocene and Miocene of Victoria, and represents the only genus so far known from the

LONG HISTORY OF FOSSIL FISH - 383

Mince =
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A

SORRY
*) NY i

x
WN

Figure 33. Osteichthyes, Actinopterygii (Tertiary, marine). A, skull of flathead Platycephalus sp. cf. P.
bassensis; B, pufferfish palate, Diodon formosus Chapman & Pritchard 1907; C-H, pharyngeal crushing teeth
from labrid fishes; C, D, Nummopalatus depressus (Chapman & Pritchard 1907). C, fragment of dentition, D,
detail of single tooth; E, F, Oplegnathus manni Chapman & Cudemore 1924; F, detail of tooth; G, H,
Labrodon batesfordiensis Chapman & Cudemore 1924; I-K, isolated lower jaws (dentosplenials) of teleosteans
from Beaumaris, Miocene, Victoria; L, M, skull of percoid fish from the Miocene Morgan Limestone, South
Australia. Scales approx. A, B, C (x 1); D (& 6); E (x2); F (« 6); G (x2); H (x6); Lk (0.75); L-M (0.3). (A,
after Corbett (1980); others after original material).

384 - LONG

Figure 34. Osteichthyes, Actinopterygii (Tertiary, marine). Otoliths (ear stones) from Miocene and
Pliocene teleosts. A, Platycephalus petilis Stinton 1958 (flathead); B, Gadus refertus Stinton 1958; C,
Coelorhynchus elevatus Stinton 1956; D, Sillago pliocaenica Stinton 1952 (whiting); E, Heterenchelys
regularis Stinton 1958; F, Plerothrissus pervetustus Stinton 1958; G, Astroconger rostratus Stinton 1958
(eel); H, Lactarius tumulatus Stinton 1958; I, Monocentris sphaeroides Stinton 1958; J, Merluccius fimbriatus
Stinton 1958; K, Trachichthyodes salebrosus Stinton 1958; L, Sebastodes fissicostatus Stinton 1963; M,
Megalops lissa Stinton 1958; N, Antigonia fornicata Stinton 1963. All specimens between 0.3 and 1 cm.
(All from Stinton 1952, 1958, 1963).

Tertiary otolith faunas not currently living around Australia (Stinton 1963). A complete
listing of the Tertiary fishes represented by otoliths, and their localities can be found in Long
& Tumer (1984).

The Redbank Plains Series, in southern Queensland, have produced well preserved
articulated fishes of early Tertiary age (Eocene-Oligocene; Hills 1934, Hill et al. 1970). The
fauna contains the osteoglossid Phaeroides queenslandicus (Fig. 35A), which is
morphologically very similar to Phaerodus (to which genus Hills first assigned the material)
from the Palaeocene of Wyoming, U.S.A., and the Eocene of Sumatra. Taverne (1973)
recognized this fish as a different genus. Phaeroides reached up to half a metre in length and is

LONG HISTORY OF FOSSIL FISH - 385

closely related to the barramundi (Scleropages) which also occurs in Tertiary freshwater
deposits, near Duaringa, Queensland (Hills 1943b). Scleropages aff. S. leichardti (spotted
barramundi) is known from an operculum of presumed Oligocene age. Today S. leichardti is
found in the Fitzroy River system of Western Australia. The gonorhynchid Notogoneus parvus
also occurs in the Redbank Plains fauna. Notogoneus is a cosmopolitan genus occurring in the
Late Eocene to Late Oligocene of Western Europe and Wyoming, U.S.A. (Hills 1934),
Interestingly the only living genus of gonorhynchid, Gonorhynchus, is exclusively marine,

=—~
=

—

Uj,

Tee eeesdd

Figure 35. Osteichthyes, Actinopterygii (Tertiary- Recent, freshwater). A, Phareoides queenslandicus (Hills
1934), Oligocene, Redbank Plains, Queensland (after Hills, 1934); B, modem Murray Cod, Maccullochella
macquariensis, which is also known from Miocene lake deposits in New South Wales. Scale approx. A (x
0.3); B (x 0.15).

living in the Indopacific regions. Other fishes in the Redbank Plains fauna are more typical of
the major group of present-day teleosteans, the Percomorpha. Percalates, the estuary perch,
lives in freshwaters and estuarics around Australia today. Its fossil representative, from the
Redbank Plains is Percalates antiquus (Hills 1934), which was about half the size of the living
species that reaches 58 cm in length. Other fishes from this fauna are indeterminate as to
genus and species but have been assigned to the Percomorpha. The Duaringa fauna contains a
preopercular assigned to the snapper Lutjanus. normally a marine genus (Hills 1943b). Other
Tertiary fishes from freshwater deposits in Queensland include the plotosid catfish Tandanus and
the living Murray cod, Maccullochella macquariensis (Fig.35B), both from a well near

386 - LONG

Brigalow, of Pliocene-Pleistocene age (Longman 1929), The Murray Cod is well known as a
fossil from the Miocene diatomaceous earths near the Wurrumbungle Ranges, New South
Wales (Hills 1946),

Crossopterygii

This term is used only in the traditional sense to denote a monophyletic group of fishes
which share lobe-fins and a kinetic braincase (divided into two moeties; Schultze 1987).
Classically the Crossopterygii contains four main groups: Onychodontiformes,
Osteolepiformes and Porolepiformes ("Rhipidistia"), and the coelacanths or Actinistia.
Onychodontiformes, Osteolepiformes and Porolepiformes are totally extinct groups which
flourished in the Devonian, with only the Osteolepiformes surviving past the Devonian until
the middle Permian. Actinistians have been around since the Devonian and were thought to be
extinct at the end of the Mesozoic until Latimeria was caught off the coast of Madagascar in
1938. Since then many coelacanths have been caught off the Comores and detailed study of
their anatomy is the basis of our knowledge of crossopterygian biology (Millot & Anthony
1958, Millot & Robineau 1978). Recently the discovery of unusual crossopterygian-like fossil
fishes has brought about a revision of the systematics of the group. It is now thought to
contain other groups of equivalent sister-group ranking to the major groups, such as
Rhizodontida (sensu Andrews 1973) and the Youngolepis group (Rosen et al. 1981, Gardiner
1984b, Long 1985a, 1985e, Schultze 1987). A recent summary of previous interpretations on
crossopterygian interrelationships, and an up-to-date phylogeny of the Sarcopterygii can be
found in Schultze (1987).

All of these crossopterygian groups are known from Australia, but Porolepiformes and
Actinistia are very poorly represented.

Osteolepiformes are well represented in the Devonian and basal Carboniferous of Australia,
with most of the material having been recently described. Gyroptychius, a common genus in
the Middle Devonian of Scotland, Canada and Russia (Jarvik 1948, Vorobjeva 1977), was
described from the Middle Devonian Hatchery Creck fauna, in New South Wales, as a new
species G. sp. cf. G. australis (Fig. 36C, E; Young & Gorter 1981). Recent study of this
material by the author and G, Young indicates that the species is highly unusual in the degree
of overhang of the snout, and grooved nares. This species may be referable to a new genus.

An isolated cosmine-covered snout from the Late Devonian South Blue Range, Victoria is
exceptionally broad and flat. Recent preparation of the braincase on this specimen showed
unusual development of strong ridges on the postero-lateral region of the ethmosphenoid. I
now believe that this species is probably a “canowindrid" allied to the genus Canowindra.

Canowindra grossi (Fig. 36B, D) was described from a single complete fish in a slab of
Upper Devonian Mandagery Sandstone, New South Wales, by Thomson (1973), who could not
decide on the relationships of the genus but suggested it was probably a porolepiform. Long
(1985b) redescribed the head of Canowindra and concluded that the genus was an unusual type
of osteolepiform probably equal in taxonomic rank to "Osteolepiformes" as currently defined.
The unusual features of Canowindra are its skull roof which has a strongly flaring parietal
shield with undifferentiated intertemporal and supratemporal bones, the cheek which has three
postorbital bones instead of the usual one, and its very small eyes. Canowindra has lost the
cosmine in the dermal skeleton and the scales are rounded with a small boss on the basal
surface. Another more primitive "canowindrid", which retains cosmine, comes from Mt
Howitt. Beelarongia patrichae (Long 1987e) is known from one skull, and (PI. 6C, E, F; Fig.
36A) several of the features on the head are not clear. An unusual feature of the genus is a
single accessory postorbital bone behind the orbit. It also shows good preservation of the
shoulder girdle and humerus, which are typically osteolepiform. Further description of
canowindrids and discussion of their affinities can be found in Young, Long & Ritchie (in

LONG HISTORY OF FOSSIL FISH - 387

Figure 36. Osteichthyes, Crossopterygii, Osteolepiformes (Devonian). A, Beelarongia patrichae Long
1987d, skull (see Fig. 39-E); B, D, Canowindra grossi Thomson 1973, new restorations of the skull (B) and
whole fish (D), Late Devonian, New South Wales; C, E, Gyroptychius sp. cf G. australis Young & Gorter 1981,
C, fronto-ethmoidal shield in side view and (E) in dorsal view; F, Gogonasus andrewsae Long 1985d, fronto-
ethmoidal shield and ethmosphenoid in side view. Scales approx. A (x0.5); B (x 0.75); C, E (x (1.5); D &
0.2); F (x 2). (C, E, after Young & Gorter 1981, all others original).

388 - LONG

press), which deals with several new genera of crossopterygians from the Middle Devonian
Aztec Siltstone of Antarctica.

The advanced osteolepid Megalichthys has recently been recovered from the basal
Carboniferous Raymond Formation near the Narrien Range, Queensland. This material was
acid prepared to yield 3-dimensional bones and scales (Pl. 5 G) and new discoveries from the
same site in early 1986 by Prof. Ken Campbell, Dr Dick Barwick and Dr Richard Fox include
well preserved braincases and one articulated fish. Megalichthys was a cosmopolitan genus in
the Carboniferous, although the various species urgently need revision at present. A superbly
preserved snout of a cosmine-covered osteolepidid, also prepared by acetic acid, is known from
the Late Devonian Gogo Formation, Western Australia ("osteolepid", Gardiner & Miles 1975).
This specimen was described as a new genus, Gogonasus andrewsae (Fig. 36F; Long 1985d)
and shows some new anatomical features of the osteolepiform ethmosphenoid. Gogonasus is
of uncertain affinities but should be regarded as a fairly primitive osteolepidid at a similar level
of organization as Thursius from the Middle Devonian of Scotland and Russia (Jarvik 1948,
Vorobjeva 1977). In 1986 a complete head of Gogonasus was recovered from Gogo (Long
1987d, 1988c, d) and description of this important specimen should reveal much new
information on the structure of the osteolepiform head.

Eusthenopterid crossopterygians have cycloid thin scales with median bosses on the basal
surface (as do rhizodontiforms). The skull roof is narrow, lacking an extratemporal bone, and
with a longer frontal division in most genera (e.g. Eusthenopteron, Eusthenodon). A primitive
eusthenopterid has been described from the Late Devonian Mt Howitt site, Victoria, which has
cycloid scales but retains the extratemporal bone in the skull roof. Marsdenichthys
longioccipitus (P|. 6A; Fig. 37B) also has a relatively long, broad parietal shield, and in many
respects is typical of an intermediate form between osteolepids and eusthenopterids (Long
1985a). Other Australian eusthenopterids are known only from isolated scales in the Late
Devonian, as at Freestone Creek (Pl. 6B). A single maxilla and isolated cleithrum of a large
eusthenopterid were collected from the Hunter Siltstone (Famennian) of New South Wales by
Dr Alex Ritchie. The unusual shape of the maxilla suggests that these remains could belong
to Eusthenodon sp., otherwise known from the Famennian of East Greenland (Jarvik 1952,
1985). Another possible eusthenopterid is a skull from Late Devonian red sandstone near
Gosses Bluff, Northern Territory, also collected by Dr Ritchie, in 1986 (cf. Nomen Nudum
1986).

Rhizodontiforms are a very poorly known group throughout the world, with nearly all
known taxa established on isolated bones, teeth and scales (Andrews 1973). Australia has one
rhizodontiform, which Woodward (1906a) first described from the Lower Carboniferous
Mansfield Group of Victoria as "Strepsodus decipiens" (Fig. 37C, E). Woodward's material of
this fish consisted of scales, teeth, gulars and some shoulder girdle bones. New material of this
fish, including two skulls and a complete pectoral fin endoskeleton indicates that it is a new
genus, Barameda (Long 1989). Vorobjeva & Obrucheva (1977) referred this taxon to the
genus "Pyctnoctenion" on scale and tooth structure, but the type species of Pyctnoctenion
differs in the nature of the dermal ornament and shoulder girdle bones. The new material from
Mansfield indicates that the rhizodontiforms are very different from eusthenopterid
osteolepiforms and probably arose from the basal group containing Osteolepiformes and
tetrapods, as all three groups share specializations of the pectoral fin skeleton, but only the
latter two groups have choanae. The Mansfield rhizodont was a large fish, with a maximum
estimated length of about 3-4 metres. Recently, Andrews (1985) described the first articulated
rhizodontiform from the Carboniferous of Scotland. The pectoral fins are large and paddle-like
but all other fins are very small. Andrews suggested that the rhizodontiforms used their large
pectoral fins and specialized shoulder girdle to instigate rolling actions in the water for tearing
off large chunks of flesh from the prey, as crocodiles do today.

LONG HISTORY OF FOSSIL FISH - 389

Figure 37. Osteichthyes, Crossopterygii (Devonian-Carboniferous): A-B, eusthenopterid osteolepiform
Marsdenichthys longioccipitus Long 1985a, head restored in dorsal (A) and lateral (B) views, Late Devonian,
Victoria; C-E, Barameda decipiens from Mansfield, Lower Carboniferous, Victoria (Long 1989); tentative
restoration of head in (C) lateral, (D) dorsal and (E) anterior views. Scales approx. x 0.75.

Other undescribed crossopterygian material from the Devonian of Australia includes many
specimens of isolated teeth and scales, but articulated or partially articulated remains are rare.
An osteolepiform parietal shield is known from the Late Devonian Pambula River site, near
Eden, New South Wales, and a partially articulated body of an osteolepiform comes from the
Amadeus Basin, Northern Territory (both G. Young, pers. comm.). Three-dimensional
osteolepiform jaws have been recently recovered from the marine Gneudna Formation
(Frasnian, Carnarvon Basin, Western Australia).

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cS

C0
BAN CS
earotae:
TR TATE ZA

Figure 38. Osteichthyes, Crossopterygii (Devonian): A, Strunius, an onychodontiform; Similar forms
occur in the Early Devonian of Victoria and New South Wales; B, G/yptolepis. a porolepiform; glyptolepid-
like scales occur in Victoria and New South Wales; C, parietal shield of Onychodus sp. from Gogo, Western
Australia. Scale A (x2); B (x 1); C (x 1.5). (A after Jessen 1966; B after Jarvik 1980; C after Andrews

1973).

Actinistia are definitely known only from fragments of palate of Triassic age from the
Knocklofty Formation, near Hobart (Dziewa 1980), belonging to an indeterminate genus of the
family Coelacanthidae. A crushed head which may belong to a coelacanth is known from the
Late Devonian Mt Howitt site, Victoria, although exact identification cannot be made, and the
scales are similar to those of the porolepiform Glyptolepis (Fig. 38B shows the
head).Porolepiformes have not been formally described from Australia, but records of their
scales are known from the Devonian around Eden (including articulated remains of
Holoptychius sp., Fergusson et al. 1979) and glyptolepid-like scales (Pl. 6D) and dermal bones
have been found from the Late Devonian Mt Howitt site, Victoria (Long 1983c). The earliest
bone of a porolepiform in Australia is that of a single isolated lower jaw from the Emsian-
Eifelian Mulga Downs Group, New South Wales. Shoulder girdle bones of Porolepiformes are
also known from the Givetian? Bunga Beds, New South Wales (Ferguson et al. 1979), and

LONG HISTORY OF FOSSIL FISH - 391

from the Famennian Jemalong Gap fauna (G. Young, pers. comm.). Young (1985b) reported
porolepid scales from the Emsian Murrumbidgee Group, New South Wales. Porolepiform
scales with complex ridge patterns are known from the Late Devonian Hunter Siltstone, near
Grenfell, New South Wales.

Onychodontids are known from the Early Devonian Fairy Beds, near Buchan, Victoria,
represented by isolated bones and tooth whorls belonging to small struniiforms. Strunius, the
only complete member of this group is shown in Fig. 38A. @rvig (1969) recognized
struniiform bones from the Taemas limestones, New South Wales. The most complete
onychodontid material in the world comes from the Late Devonian Gogo Formation, Western
Australia. Acid prepared skulls of a new species of Onychodus are currently being studied by
Dr S.M. Andrews, Royal Scottish Museum. The Gogo species (Fig. 38C) was about a metre
long, with large tooth whorls at the front of the lower jaws for stabbing prey. These anterior
teeth were so large that they almost touched the skull roof when the mouth was closed.
Onychodontids may have occupied a similar niche to moray eels on the ancient reef, lurking in
crevices ready to lunge out at passing prey.

Dipnoi

Australia has one of the most complete records of fossil dipnoans anywhere in the world,
including the most diverse Early Devonian fauna from the Taemas-Wee Jasper region of New
South Wales, and perfectly preserved material from the Late Devonian Gogo Formation,
Western Australia. Lungfish fossils are known from every geological age since the Devonian
in Australia, except the Permian, of which we have little material of any fishes. Dipnoans or
lungfishes have a skull roof primitively covered by a mosaic of small bones, the jaw
articulation is autostylic with the palate fused to the braincase. The dentition consists of tooth-
plates (or dental plates) or the denticle-shedding type (Campbell & Barwick 1983, 1987). The
most common remains of dipnoans are the hard tooth-plates, which occur in the Mesozoic and
Tertiary deposits of Australia. The living Queensland lungfish, Neoceratodus forsteri, has been
inhabiting Australia since at least the Early Cretaceous. Toothplates indistinguishable from
those of the recent lungfish have been found at Lightning Ridge, New South Wales (Kemp &
Molnar 1981). Most of the major events in dipnoan evolution occurred during the Devonian,
in which the group underwent a large radiation (Westoll 1949). Two patterns emerged from
this, one of specialization of the feeding mechanism, such as rasping feeders (e.g.
Griphognathus, a marine form), and another line of freshwater Dipnoi with tooth plates, which
show specializations towards air-breathing (lengthening parasphenoid, mobile hyoid arch and
pectoral girdle, Campbell & Barwick 1987). As the marine dipnoans became extinct at the end
of the Devonian the freshwater groups flourished, although they remained fixed to a rigid body
plan and dentition style. Today, the living genera of lungfishes show this pattern very well: a
long body with simple diphycercal tail fin, and in the most advanced forms, degeneration of the
paired fins into sensory organs (Protopterus and Lepidosiren).

The Early Devonian dipnoans from the Taemas region of New South Wales are represented
by well preserved material, including three dimensional skulls, lower jaws, isolated bones and
scales. Two genera containing four species are known. Dipnorhynchus sussmilchi (P|. 7F;
Fig. 39A, C; Etheridge 1906) is the best known form and has also been recorded from Buchan,
Victoria (Hills 1936b, Thomson and Campbell 1971). This species was first properly
described by Hills (1933, 1943a) but the exact nature of the skull roof was not made clear until
Campbell (1965) described a complete skull. Thomson & Campbell (1971) redescribed
Dipnorhynchus sussmilchi in much more detail, and this has recently been added to
byCampbell & Barwick (1982b), who redescribed aspects of the neurocranial anatomy.
Dipnorhynchus kiandrensis (Fig. 39D) was described from the slightly older Lick Hole
Limestone, near Cooma, New South Wales. It is known from a single, almost complete skull

392 - LONG

median
symphysis

nasal
capsule
cavity

meckelian
fossa

Figure 39. Osteichthyes, Dipnoi (Early Devonian). Lungfishes from Taemas and Cooma, New South Wales:
A, C, Dipnorhynchus sussmilchi (Etheridge 1906); A, palatal view of skull; C, restored skull in dorsal view;
B, Speonesydrion iani Campbell & Barwick 1983, left half of lower jaw; D, Dipnorhynchus kiandrensis
Campbell & Barwick 1982, skull roof, partially restored. Scale approx. A (x 1); B (x 1.5); C (x 0.75); D &
0.8). (A, C, after Thomson & Campbell 1971; D, after Campbell & Barwick 1982a; B, after Campbell &

Barwick 1984b).

LONG HISTORY OF FOSSIL FISH - 393

N

M

od

K

X SS
Y14 Mea
Y2

2

Figure 40. Ostcichthyes, Dipnoi (Late Devonian): A-C, Chirodipterus australis Miles 1977, head in dorsal
(A) and lateral (B) views, Gogo, Wester Australia; C, toothplate of same from the Gneudna Formation,
Westem Australia ("Dipterus" sp. cf. D. digitatus Seddon 1969); D, F, two skulls from a new genus of lungfish
from Mt Howitt, Victoria, showing variation; E, G, Griphognathus whitei Miles 1977, skull in lateral (E) and
dorsal (G) views; H, Soederberghia sp., from the Cloughnan Shale, New South Wales. Scale approx. A, B,
H (x0.75); C, D (x 1); E, G (x 0.5); F (x 1.5). (A, B, E, G after Miles 1977; C after Seddon 1969; D, F after
original material; H after Campbell & Bell 1982)

394 - LONG

Figure 41. Osteichthyes, Dipnoi (Late Devonian). Attempted reconstruction of the heads of two
undescribed genera from the Mt Howitt Fauna, Victoria (after original material). A-C, from "A". D, E, form

"B". AIL natural size.

roof (Campbell & Barwick 1982a). A massive, broad-headed species, D. kurikae, was described
from partial cranial material by Campbell & Barwick (1985). The palate of Dipnorhynchus is a

LONG HISTORY OF FOSSIL FISH - 395

broad dentine covered sheet with small tuberosities and may represent an early stage in the
evolution of dipnoan tooth-plates (Campbell & Barwick 1983). Speonesydrion iani (Fig. 39B)
1s a contemporary of Dipnorhynchus which also occurs at Taemas, New South Wales. It
differs most noticeably in that the dentine-covered sheets on the palate and lower jaws have
differentiated tooth rows and is, therefore, intermediate between Dipnorhynchus and true tooth-
plated dipnoans. Speonesydrion was diagnosed by Campbell & Barwick (1983) and then fully
described by Campbell & Barwick (1984b). It is interesting to note that the earlicst lungfish
come from the Siegennian of North America, and that both denticle shedding types
(Uranolophus) and tooth-plated forms occur at this time (Denison 1968).

The Late Devonian deposits of Australia have produced several genera of lungfish, including
complete fish from the lacustrine Mt Howitt deposit, and complete skulls from the marine
Gogo Formation, Western Australia. Other lungfish remains of this age include the skull roof
of Soederberghia (Fig. 40H) from the Famennian Jemalong Gap site, New South Wales
(Campbell & Bell 1982), a genus also known from contemporaneous deposits in East
Greenland (Lehman 1959). Isolated toothplates from the Blue Range Formation, Taggerty,
Victoria were first named as a new genus, Eoctenodus microsoma_by Hills (1929), and later
referred to the British genus Dipterus (Hills 1931). Recent re-examination of the Taggerty
dipnoan material has shown that the tooth plates differ in morphology from Dipterus and the
parasphenoid is also slightly longer and of a different shape. The material was redescribed and
referred back to Eoctenodus microsoma Hills by Long (1987a). The marine Gneudna
Formation, Late Devonian, Western Australia, has yielded isolated toothplates, which were
earlier referred to as Dipterus cf. D. digitatus (Seddon 1969). These (Fig. 40C) closely
resemble those of Chirodipterus australis (Pl. 7B-C; Fig. 40A-B), from the contemporary Gogo
Formation, and probably belong to this species, although dipterid-type toothplates occur in this
formation as well. Hills (1936a) also records dipterid toothplates from near Gingham Gap in
the Hervey Group, New South Wales, which Long (1987a) concludes could not belong to the
genus Dipterus as similar toothplates occurring in the contemporaneous Hunter Siltstone occur
with long-stalked dipnoan parasphenoids that are quite different from those of Dipterus.

The Gogo Formation has produced both tooth-plated dipnoans and denticle shedders, with
most taxa belonging to cosmopolitan genera which also occur in Northern Hemisphere faunas.
Chirodipterus, a short headed, tooth-plated form is represented by two species, C. australis and
C, paddyensis (Miles 1977). Only one partial skull is known of C. paddyensis, which differs
in the dentition having deeply grooved tooth-plates. A new chirodipterid genus was recently
found from Gogo which has elongated, narrow toothplates and dentine on the parasphenoid
(Long 1988c). The other Gogo genera are denticle shedders. The long snouted Griphognathus
(Pl. 7A-D; Fig. 40E-G) was specialized for suctorial feeding and utilized the denticles lining the
plate, jaws and gill arches for macerating food (Campbell & Barwick 1987). The homology of
Griphognathus palate bones is discussed in Campbell & Barwick (1984a). Holodipterus (PI.
7G, H), a short snouted denticle-shedder, is less well known than the other Gogo lungfish,
although recently prepared new material in the Geology Department, The Australian National
University, should greatly add to our knowledge of this genus. So far, only the cranial
anatomy and dental structure (Campbell & Smith 1987, Smith & Campbell 1987) of these
fishes has been described, although newly prepared material in the Australian National
University should contribute to our understanding of the postcranial skeleton and squamation.
One Chirodipterus specimen has been prepared showing the delicate perichondrally ossified
pelvic girdle in place (Young, Barwick & Campbell, 1990). More knowledge of the structure
of early dipnoans is urgently needed in order to place them phylogenetically within the
Osteichthyes, and to sort out relationships of Devonian dipnoan families.

The Mt Howitt lungfish (Figs. 40D, F, 41) have not been formally described, although
Long (1983c) has figured one of the skull-roofs. Two genera are present, each having short

396 - LONG

<= 2 :

Figure 42. Osteichthyes, Dipnoi (Carboniferous-Triassic): A, Gosfordia truncata Woodward 1890 from the
Triassic of New South Wales; B, attempted restoration of the skull of Delatitia breviceps (Woodward 1906) in
dorsal view (Carboniferous, Victoria.). Scale A (x0.25; B (x 1). (A, after Ritchie 1981; B, after Long &

Campbell 1985).

snouts. One genus (Figs 40D-F; 41D-E) has a primitive occipital region of the skull roof in
which the I-bones may in some cases meet mesially behind the B-bone, otherwise only seen on
Early Devonian genera (Uranolophus, Dipnorhynchus, Speonesydrion). The other genus in the
fauna (Fig. 41A-C) has a skull roof pattern similar to Scaumenacia, although the A bone is
exceptionally large. This form has true toothplates developed, whereas the other form has a

LONG HISTORY OF FOSSIL FISH - 397

denticulate palate. The body shape of these genera are similar, with a Fleurantia-like tail and
separate anterior dorsal fin.

_ The only Carboniferous lungfish known from Australia comes from the Mansfield Group,
Victoria. Ctenodus breviceps (Fig. 42B) was described from a single skull roof by Woodward
(1906a), but recently has been redescribed following preparation of the original material as a
new genus, Delatitia (Long & Campbell 1985). Delatitia differs from Crenodus by the
arrangement of the YI-Y2-Z bones and in having an occipital sensory-line canal running off the
posterior of the I bones. In these respects it is more primitive than any other Carboniferous
dipnoan and suggests that the ctenodontid group could have originated in Gondwana, a view
supported by the presence of Eoctenodus, another primitive ctenodontid, in the Late Devonian
of Victoria (Long 1987a).

From the Triassic Brookvale fauna of New South Wales comes a complete specimen of a
dipnoan, Gosfordia truncata (Fig. 42A; Woodward 1890, Ritchie 1981). Gosfordia was a deep-
bodied ceratodontid which already had achieved a body plan similar to the living lungfish by
having merged its dorsal and anal fins with the caudal fin. Jurassic and Cretaceous dipnoans are
well known by their toothplates. These, together with Tertiary dipnoans, are reviewed in a
separate section of this book by Dr Anne Kemp, and will not be considered here (Mesozoic and
Tertiary lungfish, Chap. 14).

AUSTRALIAN PALAEOZOIC FISH BIOSTRATIGRAPHY AND
BIOGEOGRAPHY

Fishes are particularly useful for dating and correlating middle Palaeozoic (Silurian-
Carboniferous) rock successions. Turner (this volume, Chap. 13) gives detailed accounts of the
use of thelodonts and other vertebrate microfossils in biostratigraphy in Australia (particularly
useful for Early - Middle Devonian correlation). Here I will briefly review the use of
macrovertebrate faunas for correlation in the Devonian and Carboniferous of Australia. It
should be remembered that combinations of both microvertebrates and macrovertebrates, when
present at single localities, enable the most precise age determinations.

Correlations between local (regional) successions are usually based on similar species
occurring in different localities (¢.g. similar Bothriolepis species at Taggerty and Mt Howitt in
Victoria, Long 1983a, Long & Werdelin 1986). Faunal zones (biozones) are based on the first
appearance or entry of a particular species or genus, or alternatively on the presence of several
species at the same time (concurrent range zones). Dating of sequences is, thus, based on the
relative time occurrences of faunas, some of which may be tied into sequences which have
volcanic rocks that have been dated using radiometric methods (absolute time dating). The
thick volcanic succession of the Cerberean Cauldron near Taggerty, Victoria has several layers
dated this way as well as a fish fauna low in the sequence, enabling correlation with other
faunas throughout Gippsland (Fig. 43) (Long & Werdelin 1986). Consideration of the
taphonomy of the fossil site is useful when using fish assemblages in biostratigraphy, as sites
with a low diversity of species may be biased in terms of preservation of the whole fauna. In
other words, the absence of a key taxon must be established as a real absence rather than as an
artifact of the size of the collection or field collecting methods (e.g. only collecting big fish
plates, etc.).

Correlations between major sedimentary basins in different states of Australia are based on
assemblages of genera, or similar faunas. A good example of this is the Wuttagoonaspis
fauna from the Mulga Downs Group, near Cobar, New South Wales. Ritchie (1973) described

398 - LONG

ZONES CENTRAL VICTORIAN § || MT. HowITT FREESTONE CK.

| VOLCANIC PROVINCE

TAGGERTY

SNOWY PLAINS FM

367-369

CERBEREAN MT.KENT CONG

VOLCANICS Spier upper Freestone
mudstone Ck Fauna

WELLINGTON

TATONG ! lower Freestone
abi Sek tr Aes Bindaree Rd Fauna Ck Fauna
upper cong.

HOLLANDS CK Wagesriy Fauna Mt Howitt Fauna RHYOLITE
RHYODACITE lower mudstone MOROKA GLEN FM

BLUE RANGE FM
lower cong

conianr aren SNOBS CK
Tatong Fauna VOLCANICS ay oso, VOleanics| ~~

. basal cong WIGHTMANS HILL CONG, U-
seers — OREMASPPIE

z
=x
Zz
o)
>
tf
fa)
Ww
5

FRASNIAN

Tabberabberan Orogeny

GIVETIAN

MOUNT HOWIT PROVINCE

MID DEV.

Figure 43. Biostratigraphic correlations between the Central Victorian Volcanic Province and the Mt
Howitt Province, Late Devonian, Victoria. Thick black lines indicate confident correlation between
stratigraphic horizons as denoted by common presence of Bothriolepis species. Wavy line represents
unconformity. Bb, B. bindarei; Bc, B. cullodenensis; Bg, B. gippslandiensis, Bt, B. tatongensis; Bw, B.
warreni. Scale approx.x4. (From Long & Werdelin 1986).

the unusual arthrodire, which is the namesake of this faunal assemblage and noted its
association with other endemic placoderm types. Since then Turner et al. (1981) have described
thelodont scales occurring with Wuttagoonaspis from New South Wales and western
Queensland (Cravens Peak Beds, Georgina Basin), thus correlating these widely-spaced
successions (Fig. 44). Similar thelodont scales have since been recognized in drill cores from
the Officer Basin, South Australia (Long et al. 1988), and the Amadeus Basin of the Northern
Territory (Young et al. 1987). The Wuttagoonaspis fauna is generally regarded as latest Early
Devonian or earliest Middle Devonian in age, based on the occurrence of conodonts in marine
intercalations within the Mulga Downs Group (Long et al. 1988).

In Australia the entry and disappearance of key placoderm genera differs from those of the
standard zonation schemes based on faunal successions in Europe, North America, East
Greenland and Russia (Young 1974). The reason for this is only recently becoming clear as
more is known of the phylogenetic and biogeographic relationships of the Australian and
Antarctic Palaeozoic fish faunas. Young (1981a) defined five Devonian faunal provinces based
on endemic populations of fishes (Fig. 45). Australia and Antarctica constitute the East
Gondwana Province, which in Devonian times was the antipodes of North America and Europe

LONG HISTORY OF FOSSIL FISH - 399

2
Oa
D2
a0
za
20

i)

2
So
33
5 &

Go
g ®
o a
] o
o
Uf

i)
5

Ss 0

oe

8

o
=|

© 4

€

°

=

400 - LONG

3

: 4
22>
/, ZS TARIM

INDOCHINA
USSR

ORS ; KAZAKHSTAN QS
EURAMERICA

* ARMORICA TIBET
ney ~=x—
A

wy

Route 1 M-Late Devonian
Groenlandaspis
Bothriolepids
Phyllolepids

Route 2 Late Devonian
Asterolepis

Route 3 Late Devonian

SOUTH AMERICA AFRICA

EURAMERICA ; :

Sone Pypvince so Route 4 ios Boone
f% ke ; Z UTH CHINA ° A

- Amphiaspid Province YW ~Galeaspid - Yunnanolepid Province siniepigs

TUVA \] EAST GONDWANA
- Tannuaspid Province - Wuttagoonaspid - Phyllolepid Province

Figure 45. Devonian vertebrate provinces, modified after Young (1981a), to show possible migration
routes for some biostratigraphically useful placoderms.

(Euramerican Province). As in modern times faunas from widely separated parts of the world
tend to differ because of differing groups having evolved in one region. For this reason, the
phyllolepid placoderms occur in Australia and Antarctica at an earlier time than for Euramerica,
and are represented by more primitive types. Thus, the likely explanation is that this group
originated in East Gondwana and did not migrate to Euramerica until a later time. Similarly,
certain fishes which are abundant in Euramerica, such as cephalaspid agnathans, are not known
to occur in Australia, not because we haven't looked hard enough for them, but probably
simply because the group was restricted to Euramerica and did not migrate from that region.
The bothriolepid antiarchs make their earliest appearance in East Gondwana (Eifelian
Monarolepis) and South China (Givetian Bothriolepis), and did not become abundant in
Euramerica until the Late Devonian. Asterolepis, a common antiarch in Europe and Russia in
the Middle Devonian, is absent in Australia but restricted to the Late Devonian in China (Pan
Kiang 1981). Groenlandaspis occurs at the very end of the Famennian in Euramerica, but is
found as early as Givetian in East Gondwana (Young 1988b).

The biostratigraphic ranges of Palaeozoic vertebrates useful for correlations in Australia and
Antarctica are shown in Figs 46, 47. Young (1988b) set up a detailed biostratigraphy for the
Devonian Aztec Siltstone, South Victoria Land, Antarctica, recognizing several taxa which also
occur on the Australian mainland. Note that these are for essentially continental facies, as
marine deposits are usually more precisely dated using microfossils such as conodonts or

palynomorphs.

LONG HISTORY OF FOSSIL FISH - 401

PLACODERMI CHONDRICHTHYES OSTEICHTHYES

myrs

AGNATHA

ACANTHODII

(No Australian

245

Perm. acanthodians)
290

CARB. PERMIAN

360

LATE DEV.

374

MID. DEV.

DEVONIAN

387

EARLY DEV.

408

SILURIAN

Isch = ischnacanthid Ant = Antarctilamna
scales Har = Harpagodens

T= Taemasacanthus Ph= Phoebodus

Ro = Rockycampacanthus McM = McMurdodus

Cul = Culmacanthus Heli = Helicoprion

Ch = Cheiracanthoides Helo = Helodus Dip = dipnorhynchid

Tur = turiniids Gy = Gyracanthides Both = Bothriolepis lungfishes
(thelodonts) H= lowittacanthus Gr= Groenlandaspis Lig = Ligulalepis scales

Ac= Acanthodes P?= Phyllolepis Rhiz =rhizodontiforms
Aus = Austrophyllolepis How = Howqualepis
Rem = Remigolepis Mars = Marsdenichthys
Sin = sinolepid Hol = Holoptychius
Sh= Sherbonaspis Ony = Onychoaontids
W= Wuttagoonaspis Meg = Megalichthys

Figure 46. Stratigraphic ranges of some Australian Palaeozoic fishes.

The Middle-Late Devonian continental deposits of southeastern Australia often contain fish
faunas typified by the presence or absence of the following key taxa: Bothriolepis (primitive
or advanced species), Groenlandaspis, phyllolepids, Remigolepis, and in one locality, a
sinolepid. Middle Devonian faunas in New South Wales and Antarctica contain Bothriolepis
associated with thelodont scales, and Groenlandaspis makes its appearance in the Givetian
(Aztec Siltstone, Antarctica). The presence of primitive Bothriolepis species with
Groenlandaspis and phyllolepids such as Austrophyllolepis (Long 1984), without thelodont
scales, is typical of a Frasnian assemblage, as seen at Mt Howitt and Freestone Creek in
Victoria. Thelodonts last on until the Frasnian in marine deposits of Western Australia (Turner
& Dring 1981), but the youngest forms from eastern Australia are probably of Middle
Devonian age (Broken River, Queensland). Famennian assemblages are characterized by the
entry of Remigolepis, with advanced Bothriolepis species, and the genus Phyllolepis probably
replaces the primitive phyllolepid genera Austrophyllolepis and Placolepis . Luse the word
probably here, because although phyllolepids are definitely known to occur in

402 - LONG

EAST GONDWANA SOUTH CHINA EURAMERICA

= nil
TZ
ZZ

sinolepid ———
Groenlandaspis

Remigolepis =<

IS
1S
1S

Phyllolepis =

Sinolep

Q
a)
2
D
2
a
<x

Remigolep

Bothriolepis

Frasnian Famennian

__
Tet

Bothriolepis

phyllolepids =
phyllolepids
Pambulaspis

Givetian

Groenlandaspis —— —
Asterolepis

Bothriolepis

Bothriolepis
Groenlandaspis

Eifelian

primitive
bothriolepid

MIDDLE DEVONIAN LATE DEVONIAN

Figure 47. Placoderm succession in Antarctica and eastern Australia (after Young 1974, 1988b, 1989b,
Long 1983a, 1984b, Long & Werdelin 1986) compared with Euramerican and Chinese faunas.

Famennian deposits in New South Wales (e.g. Jemalong Gap fauna), only fragmentary plates
have been found , which do not allow identification of the genus. Late Famennian faunas
record the entry of a sinolepid antiarch (Grenfell fauna) or the disappearance of Bothriolepis
with the entry of the osteichthyan Holoptychius (Worange Point faunas). Comparative age
ranges for East Gondwana placoderm faunas with those from South China and Euramerica are
shown in Fig. 47.

Other fishes which are not as well known may become valuable zone fossils as work on the
many undescribed Devonian faunas of Australia continues. The acanthodian Culmacanthus
from Mt Howitt, Victoria (Long 1983d), has recently been recognized from New South Wales
and Antarctica by Young (in press 1), and is easily recognized from fragmentary skull bones,
which possess a highly characteristic ornamentation.

The Carboniferous marine deposits of Western Australia, Queensland and New South Wales
are easily dated by marine microfossils, but also are rich in microvertebrates, such as sharks’

LONG HISTORY OF FOSSIL FISH - 403

teeth and scales. Such faunas have been described by Turner (1982e and in Chap. 13 this
volume) and enable age determination of samples from new localities, which may be lacking
other age diagnostic fossils. Carboniferous vertebrate macrofaunas typically lack placoderms,
which died out at the end of the Devonian, and may contain shark remains with palaconiscoid
debris (e.g. Drummond Basin, Queensland, Turner & Long 1987). Research into the
Carboniferous vertebrates of Australia, however, is only in its infancy, and biostratigraphic
schemes are only provisional at best.

REFERENCES

For a complete listing of references dealing with Australian fossil fishes see Long & Turner
(1984). The following list is of cited and useful references only.

ABELE, C., 1976. Tertiary. In: Geology of Victoria. J.G. Douglas & J.A. Fergusson, eds., Geol. Soc. Aust.
spec. Pub. 5: 177-273.

ALEXANDER, R.McN., 1967. Functional Design in Fishes. Hutchinson, London.

ie i Y., 1969. Function and Gross Morphology in Fish. Israel Program for Scientific Translations,
erusalem.

ANDREWS, S.M., GARDINER, B.G., MILES, R.S. & PATTERSON, C., 1967. Pisces. In The Fossil Record.
Harland, W. B., Gilbert-Smith, B. & Wilcock, B., eds., Geol. Soc. London: 637-683.

ANDREWS, S.M., 1973. Interrelationships of crossopterygians. In Interrelationships of Fishes. P.H.
Greenwood, R.S. Miles & C. Patterson, eds., Academic Press, London: 137-177.

ANDREWS, S.M., 1985. Rhizodontiform fish from the Dinantian of Foulden, Berwickshire. Trans. R. Soc.
Edinb. 76: 67-95.

ARCHBOLD, N., 1980. Fish scales from the Permian of Westem Australia. J. R. Soc. West. Aust. 64: 23-26.

BANKS, M.P., COSGRIFF, J.W. & KEMP, N.R., 1978. A Tasmanian Triassic stream community. Aust. Nat.
Hist. 19: 150-157.

BARDACK, D., 1962. Taxonomic status and geological position of the Cretaceous fish Ichthyodectes
marathonensis. Aust. J. Sci. 24: 387-8.

BARDACK, D., 1965. Anatomy and evolution of chirocentrid fishes. Paleont. Contr. Univ. Kans.Vertebrata
10: 1-88.

BARTHOLOMAT, A., 1969. The Lower Cretaceous elopoid fish Pachyrhizodus marathonensis (Etheridge Jr.).
In Stratigraphy and Palaeontology. K.S.W. Campbell, ed., Aust. Nat. Univ. Press, Canberra: 249-263.

BARTRAM, A.W.H., 1977. The Macrosemiidae, a Mesozoic family of holostean fishes. Bull. Br. Mus. nat.
Hist., Geol. 29: 137-234.

BENDIX-ALMGREEN, S.E., 1966. New investigations on Helicoprion from the Phosphora Formation of
south-east Idaho, U.S.A. Biol. Skr. 14 (5): 1-54.

BLIECK, A., 1984. Les hétérostraces ptéraspidiformes agnathes du Silurten-Dévonien du continent Nord-
Atlantique et des blocs avoisinants. Revision systématique, phylogénie, biostratigraphie,
biogéographie. Cahiers Paleontol. CNRS, Paris.

BLIECK, A., GOUJET, D. & JANVIER, P., 1987. The vertebrate stratigraphy of the Lower Devonian (Red Bay
Group and Wood Bay Formation) of Spitsbergen. Modern Geol. 11: 197-217.

BLIECK, A., JANVIER, P., LELIEVRE, H., MISTIAEN, B. & MONTENAT, C., 1982. Vertébrés du Dévonien
Supérieur d'Afghanistan. Bull. Mus. nat. Hist., Paris. 4 (4), C: 3-19.

BURIAN, Z. & AUGUSTA, J., 1965. Prehistoric Life on Earth. 7th Ed. Paul Hamlyn, London.

CAMPBELL, K.S.W., 1965. An almost complete skull roof and palate of the dipnoan Dipnorhynchus
sussmilchi (Etheridge). Palaeontology 8: 634-7.

CAMPBELL, K.S.W. & BARWICK, R.E., 1982a. A new species of the lungfish Dipnorhynchus from the Early
Devonian of New South Wales. Palaeontology 25: 509-527.

CAMPBELL, K.S.W. & BARWICK, R.E., 1982b. The neurocranium of the primitive dipnoan Dipnorhynchus
sussmilchi (Etheridge). J. Vert. Paleo. 2: 286-327.

CAMPBELL, K.S.W. & BARWICK, R.E., 1983. Early evolution of dipnoan dentitions and a new genus
Speonesydrion. Mem. Ass. Australas. Palaeontols. 1: 17-49.

CAMPBELL, K.S.W. & BARWICK, R.E., 1984a. The choana, maxillae, premaxillae and anterior palatal
bones of early dipnoans. Proc. Linn. Soc. N.S.W. 107: 147-70.

CAMPBELL, K.S.W. & BARWICK, R.E., 1984b. Speonesydrion, an Early Devonian dipnoan with primitive
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404 - LONG

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LONG, PA, 1986b A new Late Devonian acanthodian from Mt Howitt, Victoria, with remarks on the
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LONG, PTA 197d Aw unusual osteolepifonm fish from the Late Devonian of Victoria, Australia
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YOUNG, G.C., 1987d. Relationships between the northern and southern vertebrate faunas during the Middle
Palaeozoic. Int. Symp. Shallow Tethys 2, A.A. Balkema, Rotterdam: 79-85.

YOUNG, G.C., 1988a. New occurrences of phyllolepid placoderms from the Devonian of central Australia.
Bur. Miner. Res. J. Aust. Geol. Geophys. 10: 363-376.

YOUNG, G.C., 1988b. Antiarchs (placoderm fishes) from the Devonian Aztec Siltstone, Southem Victoria
Land, Antarctica. Palaeontographica 202: 1-125.

YOUNG, G.C., 1989a. New occurrences of culmacanthid acanthodians (Pisces, Devonian) from Antarctica and
south-eastern Australia. Proc. Linn. Soc. N.S.W. 111: 12-25.

YOUNG, G.C., 1989b. The Aztec fish fauna (Devonian) of Southern Victoria Land: Evolutionary and
biogeographic significance. In: Origins and Evolution of the Antarctic Biota, Crane, J. A., ed., Geol.
Soc. Spec. Publ. 47: 43-62.

YOUNG, G.C., 1990. New antiarchs (Devonian placoderm fishes) from Queensland, with comments on
placoderm phylogeny and biogeography. De Vis Symposium Volume, Mems. Qd. Mus.

YOUNG, G.C., BARWICK, R.E. & CAMPBELL, K.S.W., 1990 Pelvic girdles of lungfishes (Dipnoi). E£.S.
Hills Volume, R. LeMaitre, ed., Blackwell Press: 59-75.

YOUNG, G.C. & GORTER, J.D., 1981. A new fish fauna of Middle Devonian age from the Taemas/Wee Jasper
region of New South Wales. Bull. Bur. Min. Res. Geol. Geophys. 209: 85-128.

YOUNG, G.C., LONG, J.A. & RITCHIE, A., in press. Crossopterygian fishes from the Devonian Aztec
Siltstone, Antarctica: systematics and evolutionary significance. Rec. Aust. Mus.

YOUNG, G.C., TURNER, S., OWEN, M., NICOLL, R.S., LAURIE, J.R. & GORTER, J.D. 1987. A new
Devonian fish fauna, and revision of post-Ordovician stratigraphy in the Ross River Syncline, Amadeus
Basin, central Australia. Bur. Min. Res. J. Geol. Geophys. Aust. 10: 233-242.

ZHANG GUORUI, 1978. The antiarchs from the early Devonian of Yunnan. Vertebr. Palasiat. 16: 147-186,

ZHANG GUORUI, 1984. New form of Antiarchi with primitive brachial process from Early Devonian of
Yunnan. Vertebr. Palasiat. 22: 81-91.

414 - LONG

ZANGERL, R., 1981. Chondrichthyes 1. In Handbook of Paleoichthyology, H.-P. Schultze, ed., Gustav
Fischer Verlag, Stuttgart.

ZIDEK, J., 1976. Kansas Hamilton Quarry (Upper Pennsylvanian) Acanthodes with remarks on the previously
reported North American occurrences of the genus. Paleont. Contr. Univ. Kansas 83: 1-41.

APPENDIX I: CLASSIFICATION

The classification of fishes has, for some time, been the subject of dispute, and this has been compounded recently by the
addition of cladistic classifications. The use of such classifications should be restricted until agreement has been reached by
most workers on an acceptable hypothesis of piscine interrelationships. By accepting a broad dichotomy of gnathostomes
into two major groups (shark-like fishes or Elasmobranchiomorphs, and bony fishes or Teleostomes) it is relatively easy to
break down the fishes into morphotypic higher groups, despite disagreement as to their status of being classes, subclasses or
infraclasses (e.g. Moy-Thomas & Miles 1971, Andrews et al. 1967, Jarvik 1980).

In this paper the following provisional classification is used, with more subdivision of higher taxonomic groups
presented, where particularly relevant, to the Australasian record. I have avoided trying to sort out the agnathan classification
because we have so very few here in the fossil record. Tumer (1976) has provided a classification of the thelodonts.

The classification for chondrichthyans given here is modified from Zangerl (1981) and Lund (1986), but applies
primarily to Palaeozoic forms. See Kemp, this volume, for neoselachians. Agnathan classification needs revision. A
cladistic approach to this problem is given by Forey (1984), part of which is adopted here. The following arrangement gives
approximate positions of major groups only. * denotes groups absent from the Australian record, and these, therefore, are
not further subdivided.

Agnathan classification needs revision. A cladistic approach to this problem is given by Forey (1984), part of
which is adopted here. The following arrangement gives approximate positions of the major groups only.

SUBPHYLUM CRANIATA

INFRAPHYLUM MYXINOIDEA (hagfishes)
INFRAPHYLUM VERTEBRATA
Plesion: "'HETEROSTRACI"
Order Arandaspidiformes
SUPERCLASS PETROMYZONTIA
Order Petromyzontiformes (lampreys)
Order Anaspida
SUBCLASS THELODONTI
Order Thelodontida
Family Thelodontidae
Family Turiniidae
Family Apalolepididae
Order Katoporididae

SUPERCLASS GNATHOSTOMATA
Plesion: "EUGALEASPIDA"*
Plesion: "OSTEOSTRACI"*

CLASS CHONDRICHTHYES (sharks and rays)
SUBCLASS ELASMOBRANCHII (sharks)
Order Euselachii
Family Ctenacanthidae
Family Bandringidae *
Family Phoebodontidae
Superfamily Hybodontoidea

LONG HISTORY OF FOSSIL FISH - 415

Superfamily Protacrodontoidea
Order Desmiodontida *
Order Xenacanthida
Family Diploselachidae *
Family Xenacanthidae
Order Cladoselachida
Family Cladoselachidae
Order Coronodontida *
Order Symmoriida
Family Symmoriidae
Family Stethacanthidae
Order Eugeneodontida
Superfamily Caseodontoidea *
Superfamily Edestoidea
Family Agassizodontidae
Family Edestidae *
Order Orodontida
Family Orodontidae
Order Squatinactida *

SUBCLASS HOLOCEPHALI (chimaerids)
Order Chimaeriformes
Order Cocliodontiformes
Suborder Myriacanthoidei *
Suborder Cochliodontoidei
Family Cochliodontidae
Family Menaspidae *
Order Squalorajiformes *
Order Chondrenchelyformes *
Order Helodontiformes
Order Petalodontiformes
Family Petalodontidae
Family Peripristidae *
Family Pristodontidae *
Family Janassidae *

CLASS PLACODERMI (extinct Devonian armoured fishes)
Superorder Palaeacanthaspidoidei
Order Acanthothoraci
Order Rhenanida
Order Petalichthyomorpha
Superorder Dolichothoracomorpha
Order Arthrodira (Euarthrodira)
Suborder Actinolepidoidei
Infraorder Phyllolepidi
Infraorder Wuttagoonaspidi
Infraorder Actinolepidi
Suborder Phlyctaenioidei
Infraorder Brachythoraci
Family Heterosteidae
Family Buchanosteidae
Family Homosteidae
Family Brachydeiridae *
Family Leptosteidae *
Family Coccosteidae *
Family Plourdosteidae
Family Incisoscutidae

416 - LONG

Family Camuropiscidae
Family Pholidosteidae *
Family Rachiosteidae *
Family Dinichthyidae
Family Leiosteidae *
Family Trematosteidae *
Family Selenosteidae *
Family Titanichthyidae *
Family Bungartiidae *
Order Antiarcha
Suborder Yunnanolepidoidei*
Suborder Bothriolepidoidei
Superfamily Sinolepidoidea
Superfamily Bothriolepidoidea
Family Microbrachiidae
Family Bothriolepididae
Suborder Asterolepidoidei
Family Pterichthyodidae
Family Asterolepididae

CLASS TELEOSTOMI (fishes with an Internal swim-bladder)
SUBCLASS ACANTHODII (extinct spiney fishes)
Order Ischnacanthida
Family Ischnacanthidae
Order Climatiida
Suborder Climatioidei
Family Climatiidae
Family Gyracanthidae
Suborder Diplacanthoidei
Family Diplacanthidae
Family Culmacanthidae
Order Acanthodida
Family Mesacanthidae*
Family Cheiracanthidae
Family Acanthodidae

SUBCLASS OSTEICHTHYES (true bony fishes)
Superorder Actinopterygii (ray finned fishes)

Order Palaeonisciformes
Family Stegotrachelidae

Family Gonatodidae
Family Elonichthyidae
Family Tegeolepididae
Family Acrolepididae
Family Palaeoniscidae
Family Platysomidae
Family Bobasatraniidae
Family Coccolepididae
Family Urosthenidae

There are many more families of primitive actinopterygians (e.g. Kasantseva 1982), the above list contains those
families in which Australian forms can be placed.

Suborder Pholidopleuroidei
Family Pholidopleuridae
Order Redfieldiiformes

LONG HISTORY

Family Brookvaliidae
Family Redfieldiidae
Order Perleidiformes
Family Colobodontidae
Family Cleithrolepidae
Family Pholidophoridae
Order ?Acipenseriformes
Family Saurichthyidae
Order Aspidorhynchiformes
Family Aspidorhynchidae
Order Semionotiformes
Suborder Semionotidoidei
Family Semionotidae
Family Parasemionotidae
Family Macrosemionotidae

Order Pholidophoriformes
Family Archaeomaenidae
SUBCLASS TELEOSTEI
Superorder Leptolepomorpha
Order Leptolepiformes
Family Leptolepidae
Superorder Elopomorpha
Order Elopiformes
Suborder Elopoidei
Family Elopidae
Family Megalopidae
Suborder Albuloidei
Family Albulidae
Order Anguilliformes
Suborder Anguilloidei
Family Congridae
Family Heterenchelyidae
Family Muraenesocidae
Superorder Clupeomorpha
Order Clupeiformes
Family Ichthyodectidae
Family Pachyrhizodontidae
Suborder Clupeodei
Family Koonwariidae
Superorder Osteoglossomorpha
Order Osteoglossiformes
Suborder Osteoglossoidei
Family Osteoglossidae
Superorder Protacanthopterygii
Order Gonorhynchiformes
Suborder Gonorhynchoidei
Family Gonorhynchidae
Superorder Ostariophysi
Order Siluriformes
Family Plotosidae
Superorder Paracanthopterygii
Order Gadiformes
Suborder Gadoidei
Family Gadidae
Family Bregmacerotidae
Family Merlucciidae

OF FOSSIL FISH

- 417

418 - LONG

Suborder Ophidoidei
Family Ophidiidae
Family Carapidae
Suborder Macrouroidei
Family Macrouridae
(Coryphaenoididae)
Superorder Acanthopterygii
Order Beryciformes
Suborder Berycoidei
Family Caproidae
Family Trachichthyidae
Family Monocentridae
Order Scorpaeniformes
Suborder Scorpaeniodei
Family Scorpaenidae
Suborder Platycephaloidei
Family Platycephalidae
Order Perciformes
Suborder Percoidei
Family Centropomidae
Family Lutjanidae
Family Theraponidae
Family Sillaginiidae
Family Lactariidae
Family Sparidae
Family Oplegnathidae
Suborder Labroidei
Family Labridae
Family Sphyraenidae
Family Scombroidei
Order Pleuronectiformes
Suborder Pleuronectoidei
Family Pleuronectidae
Order Tetradontiformes
Suborder Tetradontoidei
Family Diodontidae

The above groups contain an enormous number of taxa and subdivision of their ranks is cumbersome. I avoid further
breakdown of these groups but refer readers to papers by Patterson (1973, 1982), Patterson & Rosen (1977) and Lauder &
Liem (1983) for recent appraisal of the classification of higher actinopterygians.

Superorder Crossopterygii
Order Onychodontiformes (Struniiformes)
Order Osteolepiformes (Osteolepidida)
Suborder Osteolepidoidei

Family Osteolepididae
Family Megalichthyidae
Family Lamprotolepidae *
Family Eusthenopteridae
Family Panderichthyidae*
Family Rhizodopsidae*
Family Canowindridae

Order Porolepiformes
Family Porolepidae
Family Holoptychiidae

LONG HISTORY OF FOSSIL FISH - 419

Order Rhizodontida
Family Rhizodontidae

Order Actinistia
Family Coelacanthidae

Superorder Dipnoi (lungfishes)

Formal classification needs revision. The following groups, known from Australia, are taken from the
classification of Miles (1977).

Family Dipnorhynchidae
Family Dipteridae

Family Chirodipteridae
Family Rhynchodipteridae
Family Ctenodontidae

Family Proteroceratodontidae
Family Neocdratodontidae

APPENDIX II: ABBREVIATIONS USED IN FIGURES

Ad adnasal bone Eth ethmoid bone
Adp accessory dermopterotic Ex extrascapular bones
AL anterior lateral plate Fr frontal ‘bones
ADL anterior dorsolateral plate fh h hysial f
AMD anterior median dorsal plate ms PEE le tae ie
Ang angular bone Hy hyomandibular bone
AOD antorbital bone he post-suborbital sensory-canal
AVL anterior ventrolateral plate D infradentary
ac.Sq accessory squamosal bones If, inf thal
alt anterolateral thickening on 8B expan

headshield IL interolateral plate
BR branchiostegal rays D infraorbital bones
bhf buccohypophysial foramen lop interopercular bone
bpt basipterygoid process ifc infraorbital sensory canal
CE central plate if.pt infranuchal pit
aa ceratohyal if.r infranuchal ridge

cleithruam Ju jugal bone
Cla clavicle IA lateral plate
CV1,2 central ventral plates lL lachrymal bone
cir circular pit-line groove LEx lateral extrascapular bone
or.PNU postorbital crista le main lateral line canal
csc central sensory-canal lda groove for lateral dorsal aorta
Den dentary aN see "Ic"
Dh dermohyal MD median dorsal plate
Dpt asnopteroué Mes mesethmoid bone

P } M.Ex median extrascapular bone

Ds dermosphenotic MG marginal plate
d.end opening of endolymphatic ML mixilateral plate

duct 5 :
dig dorsolateral sensory-line isin’ sneesalhiiar ginal plates

Mpt metapterygoid
ag foreal peg onacals , me mandibular canal
EL ee a (submarginal) mdop median depueasion
Ep epiotic bone m.gr median groove on braincase
mpl middle pit-line groove

H extratemporal bone Mx waaealla

420 - LONG

Na nasal bone pre.f precerebral fontanelle of
NU nuchal plate braincase
4 ( . pr.ot otic process of braincase
masa) opening pr-po postorbital process of
OP opercular bone braincase
oa. CE etc. overlap areas for certain plates @ ateteate bate,
orb orbit or orbital cavity Q quadsatajage? bane
P pineal plate (placod qipl quadratojugal pit-line groove
Ps paraorbital plate i rostral. plate: or bone
Par parietal bone RP rostro-pineal plate
POL posterior dorsolateral plate 4 squamosal bone
PF prefrontal bone bo rostral sensory canal
AL posterior lateral plate r.csc ridge for central canal
PM premaxilla (premaxillary) Sb suborbital bones
PMD posterior median dorsal plate Sc sclerotic bones
PMG postmarginal plate scl cleithrum
PMV posterior median ventral plate — m
FN postnasal plate SM submarginal plate
PNU paranuchal plate Mx. supramaxillary bone
PO postorbital bones Sop subopercular
POP preopercular bone SP (Sp) spinal plate (placoderms) or
re postpineal plate spiracular bone
PR (osteichthyans)
postrostral bone Ssc suprascapular bone (=PT)
PRL ecard Piste soc supraorbital sensory-canal
PRM le
PPO preorbital plate socc supraoccipital sensory-canal
PS prespiracular bone Sorc supraoral sensory-canal
PSM preoperculo-submandibur pow suborbital vault
bone sub suborbital canal
PSO postsuborbital plate : : :
: th.pp prepineal thickening
PSP parasphenoid er th.scc dermal thickening over
Psp postspiracular os labyrinth cavity
PTO postorbital plate -

Pto pterotic bone I foramen for the optic nerve
PVL posterior ventrolateral plate bite foramen for the oculomotor
ra-articular process nerve
ike i n i x 10th cranial nerve (vagus)
pmc postmarginal sensory-canal foraines

ppl posterior pit-line canal

PLATES

Plate 1. Acanthodii (Devonian). A, Howittacanthus kentoni Long 1986b (A, D, E from the Late Devonian
near Mt Howitt, Victoria.). B, scale of Cheiracanthoides from the Middle Devonian of Queensland (courtesy Dr
Sue Tumer). C, scale of Nostolepis (C, G from the Early Devonian of Buchan, Victoria.). D, E, Culmacanthus
stewarli Long 1983b. D, detail of scales, E, head in side view showing armoured cheek plates. F, G,
ischnacanthid jaw bones. F, Taemasacanthus erroli Long 1986a, Early Devonian near Taemas, New South
Wales G, Rockycampacanthus milesi Long 1986a.

Plate 2. Placodermi (Early Devonian). A, B, Buchanosteus confertituberculatus (Chapman 1916). A,

headshield and braincase in ventral view. B, trunkshield in lateral view (A-D, G from Buchan, Victoria.). C,
Wijdeaspis spinal plate. D, nuchal plate of Taemasosteus maclartiensis Long 1984c, ventral view. E, F,
complete ossified optic capsule of a palaeacanthaspidoid, from Taemas, New South Wales E, visceral view, F,
lateral view. G, detail of omament of Murrindalaspis bairdi Long 1984c.

Plate 3. Placodermi (Late Devonian). A, Remigolepis sp. headshield, Canowindra, New South Wales. B,
armour in lateral view of Bothriolepis gippslandiensis Hills 1929 from Mi Howitt, Victoria. C, headshield of
Eastmanosteus calliaspis Dennis-Bryan from Gogo, Westem Australia. Photograph of Remigolepis sp.
courtesy of Dr Alex Ritchie, Australian Museum.

.)

LONG HISTORY OF FOSSIL FISH - 421

Plate 4. Placodermi (Late Devonian). Arthrodires from Gogo. A, Eastmanosteus calliaspis (Dennis-Bryan
1987) armour in lateral view; B, Latocamurus coulthardi (Long 1988b) armour in dorsal view; C, D, new genus
of plourdosteid. C, headshield in dorsal view; D, inferognathal bone.

Plate 5. Osteichthyes, Actinopterygii (Palaeozoic). A, Liqulalepis toombsi scale, Middle Devonian,
Queensland. B, Head of Howqualepis rostridens Long (1988e) Late Devonian Mt Howitt, Victoria. C,
palaeoniscoid jaw, Lower Carboniferous, Drummond Range, Queensland. D, Mansfieldiscus sweeti (Woodward
1906), new preparation of head, Lower Carboniferous, Mansfield, Victoria. Photograph of Liqulalepis scale
courtesy of Dr Sue Tumer, Queensland Museum.

Plate 6. Osteichthyes, Crossopterygii (Devonian-Carboniferous). A, B, eusthenopterid osteolepiforms. A,
Marsdenichthys longioccipitus Long 1985a, head from Mt Howitt, Victoria. B, scale from Freestone Creek,
Victoria. C, E, F, Beelarongia patrichae Long 1987d, from Mt Howitt. C, scale. E, skull. F, cleithrum. D,
porolepiform scale cf. Glyptolepis sp., from Mt Howitt. G, scale of Megalichthys sp., Drummond Range,
Queensland, Carboniferous. From original material.

Plate 7. Osteichthyes, Dipnoi (Devonian). Fossil marine lungfishes from the Late Devonian Gogo
Formation (all except F)., Western Australia; and from the Early Devonian of New South Wales (F). A, D,
Griphognathus whitei Miles 1977. A, skull, D, scale. B, C, E, Chirodipterus australis Miles 1977. Skull in
dorsal (B) and (C) palatal views. E, scale. F, Dipnorhynchus sussmilchi (Etheridge 1906), skull in lateral
view. G, H, Holodipterus gogoensis Miles 1977. G, scale. H, anterior of skull showing endocranial cavities
lined by bone for the olfactory nerves. All specimens photographed by the author through the courtesy of
Prof. Ken Campbell, The Australian National University, Canberra.

422 6 LONG PLATE 1

PLATE 2 LONG HISTORY OF FOSSIL FISH - 423

424 - LONG PLATE

- 425

HISTORY OF FOSSIL FISH

~
x

LONC

PLATE 4

426 - LONG PLATE

PLATE 6 LONG HISTORY OF FOSSIL FISH - 427

PLATE 7

428 - LONG

CHAPTER 13

PALAEOZOIC VERTEBRATE
MICROFOSSILS IN
AUSTRALASIA

Susan Turner !

IMPMOGUCHOM 5.05. esc osiatasatdictiletloettieasete 430
INFEGH OS) 0.12 Ai eiote Bebe Mita aol 430
Identification and Taxonomy.................. 431
ASMAMNANS sce cenceence eee ep eeg eigenen ts 431
PRE TOCOMUSE fe. Ooesde Peale ences ctete hee ghee se 434
Chondrichthyans.................ceceeceeeeees 440
Elasmobranchs............cccccececeee eee 440
Xenacanths............. ccc eeee eee e ee 440
Cladoselachians..................065 440
"“CGladodontssh...tu..se mes 440
Selachians ............c. cece cee eeee eee 441
Ctenacanthoids ................. 441
Hybodontoids...............2.5. 441
Ageleodonts...............eeeee 441
Neosclachians.............0..068 441
Holocephalians................c0006 441
Helodontoids.................005 442
Menaspoids............ccceeeeees 442

Edestids? 2.2 ss. 0:58 Ses aeexhess 442
Petalodonts..............eceeeeee 442

PlaCOGENMS \....525. 0s sche wed bergen veh ee fee ced 442
TeElCOStOMES ®.....6 sce cseenes noon deer enestuvest 442
Acanthodtans....2.c.c60c:ceceesseeeceseeed 442
@limatidss « 22ss¢ct ose eedie ctalcess 443
Ischnacanthids ................eeeeees 443
Acanthodids ..............ccceeeseeeee 443

Higher Osteichthycs..............008 444
Aclinopterygians.......... sees 444

Sarcopterygians ................e eee 444
Crossopterygians .............. 444

Coelacanths ................ 445

Dipnoans ..........0..cceeeeeeeeee eee 445

SUMMARY. cctevee suds teaece canngatvedameainnanas 445
The Australasian Microvertebrate Record. .446
OrdOvician ..........c..ccceeceeceeceeeeeeeeeees 446
STUIUTIAN ete ecccct ea ceseteeticrens tas gddbtageass 446
DeVOMIAN......... cece cece cecece ee eeeeeeceees 447
Thelodonts ....5..........cc ccc ee cee eeeeeeee 447
Chondrichthyans ..................0.0065 448
PlacodermS...............ccccceeceeeeeeee 450
Teleostomes.............c cee eee cece cece ee 450
Acanthodians ................0.00ee ee 450

Higher Osteichthyans. ............. 451
Carbomniferous...............ccceceeceeese neues 451
Pe@rimb an oo .'3..) dove eeten cis do's ce Sole sivsawee ee teied'et 452
SUMMALY +5 ess renee eeeisdt Bibeeeibaeascazsd 453
Acknowledgements ..............c.ceeeeeeeeeeees 454
REFEKENCES 5 08. ce Pad red intitle ed wietiel dala as 455
Plates: ely dele shee ccoe eats feet ose 458

1 Queensland Museum, P.O. Box 300, South Brisbane, Queensland 4101 , Australia.

430 - TURNER

INTRODUCTION

Vertebrate microremains (microvertebrates or ichthyoliths) include scales, spines, teeth,
ornamented and unornamented dermal bones and endoskeletal bones of vertebrates. In
Australasian Palaeozoic rocks these remains come almost exclusively from fishes, although by
the late Palaeozoic, amphibian and reptile remains might be expected. Microvertebrates possess
all the advantages of other microfossils for use in biostratigraphical studies. They include a
range of hard parts from ancient organisms which are not well-represented in the equivalent
macrofossil record. For example, articulated fossils of agnathans, placoderms and sharks do not
always exhibit well-preserved body scales or teeth, whereas acid-prepared residues might include
such microremains.

METHODS

Microfossils, by definition, are minute and are not easily observed with the naked eye.
Thus, a range of techniques and methods can be employed to extract them from the rocks.
There follows a section outlining some of the common methods for obtaining vertebrate
microfossils. Microvertebrates can be found in, or extracted from, small or large samples of
sediment, Thelodont scales, in particular, are highly resistant to many geological processes
and preparation methods, surviving low-grade metamorphism as well as treatment with dilute
hydrofluoric acid.

When prospecting a new region for vertebrate microfossils, the various sedimentary rocks
are assessed for possible success. Searching shallow-water, or detrital, limestones often
provides the best opportunity to recover vertebrate microremains, but any lime-rich rock should
be sampled and prepared. Other rock types including dolomites, siltstones, sandstones, etc.
should be examined in the field for possible vertebrate content. Any bed which has yielded
macroremains could also contain microremains. One of the advantages of microvertebrates is
the existence of recognizable remains in very small samples of rock, such as shot-point or core
samples from boreholes, and hand specimens held in museum collections.

Any technique for preparing rock samples for phosphatic microfossils can be used to look
for ichthyoliths (see Moore 1962, Kummel & Raup 1965, Rixon 1976). The common
method of preparation is the acetic acid technique (Rixon 1976); approximately 10% or less
acetic acid (some workers use monochloracetic) solution is used to digest lime-containing
rocks. Phosphatic remains are picked from the ensuing washed residues using a fine brush or
forceps; some form of lens or binocular microscope is necessary for this process. The
vertebrate remains are then transferred to either small tubes or cavity cell slides for storage. An
alternative method is to use dilute formic acid for the "stewing-up" process. This method is
faster and can be used for dolomitic rocks, but it is highly recommended that one works in a
fume cupboard and takes greater manual care. Before placing samples into the acid solution,
larger pieces of bone should be coated with a protective coating (e.g. Bedacryl, Mowital B30 or
polyvinylacetate, see Whitelaw & Kool, this volume, on preparation techniques). This process
is repeated after each washing of the rock sample. Rock samples can remain in acetic acid
solution for about three days before the solution needs changing. Recent work on conodont
preparation has shown the need for a buffer with the acetic acid solution (Jeppson & Fredholm
1987); the easiest to use is the remaining calcium acetate solution from the previous washing,
which, therefore, should be reserved.

Where fish scales occur in sandstone, dilute hydrofluoric acid can be tried to dissolve the
rock (working in a fume cupboard, of course). Thelodont scales and possibly other, more
resistant, dentinous scales can survive this treatment, However, mechanical preparation of

PALAEOZOIC VERTEBRATE MICROFOSSILS - 431

vertebrate microremains from sandstone is probably safer unless a large, inexhaustible supply
is available. Mechanical preparation from rock, or cleaning off remaining sediment from
remains can be accomplished using fine needles and dental tools. Another technique used with
vertebrate fossils preserved in sandstone is to dissolve away the bone/dentine with dilute
hydrochloric acid, and then to make a cast from the resulting mould.

Ultrasound and freeze-thaw techniques (the latter can be done in your own backyard in colder
or extreme climates where day and night temperatures differ considerably) can be useful in
preparing microvertebrates from siltstones and sandstones. Concentrated detergent or bleach
have also been used to break up noncalcareous sedimentary rocks.

Once the loose-grained residue is obtained, this can then be further treated to concentrate the
heavy minerals, thus reducing search time under the microscope. Murray & Lezak (1977) have
summarised some of the heavy mineral separation techniques for recovery of vertebrate
microfossils. Where a fume cupboard is available, current usage favours tetrabromoethane.
Freeman (1982) devised a method which could be used by the scientist working at home.
Vertebrate remains usually occur in the heavy fractions obtained from these processes. Care
should be taken, however, not to discard any of the residue, for some scales (e.g. extremely
small ones of thelodonts, sharks and acanthodians) can occur in the light fractions.

IDENTIFICATION AND TAXONOMY

Nearly all groups of fishes can be found as vertebrate microfossils, which comprise scales,
spines (including modified forms such as pharyngial denticles, claspers and other copulation
devices),.teeth, unornamented and ornamented bones and platelets (Figs 1-7, Pls 1-5). Very
few fish are entirely naked or without hard tissues either in their exo- or endoskeleton;
examples include embryonic and juvenile fishes and some of the cartilaginous fishes, as well as
lampreys and hagfishes.

In this section each fish group will be examined to summarize what sort of remains can be
expected and how to make preliminary identification of the remains (see also Table 1). I have
followed a general grouping of fishes, and not any one specific classification. This has been
done to avoid lengthy discussion of the characteristics of fish types. Some words used in
popular parlance, "sharks" for example, have only a very general meaning in science where
some might equate "shark" with chondrichthyans, that is, the cartilaginous fishes. There are,
in fact, many "shark" fossils, particularly in the Palaeozoic, which are not completely known
and whose relationships are still obscure. For more general information and a formal
classification of fish see Long (this volume).

AGNATHANS

The agnathans comprise the "jawless" fishes and include extinct forms ranging in age from
?Late Cambrian to Late Devonian. Included in this group are heterostracans and heterostracan-
like fishes, thelodonts, cephalaspids or osteostracans, anaspids, galeaspids, as well as fossil and
modern forms of lampreys and hagfish. Recent reviews of agnathans include Janvier (1981),
Young (1981) and Archer & Clayton (1984) and Long (this volume).

The mid-Ordovician heterostracan-like fish found in North and South America and in
Australia had a dermal armour composed of small tesserae (platelets) with simple tubercular or
leaf-like dentine omament, onan acellular base (e.g. Ritchie & Gilbert-Tomlinson 1977,

Table 1. Notes to enable preliminary identification of the microfossils of the major fish
groups.

432 - TURNER

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PALAEOZOIC VERTEBRATE MICROFOSSILS - 433

Gagnier et al. 1986). These scales can appear as ichthyoliths (Fig. 1).

Apart from the strange heterostracan-like jawless fishes found in the mid-Ordovician of
Australia and North and South America, the only other agnathans known to exist in the
Palaeozoic of the Gondwanan continents, including Australasia, are the scales of Devonian
thelodonts (see e.g. Turner 1982a). Only one possible example of the strange Chinese
galeaspidiform fishes has been found in Australia (Young, pers. comm.), and, as the
identity of this fossil is still tentative, this group will not be considered here. No agnathans
have yet been found in the Silurian of Australia.

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Figure 1. Example of mid-Ordovician vertebrates yielding microremains. A, Arandaspis restoration (about
120-140 mm long); x 1; B, Arandaspis, close-up of dermal omament; x 20; C, Arandaspis scales; x 1.5; D,
Porophoraspis - dermal tubercles; x 35; all from Ritchie & Gilbert-Tomlinson 1977.

Thelodonts are worthy of a detailed discussion because their scales have proved useful
biostratigraphical indices (Turner 1973). However, recent findings from Australia and other
Gondwana countries, such as Antarctica and South America (e.g. Gagnier ef al. 1988), suggest
that in the mid-Devonian thelodonts underwent rapid speciation. Further analysis of this event
may lead to a better definition of thelodont zone fossils throughout the Devonian. Table 2
summarises current knowledge and ideas about the geographic and stratigraphical ranges of
thelodonts in Australia.

434 - TURNER
Thelodonts

I consider the Thelodonti to be a monophyletic group. It is usually placed as a subclass of
Agnatha, and its range extends from the Late Ordovician to early Late Devonian. The name
"Thelo" comes from the Greek word meaning mammal/nipple, and "dont" - tooth, in reference
to the similarity of the scales' external and internal structure to those of mammal teeth. Hence,
the name Thelodus was given to the type genus by Louis Agassiz (Turner 1976).

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Table 2. Geographic and stratigraphical distribution of thelodont scale assemblages in
Australia. Symbols: +, new turiniid species and scales comparable with Turinia polita Karatajute-
Talimaa (1978); *, Turinia fuscina Turner 1986b; dot, Turinia australiensis and close relatives;
square, new species of turiniid with body scales commonly bearing crown lappets; point-down
triangle, turiniid first described and figured by Young & Gorter 1981; triangle, Australolepis
seddoni Turner & Dring 1981.

Thelodont fish were small, with an average length around 150 mm. Some forms, however,
reached 1 metre. Thelodonts were aquatic, living in shallow-water marine to freshwater
environments. The thelodont body was covered with an external skeleton of minute (average
0.5-2.0 mm in length) dentinous scales. Thelodonts resembled modern sharks in this

Figure 2. Comparison of a selection of trunk scales from some of the known turiniid populations (from
Tumer 1986b ; for full details see her Fig. 3). A-C, L, type turiniid Turinia pagei, Gedinnian to Siegenian,
Welsh Borderland; D-E, Turinia polita, Gedinnian, England, France, Podolia; F, Turinia fuscina, Gedinnian?,
Victoria; G-H, Turinia australiensis, Emsian-Eifelian?, Western Australia, west Queensland and New South
Wales; I, K, M, new turiniid species, Cravens Peak Beds, ?Late Emsian-Eifelian? Queensland; J, Turinia
hutkensis, mid-Devonian or Early Frasnian?, Iran; N, Turinia? sp. nov., Late Eifelian - Givetian, Hatchery
Creek Conglomerate, NSW and north Queensland; O-P, Australolepis seddoni, Early Frasnian, Western
Australia. Not to scale; all turiniid thelodont scales depicted are within length range of 0.5-2.0mm.

PALAEOZOIC VERTEBRATE MICROFOSSILS - 435

436 - TURNER

covering of shagreen and in their lack of bony internal skeleton. Complete, articulated
thelodonts are rare, but the scales provide microfossils with a size range from 0.1-3.5 mm,
commonly about 1 mm long, deep and wide on average. Both overlapping and non-overlapping
scales are known. Generally, the scales from the head, mid-thorax and body are differently
shaped (e.g. Figs 2, 3). They can be recognized for each species by associating similar
morphology and histology and by comparing with scales on intact thelodont specimens. Each
scale is formed of hydroxylapatite and is made up of a crown, a neck and a base (see Fig.
4A). The crown and neck are made of orthodentine or allied dentinous tissue, penetrated by

Figure 3. Thelodont scales from Australia - types of head, transitional and special scales. A, Australolepis
seddoni with deeply dissected crown; B, Australolepis seddoni? with tripartite crown; C, Australolepis seddoni,
ventral view showing large pulp opening and shallow basal ring; D, turiniid, new species related to Turinia
australiensis, head scale ; E, turiniid, new species related to Turinia australiensis, head scale; F, turiniid, new
species related to Turinia australiensis, transitional scale. Not to scale; A-C after Turner & Dring 1981, Early
Frasnian Gneudna Formation, Western Australia; D-F after Tumer et al. 1981, Late Emsian-Early Eifelian?,
Cravens Peak Beds, westem Queensland; E after Young ef al. 1987, Early Eifelian? N'Dahla Member, Amadeus
Basin. Scale approx. 1 mm.

PALAEOZOIC VERTEBRATE MICROFOSSILS - 437

Figure 4. Thelodonts of Australia - some body scale types. A, generalized turiniid body scale - c = crown,
n = neck, b = base, o = pulp opening; B, G, turiniid new species with smooth wing-like lateral lappets; C,
turiniid new species with close double-ridged crown comparable to Turinia hutkensis of Iran; D, typical Turinia
australiensis body scale; E, F, turiniid new species related to Turinia australiensis with complex stepped
crowns and lateral posteriorly-expanded lappets - this form closely resembles turiniid body scales from
Antarctica and Bolivia; H, turiniid species possibly close to that in B & G. Not to scale; B, D-G after Tumer
et al., Late Emsian-Early Eifelian Cravens Peak Beds, western Queensland; C after Long et al. 1988, Early
Eifelian?, Munyurai, South Australia; H after Young ef al. 1987, N'Dahla Member, Amadeus Basin.

Scale approx. 1 mm.

438 - TURNER

Figure 5. Examples of shark scales: A-D, Antarctilamna prisca after Young 1982, mid Devonian, NSW,
Antarctica, Bolivia?; E, Ohiolepis after Wells 1944, a form typical of late Early Devonian in Australia; F,
Cladolepis after Wells 1944, late Early Devonian of USA; G-H, shark scales fam., gen. et sp. undet., Cravens
Peak Beds, Late Emsian-Early Eifelian?, western Queensland; I-J, shark scales fam., gen. et sp. undet., mid
Devonian, Broken River Formation, north Queensland; K, shark scales fam., gen. et sp. undet., Early
Devonian, New South Wales; L, shark scales fam., gen. et sp. undet. referred by Giffin 1980 to Skamolepis,
Zlichovian, New South Wales. Scale approx. 1 mm.

PALAEOZOIC VERTEBRATE MICROFOSSILS - 439

dentine tubules; dentine tubules merge into dentine canals, which in turn can converge into, on
average, one to three pulp canals. The base is made up of a clear bony tissue (probably akin to
aspidine), which was capable of growth, and is penetrated by the spaces occupied by Sharpey's
fibres in life. These fibres held the scale into position in the skin tissues. Characteristic of
thelodont scales is a large, single (or few) pulp canal opening in the base; this feature separates
them from the acanthodian scales which lack a pulp cavity altogether and are commonly found
together with thelodont scales. The lack of any neck canals distinguishes them from most, if
not all, shark scales. One proviso here is the lack of knowledge of primitive shark scales,
some of which seem to resemble simple thelodont scales. However, the few known, early
shark scales all have a simple, small, diamond-shaped base, unlike the growing base of
thelodonts (see Fig. 5). The style of histological growth is a feature of classification.
Common to all thelodont scales, and also a unique character for the Thelodonti, was the

erat to produce basal outgrowths, such as ‘roots' and papillae, to anchor the scales in the
lermis.

The ontogeny of the scale resembles that of human and other mammalian teeth and modern
shark scales, i.e. all simple placoid (plate-like) structures. The scale begins as a thin, cap-like
structure, and dentine is added centripetally. This means the crown and neck remain the same
size throughout the life of the scale. The inital cap is free of tubules and forms a clear outer
layer to the crown, sometimes called durodentine. The basal tissue can expand and grow
outwards and downwards to eventually obscure the basal openings in very mature scales. As in
modern sharks, scales were produced continually throughout the life of the fish, dropping out
when abraded or damaged; new scales then took their place.

Because thelodont fish were like sharks, being virtually only a soft bag of tissue covered
with scales, when they died, the scales would scatter. Complete fish are rarely preserved. As
scale variation is high, this means that there are problems in recognizing species from scales.
Recent work has done much to sort out this problem, either by bringing to light new material
with scales in situ (e.g. Turner 1986a) or by assigning scales to theoretical positions on the
body by association of morphological and histological characters (Marss 1986). The families,
genera and species are based on positioning of pulp openings, base, crown etc., and on minute
detail of crown ornament (Tumer, in press a, b).

There were three main groups of thelodont scales, based on overall morphology and
histology. These are thelodontids, katoporids and loganiids. Only the thelodontids occur in
Australasia. The thelodontids, such as Thelodus, Turinia, Australolepis, and Nikolivia, were
the longest lasting of the theiodont groups, ranging in age from the Early Silurian to Late
Devonian. They possessed relatively simple orthodentine scales with dentine tubules
converging straight into a single or few pulp canals.

In Australasia thelodontids are represented by the turiniids (Figs 2-4) and, possibly, the
nikoliviids. Australasian turiniids often had very large body scales, up to 3.5 mm, which did
not usually overlap. The head scales tend to be smaller with simple rounded shapes and a
crenulated crown rim (Fig. 3). Body scales can be simple with a smooth crown with a central
platform, but many of the body scales had more strongly ornamented, partitioned and ridged
crowns, which became deeply dissected in the later Devonian. Bases in turiniid scales tend to
be solid with a long anterior spur. The nikoliviids had more flattened crowns, which
overlapped in life. The crowns are drop- and arrowhead-shaped and sometimes had lateral
lappets with posterior points. Bases are annular and shallow around a large, oval anteriorly-
placed pulp opening. A few scales from the mid to Late Devonian in Australia appear to be
nikoliviid, but these might be highly modified turiniid scales exhibiting a parallel morphology
(Fig. 3A, B).

440 - TURNER

CHONDRICHTHYANS

The cartilaginous sharks and their relatives have a long and complex history. Only recently
have pre-Middle Devonian shark remains been recognised. Cartilage is not easily preserved but
is sometimes calcified. Teeth, finspines, claspers, eggcases and scales can all give us clues to
the shapes and lifestyles of early sharks. Most of these remains can occur as microfossils.
Shark scales come in many different designs. However, whether complex or simple in crown
structure, and some sharks apparently grew both sorts on one body, they all possess neck
canals. The teeth also come in many forms but essentially can be divided into the cutting and
grasping, pointed or cusped teeth, and the more solid-plated or rounded, crushing and gnawing
or nipping teeth. Information on many Palaeozoic sharks can be found in Zangerl (1981).

Shark scales exhibit a wide range of form and structure, depending on the group from which
they come. Many primitive sharks and most modern sharks have very simple placoid scales
(the basic lepidomorium or odontode). From the mid-Devonian onwards more complex shark
scales occur, which belong to the ctenacanth, stethacanth and other "cladodont" sharks as well
as to forms such as edestids. Examples are given here of some of the range of variation (Fig.
5). All shark scales should possess neck canals, a dentinous crown and, usually, an acellular
bony base which is non-growing and diamond-shaped. Complex shark scales may have a
bony, growing base, with fused dentine ridges forming the crown.

Shark spines, which appear to be modified and enlarged denticles, can appear as
microfossils. They tend to have a deep insertion to the body. A sheath of dentine may cover
the spine producing ribbed or tubercular omament. A large pulp cavity is present.

Palaeozoic chondrichthyan fish classification or relationships are still uncertain and so, in
this paper, the sharks are referred to only general groupings. Microfossils have come from the
following groups:

Elasmobranchs - True Sharks and Their Relatives.

A primitive group that appeared in the Silurian in Siberia is still known only from simple
scales and teeth similar to some from living primitive sharks. Australian examples from the
Devonian include Thrinacodus (Fig. 6), Mcmurdodus (Pl. 1), and scales (Fig. 5K, L).

XENACANTHS - Early Devonian to Early Triassic, youngest in Australia; advanced teeth
with characteristic diplodont (two-pronged) condition having a large "button" on the dorsal side
of base; some species possess dorsal head spines but none have been found yet in the
microfauna. (Antarctilamna(?), Xenacanthus, Orthacanthus; Phoebodus might belong
here).

CLADOSELACHIANS - Late Devonian to Late Carboniferous. Some complete specimens
of Cladoselache are known from the Cleveland Shale of the U.S.A. Cladoselachians possess
relatively simple cusped teeth; scales with large, complex, ridged crowns and generally a large
bony base; dorsal fin spines unoramented. Possible scales known in Australia.

"CLADODONTS" - ?Early Devonian to Permian. Mostly known from isolated teeth
("Cladodus"), but recent finds of complete specimens and re-examination of teeth characters are
allowing subdivision of this "bucket" group. One family includes the stethacanthids
(Symmorium, Stethacanthus, Denaea) found in the Early Carboniferous deposits of eastern
Australia. Protacrodus with ridged and crested teeth, and Phoebodus, with tricuspid teeth with
striated enamel and a large lingual extension to the base might also belong here. The scales of
some of these genera have been known for over a century but were given separate names (form

PALAEOZOIC VERTEBRATE MICROFOSSILS - 441

or organ genera); Stethacanthus includes pharyngial scales, called Stemmatodus, and strange
spines (earlier called Lambdodus) from the brush-like dorsal head apparatus; scales are generally
complex, with large bony bases.

SELACHIANS - mid-Devonian to Recent.
Ctenacanthoids - Late Devonian to Triassic? (Bandringia, Ctenacanthus,
Tristychius); complex scales. Ctenacanth scales are found in the Late Devonian in Australia.
Hybodontoids - Early Devonian? to Upper Cretaceous (Hybodus). Relatively
simple placoid scales with one or more pairs of neck canals and a simple small diamond-shaped
base.
Ageleodonts - Carboniferous (Ageleodus). Only known from slow-growing, hand
or comb-shaped teeth with a long root. The earliest record of the group comes from Australia.
Neoselachians - Early Carboniferous (Anachronistes) or earlier to Recent (most
living forms). Only placoid scales have been found in the Australasian Palaeozoic microfauna.

Figure 6. Thinacodus ("Harpagodens") ferox, shark teeth known from the Late Devonian (Famennian) to
Early Carboniferous (Early Visean), Queensland, New South Wales and Westerm Australia. Scale bar approx. 1
mm. (After Turner 1982b and see Tumer 1983).

HOLOCEPHALIANS (“undivided head") - chimaeroids (rat-fishes) and their relatives; often
called bradyodonts in the literature. Many of the Palaeozoic fossils are only known from teeth,
many of which consist of tubular dentine, producing a characteristic pitted surface.

442 - TURNER

Helodontoids - Late Devonian to Permian (Helodus, Pleurodus). Dome-like teeth
and scales occur as microfossils.

Menaspoids - Early Carboniferous to Permian (Menaspis, Deltoptychius). Teeth
and scales known as microfossils. [Menaspis also thought to be a placoderm]

Edestids - Early Carboniferous to mid-Permian (Edestus, Agassizodus, Orodus).
Teeth and scales are known as microfossils. [or could be elasmobranchs - Zangerl 1981]

Petalodonts - Early Carboniferous to Permian (Petalodus, Janassa, Polyrhizodus
(?)). Teeth and scales occur as microfossils. [or could be elasmobranchs - Zangerl 1981]

PLACODERMS

This was a group of bony, jawed, armoured fish which possessed an endoskeleton of bone
and/or cartilage (see Long 1984). These extinct fishes dominated seas and freshwaters
throughout the Devonian and became extinct in the Early Carboniferous, although their origins
are undoubtedly in or before the Early Silurian, where they have been found in China. Some
were only a few centimeters long; some grew to 6 m in total length. Most had a bony
carapace over head and thorax encompassing brain, branchial structures and well-developed
jaws. The outer surface of the armour was often ornamented, and the rest of the body often
covered in intricate body scales. Some placoderms had moveable pectoral appendages; some
had immoveable spinal plates. Each group is characterised by the style and pattern of the
bony plates and scales. For more information see Denison (1978).

Many placoderms, especially the primitive forms, had bony body scales, generally round or
rhombic, with ornament of tubercles and ridges similar to that of the main dermal plates (PI.
2). Only recently have we begun to relate dissociated body scales to genera known from
articulated plates; in much of the literature isolated scales have been referred to form genera
(e.g. Ohioaspis). Many body scales are mere nubs of bone, as in Bothriolepis (Long &
Werdelin 1986); many are tuberculated platelets ~ 2-3 mm ; some, those of an asterolepidoid
antiarch (Young 1984), for example, are very distinctive (Pl. 4F). Other microfossils include
modified jawbones with "tooth" plates called gnathals, and spines. In the ptyctodontid
placoderms the tooth plates were composed of a tubular dentine, a modification for a shell-
crushing way of life.

Placoderm fossils have been of much use in biostratigraphy in Devonian rocks, but, as yet,
little use has been made of the microfossils. In the future, placoderm body scales and jaw
elements might be useful for dating Silurian and Early Devonian sequences.

TELESTOMES - JAWED BONY FISHES

Acanthodians

The spiny sharks were primitive fusiform teleostomes (bony fishes) which possessed jaws,
paired fins, an ossified neurocranium and a separate gill skeleton. Most were small, averaging
200 mm, but forms up to 2 m are reported. Acanthodians had a body covering of small bony
scales and platelets, very tightly packed in regular criss-cross rows (Pl. 2A). The scales are
small (usually between 0.5 to 3 mm), non-overlapping, and composed mostly of bone or a
bone-like tissue called mesodentine. Special scales, or denticles, occurred on the gill arches in
a few forms. These scales have no basal pulp cavity as do thelodont and some shark scales,
but some, probably lateral line scales of acanthodians do have neck canal openings.

The scales can be differentiated by their ornament and histology. They possess a relatively
square crown, a shallow neck and, usually, a deep, rounded base. The scale is made up of

PALAEOZOIC VERTEBRATE MICROFOSSILS - 443

centrifugally-produced, concentric layers of bone and dentinal tissue. Crown ornament may be
characteristic of both species and genus. There are two main types of scale. The Nostolepis-
type has a mesodentine crown penetrated by vascular canals and enclosing cell spaces. In some
species the dentine ridges of the crown are added laterally. The base is formed of cellular bone.
The Acanthodes-type has a crown of true dentine with no cell spaces and a thick base of
acellular bone. Vascular and non-vascular (canals of Sharpey's fibres) penetrate the scale.

The scales were generally acquired at an early stage in the fish's growth and presumably
grew continuously throughout life. Each scale added successive bone and/or dentine layers in
the manner of modern fishes, but whether these were annual rings, or not, cannot be
ascertained.

Acanthodians were the only Palaeozoic fish to possess paired fin spines. Acanthodian fins
were supported by dentine-ribbed (in some), bony spines, which were usually triangular in
cross section. The fin spines were capable of some growth and were formed of three or four
layers, a superficial, or sculptured, layer of centripetal orthodentine or mesodentine, without
enameloid, which can form ribs; a middle layer of cellular bone or trabecular dentine, where
longitudinal canals can form a subcostal canal and there can be radial canals; a thin basal layer
lining the central cavity of cellular bone, or of dentine, if present. The central cavity can open
posteriorly in the lower end of the spine (see Denison 1979 for further details).

Acanthodians had jaws, some had sharp pointed teeth, and others had multicuspid piercing
and cutting teeth ankylosed (directly attached) along the jaw. Others had tooth whorls. Other
microfossils include bony, unsegmented, unbranched rays in the fins and bony vertebral arches.
For more information see Denison (1979).

Much work remains to be done on acanthodian scales and spines - their structure, variation,
and stratigraphical distribution. Many new forms are being found in Australasia. They appear
to be generally useful (Valiukevicius 1985), as they were common in marine and some
freshwater environments. They can help in the broad allocation of an age for rocks from mid-
Silurian times through to Permian.

CLIMATIIDS - Mid-Silurian to Carboniferous? Climatioids (Climatius, Cheiracanthoides,
Nostolepis) and diplacanthioids (Diplacanthus-like scales). The climatiids were probably the
most primitive acanthodians. They had short bodies with thick, high-crowned scales, well-
developed ancillary gill covers, broad, highly-sculptured fin spines and intermediate spines that
were restricted to the skin. All possessed two dorsal fins. Nostolepis—type scales and ventral
shoulder girdle platelets appear in the Australian mid- Silurian and are common in the Early
Devonian.

ISCHNACANTHIDS - Late Silurian to Carboniferous ([schnacanthus, Gomphonchus,
Poracanthodes). Ischnacanthids were long-bodied predators with Acanthodes-type scales. They
possess two dorsal fins. All fin spines deeply inserted into the body and no intermediate
spines, shoulder plates, or accessory gill covers were present. Both toothwhorls and
multicuspid teeth ankylosed to the jaws can occur.

ACANTHODIDS - Early Devonian to Permian (Acanthodes, C heiracanthus). Acanthodids
were long-bodied, filter-feeding forms with no teeth, no dermal plates, and Acanthodes—type
scales. Only one dorsal fin and a few, small intermediate fins were present. Fin spines
inserted deeply, except in primitive forms. Gill rakers were present, with an enlarged gill cover
plate in some species.

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Higher Osteichthyes

These fish have a skeleton of true bone. The upper biting edge of the mouth is formed by
two pairs of dermal bones, the maxilla and premaxilla. Teeth are typically fused to the bones.
The lateral-line canals run through dermal bones. The skull has a cranial fissure. Ossified
vertebrae and pleural ribs and an air bladder which can function as a lung are present. Included
in this group are actinopterygians, crossopterygians and dipnoans.

ACTINOPTERGYIANS - The bony ray-finned fishes have fin webs stiffened with dermal
rays called lepidotrichia. Thick ridge scales formed an upper, epichordal tail lobe.
Evolutionary tendencies in the group, including reduction in thickness of dermal bones and
scales and increase in variety of swimming, feeding and reproduction styles, have led to their
success in marine and freshwater with over 23,000 extant species.

Scales and spiny bones, which might belong to early actinopterygians appear as
microfossils in the Late Silurian of China and Baltic Europe (e.g. Lophosteus, Andreolepis).
More taxa are recorded in the Early to mid-Devonian: in Australasia there are scales called
Ligulalepis (Fig. 7), in Europe Orvikuina (Schultze 1968). These bony scales are elongate,
rectangular and ornamented. The Devonian and Carboniferous forms belong mainly to the
family Palaeconiscidae. These had ornamented, rhombic scales and bones, and teeth with a
transparent acrodin tip, which commonly occur as microfossils. The great actinopterygian
radiation began in the Carboniferous. During Permian times they began to diversify into a
wide range of ecological niches.

Actinopterygian scales are characteristically trapezoidal or diamond-shaped with one or more
longitudinal ridges on the basal surface of the scale (see Moy-Thomas & Miles 1971). The
scales have an ornamented, shiny layer on the surface, a lamellar form of dentine called
ganoine. Hence, they are known as ganoid scales. This layer is thick in early forms but
diminishes in younger (geological) forms. There was a central dentinous layer with vascular
canals and a deep layer of spongy bone. The whole scale lay beneath the skin. Growth was
periodic, and bone was added concentrically, both on outer and inner surfaces; periodic growth
rings are present. A dorsally-directed, quite pointed peg on the scale fits into a socket on the
adjacent dorsal scale producing an arrangement of sloping diagonal rows along the fish's body.

Other uscful microfossils derived from this group, and also found in acanthodians and
placoderms, are otoliths or ear stones. They are used for biostratigraphical and palaeoecological
work in the Tertiary but do also occur in the Palaeozoic (see Nolf 1985).

The primitive teleostomes (including acanthodians and Cheirolepis tend to have simple and
similar rhomboid scales.

The majority of Palaeozoic finds belong to the palaeoniscoids (e.g. Moythomasia). Most
of the early taxa were predators, but later forms were more diverse (e.g. the deep-bodied
platysomids and redfieldiids). The group existed from the Devonian to Early Cretaceous times.

SARCOPTERYGIANS - All other osteichthyans plus tetrapods have been grouped as
Sarcopterygii (with a fleshy lobe) but here I shall consider only the Crossopterygii and Dipnoi.
Crossopterygians. The bony, fringe-finned fishes include those with a stout
internal bony support for the fins, such as the famous, living fossil, the coelacanth
(actinistians), and extinct groups such as rhipidistians and onychodontids. The solid bones of
the head and the scales and fin rays were covered in enamel-coated cosmine, a dentine penetrated
by minute pores. Crossopterygian scales often have a very distinctive exterior ornament of
ridges or pustules. Some, in porolepids and osteolepids, possess rounded, dorsal pegs.
However, not much work has been done yet on distinguishing crossopterygian microfossils,
except for onychodontid teeth, with their simple orthodentine, which are readily separated from
those with more complex teeth and from actinopterygian teeth. The group probably appeared

PALAEOZOIC VERTEBRATE MICROFOSSILS - 445

in the Late Silurian but were almost extinct at the end of the Palaeozoic, except for the
Coelacanths, which survive to the present day. The earliest record seems to be scales from the
Late Silurian of China.

Primitive crossopterygians found in the Late Silurian or Early Devonian of China were
from small fishes, less than 200 mm in length. The later Devonian rhipidistians were very
large, predatory fish with thick stabbing teeth, some of which have intricate labyrinthine
structure in the dentine.

Other groups bearing cosmine-coated scales and bones include the mainly shallow-water
marine porolepiforms, which had slender, leaf-like paired fins (e.g. Porolepis, Glyptolepis,
Holoptychius) from the Late Silurian? to the end of the Devonian; the marine osteolepiforms
which had short, rounded, paired fins with a broad, muscular lobe (e.g. Osteolepis,
Megalichthys, Eusthenopteron) from the Middle Devonian to Permian; the freshwater
rhizodontids which developed complex tooth structure and thinner, lighter cycloid (rounded)
scales by losing cosmine (e.g. Rhizodus, Strepsodus), from the Late Devonian to
Carboniferous; and the marine onychodontids, which have long, striated stabbing teeth and
symphysial tooth whorls (e.g. Onychodus - Pl. 2J, Strunius) that occur commonly as
microfossils in Devonian times.

Coelacanths (hollow-spines in fin rays) or actinistians, along with some other
crossopterygians such as the large rhizodontiforms and the porolepiforms, had no choana (paired
internal nostrils). Their rounded cycloid scales lack cosmine, and they possessed a calcified or
adipose “swimbladder". The tail fin is a characteristic tassel shape, three-lobed and diphycercal.
The group is very conservative but there was a tendency to reduce neurocranial ossification
through time; they range from the Middle Devonian (or earlier) to Recent. Palaeozoic forms
occurred in shallow marine conditions, but the only remaining species, Latimeria chalumnae
(discovered in 1938), lives in deep-water in the Indian Ocean,

Pipnoans. The lungfishes are also an ancient group of fishes, which probably
began in the Late Silurian, perhaps in eastern Gondwana (China-Australia). They flourished
and underwent an adaptive radiation throughout the Devonian and Carboniferous when they
inhabited a wide range of environments. They appear to be less important in the Mesozoic and
Tertiary, and today there are only three living genera including the Queensland Neoceratodus
(see A. Kemp, this volume).

The amazing Western Australian Gogo fauna of Late Devonian age, in which three-
dimensional fish are preserved in nodules, has provided many new forms (see Long, this
volume). However, most Palaeozoic lungfish are known only from teeth and some scales,
which occasionally are found in the microfauna.

Dipnoans can be relatively large fish with a mosaic of thick bony dermal skull elements
and thick cosmoid scales, some of which in Devonian lungfish possessed short dorsal pegs.
Cosmine of skull bones and scales underwent periodic resorption and regrowth during the life
of any one individual, producing Westoll-lines. Most dipnoans had fleshy fin lobes covered
with scales. Characteristic of lungfish are two pairs of thick, ridged tooth plates and paired
vomerine blades or cones. Except for numerous denticles in early forms, dipnoans lack tooth-
bearing jawbones. Microfossils include cosmine scales and bones. Post-Devonian lungfish
have scales without cosmine. Tooth plates are the most common macro- and microfossils.

SUMMARY

Vertebrate microfossils, if not readily referable to known forms, can be identified and
handled by applying the same techniques used by conodont workers. Thelodont scales are often
placed into tentative taxa (sciotaxa, see Fredholm 1988) called "scale species" or associated into
scale "rows" based on similarity of overall scale morphology and histology (Tumer 1976,

446 - TURNER

Karatajute-Talimaa 1978, Marss 1986). These identifications can be refined later as better
specimens are found. If a scale, spine or tooth cannot be readily identified, the tendency in
recent years is to assign it to a group without giving it a species name.

Work over the past three decades has built on the foundation of the early workers such as
Orestes St John and A. H. Worthen (1875), Johannes Rohon (1893), Walter Gross (1947) and
John Wells (1944). Vertebrate palacontologists can now usually determine with confidence
both the nature of animal and the age of enclosing sediment from a small assemblage of
microvertebrates. By co-operation with workers on vertebrate macrofossils and invertebrate
fossils, many new faunas from Australasia have come to light in the last five years. These are
helping in the understanding of the evolution and biogeography of Palaeozoic vertebrates,
especially in the Southern Hemisphere. New finds in the Gondwanan realm include Early
Devonian sharks, acanthodians, and thelodonts from Australia (see below), New Zealand
(Macadie 1985), Antarctica (Young & Turner, pers. obs.), Irian Jaya (see Turner et al. 1981),
west Yunnan (Wang et al. 1986), and Bolivia (Gagnier et al. 1988) and the first Silurian fish in
Australasia (Turner & Pickett 1982, Simpson 1983).

THE AUSTRALASIAN MICROVERTEBRATE RECORD

ORDOVICIAN

Ritchie & Gilbert-Tomlinson (1977) described the first verified Ordovician vertebrates from
Australasia (see also Archer 1984). These come from the Stairway Sandstone (Middle
Ordovician, Late Arenigian to Early Llanvirnian) at Mt. Watt and two localities in the
Charlotte Range of the Amadeus Basin, central Australia. The articulated and disarticulated
remains belong to primitive heterostracan fishes of two named genera, Arandaspis and
Porophoraspis (Fig. 1). Other scale forms have not yet been described (Ritchie, pers. comm.).
Disarticulated remains are also known from the Carmichael Sandstone (Young, pers. comm.).

Arandaspis prionotolepis is represented by a complete cephalothorax with bone-like plates
covered with a scale-like ornament of presumed dentine tubercles (Fig. 1A). There are
diamond-shaped trunk scales arranged in regular rows (Fig. 1B) and others, perhaps scales,
associated with the branchial region, which have an anterior, narrow, smooth overlap zone, not
unlike those of some anaspids (Fig. 1C). Porophoraspis crenulata is known from one small
plate bearing numerous small, low, rounded tubercles with regular rows of pores opening onto
the surface (Fig. 1D).

Other distinctive scales or tubercles have also been found in the Horn Creek Siltstone in
central Australia. These dentinous scales possess an ornament resembling that on the tesserae
of the mid-Ordovician heterostracan-like fish, Sacabambaspis, from Bolivia (Gagnier et al.
1986) (Young, pers. comm.).

The known occurrences of Ordovician microfossils are restricted to shallow-water marine
deposits in central Australia, mainly in the intertidal zone. The complete specimens from
Australia, and North and South America have been found in Cruziana-zone deposits. Possible
further sources for Ordovician microfossils are the invertebrate-rich Nora Formation of western
Queensland and an area in the Molong High of New South Wales, where lagoons surrounded
volcanic islands in Ordovician times (Webby 1985).

SILURIAN

No definite Silurian agnathans have yet been found in Australasia. However, scales,
spines, jaws and tooth whorls of acanthodians appear in the Upper Silurian rocks of eastern

PALAEOZOIC VERTEBRATE MICROFOSSILS - 447

Australia. The earliest record is in mid-Ludlovian-aged limestones where they were discovered
during conodont sampling. They occur in the Mirrabooka Formation of the Orange District
(Turner 1982a, Turner & Pickett 1982), in the Queanbeyan district near Canberra (Henderson
1982), within the Laidlaw Volcanics (Long & Turner 1984), in the Silverdale Formation
(Strusz & Garrett 1991, in press), and in the Graveyard Creek Formation of the Broken River
District, north Queensland (Turner & Pickett 1982, Simpson 1983). There are scales similar
to those of the common Silurian-Early Devonian genus, Nostolepis, closely resembling
Nostolepis striata, as well as spines of climatiids (Denison 1979). Ischnacanthids are
represented by scales, including ones resembling those of Gomphonchus, and small jaws with
ankylosed teeth (Denison 1979). Some of the bony platelets might belong to early
placoderms, some of which have been found in the Silurian of China. An articulated
acanthodian tail has also been found in the Upper Silurian Baragwanathia beds near Yea,
Victoria (Long, pers. comm.).

DEVONIAN

Thelodonts

Turiniid thelodonts are common in Australian Devonian nearshore limestones. They are
well-preserved in the Mulga Downs Formation and Cravens Peak Beds (Turner et al. 1981), as
well as in other limestones from the Early Devonian of New South Wales, and more rarely,
Queensland. Rare nikoliviid thelodont scales may also occur; see Fig. 3B, C, for a nikoliviid-
like scale. In New South Wales alone, turiniids come from sediments within the Bogan Gate
and Molong Highs; the Amphitheatre Group, the Mineral Hill Volcanics, the Talingaboolba
Formation, the Yarra Yarra Creek Group, the Condobolin and Belvedere formations, the
Trundle Beds, the Garra Formation and an unusual oolitic limestone sample from Tumblong
(Turner et al. 1985). The latter contains scales of Turinia sp., T. australiensis and T. sp. cf.
T. polita (Karatajute-Talimaa 1978).

Late Early or early Middle Devonian turiniids have recently been recovered from the
Amadeus Basin and a deep borehole in South Australia (Young et al. 1987, Long et al. 1988).
The scales are closely related to those from the Cravens Peak Beds in the Georgina Basin and
the Toko Syncline (Fig. 4). Middle Devonian turiniid and nikoliviid-type scales are known
from the Hatchery Creek Conglomerate Group of New South Wales and the Broken River
Formation of Queensland. The turiniid scales from these sites closely resemble those from
west Yunnan (China), Antarctica and Bolivia (Goujet et al. 1984, Wang et al. 1986, Gagnier et
al. 1988, Young & Turner, pers. obs.). As more samples are found in Australia, it is
becoming clear that turiniids were undergoing quite intense speciation in eastern Gondwana
throughout the Devonian (Figs 2, 3). The youngest thelodonts known in Australia, and
possibly in the world, are the turiniids called Australolepis seddoni (Figs 2P, 3A) from the
Early Frasnian Gneudna Formation of the Carnarvon Basin, Westem Australia (Turner & Dring
1981). The scales of this species share characters with earlier Devonian turiniid scales in
central and eastern Australia, suggesting that some of the early forms should also be referred to
the genus Australolepis.

In western Victoria, vertebrate material from the Silverband Formation in the Grampians
includes turiniid scales of another species, Turinia fuscina (Tumer 1986b) (Fig. 2F). Frederick
Chapman had described "shark" remains from a sandstone lens in the Silverband Formation,
which were thought to be of brackish or marine origin (Turner 1986b). Talent & Spencer-Jones
(1963) examined this material and identified what they thought were shark denticles and spines.
Turner (1986b) studied these and identified the scales as turiniform and acanthodian. Shark

448 - TURNER

scales of Antarctilamna (Young 1982) may also be present. The vertebrate assemblage now
restricts the Silverband Formation to the Early Devonian (or early Middle Devonian at the
youngest), as opposed to the earlier assessments of Late Devonian to Early Carboniferous.

Whereas turiniids disappear from Laurentia (North America/Greenland/ western Europe) near
the end of the Siegenian (Early Devonian), they continue to flourish in near-shore Gondwanan
environments until the early Late Devonian. As well as the countries mentioned above,
turiniid scales have been found in the Early?-Late Devonian of Iran and Thailand (Blieck et al.
1984). Turiniids should occur in similar Devonian environments of India and South Africa.
Nikoliviids have recently been identified in Early Devonian limestones of Nevada associated
with placoderm body scales resembling those from the Early Devonian of eastern Australia
(Turner & Murphy 1988). These are the first definite Devonian thelodonts from the U.S.A.,
and they appear to be closely related to those from Arctic Canada and Europe and distinct from
the Gondwanan forms.

Australian turiniid stratigraphic occurrences are summarised in Table 2.

Chondrichthyans

The Late Silurian-Early Devonian was a time of intense experimentation in shark design.
What little we know of early sharks is almost exclusively gleaned from microvertebrate
remains. Karatajute-Talimaa (1973) has described the oldest known shark-like scales to date,
and Vieth (1980), Wang (1984) and Mader (1986) have found scales and possible teeth in the
Early Devonian of Arctic Canada, China and Spain,

In Australia some interesting scales have recently been found which cannot be referred to
thelodonts, acanthodians or placoderms. They are probably primiiive placoid scales.
Surprisingly, some apparently lack the four neck canals which Reif (1978) defined as
characteristic for the true placoid shark scale (Fig. 5K).

The Cravens Peak Beds of Queensland (possibly late Early Devonian) have yielded scales
(Fig. 5G-H, Pl. 51) similar to those called Gualepis by Wang (1984). This scale type is also
seen in the Reefton Beds of New Zealand (Macadie 1985). Associated with the Cravens Peak
scales are teeth (Pl. 1A-C, E) and prismatic calcified cartilage (Pl. 1D) from a shark which was
either related to, or convergent with, the Jurassic to Recent hexacanthid (comb-tooth) sharks
(Turner 1985). The teeth have been referred to Mcmurdodus, a genus also known from the
Middle Devonian of Antarctica (White 1968, Turner & Young 1987).

Other shark scales typical of the Early to early Middle Devonian are those called Ohiolepis
(Fig. SE) found in the U.S.A. (Wells 1944), China (Wang 1984), Europe (e.g. Friman 1983)
as well as in Australia (Giffin 1980, Turner, pers. obs.).

Some samples from the Zlichovian (latest Early Devonian) Jesse Limestone and Mt. Frome
Limestone of New South Wales have thelodont-like or quasi thelodont (probably shark) scales
(Turner 1982a, Fig. 3F, Fig. 5L). They are very different from the turiniid scales in the Early
Devonian in their slender, high crown set on a thin diamond-shaped base; they appear to
possess minute neck canals, but this has not yet been confirmed. Similar scales occur in the
Buchan Limestones of Victoria and in the Late Emsian Receptaculites Limestone Member of
the Murrumbidgee Group, Taemas district of New South Wales. The latter were referred to
"Skamolepis" by Giffin (1980). The type Skamolepis fragilis Karatajute-Talimaa (1978)
came from the Late Emsian to Early Eifelian of Latvia and Spitsbergen, and she now regards
these as shark scales (Talimaa, pers. comm.).

Another small sample of shark scales comes from the Trundle Beds of western New South
Wales. These are exceedingly simple, almost neoselachian-like scales (Fig. 5K).

One of the earliest xenacanthid sharks, Antarctilamna (Fig. 5A-D), occurs in the Middle
Devonian Bunga Beds of southeastern Australia as well as in Antarctica (Young 1982),

PALAEOZOIC VERTEBRATE MICROFOSSILS - 449

possibly in Bolivia (Gagnier et al. 1988), and in Iran (Janvier, pers. comm.). Scales and

ee of this form may also be present in the Silverband Formation of Victoria (Turner
6b).

A

Figure 7. Ligulalepis toombsi, Early Devonian (Late Lochkovian-Pragian), New South Wales and Victoria.
A, E, F, ventral view; B, C, D, dorsal view; scale approx. 1 mm. (After Schultze 1968).

Middle and Late Devonian shark scales and teeth are known from the limestones of the
Broken River district of Queensland, from New South Wales and Western Australia (Fig. SI,
J). One of the earliest phoebodont teeth appears in an early Middle Devonian site in
Queensland (Turner, pers. obs.). Ctenacanth and cladodont forms along with Phoebodus
australiensis Long (1990), and other phoebodonts and protacrodonts dominate Late Devonian
(Famennian) assemblages (Turner 1982b, and pers. obs.). Examples include limestones
in the Fairfield Group of Western Australia, the Teddy Mount Formation of north Queensland
(Pl. 2E) and at Bulga, New South Wales. These Late Devonian assemblages are remarkably
similar worldwide. Gross (1973) figured examples from Europe and the U.S.A. and a similar
assemblage occurs in China (Wang & Turner 1985), Thailand (Long 1990) and Arctic Canada

450 - TURNER

(Turner, pers. obs.). | Another useful Late Devonian (latest Famennian to Early Carboniferous
indicator is the presence of the shark teeth called Harpagodens ferox by Turner (1982b, 1983,
Fig. 6). These teeth occur in the latest Devonian to Early Carboniferous in Australia, but have
subsequently been found in Thailand, China, Europe and the U.S.A. in the Lower
Carboniferous; some were previously mistaken for conodonts. In fact, study of the St John
and Worthen collections made in the last century now convinces me that Harpagodens should
be referred to the genus Thrinacodus, which was based on very water-worn teeth from the Early
Carboniferous of the mid-West of the U.S.A.

Placoderms

Very little work has been done on placoderm microfossils. Those placoderms which
retained the primitive body covering of small bony tesserae and scales do provide, however,
valuable information, at least for the Early Devonian and probably throughout the Silurian.
Small jaws (gnathals) with teeth are also potentially diagnostic, and examples are being found
in the Early and Late Devonian of the Broken River area of north Queensland.

Body scales (PI. 4), formerly referred to Ohioaspis (Orvig 1969, Giffin 1980), are possibly
some of the commonest remains in certain relatively shallow-water marine limestones in
Victoria, New South Wales, Queensland and Tasmania. By comparison with articulated
specimens, many of these scales can be referred to Buchanosteus and related arthrodires
(Turner, Young, Long, pers. obs.) (Pl. 2H, I). Such body scales are common in some rocks
dated as Siegenian-Emsian (Pragian). Body scales of the Early Devonian acanthothoracid,
Murrindalaspis, have been described by Long & Young (1988). These and other types of
placoderm scales are also found in Early Devonian limestones in southern China, the western
U.S.A. and Europe (Goujet 1976, Poltnig 1984, Turner, pers. obs.).

In the Early Devonian Martins Well Limestone of Queensland and in some localities in
New South Wales there are placoderm tesserae very similar to those from western Europe and
identified as radotinid by Obruchev & Karatajute-Talimaa (1967) (Pl. 4D).

Telestomes

Acanthodians

Acanthodian scales are found in nearly all types of Devonian sediment. They are
predominant in the limestones of the Buchan and Taemas districts and in the Early to Late
Devonian marine sediments of eastern Australia. Complete acanthodians have also been found
in Late Devonian continental sediments in Victoria (Long 1983, 1986a, this volume) (PI. 3A).
When thelodont scales are present, they tend to be more abundant than acanthodian scales,
although in a few localities acanthodian scales predominate. The relative abundances of scales
in samples might be an artifact of sampling, or even of scale morphogenesis, rather than a
facies factor.

The first record of acanthodian scales from Australia was that made by Philip (1965) of
Nostolepis scales in the Coopers Creek Limestone of the Tyers district of Victoria. This type
of scale is common in Lower Devonian sediments. The Martins Well Limestone of the
Broken River district of Queensland and several horizons in the Early Devonian of New South
Wales (including the Trundle Beds, the Condobolin Formation, the Yarra Yarra Creek Group)
have a wide variety of scales and acanthodian platelets similar to those which Gross (1971) and
Goujet (1976) described from nostolepids in Europe. Nostolepis costata Goujet 1976 is
common in the Siegenian to Emsian limestones of eastern Australia. Nostolepid scales are

PALAEOZOIC VERTEBRATE MICROFOSSILS - 451

now known to be present in the Lower Devonian Reefton Beds (Macadie 1985) and the Baton
River Beds of New Zealand (John Simes coll., Turner, pers. obs).

Ischnacanthid and climatiid acanthodian scales, spines and jaws from many localities are
currently being studied (Long 1986b). Many are proving endemic to Australia. Some,
however, such as Cheiracanthoides comptus, which seems ubiquitous in Emsian-Early Eifelian
sediments worldwide (e.g. Wells 1944, Vieth-Schreiner 1983), and Machaeracanthus, long
known from Europe and North America (e.g. Goujet 1976, Zidek 1981, Mader 1986), are being
found in Early to Middle Devonian sites in New South Wales, Victoria, including one figured
by Philip (1965) from the Coopers Creek Beds (Pl. 3B), and Queensland.

Table 3 summarizes the stratigraphic ranges of Australian acanthodian remains.

Higher Osteichthyans

Scales, fin ray supports and teeth of crossopterygians, dipnoans, and actinopterygians are
found as microfossils in Devonian rocks (e.g. Giffin 1980, Young & Gorter 1981, Long et al.
1988). Some scales, such as Ligulalepis toombsi (Schultze 1968, Long 1982, Fig. 7), are
distinctive and seem restricted to Early Devonian limestones. Much work has yet to be done
on the use of these microremains.

CARBONIFEROUS

By the Early Carboniferous, agnathans and placoderms had disappeared, and microfossil
assemblages usually are comprised of acanthodian, palaeoniscoid, crossopterygian and shark
remains. Shark scales and teeth seem particularly useful for biostratigraphy in Carboniferous
rocks; they are now being studied by micropalaeontologists, especially in the U.S.A., and
some oil companies are making use of them for work in Alaska. There is a very complete
succession of Carboniferous to Permian shark assemblages in North America, which can
provide a standard. Michael Hansen has been studying shark scales (Hansen 1988), including
those first described by Gunnell (1933) in his conodont studics.

In Australia continental Early Carboniferous vertebrate faunas are known from Mansfield,
Victoria and the Narrien Range in Queensland (Turner & Long 1987, Turner & Hansen 1991,
in press). These faunas include the crossopterygian Megalichthys, lungfish, acanthodians
(Acanthodes), palaeoniscoids and sharks. The latter are all known as microfossils, including
teeth of xenacanthids and Ageleodus.

New finds near Rockhampton are providing information on marine chimaeroid and
cladodont sharks in the Tournaisian and Visean of Queensland (Turmer 1990). The latter
include helodont, the first Australian petalodont, psammodont and cochliodont teeth, and
Stethacanthus. In the Bingleburra, Namoi and Dangarfield formations of New South Wales
shark faunas of Tournaisian and Visean age with close similarity to those of the Rockhampton
Group are known. These assemblages include teeth of Thrinacodus ferox, stethacanthids and
helodonts as well as orodonts and caseodonts (Jones, pers. comm., Leu, pers. comm., Turner
1982b & pers. obs.). The New South Wales assemblages, as well as ones from late in the
Early Carboniferous of Queensland, are nearly all microfossils.

Good Early Carboniferous shark faunas are also present in the Tournaisian-aged Fairfield
Group and Visean Utting Calcarenite of Western Australia. These assemblages include macro-
and microfossils of Thrinacodus ferox (Turner 1983), stethacanthids (Turner 1982b), helodonts
and ctenacanths. One nodule from the Utting Calcarentite in the Bonaparte Gulf, Western
Australia, collected during geological mapping in the early 1960s, contains a specimen of
Stethacanthus exhibiting part of the jaw cartilages, a tooth battery, and external and internal
scales of the jaws. Using this specimen, the scales of Stethacanthus can be positively
identified by association with the teeth, a rare example. Rediscovery of the site of the nodules
should yield further articulated specimens. Orodus and Mesodmus remains have also been

452 - TURNER

found in the Utting Calcarenite (Long, pers. comm.). These Early Carboniferous fish faunas
contain elements in common with the classic North American and European assemblages, such
as those from the mid-western U.S.A. (St John & Worthen 1875).

The youngest Carboniferous vertebrate-bearing bed comes from the Barambah Limestone
from near Murgon, southern Queensland. This bed contains microremains (scales and teeth)
from denaeid and possible neoselachian sharks as well as palaeoniscoid teeth. The shark scales
resemble those described by Gunnell (1933) from Late Carboniferous of the U.S.A. The
Barambah Limestone is thought to be of Namurian age (Palmieri 1969). After this there are
no known microvertebrate producing sites in Australia until the Early Permian.

PERMIAN

Permian vertebrate microremains are rare, but Early Permian sites in Queensland have
yielded paleoniscoid (Turner 1982c), and rare possible shark remains (Turner, pers. obs.). The
marine Permian sequences in Western Australia and Queensland are also being investigated by
S. Turner. The interesting freshwater Permian fish assemblage from Blackwater, Queensland,
also has potential for microfossils. Michael Leu (Macquarie University) has collected new
articulated sharks from this site, which exhibit scales, spines and teeth (Leu 1990).

Table 3. Geographic and stratigraphical distribution of acanthodians in Australia. 1, Late
ploekensis-early eosteinhornensis conodont zone, Jack Limestone, north Queensland and mid-
Ludlovian, east of Trundle, New South Wales (Turner & Pickett 1982); 2, Martin's Well
Limestone, north Queensland (Turner, pers. obs.); 3, several Early Devonian limestones in New
South Wales (Tumer, pers. obs.); 4, Silverband Formation, Victoria (Turner 1986b); 5, Waratah
Bay and Coopers Creek Limestones, Victoria (Philip 1965, Turner, pers. obs.); 6, Pragian
limestones in westem New South Wales, e.g. Trungle Beds, Yarra Yarra Creek Group (Turner,
pers. obs.); 7, Mulga Downs Group, westem New South Wales (Turner, pers. obs.); 8, Mt Ida
Formation, Victoria (Turner & Long 1984); 9, Laidlaw Volcanics, Australian Capital Territory
(Turner & Long 1984); 10, Hatchery Creek Conglomerate Group (Young & Gorter 1981); 11,
Cravens Peak Beds, Queensland (Turner et al. 1981, Young & Gorter 1981) and Tandalgoo Red
Beds, Western Australia (Gross 1971 - "Gomphonchus" - probably another genus, these forms
need further study); 12, Araluen, New South Wales or Australian Capital Territory (Tumer, pers.
obs.); 13, Famennian?, Georgetown, Queensland (Turner, pers. obs.); 14, jaw, Hunter Siltstone,
Grenfell, New South Wales (Long, pers. comm.); 15, Buchan and Taemas limestones, Victoria
and New South Wales (Long 1986a); 16, Late Gedinnian to Late Emsian limestones in New South
Wales ( Turner, pers. obs.); 17, limestones in Queensland and New South Wales (Giffin 1980,
Turner, pers. obs.); 18, includes all simple, smooth-crowned Acanthodes-like scales from late
Early and Middle Devonian limestones of Queensland, New South Wales and the N'Dahla Member,
Amadeus Basin; 19, Mt Howitt, Victoria (Long 1986a); 20, new species, Pambula, New South
Wales (Young, pers. comm.); 21, Freestone Creek, Victoria (Long & Tumer 1984); 22,
Hervey'’s Range, New South Wales (Long & Turmer 1984); 23, Mt Howitt, Victoria (Long
1986); 24, Mansfield, Victoria (Long & Turner 1984); 25, Lower Carboniferous (including Upper
Telemon, Raymond and Ducabrook formations) of central Queensland (Long & Turner 1984); 26,
Acanthodes-like scales in the Coffee Hill Member, Catombal Group, New South Wales (Jones
(Australian Museum) and Turner, pers. obs.); 27, Broken River Formation, north Queensland
(Turner, pers. obs.); in addition, there are many undetermined acanthodian remains including
those from the Late Devonian Merrimbula Group, Worange Point, Eden, New South Wales;
Myrtlevale Formation and Bundock Creek Group of north Queensland.

PALAEOZOIC VERTEBRATE MICROFOSSILS - 453
SUMMARY

Vertebrate microfossils can provide us with much information about Palaeozoic fish faunas,
which cannot be gleaned from the macrofossils alone. Not only can microfossils assist in
gauging the full extent of the geographic distribution of taxa, but they can aid in the
correlation of Palaeozoic rocks and contribute to our knowledge of the early evolution of the
major fish groups. However, the need for continued research and description on the
macrofaunas is paramount if we are to understand the affinities of all the vertebrate
microfossils. For a summary of the stratigraphic ranges and relative abundance of fish groups
found both as macro- and microfossils in Australasia see Table 4.

The recent work on Palaeozoic vertebrate microfossils has led to the use of some forms as
indicator or zone fossils. The analysis of such forms in Australasia is still in its infancy.

Table 5 summarizes some of the useful indicator fossils which can assist in assessing dates
for Australasian rocks.

Early Palaeozoic agnathans, acanthodians, and sharks are all known nearly exclusively from
microfossil evidence. Major Late Ordovician-Early Silurian agnathan faunas are now known in
Siberia, North America and parts of western Europe but not as yet from the Gondwanan
continents. More intensive investigation of Australasian Ordovician, Silurian and Permian
nearshore sediments should greatly increase knowledge of the Palaeozoic vertebrates.

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UPPER
SILURIAN

454 - TURNER

Agnathans Acanthodians Sharks Placoderms Actinopterygians Crossopterygians Lungfish

PERMIAN

CARBONIFEROUS

Thelodonts

DEVONIAN

SILURIAN

ORDOVICIAN 1

Heterostracan-like
forms

Table 4. Summary of the stratigraphic ranges of fish groups based on evidence of macro- and
microvertebrate remains in Australasia. Variation in width of column gives an estimate of
abundance. E - for extinction of the group in Australasia; for crossopterygians, a Famennian
extinction for porolepiforms, and a mid-Carboniferous for osteolepiforms and rhizodontiforms.

ACKNOWLEDGMENTS

My work on Palaeozoic vertebrate microfossils of Australia was supported by grants from
the Australian Research Grants Scheme (E 8115050, 1982-1987). My thanks to the Director,
Dr Alan Bartholmai, and the Board of the Queensland Museum for the continuation of an
honorary Research Fellowship. I am especially grateful to those palaecontologists and
geologists in Australia and elsewhere who have freely given time, information and specimens
to the project; many are mentioned in the text. I particularly wish to thank Prof. Jim Warren
and Ian Stewart (Monash University), Prof. John Talent, Drs G. Bischoff and Ruth Mawson
(Macquarie University), J.S. Jell (University of Queensland), Simon Lang (Geological Survey
of Queensland), John Long (The Western Australian Museum), John Pickett (Geological
Survey of New South Wales), Tony Wright (University of Wollongong), and Gavin Young
(Bureau of Mineral Resources), who provided numerous acid-prepared residues from which
vertebrate samples were obtained.

PALAEOZOIC VERTEBRATE MICROFOSSILS - 455

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=|) 2] ai) s a| os oP lh oa Pea g a] we ® 4 8 S cle x oO
o|3| 8) 8) 5/3 2/s3|8|/e] 2/3 2 =/a|o €/|3 5 2
rt B ae 9/35 |z , |X a 13 o|2 S 9\ 82
Si o| Bl gi e]s 2/3/31 8}]e)}3 Bigiale 2 . Sle g
2 S|oo/2/8/8)elelsE = a/el\le |e/e/3 a/2=|v/z
Q vol ele! O|uea/S]/S)/8]/e2] a/& Els ® 2 =} S| =
g qa/2/a asi; sle£/2/a/2/5)28 S/eE|/z = ele | &
al 2 oil a S|£|a/| 2 o|/ 8] a = g Ql s Sle i.9a
SS|2)/o/$/s|/s2) 2/S5loele]el| sis |S/2/e\eeg 3/38 =/5/8
aS} S| °° 2); Ss] s = s 2
Ss el el e/ si selsl/5l8laleis/s]Fl/8/8] 2 s2i|e|/s|é £/ Sle \€
S8|/S)/Els|} s/s S/Z/B/P/SIS1§ | 5/2 1/8/21 28/5 S| 3 2,2
“Hl SFl|n| 2) €)02 a)/a|5 |? S/o/;/e}/e}Sleixta, asa! x ee

Table 5. Useful age-indicator fish microfossils in the Palaeozoic of Australia. For discussion
see this paper and other papers referred to in the text.

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PLATES

Plate 1. Mcmurdodus whitei, shark teeth (A-C, E) and calcified cartilage (D), late Early Devonian or early
Middle Devonian, Cravens Peak Beds, western Queensland. (From Young & Tumer 1987).

Plate 2. Devonian and Early Carboniferous examples. A, acanthodian scale in dorsal view, Cheiracanthoides
(LQ354.72) V1641 late Early Devonian Taemas Group, Wee Jasper, New South Wales; B, acanthodian scale in
dorsal view - new genus? (LQ347.20) late Early Devonian Taemas Group, Wee Jasper, New South Wales; C,
placoderm body platelet in dorsal view - rhenanid? (LQ778.33) Tullamore, Early Devonian, New South Wales
(MMMC2150); D, acanthodian toothwhorl in lateral view - (LQ777.29) Tullamore, Early Devonian, New
South Wales (MMC2151); E, “cladodont" shark tooth in lingual view - stethacanthid (LQ789.7), latest
Devonian/earliest Carboniferous? Teddy Mt. Formation, north Queensland; F, acanthodian scale -
Cheiracanthoides (LQ350.34) late Early Devonian Taemas Group, Wee Jasper, New South Wales; G, shark
tooth or cephalic denticle in dorso-lingual view - Thrinacodus ferox (LQ790.14) latest Devonian/earliest
Carboniferous Teddy Mt. Formation, north Queensland; H-I, placoderm platelet in ventral and dorsal views -
Buchanosteus (LQ356.28-27) late Early Devonian Taemas Group, Wee Jasper, New South Wales; J,
onychodontid tooth - Onychodus (LQ334.53) Lower Devonian Coopers Creek Limestone, Tyers, Victoria; not
to scale; all specimens in the 0.5-2.5 mm size range. (LQ numbers refer to ARGS-funded Project Photographic
Record housed at Queensland Museum).

PALAEOZOIC VERTEBRATE MICROFOSSILS - 459

Plate 3. Example of acanthodian scales and squamation. A, Culmacanthus stewarti Long 1983, showing
articulated squamation, ornamented cheek plates (dermal bones) and endoskeletal shoulder- girdle bones
(slender, non-omamented); B, nostolepid; C, Machaeracanthus sp. (B & C from Philip 1965).

Plate 4. Examples of placoderm body scales: A, Placoderm gen. et sp. indet., Pragian limestone low in
Trundle Beds, NSW, MMMC2146; B, Another placoderm gen. et sp. indet., Pragian limestone low in Trundle
Beds, New South Wales, MMMC2147; C, Buchanosteid- type scale, Pragian limestone in Yarra Yarra Creek
Group, New South Wales MMMC2148; D, radotinid or rhenanid scale, Lochkovian Martin's Well Limestone,
north Queensland LN561/4.; E, antiarch scale or omament, Pragian Limestone low in Trundle Beds, New
South Wales MMMC2149; F, asterolepidoid scale (CPC2258), from Young 1984. Not to scale; all in the
range of about 1-2 mm. (LN number, Australian Research Council project photographic record, Queensland
Museum; MMMC, Mineral Museum Microfossil Collection, Sydney).

Plate 5. Devonian vertebrate microfossils. A, thelodont body scale in dorsal view - Turinia pagei
(LQ771.49), Lower Devonian (Dittonian) Welsh Borderland, United Kingdom; B, thelodont body scale in
dorsal view - Turinia australiensis (LQ631.98), Belvedere Formation, Early Devonian, near Fifield, New South
Wales (MMMC2152); C, acanthodian scale in lateral view - Cheiracanthoides (LQ354.76) late Early Devonian
Taemas Group, Wee Jasper, New South Wales; D, thelodont body scale in dorsal view - turiniid n. sp.
(LQ324.34) late Early Devonian Cravens Peak Beds, western Queensland; E, placoderm body platelet in dorsal
view - Buchanosteus (LQ353.67) late Early Devonian Taemas group, Wee Jasper, New South Wales; F,
acanthodian scale in dorsal view - ?nostolepid (LQ729.24) Early Devonian (Dittonian), Welsh Borderland,
United Kingdom; G, acanthodian? scale in dorsal view - (1.Q698.53) late Early Devonian (Zlichovian) Mt.
Frome Limestone, Mt. Frome, New South Wales (MMMC2153); H, acanthodian scale in dorsal view - new
genus? (LQ349.32) late Early Devonian Taemas Group, Wee Jasper, New South Wales; I, shark scale - cf.
Gualepis (LQ623.82) late Early Devonian Cravens Peak Beds, western Queensland. Not to scale; all
specimens in the 0.5-2.5 mm size range. (LQ numbers refer to ARGS-funded Project Photographic Record
housed at Queensland Museum).

460 - TURNER PLATE 1

PLATE 2 PALAEOZOIC VERTEBRATE MICROFOSSILS - 461

462 - TURNER PLATE 3

PLATE 4 PALAEOZOIC VERTEBRATE MICROFOSSILS - 463

464 - TURNER PLATE §

CHAPTER 14

AUSTRALIAN MESOZOIC
AND CAINOZOIC
LUNGFISH

Anne Kemp!
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ETIASSIC PRECOLOS &. .cicc cose cst edocs Sudccssdsecvsseeces 467
Jurassic and Cretaceous Records.................. 472
Cainozoic Lungfish .............ccccececeeeeceeeceeeeceees 477
Palacocene and Eocene Records...............200. 477
Miocene, Pliocene and Pleistocene Records...478
COnGCIUSIONS its. 4rd... cettelsecdevesedechocctccasccest ene 484
Acknowledgements............cc.cccceseecsssceseesecsces 486
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1 Queensland Museum, P. O. Box 300, South Brisbane, Queensland 4101, Australia.

466 - KEMP, A.
INTRODUCTION

The dipnoans, or lungfish, are an ancient and often controversial group of vertebrates,
regarded by some as the sister group of the tetrapods (Rosen et al. 1981). They belong to the
Sarcopterygii or fleshy-finned fishes. In common with other Sarcopterygii (the coelacanths and
rhipidistians) lungfish have paired fins with fleshy bases and an internal skeleton which has a
single proximal bone articulating with the pectoral or pelvic girdle. The oldest known dipnoan
from Australia is Dipnorhynchus sussmilchi (Etheridge 1906) from the Spinella yassensis
limestone (Zlichovian, Early Devonian) at Taemas, New South Wales. The material includes
well-preserved skulls and mandibles. Fragments that may belong to this species also occur at
Wee Jasper and Buchan. The slightly younger Dipnorhynchus kurikae (Campbell & Barwick
1985) occurs at Wee Jasper, New South Wales and Buchan, Victoria. Members of this genus
have massive crushing palates and very thick skulls. In the same beds (at the junction of the
Bloomfield and Receptaculites limestones at Wee Jasper) the more delicate Speonesydrion
iani (Campbell & Barwick 1983) is found. This is the oldest known species in the world
with tooth plates.

The three living genera of lungfish, represented by Neoceratodus forsteri in Australia,
Lepidosiren paradoxa in South America and several species of Protopterus in Africa, form a
coherent group, although there are differences in behaviour, habitat, physiology and anatomy.
They are all plainly lungfish; the skulls and jaws, post-cranial skeleton, tooth plates and soft
tissues bear the hallmarks of the Dipnoi. They possess simple lungs, a double sac in the case
of Lepidosiren and Protopterus, single in Neoceratodus. Spawning strategies vary. N.
forsteri attaches eggs to underwater plants and shows no parental care (Kemp 1984), while the
African lungfish builds a nest and guards the eggs (Greenwood 1958). Lepidosiren and
Protopterus must breathe air to live Johnels & Svensson 1954, Burggren & Johansen 1987)
but Neoceratodus survives on gill breathing only, unless it is active, as when feeding at night
or during the breeding season (Kemp 1984, 1987a). There is no evidence that it must breathe
air to survive. Lungfish all lack a stomach, but have an intestine with a spiral valve. The
body form is eel-like, particularly in the South American lungfish, and the tail is diphycercal.
The paddle-shaped pectoral fins of N. forsteri, modified into filaments in the other living
lungfishes, move underwater like the legs of a urodele amphibian (Dean 1906, 1912) and are
used during feeding to brace the body against the substrate. In the roof of the mouth are the
controversial "nares", actually an olfactory organ with one opening just outside the mouth and
one within (Bertmar 1965). The tooth plates, made of complex dentines and ankylosed to the
jaw bones, all conform to the same basic pattern of radiating ridges. The jaw bones and skulls
are similar, despite the lighter and more streamlined structures of Lepidosiren and Protopterus,
and none of the three living groups has ossified vertebral centra. Skin and lateral line organs
are comparable in the three genera.

Fossil species, while recognizable as lungfish, show far more variation in body form,
skeletal structure and habitat, particularly in the oldest Devonian lungfish (Campbell &
Barwick 1984). Skulls of Devonian lungfish bear little resemblance to those of modern
lungfishes (Romer 1936, Thomson & Campbell 1971, Schultze 198la, Campbell & Barwick
1984, 1985), and there is little agreement on the naming of the bones, or their homology with
the skull bones of other vertebrates (Holmgren & Stensio 1936, Romer 1936, Holmgren 1949,
Lehman 1966, White 1966). Some authors refer to the bones by letters (Romer 1936,
Schultze 1981a) but the lettering systems are often inconsistent. In this paper bones of the
calvarium are labelled according to the letter system of Thomson & Campbell (1971) and
Schultze (1981a). Names of jaw bones follow Bertmar's system (1965).

There are a number of different types of dentition in the older lungfish ranging from isolated
denticles to solid crushing palates (Denison 1974, Campbell & Barwick 1983, 1985) although

AUSTRALIAN MESOZOIC AND CAINOZOIC LUNGFISH - 467

development of tooth plates from isolated cusps into radiating ridges is known in some of the
oldest dipnoans (Denison 1969, Lund 1970, 1973, Schultze 1977) as well as the living species
(Kerr 1910, Kemp 1977) and similar developmental abnormalities, e.g. bifid ridges or total or
partial absence of ridges, occur in all lungfish with tooth plates based on the radiating pattern
(Kemp 1987b).

The older lungfish lack the diphycercal tail of the modern forms (Campbell & Barwick
1984, 1988). The oldest lungfish are marine, and some of the Australian species, like
Chirodipterus australis and Griphognathus whitei, found in the Gogo Formation of Western
Australia, even lived in the deep water surrounding reefs (Campbell & Barwick 1988). Air
breathing in these fish is unlikely to have been important, and the associated anatomical
structures would have been poorly developed (Campbell & Barwick 1988).

A further dimension of confusion is added by the frequently imperfect preservation of fossil
specimens. Records of Devonian material, particularly from Australia, are good, largely
because of the heavily ossified elements of the dermal skeleton (e.g. Campbell & Barwick
1984) but many Mesozoic and Cainozoic lungfish are known only from isolated tooth plates,
which are rarely associated with jaw bones. Skull material of post-Devonian lungfish is scarce,
and skull bones associated with identifiable tooth plates almost unknown. Worse still are the
numerous isolated skull bones, skeletal elements, scales and tail impressions, often well
preserved but impossible to associate with any known species identified from tooth plates.
However, Australian Mesozoic and Cainozoic records are notable for their variety if not for
completeness, and the extraordinary radiation of species in the Cainozoic deposits of Central
and Northern Australia is unparalleled in any other continent at this time.

MESOZOIC LUNGFISH

TRIASSIC RECORDS

Gosfordia truncata (Figs 1, 2) was originally described by Woodward (1890) from five
broken specimens found in a quarry near Gosford in New South Wales (Table 1). The deposit
is from a shale horizon near the top of the Gosford Formation in the Narrabeen Group of the
Sydney Basin. This is Early Triassic in age (200-230 myBP). Ninety years after the first
description was published another specimen was found in a quarry in the Somersby Falls area,
west north west of Gosford and a few kilometres from the type locality (Ritchie 1981). The
new specimen was found in a shale lens of the Hawkesbury Sandstone, and is also of Triassic
age.

The new specimen of G. truncata is preserved as an almost perfect laterally-flattened fish.
Fin rays of the tail are delineated, and details of the post cranial skeleton are beautifully
preserved (Ritchie 1981). Some of the skull bones can be discerned (Ritchie 1981) although
they give little help in determining the relationships of the species (Fig. 1A). Although
flattened, none of the bones are out of place, and there is little distortion of the head. There is a
central series of two unpaired bones (EQ and ABC), a centrolateral series of 3 bones (M, L and
J) and a lateral series of two (Y and Z), of which the Y bone carries the jaw articulation. The
opercular and associated bones are strongly developed, as is the cleithrum. The angular is a
deep heavy bone, similar to that of Protopterus. There are no orbital bones, and no
dermosphenotic (XK bone). Skull roofing bones with this pattern indicate possible affinities
with Paraceratodus germaini (Schultze 1981a). Useful characters of tooth plates are,
unfortunately, absent from all specimens, and only the labial tips of possible ridges show in
the region of the lower jaws.

G. truncata is unusually deep-bodied for a lungfish. Like the modern Neoceratodus forsteri
it appears to have obese, rounded contours to the belly (Fig. 2), but the ratio of the depth of the

468 - KEMP, A.

tail to the length of the whole body is greater in G. truncata (0.36) than in N. forsteri (0.20).
The diphycercal tail, terminating in an obtuse angle, accentuates the deep body, and although
the specimen is preserved as a carbonised film on rock, with little surface relief, the fish was
probably lateraily compressed as well. The paired fins are similar to those of N. forsteri but
are longer and less robust (Ritchie 1981).

There is one other Triassic ceratodont in Australia known from body parts without
associated tooth plates. This is Ceratodus (Tellerodus) formosus (Fig. 2, Pl. 1), from
Brookvale near Sydney (Wade 1935). It is preserved as a whole fish in lateral view. The
detailed stratigraphy of the area is unknown (Table 1), but it is Triassic. C. formosus is small,
only 8.5 cm long and shares with juvenile specimens of N. forsteri the characteristic of a dorsal
fin reaching forward to the back of the head. Large ceratodont scales have been found in the
same deposits, and it is likely that the holotype of C. formosus is a juvenile. The depth of
tail: length of body ratio is 0.29, so in life it would have been less deep bodied than G. truncata
but not as elongate as N. forsteri.

C. formosus is not as well preserved as G. truncata, and details of the body, fins and head
are less clear (Pl. 1A). It had a diphycercal tail and typically ceratodontid paired fins. Tooth
plates show only as ridge tips sheared across (Pl. 1B) and useful characters in the dentition are
accordingly absent. Most lungfish tooth plates are indistinguishable from each other if
preserved in this way.

Preservation of the head of C. formosus is reasonably good, and much more detail shows on
the original specimen (Fig. 1B, Pl. 1B) than is visible in the photograph published by Wade
(1935). Both sides of the lower jaw (prearticular bone) are present, slightly displaced and
damaged, one bearing 4 tooth ridges and the other two, sheared across and pointing dorsally.
Immediately above these ridges a single broken ridge of the matching upper plate is visible,
with attached fragments of the pterygopalatine bone. Above this is a displaced fragment of an
oval bone, partly overlying the Q or rostral bone of the snout. Behind the rostral bones, E and
Q, is the central posterior medial bone, which is covered by numerous scales. Because of
these, the extent of this bone, and the presence or absence of centro-lateral bones cannot be
determined. An M bone above the eye may be present. Lateral to the scales, the calvarial
portion of the YZ bone and the attached jaw articulation ("quadrate"), whose distal portion
overlies the prearticular, can be distinguished. The displaced ceratohyal lies behind the
prearticular and opercular bones and branchial arch elements are also present.

Below the orbit, part of the parasphenoid is visible, but the bones of the orbit are indefinite.
A small bone 4 (the dermosphenotic of Martin 1982) is visible, close to and possibly
articulating with the calvarial YZ bone. The equivalent bone in N. forsteri articulates between
the JLM and the calvarial YZ bone (Fig. 1C). Traces of several other bones are visible in the
orbital region below the eye. The best interpretation of these structures seems to be that a
bone 4 and an orbital series were present, as in juvenile N. forsteri where an articulated XK and
a series of orbital bones, 2-4 in number and embedded in connective tissue, are found.

Figure 1. Drawings of skulls of three Australian lungfish in lateral view. A, Gosfordia truncata, freehand
drawing of F60621, from the collection of the Australian Museum, scale bar = 2.5 cm; B, Ceratodus formosus,
Camera lucida drawing of P16828 in the collection of the British Museum, scale bar = 1 cm; C, Neoceratodus
forsteri, frechand drawing of the skull of a fully grown specimen from Enoggera Reservoir, South East
Queensland; the parasphenoid and ceratohyal are obscured by more superficial bones, and the cleithrum is
omitted, scale bar = 2.5 cm; ABC, ABC or posterior central bone; A, angular; C, ceratohyal; CL, cleithrum;
D, displaced bone; E, Q, EQ, anterior central bones; I, JLM, J, L, M, bones of centrolateral series; O, orbit;
O', orbital bones; OP, opercular; PA, prearticular, PP, pterygopalatine; PS, parasphenoid; SC, scales; SOP,
subopercular; SP, splenial; V, vomer; Y, Z, YZ, XK, and 4 bones of lateral series (KK and 4 “dermosphenotic"™
and YZ, “quadrate”).

AUSTRALIAN MESOZOIC AND CAINOZOIC LUNGFISH - 469

470 - KEMP, A.

Table 1: Triassic Deposits Containing Lungfish

Species

Gosfordia truncata

Ceratodus (Tellerodus)

formosus

Ceratodus sp.

Ceratodus cf. C.
tiguidensis,

(now Arganodus cf.

A. tiguidensis)

Sagenodont

Ceratodus cf. C.
Phillipsi

Ceratodus avus

(Sagenodus laticeps)

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Locality

Railway Ballast
Quarry, Gosford,
N.S.W.

quarry near
Somersby Falls,
Gosford, N.S.W.

Brookvale, near
Sydney, N.S.W.

Brookvale, near
Sydney, N.S.W.

Blina, W.A.

Blina, W.A.

The Crater and
Duckworth Ck.,
nr. Rolleston,
cent. Qld.

Blina, W.A.

Old Beach,
Midway Point,
Conningham
locs., nr. Hobart,
Tas.

The Crater and
Duckworth Ck.,
cent. Qld.

Ashfield Sh., St
Peters, nr. Sydney,
N.S.W.

Lithologic Unit

Gosford Fm.,
Narrabeen Gp.,
near Sydney, N.S.W.

Hawkesbury Ss.,
Sydney Basin
not known

not known

Blina Shales

Blina Shales

Arcadia Fm.,
Rewan Gp.

Blina Shales

Knocklofty Fm.

Arcadia Fm.,
Rewan Gp.

Wianamatta Gp.

Material

5 incomplete
specimens
(Woodward 1890)

nearly complete
fish (Ritchie 1981)

nearly complete
fish (Wade 1935)

scales

tooth plates,
tooth plate
impressions

upper tooth
plate attached
to bone

tooth plates and
skull frags.

tooth plates

tooth plates
(Dziewa 1980)

tooth plates,
skull roof

broken palate,
body; some assoc.
scales (Woodward
1908)

AUSTRALIAN MESOZOIC AND CAINOZOIC LUNGFISH - 471

Figure 2. Artist's impression of freshwater Triassic lungfish in a natural environment. The larger deep-
bodied fish is based on Gosfordia truncata and the smaller more clongate form on Ceratodus formosus. The
fish in the foreground is the anachronistic Neoceratodus forsteri, a species which does not appear until the
Cretaceous, included for comparison of fossil and modern forms. The specimen is a juvenile. Drawing by
Rober Allen.

Unfortunately, the skull and body of C. formosus are not associated with good tooth plate
material. The dentition is consistent with those of other small Triassic lungfish but is too
poorly preserved for definite comparison. The number of ridges visible is not a useful character
as both lower tooth plates are damaged and more ridges could have been present. The body
form differs from that of G. truncata and of N. forsteri. Transferring C. formosus to Lehman's
genus Tellerodus (1975) on the grounds of a "dermosphenotic", as Martin (1982) has done is in
doubt as an equivalent bone is present in N. forsteri and may not be in Ceratodus (Tellerodus)
sturii (Kemp in prep.). Lehman (1975) and Schultze (1981a) do not recognize a
“dermosphenotic" in C.(T.) sturii, and the significant area on the specimen is damaged. It
would be better to leave C. formosus as it is until more material is available.

Other Triassic dipnoan material from Australia is incomplete and tantalizing. Three distinct
species are found, one from Western Australia, Queensland and Tasmania, one from Western
Australia and Queensland, and one which is found only in Western Australia (Table 1). The
specimens are all poorly preserved tooth plates or impressions of tooth plates with some skull
fragments that can be tentatively associated with each kind of tooth plate. The first has broad
flat tooth plates and jaw bones with primitive characteristics (Pl. 2B,C), and some similarities
to Proceratodus carlinvillensis (Romer & Smith 1934) and to sagenodont lungfish. The ridges
are short with rounded tips and originate from the back of the tooth plate, and the upper jaw
lacks a pterygopalatine process. This species is found in Blina, W. Australia, from the Blina
Shales, and at the Crater and Duckworth Creek, near Rolleston in central Queensland, a locality
in the Arcadia Formation (Rewan Group). In the latter area, some isolated skull bones are also
found, of a size comparable with the tooth plates, and having primitive characteristics as well

472 - KEMP, A.

(Kemp in prep.). The anterior skull bone has small accessory bones attached to the free edge
(PI. 2A), a feature of sagenodont lungfish but not so far of ceratodonts (Watson & Gill 1923).
It may not be justifiable to associate this skull bone and the broad, flat tooth plates as one
species, but the record of lungfish skull bones having sagenodont-like characters, previously
unknown from Australia, is important. Similarly, the presence of such a primitive tooth plate
is significant for Australian records.

The second group of poorly preserved Triassic tooth plates (Pl. 2D) has been referred to a
species described as Ceratodus tiguidensis by Tabaste (1963), which was transferred to the
genus Arganodus by Martin (1984a). It is only found at Blina, Western Australia. Despite its
poor preservation, it is plain that these tooth plates belong to a group of lungfish common in
Triassic and Cretaceous deposits in North Africa (Tabaste 1963, Martin 1984a), Russia
(Vorobyeva & Minikh 1968), and North America. One of the tooth plates was found in the
orbit of a fragmentary skull, which has no particular lungfish characteristics, and the
association of skull and tooth plate is likely to be incidental.

The third species, based on numerous small tooth plates and a tiny calvarium, is found at
Blina, Western Australia, the Crater and Duckworth Creek, Queensland and near Hobart, in
southeastern Tasmania. Specimens from the last area, found in several localities, belong in the
Triassic Knocklofty Formation and were referred to Ceratodus gypsatus by Dziewa (1980).
They are, however, closer to specimens of Ceratodus phillipsi (Agassiz 1833) than to C.
gypsatus, which has been synonymised with Ceratodus concinnus by Martin (1980). The
calvarium and the tooth plates of this species (PI. 2E, F) appear to be closer to gnathorhizids
(Carlson 1968, Olson & Daly 1972, Berman 1976, 1979, Minikh 1977) than to classical
ceratodonts. If this is the case, a record of a gnathorhizid lungfish from Triassic deposits in the
Southern Hemisphere is significant. Previously, gnathorhizids have been recorded from Late
Carboniferous deposits (Romer & Smith 1934) and Early Permian horizons (Olson & Daly
1972, Berman 1976) in North America, and Lower Triassic rocks in Russia (Minikh 1977).

The remaining dipnoan from Triassic Australian rocks (Table 1) also occurs in Cretaceous
deposits. All the material of this species is, unfortunately, badly preserved. An incomplete
and damaged specimen of a large lungfish and associated scales were described by Woodward
(1908) as Sagenodus laticeps. The material was found in the Hawkesbury Sandstone (Ashfield
Shale) at St Peters, New South Wales, in rocks that were considered at that time to be Permian
in age, and Woodward decided on the basis of its age that the specimen belonged in the genus
Sagenodus. Later work has shown that the deposits are Triassic (Wianamatta Group) and Wade
(1931) placed the specimen in Ceratodus. The St Peters material consists of a compressed and
broken head and part of the body and some isolated scales. Little is recognizable in the head
region except for part of the fan-shaped parasphenoid, fragments of the shoulder girdle and the
upper jaw bones which have been sheared across transversely. Thus nothing shows of the
occlusal punctation pattern of the tooth plates, but in shape the bony bases of the upper tooth
plates closely resemble that of the lower jaw bone of Ceratodus avus. Ceratodus laticeps is in
fact a damaged specimen of Ceratodus avus. This species was also described by Woodward
(1906), but on a fragment of a lower jaw tooth plate with a distinctive occlusal punctation
pattern (PI. 3F). The dentition of C. avus is unusually elongate and robust. The holotype of
C. avus was found in a locality at Cape Patterson in Victoria, and is Early Cretaceous in age
(Strzelecki Group, Table 2).

JURASSIC AND CRETACEOUS RECORDS

Jurassic deposits in Australia have so far produced no lungfish, and all Triassic rocks lack
the massive broad tooth plates so characteristic of Mesozoic ceratodont fossils in the Northern
Hemisphere (Agassiz 1833, Miall 1878, Haug 1904, Tabaste 1963, Schultze 1981b, Pinsof
1983 and Kirkland 1987). In this continent, typically 'ceratodont’ tooth plates do not appear

AUSTRALIAN MESOZOIC AND CAINOZOIC LUNGFISH - 473

Table 2:

Species

Ceratodus avus

Ceratodus - sp. 1

Ceratodus - sp. 2

Ceratodus wollastoni

Locality

Cape Patterson,
Vic.

Lovelle Downs St.,
near Winton, Qd.

near Manarrina,
S.A.

Chiltern Hills,
S.A.

West side of
Babbage Peninsula,
Lake Eyre, S.A.

Lightning Ridge,
N.S.W.

Grawan, N.S.W.

Walgett, Co.
Barradine, N.S.W.

Grawan, N.S.W.

Lightning Ridge,
N.S.W.

White Cliffs,

N.S.W.

Clarafield St.,
near Kynuna, Qld.

Cretaceous Deposits Containing Lungfish

Lithologic Unit

Strzelecki Gr.

Winton Fm.

Winton Fm.

Winton Fm.

Winton Fm.

Griman Creek Fm.

Griman Creek Fm.

Griman Creek Fm.

Griman Creek Fm.

Griman Creek Fm.

Coreena Fm.

Mackunda Fm.

Material

posterior frag.

of lower tooth plate,
attached jaw
(Woodward 1906)

upper and lower
tooth plates, attached
bone and other frags.

tooth frags.

tooth frags.

tooth frags.

cast of upper tooth
plate, attached bone

opalised upper tooth
plate without bone

frag. lower tooth
plate, attached bone
(Chapman 1914)

cast of two tooth
plate frags.

casts of one medium
and one large upper
tooth plate and
attached bone

cast of medium upper
tooth plate with
attached bone,
originally described
as Neoceratodus
pattinsonae (White
1926)

damaged upper tooth
plates without bone

474 - KEMP, A.

Neoceratodus forsteri

Neoceratodus nargun

Hereward St., near
Longreach, Qld.

quarry near Winton
Qld.

shore of Lk. Eyre,
S.A.

Manarrina Hill,
cent. Aust.

near Chiltern Hills,
S.A.

Lightning Ridge,
N.S.W.

shore platform,
Pt Lewis, Vic.

Dinosaur Cove,
west of Cape Otway,

Winton Fm.

Winton Fm.

Winton Fm.

Winton Fm.

Winton Fm.

Griman Creek Fm.

Otway Gr. (Zone C)

Otway Gr.

complete medium
lower tooth plate
with some attached
bone

damaged upper tooth
plate with some
attached bone

large upper tooth
plate lacking ridge
1, but having some
attached bone

lower tooth plate

upper tooth plate
frags.

one lower and two
upper tooth plate
frags. with attached
bone

damaged medium
lower tooth plate
with some attached
bone (Kemp 1983)

large, nearly
complete lower

Vic. tooth plate with
some attached bone

isolated scale
(Chapman 1912)

Kirrak, So. Gipps- Strzelecki Fm.

land, Vic.

Ceratodus sp.

Strzelecki Fm. post. frag. para-
sphenoid with cranial

ribs (Waldman 1971)

Koonwarra, So.
Gippsland, Vic.

Ceratodus sp.

EE...

until the Cretaceous in South Australia, the Northern Territory, Queensland and Northern New
South Wales (Table 2). The tooth plates of the three Cretaceous Australian species of
Ceratodus form a coherent group when analysed according to generic characters that are
independent of growth, wear and inherent variation. The most common, Ceratodus wollastoni,
was first described by Chapman (1914) from the anterior fragment of an opalised lower jaw
tooth plate. The specimen was found in Walgett, County Barradine in New South Wales,
(Griman Creek Formation) and was named after T.C. Wollaston, a well known opal dealer who
worked in the area (Table 2). Numerous other specimens have been found more recently, some
from opal beds at Lightning Ridge and the nearby village of Grawan in New South Wales, all
from the Early Cretaceous Griman Creek Formation, and some from Western Queensland and
the Northern Territory (Tables 2, 3). These localities are in the Winton Formation except for

AUSTRALIAN MESOZOIC AND CAINOZOIC LUNGFISH - 475

locality 9-71B in the Simpson Desert which may be Tertiary or Cretaceous (Mond 1974). The
upper tooth plate described by White (1926) as Epiceratodus pattinsonae fits the character

Table 3. Deposits Containing Lungfish of Uncertain Provenance
Species Locality Lithologic Material
Unit

Ceratodus sp. 1

Ceratodus sp. 2

Ceratodus wollastoni

Neoceratodus djelleh

Neoceratodus djelleh

Ceratodus wollastoni

Neoceratodus sp. 4

Neoceratodus gregoryi

Neoceratodus sp. 5

Ceratodus palmeri

Loc. 9-71B, Simpson
Desert, N.T.

Loc. 9-71B, Simpson
Desert, N.T.

Loc. 9-71B, Simpson
Desert, N.T.

Duaringa Basin, near
Duaringa, Qld.

unnamed lake 31°06'S,
140°21'E, S.A.

Billeroo Ck., S.A.
Lawson-Daily Quarry,
Lake Palankarinna,
S.A.

unnamed lake 31°06'S,
140°21'E, S.A.

Billeroo Ck., S.A.
unnamed lake 31°06'S
140°21E, S.A.

Billeroo Ck., S.A.

Ian's Prospect, S.A.

Eight Mile Plains,
Brisbane, southern

Qld.

cf. Tertiary or
Cretaceous

cf. Tertiary or
Cretaceous

cf. Tertiary or
Cretaceous

Tertiary

Eurinilla (Pleist.)

or Namba (Mio.) Fm.

Eurinilla (Pleist.)

or Namba (Mio.) Fm.

?7Mampuwordu Sands
(?Pliocene)

Eurinilla (Pleist.)

or Namba (Mio.) Fm.

Eurinilla (Pleist).

or Namba (Mio.) Fm.

Eurinilla (Pleist.)

or Namba (Mio.) Fm.

Eurinilla (Pleist)

or Namba (Mio.) Fm.

Eurinilla (Pleist)

or Namba (Mio.) Fm.

"Post-Pliocene"

tooth plate frag.
2 tooth plate
frags.

three tooth plate
frags.

one, small upper
tooth plate with
attached bone, one
medium upper tooth
plate (Kemp 1982)
tooth plate

2 tooth plates
(Kemp 1982)

tooth plate

tooth plate

4 tooth plates

8 tooth plates

16 tooth plates

tooth plate

tooth plate with
attached bone
(Jack & Etheridge
1892)

————EEE ~_ssesesees._._.___ ee

476 - KEMP, A.

analysis of C. wollastoni and is regarded as a small specimen of C. wollastoni. It was found at
White Cliffs, New South Wales, in the Coreena Formation (Table 2) and is also opalised. All
of these localities are freshwater deposits. One specimen, not well preserved but recognizable
as C. wollastoni, comes from the Mackunda Formation at Clarafield Station in West
Queensland. Other fossils that occur here are undoubtedly marine (Day et al. 1982). This does
not mean that C. wollastoni was capable of living in both freshwater and marine
environments. It is far more likely that a specimen of C. wollastoni, injured, old or sick, was
washed into the estuary of the river in which it lived and an isolated tooth plate eventually
preserved. Similar accidents have been known to overtake individuals of N. forsteri in the
Brisbane River system. C. wollastoni also occurs in the lacustrine Coli-Toro Formation of
South America (Pascual & Bondesio 1976).

C. wollastoni had heavy broad tooth plates with wide short ridges arising from an anterior
position (Pl. 3A, B). Wear on the surface has produced smoothly rounded ridge crests and
rounded inter-ridge furrows. This indicates a diet, perhaps of soft-bodied invertebrates and
aquatic plants, that produced no rough edges on the tooth plates and required minimal crushing
to prepare it for digestion. The occlusal pits on the functional surface result not from heavy
wear but from differing degrees of hardness in the underlying dentine structure (Kemp 1979,
Schultze 1981b). The species reached a considerable size, up to 4 metres in length, and is the
largest recorded in Australia.

Two new species of ceratodont, closely related to C. wollastoni but not yet described, are
also found in similar localities. The first, designated Ceratodus sp. 1, has more delicate tooth
plates (Pl. 3C,D) and is found in the Winton Formation of western Queensland, at Lovelle
Downs Station (Table 2) and in several places in South Australia and the Northern Territory,
some Winton Formation (Table 2) and some of uncertain stratigraphy (Table 3). Two of the
specimens from Lovelle Downs Station belonged to a large fish, estimated to be about 2
metres in length, and were prepared from a coprolite (Bartholomai, pers. comm.) indicating that
this lungfish existed in the company of a large carnivore. A third smaller tooth plate from a
fish estimated to be 1 m in length came out of the same coprolite.

The second undescribed Cretaceous species, designated Ceratodus sp. 2, had tooth plates that
were broader and flatter than those of C. wollastoni (Pl. 3G, H). It is known from two large
opalised tooth plates, one from Lightning Ridge and the other from Grawan in New South
Wales (Table 2), both in the Griman Creek Formation, There are also some fragments from
the Simpson Desert (Table 3) of uncertain age, either Cretaceous or Tertiary (Mond 1974). The
best specimen of this species (Pl. 3H) is unfortunately in private hands, although the
Australian Museum has a good cast. The other complete tooth plate, from Grawan, which has
no attached bone (Pl. 3G), was recently donated to the Australian Museum by Mrs C. Myer of
Grawan. The tooth plates resemble C. wollastoni in shape, but are broader with shorter ridges
and have occlusal pits on the functional surface. They are similar to the tooth plates of
Ceratodus frazieri from Jurassic deposits in North America (Kirkland 1987), and to C. kaupi
from Triassic deposits in Germany.

Two further species of Cretaceous lungfish from Australia belong to the genus
Neoceratodus. This genus is known from Triassic deposits in Russia (Chabakov 1931) and is
as old as many species of Ceratodus, so its occurrence in the Cretaceous of Australia is not
surprising. One, Neoceratodus nargun, was originally described as Ceratodus nargun by Kemp
(1983), but the tooth plates fit within the generic character analysis of Neoceratodus, and it
belongs in this genus. Since N. nargun was described from one small poorly preserved lower
tooth plate from the shore platform of Point Lewis, Victoria, another Early Cretaceous
specimen, a larger lower tooth plate, also from Victoria, at Dinosaur Cove, East Cape Otway
has been discovered (Pl. 3E). N. nargun had more delicate tooth plates than the recent N.

forsteri, with a higher crown. The Cretaceous specimens of N. nargun both came from the
Otway Group (Table 2).

AUSTRALIAN MESOZOIC AND CAINOZOIC LUNGFISH - 477

The other Cretaceous specimens have aroused some controversy (Martin 1983). They are
opalised, from the Griman Creek Formation at Lightning Ridge, New South Wales. When
analysed using characters of tooth plate and bone that are not influenced by growth, diet, wear
and inherent variation (Kemp 1990) these tooth plates are indistinguishable from those of the
Recent N. forsteri (Kemp & Molnar 1981). They cannot be referred to any other species and
should not be described as new. Martin's (1983) objection to the determination, on the basis of
the position of the pterygopalatine process, is dubious. The position of this process, which
arises from the dorsal surface of the bone and articulates with the calvarium, is inherently
variable in N. forsteri and subject to the effects of diet. Different diets in the Recent lungfish
exert different pressures on the process, and its position is not constant. The position of the
process is therefore not useful for taxonomic determination.

Two other Cretaceous lungfish records from Australia cannot be referred to particular species
(Table 3). One is a large scale found at Kirrak, South Gippsland (Strzelecki Formation). This
was described by Chapman (1912). It has a distinctive shape and ornamentation but is not
associated with tooth plates or skull material. The surface ornament differs from that on the
scales associated with the Triassic C. avus from St Peters (Woodward 1908) and on the scales
referred to C. formosus (Wade 1935). The other fragmentary record, the posterior portion of a
parasphenoid bone with cranial ribs, from Koonwarra in South Gippsland, Victoria (also
Strzelecki Formation) (Waldman 1971), is from a bone which unfortunately shows few
diagnostic characters, but is ceratodontid.

Three of the Cretaceous species, C. wollastoni, N. forsteri and N. nargun are also found in
Tertiary deposits - Miocene in the case of C. wollastoni and Miocene and Pliocene for N.
nargun and N. forsteri. The latter is also the species found still living in rivers and lakes of
southeastern Queensland.

CAINOZOIC LUNGFISH

PALAEOCENE AND EOCENE RECORDS

The dipnoan records from Palaeocene and Eocene deposits in Australia are scarce (Table 4),
and only two localities have produced lungfish material. One, at Redbank Plains, near Brisbane
in southeastern Queensland belongs in the Redbank Plains Group, regarded as Palaeocene by
Day et al. (1982) although it may be Eocene (Stephens, pers. comm.). The other, associated
with the shale oil project at Rundle in central Queensland (Henstridge & Missen 1982) is
Eocene. Oligocene deposits from this continent have so far produced no dipnoan material.

The Redbank Plains locality has yielded several poorly preserved specimens of medium-
sized lungfish. The best of these, impressions of skull bones and tooth plates from a lungfish
about 0.5 m in length, was described as Epiceratodus denticulatus by Hills (1934). However,
the tooth plates conform to the characters of tooth plates of Neoceratodus gregoryi, and
identifiable elements of the skull bones are also the same as skull bones associated with N.
gregoryi tooth plates from Miocene deposits in Central Australia. The other specimens found
recently at Redbank Plains, a tooth fragment and some impressions of disarticulated bones, are
also consistent with the characters of N. gregoryi.

Hills (1934) included some isolated caudal fin impressions of lungfish with his description
of N. gregoryi (E. denticulatus). While it is not certain that these are all from the same species,
it may be safe to assume so, in the absence of identifiable traces of other species. However,
throughout its long history, N. gregoryi occurred in other localities with other species, so the
identity of all the existing isolated tail material with N. gregoryi is by no means proved.

The Rundle locality in central Queensland has produced three fossil ceratodont tooth plates
so far. One, a small posterior fragment, is a specimen of the ubiquitous Neoceratodus sp. 1,

478 - KEMP, A.

Table 4. Palaeocene and Eocene Deposits Containing Lungfish

Species Locality Lithologic Unit Material
Neoceratodus gregoryi Redbank Plains, Redbank Plains Gr. impression of skull
nr. Brisbane, (Palaeocene of and tooth plate
SE Qld. Eocene) frags. - originally
described as
Epiceratodus

denticulatus (Hills
1934); impressions
of disarticulated
bones , one tooth

plate frag.
Neoceratodus sp. 1 Shale Oil Site, Rundle Fm. post. frag. of small
Rundle, cent. (Eocene) lower tooth plate
Qld.
Neoceratodus sp. 2 Shale Oil Site, Rundle Fm. nearly complete
Rundle, cent. (Eocene) upper tooth plate
Qld. with some attached

bone; one upper
tooth plate frag.

Se _______.____ eeeee

common in Miocene deposits of other parts of Australia (Pl. 4A). The other two specimens,
one of which is well preserved while the other is a fragment, belong to a new species,
designated Neoceratodus sp. 2 (Pl. 4B). The fossils of this species came from fish of moderate
size, less than a metre long. The tooth plates are elongate, with a complex punctation pattern
and short thick ridges. Like some of the Miocene ceratodonts described in the next section,
there are peaks of bone at the base of each ridge. This character has less diagnostic significance
than was previously thought (Kemp 1982) and seems to be correlated with the presence of
unusually short ridges, whether these occur as a normal feature of the tooth plate as in
Neoceratodus djelleh (Kemp 1982) or as a developmental abnormality (Kemp 1987).

The scale described by Hills (1943) as Epiceratodus sp. from the Narrows (Rundle
Formation) at Gladstone in Queensland is not dipnoan. The material is most likely an
impression of a fragment of turtle carapace.

MIOCENE, PLIOCENE AND PLEISTOCENE RECORDS

Miocene Formations of Central and Northern Australia were deposited in environments that
provided for an extraordinary radiation of fossil lungfish species, mainly neoceratodontid in type
with a few ceratodont specimens. The latter are all C. wollastoni, from the Etadunna
Formation, and probably represent a relict population. The remaining species, some not yet
described, belong in Neoceratodus or in one of several new related genera. Localities,
formations and species lists (with undescribed species designated by numbers) are given in
Table 5 (Miocene formations) and Table 6 (Pleistocene and Pliocene formations). The
Formations are listed in order of age where possible (Callan & Tedford 1976, Stirton et al.
1968, Rich et al. 1982).

The Etadunna and Namba formations are regarded as medial Miocene, and the Wipajiri as a
channel deposit incised into the clay of the Etadunna Formation (Stirton et al. 1968 and Rich et

AUSTRALIAN MESOZOIC AND CAINOZOIC LUNGFISH - 479

al, 1982). The records from an unnamed formation at Riversleigh in northern Queensland
(medial Oligocene to Miocene), and the Camfield Beds at Bullock Creek in the Northern
Territory (Oligocene-Miocenc) are included although they are incomplete. Riversleigh and
Bullock Creek fossils are still being prepared. Records are similarly incomplete for fossil
lungfish from the Chowilla Fauna and the Bone Gulch Fauna at Moorna Station, New South
Wales (Marshall 1972). Like the material from Chinchilla Sands (Darling Downs,
Queensland), the Malkuni Fauna and the Katapiri Formation, the specimens from New South
Wales may be Pliocene or Pleistocene (Rich et al. 1982, Stirton et al. 1968, Marshall 1972).
The Mampuwordu Sands at Lake Palankarinna, South Australia, appears to be Pleistocene
(Rich et al. 1982). The numbers in brackets after each species indicates the number of
individual tooth plates of that species at the locality. (Fossil toothplates from Australia rarely
come from the same individual. On the four occasions when matched tooth plates were found,
the specimens were counted as one individual).

During 1901 and 1902 Professor J.W. Gregory from Melbourne University led an
expedition to Central Australia. One of the results of this expedition was the discovery of a
number of lungfish tooth plates from Pliocene deposits around Cooper Creek (Gregory 1906).
These were described as two separate species by White in 1925 though in fact three species are
present. Since Gregory's expedition many palaeontologists have collected material from
Cooper Creek and other areas, and ten species are now known from Central Australia.

N. gregoryi (White 1925), present in the Redbank Plains Formation, is also found in
quantity in Central and Northern Australian Formations, and survives at least into the boundary
deposits between the Pliocene and the Pleistocene (Tables 5, 6). This is the only Tertiary
fossil ceratodont from Central Australia that has been found in association with skull material,
and is consequently the best known. N. gregoryi is similar to the Recent N. forsteri in the
shape of the head and the body. Like N. forsteri it lacked ossified centra but had double ended
ribs. The tail was probably diphycercal and elongate. There is a single EQ bone anteriorly,
which is pentagonal in shape, has only a short rostral process and carries a lunar landscape of
projections posteriorly that supported the sensory structures of the snout (Pl. 5A).

Sufficient bones and bone fragments of N. gregoryi skulls survive to show that this species
had a skull structure similar to that of N. forsteri, even in details like a small XK bone behind
the orbit, and vomerine teeth. Other skull bones, not associated with identifiable tooth plates,
have also been found in Central Australian deposits, but these cannot as yet be identified as
belonging to a particular species (PI. 5B).

Size estimation of fossil Australian lungfish is based on the dimensions of the EQ bones
and of the tooth plates. Length and breadth of the EQ bones and tooth plates in NV. forsteri are
closely correlated with body length. The EQ bone of the living species shown in Pl. 5C
comes from a lungfish 112 cm long and it measures 25 mm in breadth. The N. forsteri tooth
plate in Pl. 6G is 28 mm long and comes from a fish 104 cm long. If the same proportions
hold for closely related fossil species, the N. gregoryi EQ bone of Pl. SA would have come
from a fish close to 4 m in length, and the tooth plate in Pl. 6D from a fish about 2 m long.

N. gregoryi had large grinding tooth plates with rounded furrows and a roughened and
complex occlusal surface (Pl. 6D,E). Many of the specimens show evidence of considerable
hard wear. This is not the case with Neoceratodus species 1 (Pl. 4A), which grew almost as
large but had more elegant tooth plates, with long slender ridges and smoothly curved outlines.
Lower jaws of Neoceratodus sp. 1 were included with the upper jaw holotype of Neoceratodus
eyrensis (Pl. 4D), the second species described by White (1925). However, they do not match
this specimen, which has simple punctations, long curving ridges and a prominent procumbent
medio-lingual keel. Also, the holotype of N. eyrensis does not conform to upper jaws that
plainly match the lower tooth plates now referred to Neoceratodus sp. 1. Finally, a lower tooth
plate emerged from the medial Miocene Bob's Boulders Locality near Riversleigh Station in
northern Queensland, that has simple punctations, long slender ridges and a keel (PI. 4C). This

480 - KEMP, A.

specimen does match the upper jaw holotype of N. eyrensis. Both species can now be
described from both upper and lower tooth plates with attached bones. Neoceratodus sp. 1 is
very common, particularly in certain localities, and NV. eyrensis rather rare.

nnn ne —————————eeeeeaanaauananaeEeEE—eEee
Table 5. Miocene Deposits Containing Lungfish

Fauna/Locality

Ngapakaldi Fauna (lakes
Kanunka, Palankarinna
& Pitikanta, S.A.)
(Rich et al. 1982)

West side, Lake
Palankarinna, S.A.
(Lungfish & Tedford
locs.)

Frome Downs, Lake
Pinpa, S.A.

Mammalon Hill,
Lake Palankarinna,
S.A.

North Prospect, Lake
Frome, S.A.

Toescatter Locality,
S.A.

White Sands Basin, Lake
Palankarinna, S.A.

Pinpa Fauna (Lake Pinpa,
=Pine Lake)

Ericmas Fauna (Ericmas
Quarry, Lake Namba
& So. Prospect B,

Lithologic Unit

Etadunna Fm.?

Etadunna Fm.

Namba Fm.?

Etadunna Fm.

Namba Fm.

Etadunna Fm.

Etadunna Fm.

Namba Fm.

Namba Fm.

Species!

Neoceratodus djelleh (8)
Neoceratodus sp. 1 (16)
Neoceratodus sp. 3 (18)
Neoceratodus sp. 4 (35)
Neoceratodus gregoryi (94)
Neoceratodus sp. 5 (1)
Neoceratodus eyrensis (7)
Neoceratodus nargun (2)

Neoceratodus nargun (1)
Neoceratodus sp. 1 (5)
Neoceratodus sp. 4 (13)
Neoceratodus sp. 5 (1)

Ceratodus wollastoni (1)
Neoceratodus gregoryi (5)
Neoceratodus eyrensis (2)
Neoceratodus sp. 3 (4)
Neoceratodus sp. 4 (6)
Neoceratodus sp. 5 (1)

Neoceratodus gregoryi (1)

Ceratodus sp. 2(1)

Neoceratodus gregoryi (2)

Neoceratodus nargun (2)

Neoceratodus djelleh (15)
Neoceratodus forsteri (1)
Neoceratodus eyrensis (8)
Neoceratodus gregoryi (72)
Neoceratodus nargun (1)
Neoceratodus sp. 1 (18)
Neoceratodus sp. 3 (30)
Neoceratodus sp. 4 (38)
Neoceratodus sp. 5 (7)

Neoceratus djelleh (3)
Neoceratodus eyrensis (2)
Neoceratodus gregoryi (51)

AUSTRALIAN MESOZOIC AND CAINOZOIC LUNGFISH - 481

Lake Namba, S.A.) Neoceratodus sp. 1 (9)
& Yanda Local Fauna Neoceratodus sp. 4 (1)
(Lake Yanda, S.A.)

Tarkarooloo Local Fauna Namba Fm. Neoceratodus gregoryi (8)
(Lk. Tarkarooloo, W. Neoceratodus sp. 1 (2)
Lk. Tarkarooloo, S.A.) Neoceratodus sp. 5 (2)

Lake Namba, S.A. Namba Fm. Neoceratodus eyrensis (1)

Long Swamp, S.A. Namba Fm. Neoceratodus gregoryi (3)

Neoceratodus sp. 5 (2)

Lake east of Lake Namba Fm. Neoceratodus sp. 3 (2)
Tinko, S.A.

Kutjamarpu Local Wipajiri Fm. Neoceratodus gregoryi (114)
Fauna (Last Loc., Neoceratodus sp. 1 (604)
E. Lk. Ngapakaldi, Neoceratodus sp. 3 (4)
S.A.) Neoceratodus sp. 4 (14)

Neoceratodus sp 5 (1)

Kutjamarpu Local Wipajiri Fm. Neoceratodus djelleh (1)
Fauna (Lk. Ngapakaldi, Neoceratodus eyrensis (5)
S.A.) Neoceratodus sp. 4 (4)

Riversleigh Sites, Unnamed formation Neoceratodus eyrensis (1)
No. Qld. (Bob's Boulders, Neoceratodus gregoryi (5)
Group Site, Last Minute Neoceratodus nargun (1)
Site, Main Site, Ringtail Neoceratodus sp. 3 (4)
Site) Neoceratodus sp. 5 (2)

Riversleigh Sites, Unnamed formation Neoceratodus sp. 3 (4)

No. Qld. (Melody's Maze,
unnamed loc.)

Bullock Creek Local Fauna, Camfield Beds Neoceratodus sp. 1 (6)
Bullock Creek, N.T. Neoceratodus sp. 3 (6)
Neoceratodus sp. 4 (3)

1 Number in parenthesis indicates the number of tooth plates examined.

2The Etadunna and Namba formations are Early to Middle Miocene in age and the Wipajiri
Formation is a channel deposit within the Etadunna Formation. The unnamed formations at
Riversleigh range in age from medial Oligocene to Middle Miocene and the Camfield Beds are
thought to be of a similar age span, but the details of the stratigraphy are not entirely
resolved (see Rich, T.H. ef al., this volume).

———— Eee... ae

Neoceratodus sp. 2 from Rundle is close to N. djelleh, which was first described from bore
samples near Duaringa, also in Central Queensland (Table 3). The Duaringa deposit cannot be
dated more precisely than Tertiary as pollen in the matrix is sparse (Kemp 1982). N. djelleh
tooth plates from Duaringa (Pl. 4F) are unusual for several reasons: the ridges are short, the
occlusal surface is peculiar because of the matrix that obscures the punctation pattern and the
tooth plate is elongated and narrow. N. djelleh is one of the less common species found in the
Ngapakaldi fauna of Central Australia (Table 5 - Etadunna Formation) the Pinpa and Ericmas

482 - KEMP, A.

faunas of Central Australia (Table 5 - Namba Formation) and the Kutjamarpu Fauna of Lake
Ngapakaldi in Central Australia (Table 5 - Wipajiri Formation). These are all Miocene
localities, and the specimens were preserved in surface sand.

N. nargun, related to both N. eyrensis and N. forsteri, and first appearing in the Early
Cretaceous of Victoria (Kemp 1983) is one of the uncommon but widespread species of smaller
lungfish. The only known upper plate comes from Last Minute, another Riversleigh site in
northern Queensland. N. forsteri is also uncommon in Miocene deposits, but has been found
in material from Lake Pinpa, South Australia (Namba Formation). The remaining small
lungfish tooth plates from Central Australia, all fully formed, can be referred to one of two new
species. One, designated Neoceratodus sp. 3, has 7 to 9 short straight radiating ridges arranged
in a characteristic posteriorly directed fan (PI. 4E). This species was never large, less than a
metre in length. The tooth plates have well worn rounded inter-ridge furrows, suggesting a
rough diet and grinding action of the jaws - the signs of a small omnivore. Neoceratodus sp. 1
is relatively common in most Miocene localities and is also found in a Pleistocene deposit
(Table 6) at Lake Palankarinna in South Australia. The other, Neoceratodus sp. 5 (Pl. 6A),
slightly larger with elongate tooth plates and short thick parallel ridges, is never common and
shows peculiar wear patterns. It sometimes has smooth occlusal surfaces and notched ridges
that indicate that the animal spent a lot of time grinding its tooth plates together (Kemp 1987).
There are a number of possible explanations for this, for example, starvation or excessive
grinding of food material.

The last, relatively common Miocene species is Neoceratodus sp. 4. It had unusual modes
of growth, often developing thin ridges on spurs of bone, or growing split ridges at unusual
places on the tooth plate (Kemp 1987b). This species grew to about two metres in length (PI.
6B, C) and used its high crowned sharp ridged tooth plates for crushing and slicing rather than
grinding. One specimen, QM 10425, was included in N. eyrensis (Pl. 6B) (Kemp & Molnar
1981) but further material has indicated that it belongs in a new species.

Lungfish have little to add to the stratigraphy of Miocene localities in Central Australia and
northern Queensland. No one species is peculiar to any locality or formation (Tables 5, 6), and
there is no readily discernible evolutionary progression from one species to another. They fill
available ecological niches - large herbivore, large carnivore, small omnivore, medium
omnivore and so on. However, some conclusions can be drawn from the numbers of
specimens of each species and the numbers of each size class within species present in each of
the Miocene Formations - Namba, Etadunna, Wipajiri and unnamed sediments at Riversleigh.
In the Wipajiri Formation, the vast majority of specimens are Neoceratodus sp. 1 (80%) and
most of these are sub-adult (45% of total) or juvenile (50%). This applies also to the other
species present. There are very few large (old) specimens. The tooth plates are rounded and
worn smooth indicating a soft diet with a grinding movement of the tooth plates. In contrast,
tooth plates in the Etadunna and Namba formations are mostly large and roughly worn,
sometimes with rounded furrows (grinding), sometimes faceted (crushing). There are few
juvenile or sub-adult specimens, and some of the plates are affected by disease (Kemp 1987b).
The two recent localities of Enoggera Reservoir and the Brisbane River produce populations
with characteristics that parallel these two Miocene formations. Fish from Enoggera Reservoir
are large and old, with worn, faceted tooth plates (Pl. 6H) and frequent evidence of dental
disease. There are few juveniles in this locality. Fish from the richer environment of the
Brisbane River belong in all possible age groups and have healthy tooth plates with a rounded
wear pattern (PI. 6G). Dietary items available in Enoggera Reservoir are comparatively scanty
and harsh, prawns and Hydrilla verticillata (Kemp et al. 1981). Far more soft and easily
available food can be found in the Brisbane River - worms, prawns, algae, eel grass (Vallisneria
spiralis), small bivalves, snails and tadpoles. It appears possible that the areas represented by
the Namba and Etadunna formations were large, deep freshwater lagoons and that the Wipajiri
Formation developed from a river. This supports the previous ideas, based on the mammalian

AUSTRALIAN MESOZOIC AND CAINOZOIC LUNGFISH - 483

fauna, that the Wipajiri Formation is a channel cut into the Etadunna Formation (Stirton ez al.
1968 and Rich et al. 1982).

Table 6. Pliocene and Pleistocene Localities Containing Lungfish

Fauna/Locality Lithologic Unit? Species!

Kanunka Fauna (Stirton Quarry, Katipiri Sands Neoceratodus nargun (1)
Lk. Kanunka; Murrapatterina Neoceratodus gregoryi (20)
St., near Cooper Creek; Neoceratodus sp. 1 (4)
Cooper Creek; Green Bluff, Neoceratodus sp. 4 (3)
Cooper Creek, S.A.)

Malkuni Fauna (Pateramordu and ?Katipiri Sands Neoceratodus eyrensis (15)
Lower Cooper Creek, S.A.) (White 1926)

Neoceratodus gregoryi (5)
(White 1926)
Neoceratodus sp. 1 (3)
(White 1926)

Chinchilla Fauna Chinchilla Sands Neoceratodus forsteri (11)
(Chinchilla, Chinchilla Ceratodus palmeri (8)
Rifle Range, Darling (Krefft 1874)

Downs, S.A.)

Chowilla Fauna Moorna Formation Neoceratodus forsteri (3)
(Chowilla Project and Neoceratodus gregoryi (4)
unknown loc., N.S.W.)

Bone Gulch Local Fauna Blanchetown Clay, Neoceratodus forstert (2)
(Bone Gulch, N.S.W.) ?Moorna Fm. Ceratodus palmeri (1)

Palankarinna Fauna, Mampuwordu Sands Neoceratodus sp. 3 (3)

Lake Palankarinna, S. A.
Billeroo Creek, S.A. Eurinilla Fm. Neoceratodus gregoryi (1)

1 Number in parenthesis indicates the number of tooth plates examined.
2 See Rich, T.H. et al., this volume, for a discussion of the stratigraphic units mentioned in
this chart.

oOo OOOO

The unnamed Riversleigh formations appear to be different from the Namba, Etadunna and
Wipajiri formations. The detailed stratigraphy of the area has not been completely worked out
(Archer, e¢ al. 1989) but the deposits that contain lungfish are regarded as medial Miocene
(Tedford 1967, Smart et al. 1980, Archer pers. comm.). Numbers are still too low to be
certain and material is still being prepared from this locality, but the composition of species is
different, some species e.g. N. djelleh being notably absent. With the exception of the large
lower tooth plate of N. eyrensis, the specimens are small, less than 30 mm in length. The
Camfield Beds at Bullock Creek have so far produced small specimens only, some belonging to
the ubiquitous Neoceratodus sp. 1, although Neoceratodus sp. 3 and 5 and N. gregoryi are also
represented.

484 - KEMP, A.

Pliocene and Pleistocene localities (Table 6) have far fewer lungfish than Miocene
Formations, even though these localities produced the first fossil Australian ceratodont tooth
plates, belonging to Ceratodus palmeri from Chinchilla in Queensland. These were described
in 1874 by Krefft, who also published the first description of the living Australian lungfish in
1870. C. palmeri was one of the larger species, with broad flat cuspless ridges (C. palmeri was
synonymised with N. forsteri by de Vis (1884), but the species is, in fact, quite distinct (Kemp
& Molnar 1981). Most specimens are moderate in size (Pl. 6F). The holotype is a fragment
of a large tooth plate now in the British Museum of Natural History. It was found with most
other specimens of C. palmeri in the Pleistocene or Pliocene localities in the Darling Downs,
southeastern Queensland. One specimen was collected near Brisbane, in a possible Pliocene
locality at Eight Mile Plains, but information on this has since been lost (Jack & Etheridge
1892). Some tooth plates referrable to C. palmeri, mentioned by Marshall (1972) have also
been found in New South Wales in the related Blanchetown Clay, with N. forsteri. N. forsteri
and N. gregoryi have also been found in the related Moorna Formation, (Table 6) also in New
South Wales.

Most of the lungfish species from the Pliocene and Pleistocene deposits, or from the
boundary between the two epochs, are relict populations. Only five of the nine species found
in the Miocene occur in the Pleistocene, two on the Pliocene-Pleistocene boundary and two in
the Pleistocene. C. palmeri is known as yet only from Pliocene or Pleistocene localities.
Natural populations of the one surviving species, N. forsteri, are now restricted to the Burnett,
Mary and possibly the Brisbane rivers in southeastern Queensland (Kemp 1987a) but the
distribution of this species in the Pliocene/Pleistocene boundary deposits ard the Pleistocene
was wider, from New South Wales to the Darling Downs in Queensland, an area much larger
than its present distribution. Localities suitable for lungfish are becoming scarcer in this
continent and we can only hope that lungfish in Australia do not soon become extinct.

CONCLUSIONS

The composition of the dipnoan faunas from Mesozoic and Cainozoic deposits of this
continent differs from equivalent faunas in other parts of the world. The major groups of
Mesozoic dipnoans from other parts of the world are represented in tooth plates from Australian
deposits, but are not known in equivalent numbers of specimens or of species. Few of the
massive, broad-tooth-plated forms so common in European deposits e.g. Ceratodus latissimus,
occur in Australia, and only Ceratodus wollastoni and Ceratodus sp. 2 have massive, broad
tooth plates. One Australian Triassic lungfish belongs in the Arganodus/Asiatoceratodus
lineage, cf. A. tiguidensis. One small lungfish possibly represents the gnathorhizids,
previously a Northern hemisphere genus, and there is one species representing the
proceratodontid - sagenodontid group previously known only from Europe and North America.
There appear to be none of the European ptychoceratodonts (Schultze 1981a) in Australia.
Lungfish with large heavy cutting tooth plates represented by Ceratodus dorotheae (Case 1921)
from the Triassic of Texas and by Ceratodus szechuanensis (Young 1941, Liu & Yeh 1960) and
Ceratodus youngi (Liu & Yeh 1957) from Mesozoic deposits of China are also absent from
Australia. Two of the other Mesozoic lungfish, N. forsteri and N. nargun, represent the
neoceratodontids which also appear in Mesozoic deposits in Europe (N. facetidiens, Chabakov
1931) and in South America (Neoceratodus sp. cf. N. forsteri Pascual and Bondesio 1976). It is
unlikely therefore that neoceratodonts initially evolved in Australia. One Mesozoic species
described mainly on tooth plate material, Ceratodus avus, is of a different and unusual type.
The tooth plates are elongate with short cuspless ridges. Of the two species described mainly
on body form, C. (T.) formosus has the elongate shape and diphycercal tail common to other
known post-Palacozoic dipnoans e.g. Megapleuron zangleri (Schultze 1977) and N. forsteri.

AUSTRALIAN MESOZOIC AND CAINOZOIC LUNGFISH - 485

The other, Gosfordia truncata is unusual and unique - no other known dipnoan has a comparable
deep square tail.

Cainozoic Australian lungfish are remarkable in several ways. The broad-tooth-plated
ceratodonts finally died out in the Miocene of Australia, although they survived later here than
elsewhere. Nowhere in other parts of the world are neoceratodonts found in such variety and
abundance as in Central and northern Australia. The radiation of neoceratodontid and related
genera in the Miocene was extensive, and most of these fish died out in the Pliocene. N.
forsteri, which survives to the present day in southeastern Queensland and which first appeared
in Cretaceous deposits, played only a minor role in this radiation. The unusual variety of
elongate and short-ridged tooth plates represented by the Triassic and Cretaceous C. avus
appears in several related small forms in the Miocene. The diversity of fossil lungfish lasted
much longer in Australia than in other parts of the world. Fossil dipnoans of ceratodont stock
are unknown in Europe after the Jurassic and in Asia, and America after the Cretaceous (Romer
1966, Kirkland 1987). The African ceratodont which survived to the Palaeocene, Ceratodus

Figure 3. Mesozoic and Cainozoic lungfish of Australia are not the direct ancestors of land animals. Apart
from being on the scene much too late, the size and weight of the body, and the relative weakness of the
limbs means that no ceratodont lungfish, to the best of our knowledge, could have crawled out of the water, or
maintained itself on land. The statement, commonly encountered and repeated occasionally in popular
accounts of lungfish, that the Recent N. forsteri is capable of leaving the water deliberately to rest or to evade
predators, is untrue. Whether the lungfish as a group are close to the ancestry of the tetrapods is still open to
question. Drawing by Robert Allen.

486 - KEMP, A,

protopteroides, is now regarded as Protopterus protopteroides (Martin 1984b), so ceratodonts
presumably also died out in Africa in Cretaceous times. Fossil species of Lepidosiren are
present in Miocene deposits in South America, where Lepidosiren paradoxa, the living South
American lungfish, is still found. Several fossil species of Protopterus survived into Eocene
times in Africa (Romer 1966), and several species survive there today. Both of these groups are
unknown in Australia.

A number of recent publications (Rosen et al. 1981, Forey 1987) have claimed that the
Dipnoi is the sister group of the tetrapods. This idea is difficult to justify, as several authors
have pointed out (Campbell & Barwick 1987, Panchen & Smithson 1987). In reaching these
conclusions Rosen et al. (1981) depended heavily on the structure of N. forsteri, which is
regarded as the most primitive of the three living lungfish genera. My investigations of this
genus do not lend any support to their views of lungfish/tetrapod relationships (Fig. 3).

ACKNOWLEDGEMENTS

Most of the material described in this paper was provided from the collections of the
Museum of Victoria, the South Australian Museum, the Queensland Museum, the Australian
Museum, the Museums of Vertebrate Palaeontology at Berkeley and at Riverside, the American
Museum of Natural History, the Hunterian Museum and the British Museum of Natural
History. I am particularly grateful to Dr T. Rich and Mr N. Pledge who sent me material that
they collected, and to Dr M. Archer, who sent me material and gave me considerable assistance
at Riversleigh in 1987. The Joyce W. Vickery Scientific Research Grants and the Ethel Mary
Read Grants also provided financial assistance to visit Riversleigh. The Australian Research
Grants Scheme supported the work with a travel grant to America in 1984.

The illustrations in Figs 2, 3 were drawn by Robert Allen, of the Queensland Museum, and
the photographs in Pl. 1 were prepared by Peter Forey of the British Museum. Gary Cranitch
of the Queensland Museum took the remaining photographs. Dr D.H. Kemp and A. Rozefelds
read and commented on the text. Dr R.E. Barwick and Prof K.S.W. Campbell provided
additional information on Devonian dipnoans, and made helpful comments on the text, together
with an anonymous referee. Thanks are also due to Peta Woodgate, who typed the manuscript.
Draga Gelt helped with the tables.

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WOODWARD, A.S., 1906. On a tooth of Ceratodus and a dinosaurian claw from the Lower Jurassic of
Victoria, Australia. Ann. Mag. nat. Hist. 18: 1-3.

WOODWARD, A.S., 1908. The fossil fishes of the Hawkesbury Series at St Peters. Mem. Geol. Surv.
N.S.W., Paleontology 10: 1-29.

YOUNG, C.C., 1941. On two new fossil fishes from Southwestern China. Bull. geol. Soc. China 21: 91-96.

490 - KEMP, A.

PLATES

Plate 1. The Australian Triassic species, Ceratodus formosus. A, photograph of the holotype P16828, in
the British Museum of Natural History, scale bar = 3 cm; B, Photograph of the head, scale bar = 1 cm.
Specimen photographed by Peter Forey of the British Museum of Natural History.

Plate 2. Photographs of tooth plates and skull bones of Australian Triassic lungfish. A, undescribed
sagenodont skull bone from the Arcadia Formation of Western Queensland; the bone is from the anterior part
of the calvarium and has two accessory bones (arrowed) on one side; B, upper jaw bone and attached tooth
plate of possible sagenodont from the Blina Shales of Westem Australia; C, lower tooth plate of the same
species from the Arcadia Formation of Western Queensland; D, Ceratodus cf. C. tiguidensis (Arganodus cf. A.
tiquidensis) from the Blina Shale of Westem Australia; E, F, Tooth plates of Ceratodus ef. C. phillipsi from the
Arcadia Formation of Western Queensland; E, upper tooth plate with attached bone; F, lower tooth plate.
Scale bar = 2 cm.

Plate 3. Photographs of tooth plates of Australian Cretaceous lungfish. The specimens are shown in
occlusal view. A, upper tooth plate of Ceratodus wollastoni, showing breadth of the tooth plate, occlusal pits
and short thick posterior ridges; ridge 1, long and slender in upper plates of this species, is broken at the
base; Winton Formation, Queensland; B, lower tooth plate of C. wollastoni, showing occlusal pits and the
width of the first ridge of the lower jaw in this species; Winton Formation, Central Australia; C, D, upper
tooth plates and some attached bone of Ceratodus sp., 1; this species also has occlusal pits, but ridge 1 in the
upper jaw is broad; Winton Formation, Queensland; E, lower tooth plate of Neoceratodus nargun; this
specimen has a broken posterior end; Otway Group, Victoria; F, lower tooth plate fragment of Ceratodus avus
from Cape Patterson, Victoria; Strzelecki Group; G, H, upper tooth plates of Ceratodus species 2; this species
closely resembles Ceratodus kaupi from Europe; Griman Creek Formation, New South Wales. Scale bar = 2
cm.

Plate 4: Photographs of Cainozoic neoceratodont lungfish tooth plates from Australia. A, Neoceratodus
sp.1 from the Etadunna Formation, Central Australia; occlusal view of lower tooth plate and attached bone; B,
Neoceratodus sp.2 from the Rundle Formation, Queensland; occlusal view of upper tooth plate and some
attached bone; C, Neoceratodus eyrensis, lower tooth plate from an unnamed formation at Riversleigh, north
Queensland; occlusal view; D, Neoceratodus eyrensis, upper tooth plate from the Katapiri Formation, Central
Australia; occlusal view; E, Neoceratodus sp.3, from the Etadunna Formation, Central Australia; upper tooth
plate showing the posterior fan of ridges characteristic of but not universal to this species; occlusal view; F,
Neoceratodus djelleh from Duaringa in Queensland; upper tooth plate and bone showing the short ridges found
in this species, and the peaks of bone on the ridge base; occluso-labial view. Scale bar = 2cm.

Plate 5. Photographs of EQ (rostral) bones of Australian neoceratodont lungfish. A, the rostral bone
associated with large Neoceratodus gregoryi tooth plates from the Namba Formation, South Australia; dorsal
surface; B, A smaller isolated rostral bone, from the Etadunna Formation, South Australia; dorsal view; C,
D, rostral bones of the Recent lungfish, Neoceratodus forsteri showing variation common in this species; C,
From an adult lungfish, 112 cm long, caught in Enoggera Reservoir, near Brisbane; D, from a sub adult
lungfish, 58 cm long, caught in the Brisbane River; dorsal view. Scale bar = 2 cm.

Plate 6. Photographs of Cainozoic lungfish tooth plates from Australia. A, Neoceratodus sp.4, an upper
plate, from the Namba Formation, Central Australia showing the polished striated occlusal surface and the
notches (arrowed) in the ridges; B, upper tooth plate, Neoceratodus sp.5, from the Etadunna Formation,
Central Australia, previously included in N. eyrensis (Kemp & Molnar, 1981); occlusal view; C, lower tooth
plate, Neoceratodus sp.5, from the Namba Formation, Central Australia; occlusal view; D, E, tooth plates of
Neoceratodus gregoryi from the Namba Formation, Central Australia; D, upper; E, lower tooth plate, both in
occlusal view; F,complete upper tooth plate, Ceratodus palmeri, from the Chinchilla Sands, Queensland,
showing the chevron pattern of the punctations on the occlusal surface and the curved, parallel ridges;
occlusal view; G, occlusal view of a lower tooth plate of Neoceratodus forsteri from the Brisbane River, South
East Queensland; H, occlusal view of an upper tooth plate of Neoceratodus forsteri from Enoggera Reservoir
South East Queensland. Scale bar = 2 cm.

491

NOZOIC LUNGFISH

AUSTRALIAN MESOZOIC AND CAI

PLATE 1

492 - KEMP, A. PLATE

PLATE 3 AUSTRALIAN MESOZOIC AND CAINOZOIC LUNGFISH - 493

PLATE 4

494 - KEMP, A.

PLATE § AUSTRALIAN MESOZOIC AND CAINOZOIC LUNGFISH - 495

496 - KEMP, A. PLATE 6

CHAPTER 15

CHONDRICHTHYANS IN
THE CRETACEOUS AND
TERTIARY OF AUSTRALIA

Nocl R. Kemp!

IMtOGUCTION: 620s is Bie ee eens na uns 498
Systematics of Chondrichthyans ............ 500
Nomenclature of Dentitlion................6 501
MOGUL Y DSS: 0, sarod oust oR otteae este sed Samesatt 502
ND DLO VIAUONS Kos cones ast Bek ensgeedarepeideecanae 503
The Chondrichthyan Record in Australia.. 504
(GLETACCOUSK Bites schege colo sis hecho sesh hd neasielsee 504
Depositional Sctting.............ceeeeeeee 504
CUCENSIANG, por2.5.. Aogedeoseeder es oeens 504
Western Australia.............eeeeeee eee 506
Biostratigraphy and Biogeography...... 506
Aptian, Early Cretaccous ..........+- 506
Albian, Early Cretaceous ............. 506
Coniacian - Santonian, Late
GCrEfACCOUS 8.5.25 Mieke oy pose stele 508
Santonian - Campanian, Late
GrétaCeOusiw has: ei Se: 508
Maastrichtian, Late Cretaccous...... 509
ABELLA LY? cheba scat tac aes Ger yee soe 509

nnn nn aE E EEE

Depositional Setting ...............:.eeeeee 509
Biostratigraphy and Biogeography...... 510
Palaeocene - Eocene ...............2065+ 510

OUP OCENE? oie cpereqesncenveseese'ssehees S11

Early - Middle Miocene................ 513

Late Miocene - Pleistocene........... 515
SUMMALY........... ce eee eee eee ee ee cece eeeeeees eens 518
Acknowledgements ............:.:eeeneeeeeee eres 518
REfENENCES 4:2 Bice tetAvcawesteesipe Tene seses oa 519
PL Ate Si 6 sc: Se aavers Bechet Wieck svt eectece« ots ab 524

1 Tasmanian Museum, G.P.O. Box 1164 M, Hobart, Tasmania 7001, Australia.

498 - N. KEMP

INTRODUCTION

The chondrichthyans embrace all those fish having well developed jaws, paired fins, and an
internal skeleton composed entirely of cartilage. In more familiar terms the class includes
sharks, rays, skates and the chimaeroids or holocephalians some members of which are
commonly known, in Australia at least, as elephant fish, spook fish and ghost sharks. In an
endeavour to clarify the broad and varied usage of terms for cartilaginous fishes Maisey (1984b)
defined the following: In chondrichthyans he included all modern and fossil sharks, rays and
holocephalians; "Shark" he applied as a purely descriptive term to all non-holocephalian
chondrichthyans including batoids i.e. rays and skates. However, Compagno's (1984) use of
"shark" excludes not only holocephalians, but also batoids. The meaning then of "sharks"
should be read in the context of the particular author involved. When used in the sense of
Compagno (1984) "sharks and rays" does serve to emphasize a point which cannot be made as
readily as with Maisey's (1984b) definition. Neoselachians Maisey (1984b) considered as a
monophyletic group of "higher" elasmobranchs. This group includes all living sharks (and
thus rays and skates) and some Mesozoic sharks. The term was coined by Compagno (1977)
and equates with Reif's (1977) Euselachii and Maiscy's earlier (1975) term Euselachiformes.
Selachians is a term which should also be interpreted carefully, again, in the context of the
author, and the date of publication involved. It has been used at an ordinal level e.g. Romer
(1962, 1966) for all "typical sharks, Palaeozoic to Recent, with claspers, narrow-based fins..."
but excluding many Palaeozoic sharks, and all holocephalians and batoids (Romer 1962: 423);
or at a class level e.g. Patterson (1967) for all non-holocephalian chondrichthyans; or,
generally e.g. Cappetta (1987) also for non-holocephalian chondrichthyans but in this context
referring to only Mesozoic, Teriary and Recent taxa.

Compared with other groups of fishes, such as the placoderms and the Osteichthyes, the
chondrichthyan fossil record is markedly incomplete. Bone, be it in the form of an exo- or
endoskeleton, has a much greater chance of preservation and survival than does cartilage. It is
only in exceptionally favourable circumstances that (articulated) remains of the cartilaginous
endoskeleton of chondrichthyans are found, e.g. Dean (1902, 1909), Jaekel (1906), Zangerl
(1973), Capetta (1975), Maisey (1986). However, many parts of the body of chondrichthyans
are hardened by calcium, for example, dermal denticles, spines, vertebrae and teeth, and so the
chances of their preservation are enhanced by this mineralization. The range of hard parts
within the class is not uniform and varies between and within families. Applegate (1967)
describes this variation within some fossil and most living sharks (not batoids).

Dermal denticles, described as microfossils, because they are difficult to see with the naked
eye, are reasonably common in the fossil record. They form an important part of the
Palaeozoic chondrichthyan fauna, e.g. Zangerl (1981), Turner (this volume). Many
contemporaneous teeth are also in the microfossil size range. The methods employed for
recovering teeth, such as acid treatment of limestone (Whitelaw & Kool, this volume),
breaking down and washing and sieving of sediment (Turner, this volume) obviously will
reveal denticles, and other hard parts as well.

In comparison, records of dermal denticles in the Tertiary especially are rare. With the
relative abundance of macro-teeth (greater than 5 mm) techniques used for finding denticles are
usually not necessary. However, when they are employed, dermal denticles are found, ¢.g.
Ledoux (1972), Case (1978). As Cappetta (1987) points out, it might be because of the small
size of dermal denticles that palaecontologists are often not interested in them.

Spines are well mineralised and also form an important component of the chondrichthyan
fossil record, especially in the Palaeozoic and Mesozoic, e.g. ctenacanthids, hybodontids.
Recent publications by Maisey (1982a, 1982b, 1987) and Reif (1982) and references therein
cover aspects such as structure and morphology, occurrences and taxonomy of this field. In

FOSSIL CHONDRICHTHYANS - 499

Australia, however, spines are rare. Palaeozoic records include a xenacanthid from the
Devonian of New South Wales (Young 1982), a ctenacanthiform from the Permian of
Queensland (Leu 1987) and tentative xenacanthiform remains from the Devonian of Victoria
(Turner 1986). There is a paucity of records from the Mesozoic (Long & Kemp 1982), while

Ae Tertiary has apparently yielded only a few specimens (Chapman & Cudmore 1924, Pledge
1985).

Vertebrae are calcified to varying degrees. Many examples of isolated, or only a few
articulated vertebral centra have been described, e.g. Leriche (1910), Chapman (1918), Casier
(1946), Applegate (1964), Cappetta (1980), Itoigawa et al. (1985), but these are often not
readily identifiable, especially down to the specific level.

Springer & Garrick (1964) note that X-radiographs of nearly all living sharks clearly show
the number of vertebrae present. The exceptions are two hexanchoid genera, Hexanchus and
Notorynchus (the six- and seven-gill sharks, respectively) whose vertebrae are insufficiently
calcified to produce and image. The third extant hexanchoid Heptranchias, another sevengill
genus, shows partial calcification of the vertebral centra precaudally, but most caudal vertebrae
are well calcified. The frilled shark, Chlamydoselachus, is an exception, too, having only
some vertebrae, both pre- and caudal, calcified. This notochordal condition, i.e. where the
vertebral column is not well calcified and the notochord is consequently not segmented, is also
found in a number of extant squaloids (Echinorhinus, Aculeola; Compagno 1977, pers.
comm.). In living hexanchoids, it is regarded as a secondary character being derived from
ancestors with well developed centra and a distinctly segmented notochord, e.g. the Late
Jurassic hexanchoid Notidanoides muensteri (Compagno 1977, Maisey 1986) and the
Cretaceous Hexanchus gracilis, which shows a strongly calcified vertebral column, at least
(Maisey & Wolfram 1984).

Notwithstanding the cartilaginous nature of the class, bone does exist in chondrichthyans.
The base of teeth and the pedicle of dermal denticles or placoid scales possess an acellular bone
(Halstead 1974, Moss 1977, Reif 1978b), while the endoskeleton has a thin layer of what may
be considered bone. The flexible cartilaginous endoskeleton is given some rigidity by the
presence of a mosaic of prisms of calcified cartilage, "tesserae" of Applegate (1967), but Moss
(1977) points out that this term should be restricted to the dermal armour of ostracoderms and
placoderms. On the outer surface of these prisms a "cap zone" is laid down by the inner surface
of the perichondrium (a layer of connective tissue surrounding cartilage). On histological
grounds, this cap zone may be interpreted as a thin veneer of bone (Kemp & Westrin 1979).
The fact that osteocytes - the cells which govern the laying down of bone tissue - have never
been found in any chondrichthyans (Applegate 1967) " is accounted for by the transitory nature
of these cells in what is an acellular bone tissue" (Moss 1977: 337).

Teeth are heavily calcified in keeping with their mechanical role in feeding. As sharks have
the ability to constantly shed and replace teeth throughout their life (see below) and as a
consequence of the heavily mineralized nature of the teeth, they are the most common of the
hard parts of macrofossil found in the fossil record. This is especially so in Australia, where
relatively little systematic searching has been conducted. Even with this relative abundance of
teeth compared with the relative dearth of other hard parts - dermal denticles, spines, vertebrae
and chimaerid dental plates - until the 1980's surprisingly little work had been done on any of
these fossils. Their presence in a formation or certain horizon may be mentioned in very
general terms, e.g. Abele (1988: 281, 316), or in faunal lists, e.g. Tate (1888: 169, 1893:
247). Recently Long (1985) and Lees (1986) have published descriptions of two new
Mesozoic chimaeroids from the Cretaceous of South Australia and Queensland, respectively.

Descriptions of Tertiary chondrichthyans (mostly elasmobranchs) are limited to only six
workers in more than a century: McCoy (1875), Chapman & Pritchard (1904, 1907),
Chapman (1913, 1917); Chapman & Cudmore (1924), Pledge (1967, 1985) and Kemp (1970,

500 - N. KEMP

1978, 1982). Papers by these workers included the descriptions of more than SO species, two-
thirds of which are attributable to Agassiz (1833-1844) and Chapman and his co-authors.

Studies subsequent to Agassiz's Poisson Fossiles (1833-1844), e.g. Leriche (1902, 1905,
1910, 1926), revealed that many of Agassiz's species were erected on the basis of dental
morphotypes only. Work by Kemp (1970, 1978, and this paper), Pledge (1985) and to a lesser
extent Keyes (1982) has clarified many of the misidentifications and nominal species included
in the earlier local descriptions. This has been done on the basis of published synonymies and
also by direct comparison with jaws of extant chondrichthyans, as initiated by earlier workers,
e.g. Leriche (1902, etc.) and Casier (1946).

SYSTEMATICS OF CHONDRICHTHYANS

The classification of chondrichthyans used here is adapted from that of Smith & Heemstra
(1986), Compagno (1984) (which do not include extinct taxa), and Cappetta (1987), with
respect to the fossil record.

A radically different classification of sharks and rays proposed by Glikman (1964, 1967,
fide Cappetta 1987) although not accepted for various reasons (see below) must be mentioned.
Glikman divided all sharks and rays into two infraclasses: Orthodonta and Osteodonta, based
on the histological structure of their teeth.

Orthodont teeth have a pulp cavity surrounded by a form of dentine known as orthodentine.
Osteodont teeth have no pulp cavity, with the crown and root being formed from another
form of dentine, osteodentine (=trabecular dentine). The crowns of both types have an outer
layer of enameloid.

Patterson (1966), Compagno (1973), Zangerl (1981) and Cappetta (1987) all reject
Glikman’s classification. Reasons such as: the extant frill shark Chlamydoselachus having
teeth which could be classed as either orthodont or osteodont (Pfiel ((1983); fide Maisey
1987) regards Chlamydoselachus as having orthodont teeth); the common Tertiary carcharhinid
Hemipristis serra having orthodont teeth while its extant relative H. elongatus possesses teeth
of the osteodont type (Compagno 1973, 1984); Glikman's inconsistency of including the
Hexanchidae, the 6- and 7-gill sharks, which have osteodont teeth, in the Orthodonta; the
necessary separation of Pucapristis, with osteodont teeth from Dalpiazia with orthodont teeth,
both being Late Cretaceous batoids of the family Sclerorhynchidae (Cappetta 1987), all
demonstrate the artificial nature of this classification. Additionally, tooth size appears to have
some bearing on the histological nature of the tooth. Larger teeth, such as those of lamnoids
and many carcharhinoids are of the osteodont type. Smaller teeth e.g. in Pristiophorus,
pristids, and many orectoloboids are of the orthodont type (Compagno 1977, Zangerl 1981).

Maisey (1987: 28) while pointing out “at face value, Glikman's suggestions amount to no
more than a gross oversimplification of reality" does add qualifying remarks with respect to
Patterson's (1966) and Compagno's (1973) arguments, suggesting perhaps some ambivalence
in his own mind; however he does not adopt Glikman's classification.

Compagno's (1977) term Neoselachii was proposed to encompass all extant sharks (and
batoids) and certain groups of Mesozoic sharks of modern design e.g. palaeospinacids,
orthacodontids, anacoracids. Work by Reif (1977) on the ultrastructure of teeth of Recent
sharks and the above Mesozoic sharks revealed the presence of three layers in the enameloid: a
shiny layer enameloid (SLE) which prevents the formation of cracks in the tooth; a parallel
fibred enameloid (PFE) which increases the bending strength, and an underlying layer of tangled
(or haphazardly) fibred enameloid (TFE). SLE and PFE are not known from any other group
of fossil sharks nor from any living or fossil batoids (Reif 1977). The PFE then is an
autapomorphy for neoselachian sharks. In neoselachian batoids the PFE layer appears to have
been secondarily lost (Thies & Reif 1985).

FOSSIL CHONDRICHTHYANS - 501

Several differences in classification from that of Cappetta (1987) may be noted. The use of
the genus Carcharodon in keeping with the decisions of Keyes (1972) and Welton &
Zinsmeister (1980) in rejecting Casier's (1960) splitting of Carcharodon into three genera.
The use of the genus Carcharias for Tertiary odontaspidid teeth of similar morphology to that
of the extant species C. taurus (Rafinesque). My tentative thoughts on this subject were
confirmed by the assignation of fossil odontaspidids to either Carcharias or Odontaspis in a
paper in press by Thies (pers. comm.). Cappetta admits (1987: 86) that "The systematics of
the family is far from clear, mainly at the generic level...and their attribution to a genus may
be very difficult." Thies (pers. comm.) suspects that the family contains many generic
synonyms and so has adopted the "lumping" approach. Applying the criteria for the various
genera (e.g. Herman 1975, Cappetta 1987) to the Tertiary teeth at hand has certainly not
enabled their generic differentiation.

Following the recent decision by the ICZN (1987), wherein the name Carcharias is
conserved, the confusion arising from the use of many names for this extant genus, especially
over the past decade e.g. Odontaspis (part.), Eugomphodus, Synodontaspis should now be
removed.

I have retained the Mid-Tertiary species of Le Hon in /surus viz. I. benedeni as I consider
its similarities to other robust species of that genus e.g. J. retroflexus, I. planus strong
enough to warrant its retention. Cappetta (1987) includes the species in his new genus
Parotodus Cappetta (1980) in Glickman's family Otodontidae (Lamniformes, also).

NOMENCLATURE OF DENTITION

Two or three hundred teeth in any one set of jaws of a shark is not abnormal in many
species e.g. Carcharias, Isurus. However, not all of these teeth are in use at any one time.
The teeth of an elasmobranch are being continuously replaced throughout the life of an
individual. The immature teeth, which lie beneath the fold of buccal membrane against the
jaw, are known as replacement teeth. Those which have broken through this membrane and are
in use are termed functional teeth.

A row of teeth comprises the functional tooth, or teeth, plus all the replacement teeth
arising from one dental germinal area of the jaw. A row of teeth lies at approximately right
angles to the jaw line. This and other terms dealing with teeth position and placement in the
jaw are illustrated in Pl. 1 (A-C).

A series of teeth is the line of functional teeth, all in different rows, around the upper and
lower jaw. There may be a single series of teeth, e.g. Squalus, or one or two series of teeth
along the sides of the jaws with two or three series in the anterior part of the mouth e.g.
Isurus. When a number of series are closely juxtaposed they present a continuous "pavement"
of teeth, e.g. Heterodontus, Myliobatis. A sharp boundary often cannot be drawn between
functional and replacement teeth and so a series of teeth does not necessarily form a
continuous line around the jaw.

In those elasmobranchs with a single series presenting a continuous cutting edge e.g.
Dalatias, the replacement of that series always moves forwards as a whole into the functional
position (Bigelow & Schroeder 1948) Where the series does not form a continuous cutting edge
functional teeth are lost - commonly accidently - or shed irregularly and replaced individually
by the next tooth in that row. The rate of replacement varies. In Carcharias for example, a
functional tooth may take from two days to a week to move, after rupturing the gum, to the
outside of the mouth and be shed (Breder 1942). In contrast, in Heterodontus for example, it
takes about four weeks (Reif 1976). This shedding of teeth throughout the life of
elasmobranchs is significant as many teeth found in the fossil record represent teeth lost in
this way. Reif (1976) estimates that during the 10 year life span of the Port Jackson Shark
Heterodontus portusjacksoni (Meyer) it produces somewhere between 2000-4000 teeth.

502 - N. KEMP

As the last part of a functional tooth to calcify is the tip of the root, complete fossil teeth
thus indicate their mature nature (Applegate 1965). If the root tips are incomplete it can mean
it is either a replacement tooth, or there has been post mortem wear. Occasionally a
concentration of teeth is found obviously coming from one individual, but these finds are rare,
e.g. a block of Miocene limestone from Batesford Quarry, Victoria (NMV P12984) has 32
visible associated teeth.

TOOTH TYPES

Heterodonty is the term used to describe the differences in tooth size and shape within a set
of jaws (Applegate 1965).

Monognathic heterodonty (Compagno 1970) is the condition where the morphology of
teeth in one jaw differs along the series.

Dignathic heterodonty (Compagno 1970), the more common condition, is a difference in
morphology of teeth between the two jaws. The hexanchids e.g. Heptranchias and
Notorynchus (P|. 6 A and B respectively) clearly demonstrate dignathic heterodonty, with
differences in tooth morphology within the upper jaws and between the upper and lower jaws.
Obviously in the fossil record this gives rise to individual teeth of many different
morphotypes.

Ontogenic heterodonty (Compagno 1970), is where the morphology of a tooth in the
functional position changes gradually with replacement teeth during growth e.g. in Hexanchus
griseus a lower antero-lateral tooth may have from six to eight or nine crownlets in a shark
about a metre long. In a mature adult, more than two metres in length, a tooth from the same
jaw position may have up to 12 crownlets (Kemp 1978). Other such ontogenic changes e.g.
relative stoutness of tooth (compare the teeth of the juvenile Jsurus oxyrinchus with those of
the mature specimen, P1.17 A and B respectively) proportion of height to width both of tooth
and of crown, degree of serration of margins, may be barely discernible between a functional
and a replacement tooth at a given stage of growth, but as with the number of crownlets, teeth
shed by an individual as a juvenile and as a mature adult would present two different
morphotypes in the fossil record.

Dental sexual dimorphism (gynandric heterodonty of Compagno 1970) is more apparent in
mature specimens of a species exhibiting differences in tooth morphology between the male
and female (Garrick 1967), e.g. teeth of immature males in Carcharhinus brachyurus are similar
to mature females, but the crowns of mature males are distinctly more hooked apically (Bass er
al. 1973). Gynandric heterodonty is common in batoids (Cappetta 1987). In the eagle ray
Myliobatus, for example, there is a marked sexual difference in tooth size, the males having
labio-lingually shorter teeth than the females. As Welton and Zinsmeister (1980) point out,
this is a significant feature as such length-width ratios have been employed by palaeontologists
in order to differentiate taxa.

The salient morphological features of elasmobranch teeth are shown in Pl. 2 (E-H).

As heterodonty is common in elasmobranchs a number of terms have been proposed to
cover the variety of tooth morphotypes which share certain characteristics. Pl. 1C illustrates
some of the terms in use, in this case applicable to Carcharias taurus as proposed by Leriche
(1905) and later modified by Applegate (1965). These terms cover most Lamniformes but
there are variations e.g. Odontaspis has symphyseals in both jaws, Isurus has none.

In hexanchoids on the midline of the lower jaw there is a single tooth termed a medial. In
the upper jaw there are a number of rows around the symphysis, on either side and/or on the
symphysis (Kemp 1978), these are termed symphyseals (Ward & Thies 1987). The remaining
larger teeth of both jaws are termed antero-laterals, the much smaller rows near the jaw
articulation are known, as in Lamniformes, as posteriors.

FOSSIL CHONDRICHTHYANS - 503

Antero-laterals is also applied to the teeth not on or near the symphyseal area of many
Carcharhiniformes e.g. Carcharhinus plumbeus and C. brachyurus (Pl. 33 A and B
respectively). The small teeth on or near the midline are termed medials.

Reif (1976) was unable to apply Applegate's (1965) tooth terminology to the tecth of
Heterodontus as the morphological transitions in their dentition are completely continuous.
This, in addition to ontogenic changes in tooth rows, makes it possible to consistently
identify, with certainty, only one row of teeth in each jaw quadrant. This row consists of the
main molariform teeth and is termed Go. Molariform teeth distal to it are designated G+1,
G42, etc. and mesially, G_1, G_2, etc. Symphyseal and post-symphyseal teeth lie on and near
the midline respectively. A dental formula can be used for comparative purposes by stating the

numbers of each tooth type present in a jaw e.g. Carcharias taurus (P1. 1C) showing the right
side of the jaw only:

Po Lg In A3
PiiL5 A3 S81

Holocephalians do not replace their teeth during their life-time. They possess continuously
growing dental plates which wear on the oral/apical surface and which are replaced on the basal
surface. There are two pairs of these plates in the upper jaw, the anterior vomerines and the
posterior palatines, and one pair, the mandibulars, in the lower jaw (Pl. 2A-D, which also
illustrates the other terminology associated with the dental plates of chimacroids). The
conspicuous areas e.g. Callorhinchus milii (Pl. 2 A, C) and ridges e.g. Hydrolagus sp. (PI. 2
B, D) are known as tritoral surfaces (Bigelow & Schroeder 1948). The term pearlstring dentine
(Zangerl 1981) has been applied to material forming the type of tritors as seen for example in
Hydrolagus sp. Reif (1982) found that the mandibular plate of the Late Jurassic chimaeroid
Pachymylus consisted mostly of osteodentine. Australian taxa, both extant and extinct,
mentioned in the text below are illustrated in Pls 3-40. The localities where all Australian taxa
were found are shown in Fig. 1.

ABBREVIATIONS

Institutions and other collections: AM, Australian Museum, Sydney, N.S.W.;
BMcD, teeth in the private collection (pc) of R. McDonald, Foster, Victoria; CMcC, pc, C.
McCrae, Beaumaris, Victoria; CY, pc, C. Yee, Hamilton, Victoria; DE, pc, D. Evans,
Melbourne, Victoria; DJT, pc, D.J. Taylor, Sydney, N.S.W.; GPe, pc, G. Pedrina, Melbourne,
Victoria; GPi, pc. G. Pitt, Melbourne, Victoria; GSV, Geological Survey of Victoria; ILG,
pe, ILL. Gill, Melbourne, Victoria; JAL, pc. J.A. Long, Western Australian Museum, Perth;
JP, pe. J. Pelchen, Hamilton, Victoria, MA, pc of M. Archer, University of N.S.W.; MM,
Geological Survey of N.S.W; MP,. pc of M. Parker Australian Portland Cement Limited,
Geelong,Victoria, MUGD, Department of Geology, University of Melbourne, Victoria;
NMV, Museum of Victoria; NRK, pc. N.R. Kemp, Tasmanian Museum; OPS, pc.
O.P.Singleton, University of Melbourne, Victoria; PS, pc, P. Simmons, Caulfield, Victoria;
QM, Queensland Museum; RB, pc, R.F. Baird, Monash University, Melbourne, Victoria;
RJFJ, pc, R.J.F. Jenkins, Adelaide, South Australia; SAM, South Australian Museum;
SAMD, South Australian Department of Mines; SW, pc, S. Wright, Portland, Victoria; TFF,
pe. T.F. Flannery, Australian Museum, Sydney, N.S.W.; TM, Tasmanian Museum; UAGD,
Department of Geology, University of Adelaide, South Australia; WAM, Western Australian
Museum. Anatomical: a.d.(a.), angle of divergence of root, acute; a.d.(o.), angle of
divergence of root, obtuse; an., anterior teeth; ant., anterior; bas., basal surface; b.de., basal
denticles (or, lateral denticles or lateral cusplets); cr., crown of tooth; cu., cusp, or apex, of

504 - N. KEMP

crown; di.n., distal notch of margin; d.m., distal margin; (en.), entire margin, i.e. not serrated
or denticulated; fu., functional tooth; ?fu., tooth transitional between replacement series and
functional series; in., intermediate teeth; j.ar., jaw articulation; la.f., labial face of tooth; lat.,
lateral teeth; li-f., lingual face of tooth; 1.j., lower jaw; I.m., left mandibular dental plate; I.r.1,
first row of teeth of left side of jaw; l.r.2, second row of teeth of left side of jaw; l.st.,
longitudinal striations; m.m., mesial margin; n.f., nutritive foramen; o., oral surface; occ.,
occlusal surface; p., palatine dental plates; po., posterior teeth; post., posterior; re.,
replacement tooth; r.m., right mandibular dental plate; r.p., right palatine dental plate; r.r.1,
first row of teeth of right side of jaw; r.r.2, second row of teeth of right side of jaw; rt., root
of tooth; r.v., right vomerine dental plate; s., symphyseal tooth; (s.e.), serrated margin; ser.,
one series of teeth; sy., symphysis of jaw; t.j.ar., towards jaw articulation; t.j.sy., towards jaw
symphysis; tr., tritoral surface; u.j., upper jaw; v., vomerine dental plates; XY, oral margin;
XZ, symphysial margin.

THE CHONDRICHTHYAN RECORD IN AUSTRALIA

CRETACEOUS

Except for a handful of published descriptions e.g. Long (1985), Lees (1986) and see Turner
(1982) little work has been done on the Australian Cretaceous fauna. This is obviously due
in some part to the relatively small number of specimens which have come to light and their
mostly fragmentary nature. An attempt has been made here to figure most Australian
Cretaceous elasmobranch teeth. Plates 3-5 represent virtually all the identified taxa, and about
half the known specimens held in museums in Australia. As can be seen, this then does not
represent a (numerically) large collection. Notwithstanding this, at least 18 taxa representing
about a dozen families have been recognized.

Depositional Setting

Queensland

All the teeth, and chimaeroid dental plates, come from the Early Cretaceous Rolling Downs
Group of the Eromanga Basin. All the units were more or less conformably Geposited with the
Early Cretaceous (Aptian-Albian) Wallumbilla Formation being overlain, in stratigraphic
order, by the Toolebuc Formation, the Allaru Mudstone and the Mackunda Formation, all of
which are Albian in age (Smart & Senior 1980). All four units were deposited in a shallow
marine environment with minor local variations accounting for lithological changes.

The upper part of the Wallumbilla Formation was deposited under paralic environmental
conditions in the southern part of the basin, while to the north there appeared to be limited
communication with the sea resulting in restricted marine and lagoonal environments (Day ef
al. 1983).

The overlying Toolebuc Formation, interbedded black shale and limestones, was deposited
following the establishment of the northern seaway. The Allaru Mudstone resulted from the
deposition under normal marine conditions during this marine transgression. The Mackunda
Formation represents the regressive cycle which began in late Albian times and which closed
sedimentation of the Basin (Smart & Senior 1980, Day et al. 1983). A wide range of fossils -

FOSSIL CHONDRICHTHYANS - 505

SY Cairn
/ : ie Nel - a ‘
ly Richmond
DY 7
Toll wh. 3 Mckinley! —@ — Hughenden
Cape Range gf” Boulia +—@ Aramac
‘ Cardabia H ea ¢ Dartmouth
—--— Pg
ws | Waternal
\ ! 6 Brisbane
Gantheaume Bay e S.A i ~~~ gouthport
Gingin& _ Perth ek ! /
aa ~, | }
Molecap Hill landakot \ A, N.S.W. ff
¢ fing og ~. cs dney
oe Adelaide ak \ y
r= ! vic a) cater
Sry ee
> Melbourne
Cy Tas.
“Hobart
@ Broken Hill
| ® Buckalow
SOUTH AUSTRALIA
Morgan NEW SOUTH WALES
Blanchetown @ ren Ne ole :
\Abattoirs YW \f I % Sydney
\ Bore, % : $
3505 rennants ae
1 Adelaide ° sp Murray | \
Marion No.2 Bore Brg es hy miseINdee: L
Noarlunga Talem *ioBores
e < 35°S
J Wellington | * ues =
fl vi I * ee
Strathalbyn | SE TORIA Nacl: ek Deca -
Blanche i Geelong ~~
Point | Batesford \
| Naracoorte, E & Curlewis '
Robe @1 WS No 5 Bore Native Hut _| Fyansford H
Peels : | Hamilton creen _--~| Waurn Ponds —~
Bore | 4 Grange Burn ! Keilor "Ts
/\Muddy Creek / / a . ™Eden
i Byaduk @ / lelbourne
Mount Gambier yaeus Birregurra / South Yarra Bairnsdale
Portland Kawarren\y@ Beaumaris Lakes Entrance
Pt Fairy e \ Jemmys Point
Princetown ~~~ / \ Meromans:
< iter A Aireys | | Torquay reek
Gellibrand River J / Inlet. | |Bird Rock
eA0SS. Rivernook Jan Juc | Mornington
Aire River Point Balcombe Bay
Browns Creek D Addis Fossil Beach
Fishing Point 5 Grices Creek Flinders Island
Hordem Vale Cart Manyung Cameron Inlet 400
8G, Rocks yy 40°95
Cape Grim Fossil Bluff
Table Cape

TASMANIA

Macquarie Harbour

Hobart

1400E | 1500E

Figure 1. Localities producing Cretaceous and Tertiary chondrichthyans in Australia.

506 - N. KEMP

reaper ae invertebrate and vertebrate - has been found in the Rolling Downs Group ( Day
etal. FE

Except for the Aramac and Barcaldine (and the Pulchera Waterhole and Elizabeth Springs,
see below) localities, all material is of Albian age. The former two localities could be from
the Wallumbilla or Toolebuc formations or Allaru Mudstone and so the age cannot be more
precise than Aptian-Albian (A.C. Rozefelds, pers. comm.)

The Pulchera Waterhole and Elizabeth Springs localities are associated with mound springs
and are mapped as Pleistocene in age. The sharks’ teeth and associated belemnites are
presumably being reworked from the underlying Lower Cretaceous Beds (A.C. Rozefelds, pers.
comm.).

Western Australia

The Molecap Greensand and the Gingin Chalk, where they outcrop at Gingin, north of
Perth, are the two main sources of sharks’ teeth.

The Late Cretaceous (Coniacian-Santonian) Molecap Greensand, rich in microflora but rare
in macrofossils, was deposited in a shallow sea over an irregular topography, with conditons
apparently unfavourable for the development of a normal benthic fauna (Playford et al. 1976).

Overlying the Molecap Greensand, with apparent conformity, is the Gingin Chalk, also of
Late Cretaceous (Santonian-Campanian) age. It was deposited in a shallow, warm sea, and is
characterised by an abundant coccolith assemblage which is little diluted by terriginous detritus
(Playford et al. 1976).

A third source of teeth is the Ascot Beds, a thin sequence of Pliocene marine sediments
unconformably overlying the Cretaceous and Eocene-Palaeocene deposits in the Perth Basin
(Baxter & Hamilton 1981). Black, very worn and polished fossils and phosphatic nodules are
found in the Ascot Beds. These are thought to be reworked from the underlying Cretaceous
sediments (Baxter & Hamilton 1981; Kendrick 1981, and pers. comm.). More recent studies
on the sharks’ teeth from this source (Kemp 1982) have proven to be unfruitful as no firm
identifications have been made due to the very worn and polished nature of the specimens.

Biostratigraphy and Biogeography

Aptian, Early Cretaceous

The chimaerid Edaphodon eyrensis Long from the Aptian, Lower Cretaceous Bulldog Shale,
south of Lake Eyre, South Australia is one of only two chimaeroids known from the
Australian Mesozoic. It is also one of the earliest known species of this genus (Long 1985,
Lees 1986). The confirmed Aptian age makes it probably the earliest Australian Cretaceous
chondrichthyan taxon. The Queensland material which may be of Aptian age (see above) is
included in the following, Albian, section.

Albian, Early Cretaceous

From the Toolebuc Formation of western Queensland comes the other Australian Mesozoic
chimaeroid, a callorhinchid, Pryktoptychion tayyo Lees (Lees 1986). The chimaeroids, which
reached their maximum diversity during the Cretaceous were widespread during the Mesozoic
and have been recorded from North America, Russia, Europe and New Zealand (Hussakoff
1912, Ward & McNamara 1977, Keyes 1981, Lees 1986).

Two rostral teeth of the sawshark Pristiophorus from the Toolebuc Formation of central
Queensland are referred tentatively to P. tumidens (Woodward 1932). This is the first record of
Pristiophorus from the Cretaceous of the southern hemisphere (Keyes, pers. comm.) and the
oldest Pristiophorus in the fossil record. Cappetta (1980) described complete skeletons of P.
tumidens from the Upper Santonian, Upper Cretaceous of Lebanon, the only other known
Cretaceous occurrence of Pristiophorus (Cappetta 1987).

FOSSIL CHONDRICHTHYANS - 507

The cretoxyrhinid, Cretolamna appendiculata (Ag.) was a cosmopolitan, long-ranging
species, Early Cretaceous-Early Eocene (Cappetta 1987), and so it is not surprising to find it in
both the Queensland fauna and also the younger (Late Cretaceous) Western Australian fauna.
Many broken teeth in the material to hand are probably also attributable to this species.

Five teeth collected from the claypans of the Simpson Desert, Northern Territory, near its
borders with Queensland and South Australia (24° 12' S 136° 40' E, Mond, 1974; J.W.
Warren, pers. comm.) are assigned tentatively to this species. Photocopies of the teeth (N.S.
Pledge, pers.comm.) show their morphology to be similar to teeth from the Queensland Albian
(e.g. Pl. 4 A, B) and the Upper Cretaceous of the Belgian Basin (Herman 1975). Tentative
thoughts about the outcropping material of the clay pans being of Cretaceous age (J.W.
Warren, pers. comm.; A. Mond, pers. comm.) and not Etadunna Formation and of ?Oligocene,
possibly Miocene age (Mond 1974) are confirmed by the presence of this Cretaceous-Early
Eocene species.

Another cretoxyrhinid, Paraisurus macrorhiza (Pictet & Campiche 1858) occurs in the
Queensland Toolebuc Formation. The genus is restricted to the Albian, Lower Cretaceous,
although its geographical distribution is wide e.g. France, England, North America, USSR
(Cappetta 1987).

A number of fragmentary teeth, mostly represented by only crowns, are referred tentatively
to Cretoxyrhina on the basis of their general morphology, which is very like that of C.
mantelli (Ag.), and especially the thick (labio-lingually) root, when present.

Teeth of a form relatively common in the Allaru Mudstone of Queensland are referred
tentatively to “Lamna" arcuata Woodward. This species occurs in the Upper Cretaceous of
Europe e.g. England, Holland, France, Belgium (Herman 1975). The status of the genus of
this species is unclear. Herman (1975) included it in his new genus Plicatolamna. Cappetta
(1987) however, while placing a number of species of Herman's Plicatolamna in Sokolov's
Cretodus, a cretoxyrhinid, hesitates to include Woodward's species, which he (Cappetta 1987)
leaves as "Lamna" arcuata. As this name is adopted here no further geographical or
stratigraphical comparisons are made.

The most common elasmobranch taxon in the Qucensland Albian is the anacoracid
Pseudocorax australis (Chapman 1909). This occurrence is the oldest record of the genus in the
fossil record. Pseudocorax is widespread geographically, being found in Europe, North
America and North Africa, but restricted to the Turonian-Maastrichtian, Upper Cretaceous of
these continents (Cappetta 1987)

Similarly, another anacoracid, Microcorax - whose presence is based on a single, very well
preserved tooth from the Queensland Albian, identified only as Microcorax sp. - also found
only in the Upper Cretaceous (Cenomanian-Campanian ) of Europe, North America and North
Africa (Cappetta 1987), represents the earliest record for this genus.

The palaeospinacid Synechodus (or Palaeospinax, the two genera may be synonymous,
(Cappetta 1987, Thies, pers. comm.)) from an unknown horizon associated with Pleistocene
mound springs (see above ) in western Queensland, is the earliest of the two Australian
occurrences of this genus, which is also found in the Upper Cretaceous of Western Australia.
The genus is wide ranging both geographically e.g. Europe (Herman 1975, Thies 1983), North
America (Case 1978), New Zealand (Chapman 1918, Keyes 1981) and stratigraphically, from
the Early Jurassic (Thies 1983) to Palacocene (Keyes 1981).

Two teeth are referred to the mitsukurinid genus Anomotodon. There are a number of
specimens to hand of this general form but in many case they are more eroded and so it is
difficult to tell if basal denticles were, or were not present. If not, they may belong to another
mitsukurinid genus, Scapanorhynchus. Both genera are known from the Lower and Upper
Cretaceous and their geographical and stratigraphical distribution is widespread (Cappetta
1987). A tooth referred tentatively to S. subulatus (Ag.) (PI. 3M) may belong to the
odontaspidid genus Palaeohypotodus (Thics, pers. comm., Herman 1975). Cappetta (1987)

508 - N. KEMP

considers "Scapanorhynchus" subulatus to be an odontaspidid. In light of these opinions,
geographical and stratigraphical comparisons will not be made for this specimen,

Coniacian-Santonian, Late Cretaceous

The teeth referred to Notorynchus sp. from the Western Australian Molecap Greensand are
the only Late Cretaceous Notorynchus known. The genus is represented by a single species,
N. aptiensis (Pictet 1864) from the Lower Cretaceous of France, Germany and England. The
Western Australian specimens differ from this taxon and is probably a new species (Ward &
Thies 1987, Cappetta 1987, Thies, pers. comm.). Notorynchus is well known from the
Tertiary (see below).

Teeth identified as representing two genera Centrophoroides and Protosqualus indicate the
presence of these two squalids in the shallow Late Cretaceous seas of (now) Western Australia.
A number of fragmentary tecth may also be referrable to these taxa whose characteristics do not
yet seem to be clearly defined (Herman 1975, Thies 1979, Cappetta 1987, Thies, pers.
comm.). Both genera are known from the Cretaceous and Upper Cretaceous respectively of
Europe (Cappetta 1987).

The angel shark Squatina is a long ranging - Late Jurassic of England (Thies 1983), to the
present day - and geographically widespread genus, both extinct and extant (Compagno 1984,
Cappetta 1987). Several teeth and a number of fragments, identified only as Squatina sp., are
represented in the Molecap Greensand. Teeth from modern Squatina differ little from
Cretaceous and Jurassic examples (pers. observ., Thies 1983) and thus identification of a few
isolated teeth to a specific level is virtually impossible.

As mentioned above, the mitsukurinid genus Scapanorhynchus is both geographically and
stratigraphically widespread. Several teeth, and crowns, from the Molecap Greensand are
referred to Scapanorhynchus sp.

Another cosmopolitan taxon, Cretolamna appendiculata, found in the Albian, Lower
Cretaceous of Queensland (see above) also occurs in the Molecap Greensand of Western
Australia,

A small number of incomplete teeth are referred tentatively to another cretoxyrhinid genus,
Protolamna. Preservation of the specimens is such that it is difficult to judge if there were
basal denticles present or not. Protolamna ranges from the Aptian, Early Cretaceous, to the
Cenomanian, Late Cretaceous (Cappetta 1987).

Synechodus is another of only a few taxa which are common to both the Queensland Lower
Cretaceous strata (see above) and the Western Australian Cretaceous, once again the Molecap
Greensand. The Western Australian specimens referred to Synechodus, also can not be readily
equated to any known species from Europe (Thies, pers. comm.) nor New Zealand (Chapman
1918). When compared with the New Zealand species they lack the marked labial striations of
S. sulcatus (Davis 1888) and possess a more attenuated crown than S. validus (Chapman
1918). The Australian specimens are left as Synechodus sp. until more, and hopefully better,
material becomes available.

An unnamed chimaeroid from the Molecap Greensand, possibly [schyodus, is currently
under study (K.J. McNamara, pers. comm.).

Several isolated vertebral centra, some complete, some broken, have been recovered from
the Molecap Greensand. They are only identified as Lamniformes, which is in keeping with
the presence of teeth of a number of taxa of this order in the horizon.

Santonian-Campanian, Late Cretaceous

The odontaspidid genus Hispidaspis occurs in the slightly younger Gingin Chalk, from
Western Australia. Hispidaspis is found in beds of from Early-Late Cretaceous age in Europe
and USSR (Cappetta 1987). Several more incomplete specimens are referred tentatively to
Hispidaspis.

FOSSIL CHONDRICHTHYANS - 509

The Toolanga Calcilutite, in the Carnarvon Basin, does actually extend into the
Maastrichtian, as shown by drilling. However, in the southern part of the Basin it is of
Santonian-Campanian age, with the Santonian fauna from the base of the formation being
essentially the same as the Gingin Chalk of the Perth Basin (see above) (Playford et al. 1975,
G.W. Kendrick, pers. comm.). It is from this part of the formation that "Lamna" cf venusta
os This species is present in the Upper Cretaceous of Belgium and France (Herman

A well preserved crown, identified as Cretoxyrhina cf mantelli, from the Gingin Chalk, is
in keeping with the stratigraphic limits of the species, which, while present in the Santonian
of many continents e.g. Europe (Herman 1975), Africa (Dartevelle & Casier 1943), U.S.A.
(Cappetta & Case 1975) does not seem to reach the Campanian (Cappetta 1987).

Maastrichtian, Late Cretaceous

Sharks’ teeth have been recorded from the Miria Formation, of Maastrichtian age (Playford
et al. 1975, A.E. Cockbain, pers. comm.), from Cardabia Station in north-western Western
Australia. A large, reasonably well preserved tooth identified as Cretoxyrhina mantelli extends
the range of this species from the previously oldest occurrence of Santonian age (Cappetta
1987).

TERTIARY

Unlike the Western Australian and Queensland Cretaceous chondrichthyan remains, which
come from a small number of horizons, the Tertiary fauna occurs in a (relative) multitude of
formations. These then will not be dealt with in an initial description as were the Cretaceous
beds, but in stratigraphical order, in relation to the taxa found therein.

Depositional Setting

The depositional settings of the various strata in which south-eastern Australian
chondrichthyans occur have been taken from only a few references viz. southern Australia,
McGowran (1989); South Australia, Pledge (1967, 1985), Abele et a/. (1988); Victoria,
Darragh (1985, 1986), Abele et al. (1988) and Tasmania, Sutherland & Kershaw (1971) and
Quilty (1972). References for Western Australian localities are more diverse and along with
relevant personal communications for all States, are mentioned in the text (below).

In south-eastern Australia there are perhaps 50-60 sites which have yielded chondrichthyan
remains. Compared with many overseas localities e.g. the Eocene London Clay, England, the
Miocene Shark Tooth Hill locality, California, U.S.A., the Tertiary Paris and Belgium Basins,
our deposits are, on the whole, relatively barren. The few productive sites include the
winnowed deposits of the nodule beds: the Late Miocene (the nodule bed is at the base of the
Black Rock Sandstone, which extends into the Early Pliocene; unfortunately it is not always
possibie to say whether the fossils come from the nodule bed or higher up in the formation)
Beaumaris and the latest Miocene-earliest Pliocene Grange Burm localities where fossils are
usually very worn. Many sites, however, yield specimens the numbers of which may just run
into double figures. The record is further fragmented by the lack or paucity of fossiliferous
outcrops of certain ages e.g. Early Palaeocene, Early and Middle Eocene, Early Oligocene and
Middle Miocene. Little new material has come to light over the last seven to eight years to
warrant changing this description (Kemp 1982).

The inclusion of elasmobranch material from the Western Australian Museum collections
in the present study also, unfortunately, does little to enhance this basic picture. A few more
taxa have been added to some localities in the southeast of the continent (see below); it seems
then that the west also yields only desultory finds, more often fragmentary than not.

510 - N. KEMP

As mentioned above there has been little systematic collecting done in Australia, and the
collections in the Museum of Victoria demonstrate this influence of a collecting bias, which
obviously has some bearing on the faunal lists. In the case of /surus hastalis (Ag.), the extinct
mako or blue pointer, more than 1500 teeth come from the Beaumaris nodule bed. The teeth
of this species are more or less flat and triangular, the uppers more so than the lowers, the
anteriors more so than the smaller laterals and posteriors (PI. 20A). About 60% of the teeth of
I. hastalis in the collection are from the upper jaw. The majority of these are anteriors or large
laterals; very few are posteriors and small laterals. Similarly most teeth from the lower jaw are
anteriors and large laterals. There are more teeth of J. hastalis in the collection than all the
other taxa of elasmobranchs combined. This may reflect the relative abundance of this species
in the late Tertiary seas, but it is more likely that it is a reflection of collectors seeing and
picking up large, more noticeable teeth. It seems unlikely that there was a dearth or total lack
of many smaller species of the elasmobranchs found in contemporaneous deposits outside
Australia e.g. Triakis, Dasyatis, Oxynotus, Scylliorhinus, Rhina, Galeus, Squalus,
Centrophorus , Dalatias, (Leriche 1910, Ledoux 1972, Cappetta 1976, Welton 1981, Keyes
1984), and indeed in southern Australian waters today.

In order to test this apparent lack of teeth of smaller species field work in 1984 resulted in
the collection of both bulk samples and sieve concentrates from the major, and a number of
minor, Victorian localities, as well as several Tasmanian localities. At the Tasmanian
Museum further sieving and binocular microscopic examinations of tens of kilograms of the
concentrates revealed not a single elasmobranch remain. In addition to the limestones a
number of other rock types were sampled e.g. the highly fossiliferous (predominately
gastropods and foraminifera) calcareous, clayey silt of the Middle Miocene Fyansford
Formation in Balcombe Bay, Victoria, this too proved barren (of sharks' teeth and denticles).
No satisfactory answer has yet been found to explain this situation.

Environments of deposition are not easily ascertained on the basis of shark faunal
assemblages alone. Teeth shed by pelagic sharks are going to drop to the bottom be it in deep
or shallow water. Coastal dwellers however, will give an indication of their habitat, by their
remains. The fortuitous occurrence of one or a number of species in a certain horizon may lead
to erroneous conclusions. This is especially so with a fragmentary record such as exists in
Australia; usually the associated molluscan assemblages give an accurate indication of the
conditions at the time of deposition.

Biostratigraphy and Biogeography

Palaeocene-Eocene

The dominant shark of the Early Tertiary in Australia was the odontaspidid Carcharias. The
teeth ascribed to this genus practically all show strong, or at least incipient, lingual striations.
C. sp. and C. macrotus (Ag.) are found in the Western Australian Boongerooda Greensand of
the Cardabia Group, which unconformably overlies the Maastrichtian Miria Marl, in the
Carnarvon Basin, in the northwest of the State (Playford et al. 1975). Present too in the
Boongerooda Greensands was a hexanchid which is currently under study (J.A. Long, pers.
comm.).

In southeastern Australia, in the shelf seas and shallow bays of the deepening Otway Basin
a number of taxa are represented. From the Middle Paleocene Pebble Point Formation, near
Princetown, Victoria, has been found a single tooth of the large lamniform Otodus obliquus
Ag. This species occurs in Europe, Africa and North America e.g. England (Casier 1966, Gurr
1963), Belgium (Leriche 1905), U.S.A. (Leriche 1942) but is restricted to the Late Palaeocene
to Early Eocene horizons. The Victorian specimen, then, extends its range back to the Middle
Palaeocene.

FOSSIL CHONDRICHTHYANS - 511

Carcharias sp., C. macrotus and a more slender form of Carcharias, here referred to C. cf.
acutissima, and a hexanchid, Hexanchus ?sp. 1, are also represented in the Otway Basin, being
found in the Trochocyathus Bed at Rivernook, near Princetown. The Carcharias species are the
most common shark taxa; only one hexanchid tooth has been found so far. C. macrotus is a
species common in the Palaeocene-Eocene of England (White 1931, Ward 1980), France and
Belgium (Leriche 1905, Casier 1946) and Africa (Casier 1946).

Following the marine regression of the Early Eocene the sea advanced diachronously during
the Middle and Late Eocene. In the western part of the basin - the Gambier Embayment - from
the middle Eocene horizons, are found Otodus obliquus, Carcharias sp., C. macrotus, C. cf.
acutissima and a mitsukurinid, Mitsukurina maslinensis (Pledge) known only from South
Australia to date. Also preserved in these pelletal chamositic greensands e.g. in Peel's Bore,
near Robe, South Australia, are Myliobatis sp., a cosmopolitan genus of ray which is found
throughout most of the Tertiary (Cappetta 1987), Hexanchus agassizi Cappetta and another
Hexanchus, H. sp. 1 which may be conspecific with the specimen from Rivernook.

The majority of specimens from Peel's Bore are well worn crowns but there are some
complete teeth preserved e.g. Mitsukurina maslinensis, which appear to be restricted to
horizons representing shallow water.

Further to the west, still in South Australia, Carcharias sp., C. macrota and Mitsukurina
maslinensis are also present in the Late Eocene deposits e.g. Tortachilla Limestone.

Contemporaneous beds from two basins in southern Western Australia viz. the Nanarup
Limestone of the Bremer Basin and the Toolinna Limestone of the Eucla Basin (McGowran
1989) have yielded a number of odontaspidid teeth here referred to Carcharias sp.; most
specimens being preserved as only crowns - usually striated lingually - or with only portions
of roots and rarely a basal denticle.

In the Late Eocene Blanche Point Marl of the South Australian St Vincent Basin
(McGowran 1989) are found, again Hexanchus agassizi and the first record in Australia of the
other seven gill hexanchid, Heptranchias, as H. howellii (Reed), Myliobatis spp., Carcharias
sp., C. macrotus, and Mitsukurina maslinensis. The type specimen of Heptranchias howellii
is also from the Eocene, of the U.S.A. (Reed 1946) and the species occurs in other Eocene and
also Oligocene localities from that continent (Welton 1974) as well as the Oligocene of Japan,
as H.ezoensis (Applegate & Uyeno 1968).

On the west coast of Tasmania ?7Eocene (Upper Cretaceous-Lower Tertiary) strata are
predominantly continental but there were short transgressive intervals which resulted in the
deposition of thin calcareous layers rich in bivalves. These deposits, which are very leached
carbonaceous siltstones, have yielded a single tooth of Carcharias sp.

The last appearance of Otodus obliquus is in the Upper Eocene/Lower Oligocene from the
Olney No. 1 Bore in northwestern Victoria. The single tooth is from the siltstones and
claystones of the Olney Formation deposited under marginal marine conditions of the Murray
Basin. In Europe and North America O. obliquus is restricted to the Paleocene and Eocene
(Gurr 1963, Casier 1966, Leriche 1942).

Oligocene

During the Early Oligocene the continental shelf prograded resulting in little marine
deposition. The sea advanced strongly towards the basin margins during the mid-Late
Oligocene and the littoral to shallow, high energy environments supported a large invertebrate
and - judging by the reasonable numbers of sharks' teeth - a moderate vertebrate community as
well.

Typical of these deposits are the limestones found at Mt Gambier, South Australia (Clifton
Formation), Waurn Ponds and Airey's Inlet (Waurn Ponds and Point Addis Limestone
Members, respectively, of the Jan Juc Formation) all being bryozoal calcarenites with the latter
two being exceptionally coarse grained. The marly horizons, ¢.g. at Bird Rock near Torquay,

512 - N. KEMP

of the Jan Juc Formation, and the Ettrick Marl, the contemporaneous beds in the Murray
Basin, to the west, were deposited in shallower water, subjected to less wave and current
action.. Further west again, in the eastern part of St Vincent Basin, the sediments of the Port
Willunga Formation were deposited during the late Eocene, and Oligocene in a basin with
restricted access to the open seas (Lindsay & McGowran 1986).

The last Australian Tertiary appearance of Hexanchus, as H. agassizi is in the Ettrick Marl,
near Wellington, South Australia. Overseas H. agassizi continues until the Miocene, in
Europe (Cappetta 1987).

The cosmopolitan, very long ranging, Jurassic-Recent, Heterodontus makes its first
appearance in the limestones of the Maude Formation of Victoria as H. cainozoicus. It is not
clear if the specimens are from the Lower, or Upper Maude Limestone, and so they could be
either Late Oligocene, or early Miocene in age. Carcharias macrotus is still present, being
found in the Waurn Ponds Limestone, but now, as with the Miocene representatives, the teeth
are very much larger. Dartevelle & Casier (1943) and Casier (1946, 1966) maintained the same
name for both the early Tertiary (Palacocene) and younger (Late Oligocene-Early Miocene)
forms in Europe, noting a significant increase in size over this time.

Another Carcharias form occurs in the Jan Juc Formation, both in the marls at Torquay,
and in the Waurn Ponds Limestone Member, and also in the Kawarren Limestone. This form
is indistinguishable from the extant C. taurus Raf. Cappetta (1987: 91) notes that "Pliocene
specimens (of Synodontaspis acutissima) are hardly separable from the Recent S. taurus”

In Australia this problem then arises earlier. Rather than call teeth of this morphotype, of
the common European Miocene-Pliocene taxon, C. acutissima, I refer them to the extant
species until suitable material becomes available to differentiate them on morphological rather
than stratigraphical grounds.

The genus /surus which ranges from the Late Palaeocene to Recent (Leriche 1905) makes
its first appearance in Australia with two forms, /. desori (Ag.) and J. planus (Ag.). Overseas
neither species appears until the Miocene e.g. J. desori in the Miocene of Africa (Dartevelle &
Casier 1943) and Europe (Cappetta 1970), and J. planus, in the Miocene of California, U.S.A.
(S.P. Applegate, pers. comm.). Both species are found in the marls of the Jan Juc Formation,
and J. desori also occurs in the Kawarren Limestone, and the Waurn Ponds Limestone, where it
is probably the most common taxon.

Carcharodon angustidens (Ag.) is the first species of Carcharodon to appear in Australia. It
is known from a number of localities and formations including, in Victoria, the limestones at
Point Addis, the Jan Juc Formation at Torquay, and the Waurn Ponds Limestone Member at
Waurn Ponds; South Australia, the Port Willunga Formation, in Aldinga Bay, south of
Blanche Point, the Gambier Limestone at Mount Gambier (Late Oligocene-Early Miocene). It
also occurs in the Oligocene overseas e.g. Belgium (Leriche 1910), Frarice (Priem 1906), New
Zealand (Chapman 1918, I.W. Keyes, pers. comm.).

The relatively uncommon genus Carcharoides is represented in Europe, in the Oligocene
and Miocene, by a form with entire margins, C. catticus (Philippi) (Antunes 1969) but in the
South American Lower Miocene of Patagonia (Ameghino 1906), and the Australian Upper
Oligocene-Upper Miocene (Chapman 1913, 1914, 1917), by a form with serrated margins, C.
totuserratus (Ameghino). As argued by Cappetta (1987) the general morphology of
Carcharoides seems sufficiently distinct to warrant separating it from Lamna, in which genus it
has been usually included. The holotype of Chapman's C. tenuidens is an anterior tooth,
which is here considered conspecific with Ameghino's C. totuserratus, which is based on a
lateral tooth.

A group very widespread in modern seas is Carcharhinus (Garrick 1982, Compagno 1984)
which makes its first appearance in the Middle Eocene of Egypt (Cappetta 1987). Australia's
earliest Carcharhinus comes from the Limestone at Waurn Ponds, it is referred to Carcharhinus
sp. only. The ray, Myliobatis also occurs in the Waurn Ponds Limestone.

FOSSIL CHONDRICHTHYANS - 513

Early-Middle Miocene

In the Otway and Murray Basins during the Early Miocene the sea continued to advance
towards the margins depositing bryozoal limestones and calcarenites of the Morgan Limestone
and Mannum Formation e.g. at Morgan, Mannum, Murray Bridge, Strathalbyn in South
Australia, and the Port Campbell Limestone, including its marginal members, the Bokhara
Limestone and Muddy Creek Marl, e.g. at Port Campbell, Hamilton and Grange Burn, in
Victoria, in shallow (and for the limestones) high energy environments, while in the deeper
parts of the Otway Basin, fined grained sediments, calcareous marl, silt and clay, é.g.
Gellibrand Marl and Fishing Point Marl were being laid down. To the east in the deepening
Torquay Basin the limestones were replaced by calcareous clay and silts of the Puebla
Formation. Around the (then) Dog Rocks Islands, near Geelong the Batesford Limestone was
deposited in less than 30 metres of water in which strong currents prevailed. The Batesford
Limestone conformably passes upwards into the calcareous silt and clay of the markedly
diachronous Fyansford Formation, which was laid down well offshore in water depths of 80-
100 m. The marine transgression continued and further to the east, in the Mornington area the
deposition of the marl and clay of the Fyansford Formation commenced late in the early
Miocene. During the middle Miocene the sea started to retreat again depositing the grey
calcareous clay and silt at Fossil Beach, Mornington, while to the west the uppermost part of
the Fyansford Formation was deposited around Geelong in water of less than 30 m depth.
Further to the west, at Grange Burn, the Muddy Creek Marl overall conformably overlies the
Bokhara Limestone but appears to be integrated with it locally.

To the south, in northern Tasmania, the shallow temperate seas of the early Miocene are
preserved as the richly fossiliferous - again with a predominantly molluscan fauna - beds at
Fossil Bluff, near Table Cape. The deposition of the Freestone Cove Sandstone commenced in
a shallow bay, initially preserving the intertidal zone but deepened to 4-6 m during the marine
transgression. The conformably overlying Fossil Bluff Sandstone contains lithologies varying
from fine siltstones and shales to glauconitic sandstone, calcareous sandstone and calcarenites.
Towards the top of the Fossil Bluff Sandstone deposition was taking place in 10-20 m of
water.

To the west, at the northwestern tip of Tasmania, are preserved the Cape Grim beds which
were deposited in a channel cut into the underlying Tertiary basalt during the early Miocene.
The tuffaceous sandstones and calcarenites were laid down in water of about 20 m depth, as
indicated by the molluscan fauna. However the foraminifera could be interpreted as indicating
water between 20-40 m. Quilty (1972) also suggests that due to the channel in which the
sediments were deposited the winnowing effects of local currents could have influenced the
foraminiferal fauna by removing the lighter species. In this way, too, the smaller taxa of
elasmobranchs could have been removed, as may have been the case at Dog Rocks in Victoria
during the deposition of the Batesford Limestone.

Three teeth have been found in the Miocene Cape Range Group of the Carnarvon Basin in
northwestern Western Australia. The earliest Miocene Mandu Calcarenite is conformably
overlain by the Middle Miocene Talki Limestone, of the East Indian Letter Classification ©,
and f, respectively (Playford et al. 1975, Chaproniere 1981).

Notorynchus primigenius (Ag.) is the sole hexanchoid present in the Miocene of Australia.
It is moderately common in the Batesford Limestone and occurs in the Middle Miocene Muddy
Creek Marl, near Hamilton.

The sawshark Pristiophorus, as P. lanceolatus (Davis 1888) makes its first Australian
Tertiary appearance, again from Batesford and Hamilton.

Heterodontus cainozoicus is moderately widespread in the Early Miocene being found in
Tasmania at Cape Grim, and Fossil Bluff, in both the Freestone Cove and Fossil Bluff
Sandstones, and in Victoria, from 64-66 m in Mallee Bore No. 8, and from the Fyansford

514 - N. KEMP

Formation at Curlewis. Included in H. cainozoicus are a number of nominal species described
and/or recorded by Chapman and other authors e.g. Chapman & Pritchard 1904, Chapman
1918, Chapman & Cudmore (1924) which all fall into what is considered a normal
morphological range for this extinct Port Jackson Shark,

Orectolobus - the extant Wobbegong of Australia and Japan (Compagno 1984) - was until
recently unknown from the fossil record. Pledge's (1985) record of O. gippslandicus (Chapman
& Cudmore 1924) from the Lower Pliocene of South Australia constitutes the first record.
Several incomplete teeth from Batesford, which are here referred to Orectolobus sp., extend the
time range of this genus back to the Early Miocene.

A tooth referred tentatively to Odontaspis sp. comes from the Lower Miocene Cape Grim
beds, in northwest Tasmania. It is the only representative of this genus recognised in the local
Tertiary. The lingual striations of all other odontaspidids make them assignable to Carcharias.

Carcharias taurus is a common species, being found in the Limestone at Batesford, the clay
of the Puebla Formation at Torquay as well as at Fossil Bluff, Tasmania, in the Fossil Bluff
Sandstone. In the Middle Miocene it occurs in the Muddy Creek Marl near Hamilton and in
the blue clays of the Fyansford Formation at Balcombe Bay and Grice's Creek.

The Fyansford Formation at Balcombe Bay has also yielded the youngest - Middle Miocene
- example of C. macrotus, a species which lasted until the Late Eocene in Europe (Cappetta
1987).

The presence of Mitsukurina maslinensis in the Mannum Formation equivalent, at
Strathalbyn is based on three poorly preserved specimens. These may be reworked from the
underlying Eocene (Pledge 1967).

The two species of Jsurus, I. desori and I. planus, which appeared in the Late Oligocene
continue into the Miocene, both being found in the Batesford Limestone at Batesford, Muddy
Creek Marl at Hamilton, the Puebla Formation at Birregurra (/. planus only), Morgan
Limestone at several localites in South Australia and the Fyansford Formation at Balcombe
Bay (J. desori only). The Middle Miocene occurrences are the last of J. planus. It does not
extend past the Miocene of California (S.P. Applegate, pers. comm.) and Japan (Itoigawa et al.
1985).

The uncommon taxon J. benedeni (Le Hon 1871) appears in the middle Miocene, in the
Fyansford Formation at Balcombe Bay. Overseas it occurs in the Miocene, of Japan (Itoigawa
et al. 1985), Europe, e.g. Italy (Menesini 1969) and Belgium (Leriche 1926), Zululand, South
Africa (Davies 1964) and the Pliocene of North America and Italy (Cappetta 1987) and of
Angola (Antunes 1978).

The cosmopolitan species /. hastalis (Ag.), one of the most common sharks of the Tertiary
is found from innumerable localities from southeastern Australia. Interestingly enough the
genus has yet to be recorded from the west of the continent. In Victoria it occurs in the
Batesford Limestone, Bockhara Limestone, the Gellibrand Clay, the Ironstone beds, Keilor, the
Holey Plains Marl Member of the Seaspray Group at Merriman's Creek, Muddy Creek Marl,
near Hamilton, Fyansford Formation, Balcombe Bay; in South Australia, in the Mannum
Formation at Mannum, the Morgan Limestone at Morgan and the Mt Gambier Limestone at
Mt Gambier; in Tasmania in the Freestone Cove Sandstone of Fossil Bluff.

I. retroflexus (Ag.) makes its first appearance in the Early Miocene, in the Batesford and
Morgan Limestones and also in the Freestone Cove Sandstone. It occurs in the Miocene of
Europe e.g. Belgium (Leriche 1926), France (Cappetta 1970).

There are two extant species of Jsurus, J. oxyrinchus Raf. and I. paucus Guitart Manday
(Garrick 1967, Compagno 1984) Teeth referred to I. cf. oxyrinchus and J. cf. paucus are
reported from the Miocene of Japan (Itoigawa et al. 1985) and Portugal (Antunes et al. 1981).
|. oxyrinchus, I. cf. oxyrinchus and I. cf. paucus are recognised for the first time from the
Australian Tertiary, both from the Batesford Limestone at Batesford, and /. cf paucus
additionally from the Puebla Formation at Birregurra, and Muddy Creek Marl at Hamilton.

FOSSIL CHONDRICHTHYANS - 515

Carcharodon angustidens makes its final appearance in the Early Miocene, in the Mt
Gambier Limestone, South Australia, and Freestone Cove Sandstone, Fossil Bluff, Tasmania,
and also in the Cape Range Group, in northwestern Western Australia. Three specimens of
Carcharodon (two broken) - one of which is C. angustidens (and one C. megalodon)- have been
found at this Western Australian locality but it is unclear as to which came from which
horizon: the older Early Miocene Mandu Calcarenite or the conformably overlying Middle
Miocene Tulki Limestone. Overseas it is recorded from the Early Miocene of Belgium and
France (Leriche 1926). The New Zealand records of C. auriculatus (Blainville) (Keyes 1972)
which do extend to the Early Miocene may include C. angustidens, as recognised here.

Keyes (1972) also records another species of Carcharodon, C. megalodon Ag. from the
earliest Oligocene, to the Early Pliocene. In Australia, like practically all other occurrences
outside New Zealand, C. megalodon does not appear until the Early Miocene. Here it occurs,
again, in the Cape Range Group of Western Australia, in a number of localities in southeastern
South Australia, including Mt. Gambier, Point McDonnell, Lake Bonney, and Morgan and
Blanchtown, and from Victoria at Torquay, Batesford and in the lower Middle Miocene
Gippsland Limestone at Orbost, east of Lakes Entrance.

A unique association of some 30-odd large teeth from the Batesford Limestone is referred to
Lamnidae incertae sedis. The teeth are very close to /surus benedeni but the presence of two
teeth, both of different morphology, one of which is definitely, and one tentatively referrable to
a symphysial position, excludes it from this genus. A tooth from the blue clays of the
younger - Middle Miocene - Fyansford Formation at Balcombe Bay is also of this taxon.

Hemipristis serra Ag. is a cosmopolitan species, more common in deposits representing
conditions of warm water (Cappetta 1987). Strangely, its presence in Australia is indicated by
just two teeth (Pledge1967), both from the Lower Miocene Morgan Limestone, near Morgan,
South Australia.

Individual teeth of the upper jaw of species of Carcharhinus can - sometimes - be
differentiated; the lower teeth only rarely, with confidence. For this reason a number of upper
teeth from Batesford, are compared with the extant species C. brachyurus (Gunther 1870).
Carcharhinus teeth of the lower jaw, and uppers of a morphotype not immediately comparable
with those of C. brachyurus are referred to Carcharhinus sp.; they are found at Batesford, at
Fossil Bluff, in the Freestone Cove Sandstone, in the Muddy Creek Marl near Hamilton, and
the blue clays of the Fyansford Formation near Mornington.

An extant cosmopolitan species of Tiger shark Galeocerdo aduncus Ag., which occurs from
the Early Oligocene e.g. France (Priem 1906) through to the Early Pliocene e.g. Australia
(Pledge 1985) makes its first appearance in Australia in the Early Miocene, It is found in the
Morgan Limestone of the River Murray cliffs, near Morgan, at Batesford, at a number of early
Miocene localities in western Victoria e.g. Hordern Vale, Fischers Point, near the Gellibrand
River, and the Middle Miocene Muddy Creek Marl near Hamilton.

Teeth very similar to one genus of hammerhead, Sphyrna, and referred to Sphyrna sp. 1 and
Sphyrna sp. occur in the Cape Grim beds in Tasmania and the Batesford Limestone at
Batesford, respectively.

The first chimaeroid in the Tertiary of Australia is represented by the holotype of Ischyodus
mortoni (Chapman & Pritchard 1907), which is also the sole occurrence of the taxon, from the
Fossil Bluff Sandstone at Fossil Bluff, Tasmania.

Late Miocene-Pleistocene

In the Murray Basin the very Late Miocene or Early Pliocene marine transgression was
followed by the deposition of the glauconitic marl - including clayey marl-clayey sand and sand
- of the Bookpurnong Beds, in shallow water conditions. These Beds cover much of
northeastern Victoria and may represent the fossiliferous horizon intersected at 97 m in the
Buckalow Bore, New South Wales (T.A. Darragh, pers. comm.). This bore, drilled in the

516 - N. KEMP

1920's, yielded, in addition to many molluscan specimens, a variety of sharks’ teeth "in a fine
state of preservation" (Kenny 1934: 96), which unfortunately are now lost. The Buckalow
material to hand is very worn and polished.

The Early Pliocene saw the deposition of the Loxton Sands (in South Australia) under
shallow estuarine conditions. The particular horizon from which the sharks’ teeth come may
even represent an inter-tidal aspect. To the west, in St Vincents Basin in the late Pliocene, the
Dry Creek Sands were laid down in a littoral, near-shore environment .

In Western Australia, in the Perth Basin, Pliocene sediments are represented by the richly
fossiliferous Ascot Beds. These calcarenites were deposited in a sub-littoral inner shelf
environment at a time of low terrigenous sediment. The presence in the included phosphate
nodules of the Cretaceous bivalve Inoceramus indicates that the phosphatisation occurred in the
Cretaceous, prior to the development of the Ascot Beds (Baxter & Hamilton 1981, G.W.
Kendrick 1981, pers. comm.). As mentioned above, no black, eroded and polished teeth, which
presumably represent the Cretaceous fauna, have been identified.

The Jandakot beds are encountered in shallow bore holes around Perth (Mallett 1982).
These Late Pliocene-Early Pleistocene clays, silty clays and sands were deposited in shallow
water conditions in temperatures similar to those found in southern Western Australia today
(Mallett 1982, pers. comm.).

The Peppermint Grove Limestone of Early to Middle Pleistocene age was deposited in the
sheltered seaward part of an estuary during the marine intercalation of a transgressive phase; the
beds are generally only about one metre thick (G.W. Kendrick, pers. comm). Teeth from
Strongs Cave near the south west of Western Australia are probably Pleistocene in age too
(G.W. Kendrick, pers. comm.).

The greatest variety of taxa comes from the nodule beds at Beaumaris (part of the Black
Rock Sandstone) and Grange Burn (part of the Grange Burn Formation) in Victoria. The
majority of these specimens are usually worn. The small but varied fauna from the Loxton
Sands in South Australia are often fragmented but are not so worn.

The first fossil record of the extant species of Notorynchus, N. cepedianus is from the
Lower Pliocene Jemmys Point Formation in eastern Victoria, based on a single tooth.
Pristiophorus lanceolatus makes its final appearance in the Early Pliocene, at Beaumaris. In
New Zealand it survives until the Late Pliocene (Keyes 1982). Heterodontus cainozoicus,
common at Beaumaris is not known in beds younger than the Grange Burn Formation at
Hamilton. Orectolobus gippslandicus is recorded from the Lower Pliocene Loxton Sand. As
mentioned above this is the first fossil record of the genus. The Recent species of Grey Nurse
shark Carcharias taurus continues its Tertiary record, being found at Beaumaris, Victoria, from
97.5 m in the Buckalow Bore, New South Wales and in the Loxton Sands (as Odontaspis cf.
acutissima and Lamna cf. cattica) South Australia, from the Cameron Inlet Formation, Flinders
Island, Tasmania, from a number of localities from the Jandakot beds and the Peppermint
Grove Limestone in Western Australia. Carcharias sp., which includes fragmented teeth,
usually crowns, or very worn and polished specimens, is also found at most of these localities.
Antunes (1978) also recognizes C. taurus from the fossil record, in the Pliocene of Angola.

One well preserved tooth from the Black Rock Sandstone at Beaumaris is referred
tentatively to Lamna. The two pairs of basal denticles suggest an odontaspidid but the uneven
nature of the crown, the convex labial face, the shape of the root and the placement of the basal
denticles well separated from the crown compares favourably with Lamna. Many teeth from
the Tertiary of Australia, previously referred to species of Lamna e.g. Lamna sp. A (Kemp
1970, 1982), Lamna cf. cattica (Pledge 1985) have, with access to more and better material,
proved to be the laterals of Carcharias taurus, a species which, previously, not surprisingly,
was deficient (numerically) in examples of lateral teeth.

A number of species of Jsurus continues on into the Late Miocene-Early Pliocene in
southeastern Australia viz. /. hastalis, being found in the nodule beds at Beaumaris and Grange

FOSSIL CHONDRICHTHYANS - 517

Burn in Victoria, in the Loxton Sands, South Australia, and the Late Pliocene Cameron Inlet
Formation of Flinders Island; /. desori , J. benedeni and /. retroflexus all occurring at Beaumaris
and the rare serrated species, J. escheri (Ag.) being known from one well preserved specimen
from the Grange Burm Formation near Hamilton. This extends the range of a number of
species: J. desori, from the Middle Miocene, of France (Cappetta 1970) to the late Miocene; /.
escheri, from the Late Miocene e.g. Belgium (Leriche 1926) to the Early Pliocene; J. benedeni,
I, hastalis and I. retroflexus also become extinct in the Pliocene of Europe (Cappetta 1987).
The extant species of J. oxyrinchus is recorded from the Lower Pliocene Cameron Inlet
Formation.

The extant White Pointer Carcharodon carcharias (Linn.) occurs in the Early Pliocene of the
Grange Burn Formation at Forsythes Bank, near Hamilton, in the Jemmys Point Formation at
Lakes Entrance, the Whalers Bluff Formation at Duttons Way near Portland in Victoria. In the
Late Pliocene in Tasmania it occurs in the Cameron Inlet Formation of Flinders Island and in
South Australia is known from the Dry Creek Sands in the Abattoirs Bore near Adelaide.
Specimens of Pleistocene age have been recovered from the West Melbourne swamp, earlier
this century. Overseas C. carcharias makes its first appearance in the late Early Miocene of
Switzerland (Leriche 1927). These teeth, up to 70 mm high, are extraordinarily large for C.
carcharias. Antunes (1978) and Cappetta (1987) question this occurrence. C. carcharias
appears in the latest Miocene (Cappetta 1987) and is common in the Pliocene, e.g. North
America (Leriche 1942), Angola (Antunes 1978), New Zealand (Chapman 1918, Keyes 1972),
Japan (Ishiwara 1921). Several broken crowns from Strongs Cave in Western Australia may
represent the extant C. carcharias.

Carcharodon megalodon continues to be recorded in younger beds, from the Black Rock
Sandstone, at Beaumaris, and the Grange Burn Formation at Forsythes Bank, near Hamilton.
Part of a tooth referred to C. cf. megalodon is recorded from the Lower Pliocene Loxton Sands,
while C. megalodon is definitely known from the Upper Pliocene Cameron Inlet Formation,
Flinders Island. It is believed that C. megalodon survived until the Pleistocene (fide Cappetta
1987).

The scyliorhinid Megascyliorhinus is recognised from the Black Rock Sandstone at
Beaumaris, being recorded originally as an oral tooth of the sawshark Pristiophorus lanceolatus
by Chapman & Cudmore (1924) (Keyes 1984). Cappetta (1987) disputes Keyes's (1984)
specific identification of Megascyliorhinus cooperi Cappetia & Ward. It is the only record of
this genus in Australia.

Well preserved teeth from the Loxton Sands represent the first occurrence in the Australian
fossil record of Galeorhinus. Pledge (1985) compares the teeth to the extant Tope, or School
shark of Australian waters, G. australis (Macleay)

Upper teeth compared with Carcharhinus brachyurus are recorded from Beaumaris and
Grange Burn, and from the Lower Pliocene Loxton Sands. Carcharhinus sp., referring to many
upper teeth and all lower teeth of Carcharhinus occur over a wide area: Cameron Inlet
Formation, Flinders Island, Tasmania; Beaumaris and Grange Burn in Victoria, from the
Buckalow Bore in New South Wales, from the Loxton Sands in South Australia (e.g. the
lower teeth and some upper teeth of Pledge’s (1985) C. cf. brachyurus), and from the Ascot
Formation from Rando's Bore and the Jandakot beds from Paulik's Bore, both in Perth,
Westem Australia.

The cosmopolitan extinct Tiger shark Galeocerdo aduncus continues in Australia, in the
Late Miocene and Early Pliocene at Beaumaris, Forsythes Bank at Grange Burm and in the
Loxton Sands. It is known as late as the Pliocene in Japan (fide Cappetta 1987), A small
number of teeth from the Loxton Sands, referred to cf Sphyrna sp. (Pledge 1985), and the first
Australian record of Sphyrna, appear to be of that genus.

The stingray Dasyatis is known from a single tooth from 49-52 m in Mallee Bore No. 5,
probably from the Upper Miocene Bookpurnong Beds of the Murray Group; it is referred to

518 - N. KEMP

Dasyatis sp. only. A tooth figured by Pledge (1985) as Mustelus sp., could possibly be
Dasyatis; it comes from the Lower Pliocene Loxton Sands.

The eagle ray Myliobatis, as Myliobatis spp. is known from several localities: the Black
Rock Sandstone at Beaumaris, the Grange Burn Formation at McDonalds Bank, Muddy Creek,
near Hamilton, the Loxton Sands, and from the Pleistocene in West Melbourne swamp, at
Fisherman's Bend. A very worn portion of a caudal spine from the Buckalow Bore is referred
to ?Myliobatis.

A number of chimaerids make a brief appearance in the Late Miocene-Early Pliocene.
Edaphadon sweeti Chapman & Pritchard is found at Beaumaris and Grange Bum, E. mirabilis
Chapman & Cudmore, at Beaumaris, and several dental plates, referred to Ishchyodus cf. dolloi
are also found at Beaumaris and Grange Burn. This latter form is referred only tentatively to
Leriche's species as it is only known from the early Tertiary in Europe - middle Palaeocene of
the London Basin (Ward 1980) and Late Palaeocene in Belgium (Leriche 1902).

SUMMARY

Very little has been published on the Cretaceous chondrichthyan fauna of Australia. The
present study figures nearly all of the 18-odd identified taxa, representing a dozen families, held
in Australian museums. The Cretaceous fauna is restricted to the Lower Cretaceous (Albian-
Aptian portion) Rolling Downs Group of Queensland, while in Western Australia it mainly
occurs in the Upper Cretaceous Molecap Greensand (Coniacian-Santonian) and Gingin Chalk
(Santonian-Campanian). All genera of sharks are known in overseas Cretaceous deposits. The
most common Australian genus, Pseudocorax, also is the earliest occurrence in the fossil
record, extending its range from the Late Cretaceous (Turonian) to the Early Cretaceous
(Albian). Several other ranges of genera represent earliest or latest records.

The Tertiary fauna which has been relatively more widely published is figured almost in
toto for the first time. The predominant group of the early Tertiary is the odontaspidid
Carcharias, represented by several species. The extant C. taurus is recognized from the Late
Oligocene-Early Miocene. The lamnid /surus radiates during the middle Tertiary with up to
eight species recorded. Several earliest and latest records of taxa are noted, e.g. Otodus obliquus
Ag. extended from Middle Palaeocene back to Late Palaeocene, Isurus desori (Ag.) from Middle
Miocene to Late Miocene. The Australian fauna is noteably lacking the smaller taxa (teeth 5
mm or less) of Recent waters, and overseas Tertiary deposits. This may be due to
preservational factors rather than impoverished faunas.

ACKNOWLEDGMENTS

1 am indebted to the following people for the loan of their teeth (sharks'), or teeth in their
care, for this study: S. P. Applegate, Instituto de Geologia, Ciudad Universitaria, Mexico
City, Mexico; M. Archer, University of New South Wales; R.F. Baird, Monash University;
T.A. Darragh, Museum of Victoria; D. Evans, Melbourne; T.F. Flannery, Australian
Museum, Sydney; I.L. Gill, Melbourne; G.W. Kendrick, Western Australian Museum; P.A.
Jell, Queensland Museum; R.J.F. Jenkins, University of Adelaide; J.A. Long, Western
Australian Museum; C. McCrae, Beaumaris; B. McDonald, Foster; KJ. McNamara, Western
Australian Museum; R.E. Molnar, Queensland Museum; M. Parker, Australian Portland
Cement Limited, Geelong; G. Pedrina, Melbourne; J. Pelcher, Hamilton; G. Pitt, Melbourne;
N.S. Pledge, South Australian Museum; A. Ritchie, Australian Museum, Sydney; A.C.
Rozefelds, Queensland Museum; P. Simmons, Caulfield; O.P. Singleton, University of
Melbourne; D.J. Taylor, Sydney; E.M. Thompson, Museum of Victoria; S. Wright, Portland
and C.J. Yee, Hamilton. I gratefully acknowledge correspondence and/or discussions in many
aspects of this work with I.W. Keyes, New Zealand Geological Survey; D. Thies, Institut fiir

FOSSIL CHONDRICHTHYANS - 519

Geologie und Palaontologie der Universitat, Hannover, West Germany; G.R. Case, New
Jersey, U.S.A.; A.E. Cockbain, Western Australian Geological Survey; L.J.V. Compagno,
J.L.B. Smith Institute of Ichthyology, Grahamstown, South Africa; T.A. Darragh, Museum of
Victoria; B. Hutchins, Western Australian Museum; P.A. Jell, Queensland Museum; G.W.
Kendrick, Western Australian Museum; P. Last, C.S.I.R.O., Tasmania; K.J. McNamara,
Western Australian Museum; C.W. Mallett, C.S.I.R.O., Victoria; A. Mond, Bureau of
Mineral Resources, Canberra; C. Patterson, British Museum (Natural History), London; J.W.
Pickett, New South Wales Geological Survey; N.S. Pledge, South Australian Museum; S.
Pritchard, New South Wales Department of Water Resources; P.G. Quilty, Antarctic Division,
Hobart; W.-E. Reif, Institut fiir Geologie und Paldontologie der Universitat, Tiibingen, West
Germany; J. Stevens, C.S.I.R.O., Tasmania; E.M. Thompson, Museum of Victoria; S.
Turner, Queensland Museum; J.W. Warren, Monash University, Victoria; and D.J. Ward,
London, U.K. and B.J. Welton, Texas, U.S.A. both of whom introduced me to the practical
aspects of bulk sampling and sieving techniques. I also thank T.H. Rich, Museum of Victoria
for providing financial assistance for field work in Victoria, and E.M. Thompson for her help
once there; P.V. Rich, Monash University for her patience and tacit encouragement; A.A.
Cupit, Tasmanian Museum for her unsolicited help in typing all the references and R.E.
Buttermore, Tasmanian Museum without whose "Mac" this m/s would still be sitting in a
manual typewriter. I especially thank my daughters, Nenagh and Zoé for their patience and
domestic help, and their (proven) resilience which will enable them to recover from their recent
prolonged period of parental deprivation.

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PLATES

Plate 1: A, Isurus oxyrinchus, lower jaw, (NRK) x0.5; B, Jsurus oxyrinchus, cross-section of lower jaw
showing first row of teeth of right side of jaw (SAM), x1; C, Carcharias taurus, one series of teeth from right
side of upper and lower jaws (MUGD F3896), x0.5.

Plate 2. A-B, Callorhinchus milii, dental plates (TM Z2341), x1: A, upper jaw, vomerines and palatines,
not in situ; B, mandibular plate, in situ; C-D, Hydrolagus sp., dental plates (TM Z2342), x1.5; C, upper jaw,
vomerines and palatines, in situ; D, mandibular plates, in situ; E,Galeocerdo aduncus, anterior tooth of upper
jaw, labial view (NMV P26770), x2; F, Carcharias macrotus, first anterior tooth of upper jaw, lingual view
(NMV P27451), x1; G, Isurus retroflexus, lateral tooth (second or third row) of upper jaw, labial view (NMV
evign x1; H, Myliobatus sp., tooth of dental pavement, H1, occlusal (biting) surface, H2, basal surface,
CY16), x1.5.

Plate 3. A,B, Notorynchus sp.: A, lower anterolateral, labial and lingual views (MA 8.1), x4; B, upper
symphyseal, labial and mesial(?) views (MA 8.5), x4; C, Centrophoroides sp., labial and lingual views (MA
4.1), x3; D, E, Protosqualus sp.: D, labial and lingual views (MA 4.2), x3; E, labial and lingual views (MA
14.1), x4; F, G, Pristiophorus cf. tumidens , rostral teeth, Toolebuc Fm., Yambore Creek, Nelia, central
Queensland, Early Cretaceous (Albian): F, dorsal (note attached portion of rostrum), posterior views (QM
F14644), x3; G, dorsal, posterior views (QM F14643), x2; H, Squatina sp., labial, incisal and lingual

views (MA 7), x4; I, Hispidaspis sp., anterior(?), labial, mesial and lingual views (WAM 8248), Gingin

Chalk, Molecap Hill, Western Australia, Late Cretaceous (Santonian-Campanian), x3; J, Hispidaspis(?), labial
and lingual views (WAM 75.12.22), Gingin Chalk, Molecap Hill, Gingin, Western Australia, Late Cretaceous
(Santonian-Campanian), x2; K, L, Anomotodon sp., Aramac, central Queensland, Early Cretaceous (Aptian-

Albian), both x1.5: K, labial, mesial and lingual views (QM F1311); L, labial, mesial and lingual views
(QM F14647); M, Scapanorhynchus cf subulatus, labial view (QM F17374), Allaru Mudstone, or Mackunda
Fm, Dartmouth, central west Queensland, Early Cretaceous (Albian), x1; N, O, Scapanorhynchus sp. N,
labial, mesial and lingual views (WAM 74.5.58), x1.5; O, anterior, labial, mesial and lingual views (WAM
68.5.42.1), x3; P-U, "Lamna" cfarcuat. P, labial view (QM F12174), Allaru Mudstone(?), O'Connell Creek,
near Richmond, north central Queensland, Early Cretaceous (Albian), x3 (holotype of Hybodus incussidens de
Vis 1911); Q, labial view (QM F1900), Allaru Mudstone(?), 80 km south of Richmond, north central
Queensland, Early Cretaceous (Albian), x1.5; R, labial and lingual views (QM F17385), Allarw Mudstone,
Dinga Ding Station, McKinley, central Queensland, Early Cretaceous (Albian);, x1.5 S, labial and lingual
views (QM F17373), Allaru Mudstone, or Mackunda Fm., Dartmouth, central west Queensland, Early
Cretaceous (Albian), x1.5; TT, labial and lingual views (QM F17354), Allaru Mudstone(?), Iona Station,
Hughenden, central Queensland, Early Cretaceous (Albian), x5; U, labial and lingual views (QM F17386),
associated with Pleistocene mound springs, presumably from reworked Lower Cretaceous beds, north end of
Pulchera Waterhole, Mulligan River, central westem Queensland, Early Cretaceous, x1.5; V, “Lamna" venusta
Leriche, labial, mesial and lingual views (WAM 7195), Toolanga Calcilutite, White Cliff, Gantheaume Bay,
Westem Australia, Late Cretaceous (Santonian-Campanian), x1.5; W, Lamniformes, indeterminate, lingual

view (QM F14880), Mackunda Fm., Stamford, Hughenden, central Queensland, Early Cretaceous (Albian),
x1.5. A-E, H, N, and O, Molecap Greensand, Molecap Hill, Gingin, Westem Australia, Late Cretaceous

(Coniacian-Santonian).

Plate 4. A-E, Cretolamna appendiculata: A, labial, mesial and lingual views (QM F12243), Toolebuc Fm.,
Booree Park, near Richmond, north central Queensland, Early Cretaceous (Albian), x1; B, lateral(?), labial
and lingual views (QM F15548), Toolebuc Fm., Elizabeth Springs, south of Boulia, western Queensland,
Early Cretaceous (Albian), x1; C, upper lateral, labial view (QM F14648), Allaru Mudstone, or Mackunda
Fm., Dartmouth, central westerm Queensland, Early Cretaceous (Albian), x1.5; D, lower lateral, labial, mesial

FOSSIL CHONDRICHTHYANS - 525

and lingual views (WAM 63.9.25), Molecap Greensand, Molecap Hill, Gingin, Westem Australia, Late
Cretaceous (Coniacian-Santonian) x1.5; E, lateral(?), labial, mesial and lingual views (QM F17384),
associated with Pleistocene mound springs, presumably from reworked Lower Cretaceous beds, north end of
Pulchera Waterhole, Mulligan River, central western Queensland, Early Cretaceous, x1; F, Cretolamna cf
appendiculata, lingual and mesial views (QM F14646), Huntingdon Station, Queensland (no other data
available), x1; G, Cretoxyrhina mantelli, labial, mesial and lingual views (WAM 60.9.1), Miria Marl,
Toothawarra Creek, Cardabia Station, Western Australia, Late Cretaceous (Maastrichtian), x1; H, Cretoxyrhina
cf mantelli, labial and lingual views (WAM 62.8.29), Gingin Chalk, One Tree Hill Quarry, Gingin, Western
Australia, Late Cretaceous (Santonian-Campanian), x1.5; I, Cretoxyrhina(?), labial, mesial and lingual views
(QM F17370), Aramac, central Queensland, Early Cretaceous (Aptian-Albian), x1; J, K, Paraisurus macrorhiza:
J, labial, mesial and lingual views (QM F10607), Toolebuc Fm., northeast paddock, Warra Station, near
Boulia, western Queensland, Early Cretaceous (Albian), x1; K, labial, mesial and lingual views (QM F17375),
x1; L, Protolamna(?), labial, mesial and lingual views (WAM 75.8.11.1), Molecap Greensand, Molecap Hill,
Gingin, Westem Australia, Late Cretaceous (Coniacian-Santonian), x3; M-W, Pseudecorax australis: N, O,
from Toolebuc Fm., northeast paddock, Warra Station, near Boulia, western Queensland, Early Cretaceous
(Albian); P-W from Toolebuc Fm., Iona, Hughenden, central Queensland, Early Cretaceous (Albian). M,
labial and lingual views (QM F17477), Toolebuc Fm(?), Cambridge Downs, near Richmond, north central
Queensland, x3; N, labial and lingual views (QM F17424), x3; O, labial and lingual views (QM F17394), x3;
P, labial and lingual views (QM F17358), x6; Q, labial and lingual views (QM F17359), x6; R, labial and
lingual views (QM F17356), x6; S, labial and lingual views (QM F17361), x4; T, labial and lingual views
(QM F17357), x6; U, labial and lingual views (QM F17355), x6; V, labial and lingual views (QM F17362),
x4; W, labial and lingual views (QM F17360), x6; X, Microcorax sp., labial, mesial and lingual views (QM

F17391), Toolebuc Fm., northeastem paddock, Warra Station, near Boulia, western Queensland, Early
Cretaceous (Albian), x3.

Plate 5. A-C, Lamniformes, vertebral centra: A, articular surface, profile (WAM 10497), x1; B, articular
surface, profile (QM F14653), Allaru Mudstone, Dinga Ding Station, McKinley, central Queensland, Early
Cretaceous (Albian), x1; C, articular surface, profile (MA 1), x2; D-F, Synechodus sp.: D, labial, mesial and
lingual views (WAM 65.10.8), x1.5; E, labial view (QM F17383), associated with Pleistocene mound
springs, presumably from reworked Lower Cretaceous beds, north end of Pulchera Waterhole, Mulligan River,
central westem Queensland, Early Cretaceous, x2; F, labial and lingual views (WAM 10500), x2; G, batoid(?)
dermal denticle, posterior, lateral and apical views (MA 5), x3; H-J, elasmobranch(?), dermal denticles, all
x10; H, opposite views (QM F17353); I, opposite views (QM F17352); J, opposite views (QM F17351). A,
C, D, F, G are all from the Molecap Greensand, Gingin, Western Australia, Late Cretaceous (Coniacian-
Santonian).

Plate 6. A, Heptranchias perlo: Teeth of 887 mm female from New South Wales, right side of upper and
lower jaws including lower medial tooth; posteriors, except for first upper, are not shown (TM D1247): Al,
labial view, x1.8; A2, mesial view (note, printed from reversed negative), x3; B, Notorynchus cepedianus.
Teeth of right side of upper and lower jaws including upper central and lower medial teeth; posteriors, except
for first upper, are not shown (NMV, from dried jaws, no data): B1, labial view, x1; B2, mesial view, x1.5.

Plate 7. A-D, Heptranchias howelli, lower anterolateral teeth, labial and lingual views, Blanche Point Marl,
Blanche Point and Noarlunga, South Australia, Late Eocenc, x2: A, (SAM P19573); B, (UAGD F17284a); C,
(UAGD F17284b); D, (SAM P19572). E-M, Hexanchus agassizi: E-H, M, Renmark Group (lower part),
Naracoorte No. 5 Bore, 135-145 m, South Australia, Middle Eocene; E, upper anterolateral, labial and lingual
views (SAM P19552b), x2; F, upper anterolateral, labial and lingual views (SAM P19552c), x2; G, upper
anterolateral, labial and lingual vicws (SAM P19552a), x2; H, upper(?) anterolateral, labial and lingual views
(SAM P19552d), x2; I, lower anterolateral, labial and lingual views (SAM P10867), Ettrick Marl, Murray
Group, River Murray cliffs near Wellington, South Australia, Late Oligocene, x1.5; J, lower anterolateral,
labial view (SAM P19643), Banded Marl Member, Blanche Point Marl, Blanche Point, South Australia, Late
Eocene, x1.5; K, lower anterolatcral, labial and lingual views (RJFJ 121b), Blanche Point Marl, Blanche
Point, South Australia, x2; L, lower anterolateral, labial and lingual views (UAGD F17262), Blanche Point
Marl, Blanche Point, South Australia, x2; M, lower anterolateral, labial and lingual views (SAM P19552e),
x2; N, Hexanchus sp. 1, lower anterolateral, labial and lingual views (SAMD ¥V34), x2, Renmark Group
(lower part), Naracoorte No. 5 Bore, 135-145 m, South Australia, Middle Eocene; O, Hexanchus ?sp. 1,
lower anterolateral, labial and lingual views (JAL), Trochocyathus Bed, Rivernook Mbr., Dilwyn Fm.,
Rivernook, Victoria, Late Palaeocenc-Early Eocene, x2; P-U, Notorynchus primigenius, Batesford Limestone,
Batesford, Victoria, Early Miocene: P, upper(?) anterolateral, labial and lingual views (MP 45), x1.5; Q,
lower anterolateral, labial and lingual views (MP 38), x1.5; R, lower anterolateral, labial and lingual views
(MP 37), x1.5; S, lower anterolateral, lingual and lingual views (MP 44), x1.5; T, lower anterolateral, labial
and lingual views (MP 39), x1.5; U, lower anterolateral, labial and lingual views (NMV P27411), x2; VY,

526 -N. KEMP

Notorynchus cepedianus, lower anterolateral, labial and lingual views (TM Z1991), Jemmys Point Fm.,
Jemmy's Point, Victoria, Early Pliocene, x1.5.

Plate 8. A, Heterodontus portusjacksoni, jaws from specimen from Bass Strait, Victoria (no other data
available) (NRK): A1, teeth of upper jaw, x0.9; A2, teeth of lower jaw, x0.9; A3, replacement teeth of four
most distal rows (Gi2-G,1-Go-G_; of Reif (1976)) of Tight side of upper jaw, x1.4; Note strongly pitted

occlusal surfaces and longitudinal ridges; these features are quickly eroded in functional teeth. The ridges on
the teeth of row G_ (row on right side of photo) are medianly placed in the lower (in the photo) tooth and
laterally placed in the succeeding tooth (above it in the photo); B-L, Heterodontus cainozoicus: B, H, Fossil
Bluff Sandstone, Fossil Bluff (Table Cape), Tasmania, Early Miocene; C-E, Black Rock Sandstone, Beaumaris,
Victoria, Late Miocene-Early Pliocene; F, G, Cape Grim, Tasmania, Early Miocene; I-L, Maude Formation,
Moorobool Valley limestone, near Geelong, Victoria, Late Oligocene-Early Miocene; B, occlusal and labial
views (NMV P13386), x2; C, occlusal and labial views (NMV P13387), x1; D, occlusal and labial views
(NMV P13388), x1 (holotype of Cestracion longidens); E, occlusal and labial views (NMV P13380), x1.5;
F, occlusal view (NRK), x1.5; G, occlusal view (NRK), x1.5; H, occlusal and labial views (MV P13389),
x1; I, occlusal and labial views (NMV P5382), x1; J, occlusal and labial views (NMV P5379), x1; K,
occlusal and lingual views (NMV P5378), x1; L, occlusal and labial views (NMV P5380), x1.

Plate 3%. A, B, Pristiophorus lanceolatus, rostral teeth, both from Black Rock Sandstone, Beaumaris,
Victoria, Late Miocene-Early Pliocene, x1.5: A, dorsal and ventral views (NMV P160488a); B, dorsal and
ventral views (NMV P160488b).

Plate 10. A-H, Heterodontus cainozoicus, Black Rock Sandstone, Beaumaris, Victoria, Late Miocene-Early
Pliocene: A, occlusal view (NMV P5369), x1.5 (holotype of Cestracion cainozoicus ); B, occlusal and labial
views (NMV P5372), x1.5; C, occlusal and labial views (NMV P5374), x1.5; D, occlusal and labial views
(NMV P13381), x1.5; E, occlusal view (NMV P13385), x2; F, occlusal view (NMV P13383), x2; G,
occlusal view (NMV P13384), x2; H, occlusal view (NMY P13382), x2. I-K, Orectolobus sp., Batesford
Limestone, Batesford, Victoria, Early Miocene, x3; I, labial, incisal and lingual views (NRK); J, labial,
incisal and lingual views (NRK); K, labial, incisal and lingual views (NRK).

Plate 11. A, B, Carcharias taurus, teeth, not including posteriors, of 2.76 m male taken in 6 m, off
Newcastle, New South Wales, right side of upper and lower jaws (MUGD 3896): A, labial view, x0.75; B,
mesial view (note, printed from reversed negative), x1.

Plate 12. A-O, Carcharias cf acutissima , from Trochocyathus Bed, Rivernook Mbr., Dilwyn Fm.,
Rivernook, Victoria (GSV AW7 locality), Late Palaeocene-Early Eocene, x1.5: A, upper(?) anterior, labial,
mesial and lingual views (NMV P27934): RB, upper lateral, labial, mesial and lingual views (ILG); C, upper
lateral, labial, mesial and lingual views (ILG); D, upper lateral, labial view (ILG); E, upper lateral, labial,
mesial and lingual views (NMV P27933); F, upper lateral, labial, mesial and lingual views (ILG); G, lower
second anterior, labial and lingual views (ILG); H, lower second anterior, labial, mesial and lingual views
(ILG); I, lower lateral, labial, mesial and lingual views (ILG); J, lower lateral, labial, mesial and lingual views
(NMV P27932); K, lower lateral, labial, mesial and lingual views (NMV P27931); L, lower lateral, labial and
lingual views (ILG); M, lower lateral, labial and lingual views (GSV); N, lower lateral, labial, mesial and
lingual views (ILG); O, lower(?) lateral, labial and mesial views (NMV P27929); P, Carcharias sp-, labial
and lingual views (ILG).

Plate 13. A-T, Carcharias macrotus; A,B, H, N, Blue clay, Fyansford Fm., Beaumaris, Victoria, Middle
Miocene; C-G, I, J, L, O-T, Trochocyathus Bed, Rivernook Mbr., Dilwyn Fm., Rivernook, Victoria, Late
Palaeocene-Early Eocene: A, upper first anterior, labial, mesial and lingual views (NMV P27451), x1; B,
upper second anterior, labial, mesial and lingual views (NMV P27450), x1; C, upper (second?) anterior,
labial, mesial and lingual views (ILG), x1.5; D, upper anterior, labial and lingual views (NMV P27922), x1;
E, upper anterior, labial and lingual views (ILG), x1.5; F, upper anterior, labial and lingual views (ILG), x1;
G, upper lateral, labial and lingual views (ILG), x1.5; H, upper lateral, labial, mesial and lingual views (NMV
P27452), x1; I, upper lateral, labial and lingual views (NMV P27925), x1.5; J, upper lateral, labial, mesial
and lingual views (NMV P 27924), x1; K, upper lateral, labial and lingual views (WAM 71.2.39),
Boongerooda Greensand, Cardabia Gr., Toothawarra Creek, Cardabia Station, Western Australia, Late
Palaeocene, x1.5; L, upper lateral, lingual and labial views (ILG), x1.5; M, lower first anterior, labial,
mesial and lingual views (NMV P10968), Waum Ponds Limestone, Jan Juc Fm., Waum Ponds, Victoria, Late
Oligocene-Early Miocene, x1; N, lower first anterior, lingual and labial views (NMV P27449); O, lower first
anterior, labial and lingual views (ILG), x1.5; P, lower second anterior, labial, mesial and lingual views
(ILG), x1.5; Q, lower second(?) anterior, labial and lingual views (NMV P27927), x1.5; R, lower anterior,
labial and lingual views (ILG), x1; S, lower lateral, labial and lingual views (NMV P27933), x1; T, lower

FOSSIL CHONDRICHTHYANS - 527

lateral, labial and lingual views (ILG), x1.5.

Plate 14. A-U, Carcharias taurus, teeth of upper jaw, x1.5; A, N, Q, Peppermint Grove Limestone,
Peppermint Grove, Perth, Western Australia, late Middle Pleistocene; C, R, Muddy Creek Marl Mbr., Pon
Campbell Limestone, Muddy Creek, Hamilton, Victoria, Middle Miocene; D, F, G, I, K-M, O, P, S, U,
Batesford Limestone, Batesford, Victoria, Early Miocene; H, T, formation unknown (?Bookpumong Beds),
Buckalow Bore No. 9730, 97.5 m, 80 km southsouthwest of Broken Hill, New South Wales, Late Miocene(?):
A, first anterior, labial, mesial and lingual views (WAM 60.10.29); B, first (?second) anterior, labial,
mesial and lingual views (NMV P27437), Jan Juc Fm., Torquay, Victoria, Late Oligocene-Early Miocene; C,
second anterior, labial and lingual views (CY 7); D, third anterior, labial, mesial and lingual views (MP 9);
E, third anterior, labial and lingual views (NMV P27401), Black Rock Sandstone, Beaumaris, Victoria, Late
Miocene-Early Pliocene; F, third anterior, labial, mesial and lingual views (MP 1); G, intermediate, labial
and lingual views (MP 28); H, lateral, labial and lingual views (MM F31317); I, lateral, labial, mesial and
lingual views (MP 11); J, lateral view (NMV P27404), Fossil Bluff Sandstone, Fossil Bluff (Table Cape),
Tasmania, Early Miocene; K, lateral, labial, mesial and lingual views (MP 10); L, lateral, labial, mesial and
lingual views (MP12); M, lateral, labial and lingual views (NMV P27433); N, lateral, labial, mesial and
lingual views (WAM 65.9.2); O, lateral, labial mesial and lingual views (MP 24); P, lateral, labial and
lingual views (MP 16); Q, lateral, labial and lingual views (WAM 66.1.12); R, lateral, labial, mesial and
lingual views (JP 36); S, lateral, labial and lingual views (MP 15); T, lateral, labial and lingual views (MM
F31318); U, lateral, labial and lingual views (MP 26).

Plate 15. A-K, Carcharias taurus, teeth of lower jaw: A, symphysial, labial and mesial views (MM
F31320); B, symphysial, labial and lingual views (NRK); C, first anterior, labial view (NMV P27405),
Fossil Bluff Sandstone, Fossil Bluff (Table Cape), Tasmania, Early Miocene; D, first (?second) anterior,
labial, distal and lingual views (AM F37185), Botany, Sydney, New South Wales (no other data available);
E, second anterior, labial and lingual views (NMV P27400), Black Rock Sandstone, Beaumaris, Victoria, Late
Miocene-Early Pliocene; F, third anterior, labial and lingual views (CY 6), Muddy Creek Marl, Clifton Bank,
Muddy Creek, Hamilton, Victoria; G, third anterior, labial and lingual views (MM F31315); H, lateral,
labial, mesial and lingual views (MP 2); I, lateral, labial and lingual views (WAM 76.6.43), Jandakot beds,
Paulik's Bore, 34 m, Semple Road, Jandakot, Perth, Western Australia, Pliocene-Pleistocene; J, lateral,
labial, mesial and lingual views (MP 27); K, lower(?), lateral, labial and lingual views (MP 14); L,
Carcharias cf taurus Rafinesque, upper lateral, lingual and labial views (MM F31316). A, G, L, formation
unknown (?Bookpurnong Beds), Buckalow Bore No. 9730, 97.5 m, 80 km southsouthwest of Broken Hill,
New South Wales, Late Miocene(?); B, H, J, K, Batesford Limestone, Batesford, Victoria, Early Miocene;
all teeth x1.5, except B, x2.

Plate 16. A, ?Odontaspis, upper lateral, labial, mesial and lingual views (NRK), Cape Grim, Tasmania,
Early Miocene, x1.5; B, ?Lamana , upper lateral, labial, mesial and lingual views (NMV P27935), Black Rock
Sandstone, Beaumaris, Victoria, Late Miocene-Early Pliocene, x1.5. C, Lamna nasus, teeth from specimen
from Bass Strait, Victoria (no other data available), right side of upper and lower jaws (MUGD F3897): Cl,
labial view, x0.8, C2, mesial view (note, printed from reversed negative), x1.5; D, E, Otodus obliquus: D,
upper(?) anterior, labial, mesial and lingual views (DJT), Pebble Point Fm., Wangerrip Gr., near Princetown,
Victoria, Middle Palaeocene, x1; E, upper lateral, labial, mesial and lingual views (GSV), Olney No. 1 Bore,
Murray Basin, 262 m, Olney Fm., Renmark Gr., near Murray River, South Australian/Victorian border, Late
Eocene-Early Oligocene, x1.

Plate 17. A, B, /surus oxyrinchus: A, teeth of 1.5 m specimen from Eden, New South Wales, right side of
upper and lower jaws, labial view (MUGD F3898), x1; B, teeth of 3.3 m specimen from Bass Strait,
southeastem Victoria, right side of upper and lower jaws. Note more robust nature of teeth, less sinuous
anteriors and more curved apices, distally, of laterals compared with smaller specimen (above) (MUGD
F3899), x0.6; B1, labial view; B2, mesial view.

Plate 18. A,B, /surus benedeni, Black Rock Sandstone, Beaumaris, Victoria, Late Miocene-Early Pliocene,
xl: A, upper lateral, labial, mesial and lingual views (NMV P42536); B, lower lateral, labial, mesial and
lingual views (NMV P14648); C-K, /surus desori; C, D, E, Waum Ponds Limestone Mbr., Jan Juc Fm.,
Waum Ponds, Victoria, Late Oligocene-Early Miocene, x1; F, G, Muddy Creek Marl, Clifton Bank, Muddy
Creek, Hamilton, Victoria, Middle Miocene, x1; H, I, K, Batesford Limestone, Torquay Gr., Batesford,
Victoria, Early Miocene; C, upper first anterior, labial, mesial and lingual views (NMV P27174); D, upper
second anterior, labial, mesial and lingual views (DE); E, upper second anterior, labial, mesial and lingual
views (NMV P27171); F, upper lateral, labial, mesial and lingual views (CY 12); G, upper lateral, labial,
mesial and lingual views (CY 11); H, upper lateral, labial and lingual views (RB), x1; I, upper lateral, labial
and lingual views (MP 32), x1; J, upper lateral, labial and lingual views (NMV P27203), Blue clay,

528 - N. KEMP

pyees ions say hele Beach, Balcombe Bay, Middle Miocene, x1; K, upper posterior, labial and lingual

Plate 19. A-G, Jsurus desori, all x1; B, D, Jan Juc Fm., Torquay, Victoria, Late Oligocene-Early Miocene;
C, E, F, Waurn Ponds Limestone Mbr., Jan Juc Fm., Waurn Ponds, Victoria, Late Oligocene-Early Miocene:
A, lower first anterior, labial and lingual views (NMV P27198), Black Rock Sandstone, Beaumaris, Victoria,
Late Miocene-Early Pliocene; B, lower second anterior, labial, mesial and lingual views (NMV P27181); C,
lower third anterior, labial, mesial and lingual views (NMV P27183); D, lower lateral, labial, mesial and
lingual views (NMV P27189); E, lower lateral, labial, mesial and lingual views (NMV P27177); F, lower
lateral, labial, lingual and mesial views (GPi); G, lower lateral, labial and lingual views (MP 30), Batesford
Limestone, Batesford, Victoria, Early Miocene; H, Jsurus escheri, upper anterior, labial, mesial and lingual
views (JAL), Grange Bum Fm. (upper unit), Hamilton, Victoria, Early Pliocene, x1.

Plate 20. A-L, Isurus hastalis, Black Rock Sandstone, Beaumaris, Victoria, Late Miocene-Early Pliocene:
A, artificial tooth set representing teeth of left side of upper and lower jaws, labial view (intermediate tooth
not represented); upper jaw (NMV P26812, OPS, NMV P26813-26818, NMV P5430, OPS); lower jaw (OPS,
NMV P26820-26821, OPS, NMV P26822-26824, OPS, NMV P26826-26827); x0.3; B, upper first anterior,
labial and lingual views (NMV P26847), x1; C, upper first anterior, labial, mesial and lingual views (NMV
P26851), x0.5; D, upper second anterior, labial and lingual views (NMV P26852), x1; E, upper second
anterior, labial and lingual views (NMV P27196), x0.5; F, upper first lateral, labial and lingual views (NMV
P26856), x1; G, upper first lateral, labial, mesial and lingual views (NMV P26858), x1; H, upper second
lateral, labial, mesial and lingual views (NMV P26859), x1; I, upper fourth or fifth lateral, labial, mesial and
lingual views (NMV P26867), x1; J, upper posterior, labial and lingual views (NMV P26868), x1; K, upper
posterior, labial and lingual views (NMV P26869), x1; L, upper posterior, labial, mesial and lingual views
(NMV P26870), x1.

Plate 21. A-L, Jsurus hastalis; A-G, I-L, Black Rock Sandstone, Beaumaris, Victoria, Late Miocene-Early
Pliocene: A, lower first anterior, labial and lingual views (NMV P26828), x1; B, lower first anterior, labial,
mesial and lingual views (NMV P7586), x0.5; C, lower second anterior, labial, mesial and lingual views
(NMV P26830), x1; D, lower second anterior, labial, mesial and lingual views (OPS), x0.5; E, lower third
anterior, labial and lingual views (NMV P26832), x1; F, lower third anterior, labial and lingual views (NMV
P26834), x0.5; G, lower lateral, labial and lingual views (NMV P26837), x1; H, lower lateral, labial, mesial
and lingual views (NMV P27197), Grange Bum Fm., Forsythes Bank, Hamilton, Victoria, Late Miocene-Early
Pliocene, x1; I, lower lateral, labial, mesial and lingual views (NMV P26841), x1; J, lower posterior, labial
and lingual views (NMV P26843), x1; K, lower posterior, labial and lingual views (NMV P26844), x2; L,
lower posterior, labial, mesial and lingual views (NMV P26845), x2; M, /surus oxyrinchus, lower lateral,
labial, mesial and lingual views, (NRK), Batesford Limestone, Batesford, Victoria, Early Miocene, x1; N-Q,
Isurus cf oxyrinchus; N, P, Q, from Batesford Limestone, Batesford, Victoria, Early Miocene; N, upper
second anterior, labial and lingual views (NRK), x1.5; O, upper lateral, labial, mesial and lingual views
(TFF), Jan Juc Fm., Jan Juc, Victoria, Late Oligocene-Early Miocene, x1.5; P, lower lateral, labial and
lingual views (MP 33), x1.5; Q, lower lateral, labial and lingual views (CY 44), x1.5.

Plate 22. A-E, Isurus cf. paucus: A, lower first anterior, labial, mesial and lingual views (JP 20), x1; B,
lower first anterior, labial, mesial and lingual views (NRK), x1; C, lower(?) lateral, labial, mesial and lingual
views (JP 21), x1; D, lower lateral, labial and lingual views (MP 31), x1.5; E, upper(?) lateral, labial,
mesial and lingual views (NMV P27188), Puebla Fm., Torquay Gr., Birregurra, Victoria, Early Miocene, x1;
F-L, /surus planus: F, upper anterior, labial, mesial and lingual views (NMV P27206), x1; G, upper
anterior(?), labial, mesial and lingual views (NMV P27168), x1; H, upper lateral, labial, mesial and lingual
(CY 12), x1; TU, upper lateral, labial and lingual views (NMV P27193), x1; J, upper lateral, labial, mesial, and
lingual views (MP 29), x1; K, upper posterior, labial and lingual views (JP 5), x1.5; L, lower first anterior,
labial, mesial and lingual views (JP 6), x1. A, C, F, H, I, K, L, Muddy Creek Marl Mbr., Port Campbell
Limestone, Muddy Creek, Hamilton, Victoria, Middle Miocene; B, D, F, I, J, Batesford Limestone,
Batesford, Victoria, Early Miocene.

Plate 23. A-C, Jsurus planus, all x1: A, lower lateral, labial, mesial and lingual views (NMV P27184),
Puebla Fm., Torquay Gr., Birregurra, Victoria, Early Miocene; B, lower lateral, labial, mesial and lingual
views (NMV P27186), Jan Juc Fm., Torquay, Victoria, Late Oligocene-Early Miocene; C, lower lateral, labial
and lingual views (NRK), Batesford Limestone, Batesford, Victoria, Early Miocene, x1; D-L, /surus
retroflexus, Black Rock Sandstone, Beaumaris, Victoria, Late Miocene-Early Pliocene, x1: D, upper first
anterior, labial, mesial and lingual views (NMV P27207); E, upper first anterior, labial, mesial and lingual
views (NMV P26799); F, upper second anterior, labial, mesial and lingual views (NMV P26793); G, upper
first lateral, labial, mesial and lingual views (NMV P26794); H, upper second or third lateral, labial, mesial
and lingual views (NMV P26790); I, upper second lateral, labial, mesial and lingual views (NMV P 13406);

FOSSIL CHONDRICHTHYANS - 529

J, upper lateral, labial, mesial and lingual views (NMV P26807); K, upper posterior, labial, mesial and
lingual views (NMV P26802); L, upper posterior, labial, mesial and lingual views (NMV P27936).

Plate 24. A-G, Isurus retroflexus, Black Rock Sandstone, Beaumaris, Victoria, Late Miocene-Early Pliocene,
all x1: A, lower first anterior, labial, mesial and lingual views (NMV P26777), B, lower first anterior,
labial, mesial and lingual views (NMV P5507); C, lower second anterior, labial, mesial and lingual views
(NMV P26778); D, lower first or second lateral, labial, mesial and lingual views (NMV P26798); E, lower
second or third lateral, labial, mesial and lingual views (NMV P26805); F, lower third or fourth lateral,
labial, mesial and lingual views (NMV P26791); G, lower posterior, labial, mesial and lingual views (NMV
P26801). H-J, /surus sp.: I, J, Batesford Fm., Batesford, Victona, Early Miocene, x1.5: H, lower lateral,
labial, mesial and lingual views (CY 33), Whalers Bluff Fm., Portland, Victoria, Early Pliocene, x1; I, labial,
mesial and lingual views (MP 35); J, aberrant tooth, labial and lingual views (MP 36).

Plate 25. A,B, Carcharodon carcharias: A, teeth of 1.39 m juvenile from Eden, New South Wales, left
side of upper and lower jaws, labial view. Note basal denticles on nearly all teeth and entire (non-serrated)
mesial margin on first lower anterior tooth (MUGD F3901), x1.2; B, teeth of 3.5 m adult from Port Fairy,
Victoria, left side of upper and lower jaws, labial view. Note lack of basal denticles and more robust nature of

teeth compared with juvenile (above). Mesial branch of root of both first and second lower anteriors slightly
damaged (MUGD F3902), x0.6.

Plate 26. A-J, Carcharodon angustidens, all teeth x0.5; A, D, F-H, Freestone Cove Sandstone, Fossil
Bluff (Table Cape), Tasmania, Early Miocene: A, upper first anterior, labial and lingual views (NMV
P13218); B, upper second anterior, labial, mesial and lingual views (WAM 68.9.1), Cape Range Gr., Cape
Range, Westem Australia, Early-Middle Miocene; CC, upper lateral, labial and lingual views (NMV P27417),
Kawarren Limestone, Clifton Fm., Kawarren, Victoria, Late Oligocene; D, upper posterior, labial and lingual
views (NMV P5467); E, upper lateral, labial, mesial and lingual views (NMV P5465), Jan Juc Fm., Torquay,
Victoria, Late Oligocene; F, lower first anterior, labial, mesial and lingual views (NMV P5385); G, lower
second anterior, labial, mesial and lingual views (NMV P5384); H, lower third anterior, labial, distal and
lingual views (NMV P27413); I, lower lateral, labial and lingual views (NMV P27412); J, lower lateral,
labial, mesial and lingual views, Jan Juc Fm., Torquay, Victoria, Late Oligocene (NMV P5466).

Plate 27. A-M, Carcharodon carcharias; A, B, E, G, I, J, M, Grange Bum Fm., Hamilton, Victoria, Late
Miocene-Early Pliocene; C, K, L, Whaler's Bluff Fm., Dutton's Way, Portland, Victoria, Early Pliocene: A,
upper anterior, labial, mesial and lingual views (NMV P27423), x1; 3B, upper anterior, labial and lingual
views (JP), x1; C, upper anterior, labial and lingual views (SW), x1.5; D, upper lateral, labial and lingual
views (WAM 68.9.129), Cameron Inlet Fm., east coast, Flinders Island, Tasmania, Late Pliocene, x1.5; E,
upper lateral, labial, mesial and lingual views (JP 35), x1; F, upper lateral, labial, mesial and lingual views
(MUGD F3892), Jemmys Point Fm., Jemmys Point, Victoria, Early Pliocene, x1; G, upper lateral, labial and
lingual views (JP), x1; H, upper lateral, labial and lingual views (CY 31), Black's Pit quarry, Byaduk,
Victoria, ?Pliocene, x1; I, lower lateral, labial and lingual views (GPe), xl; J, lower lateral, labial and
lingual views (WAM 79.5.93), x1; K, lower lateral, labial and lingual views (CY 62w), x1; L, lower lateral,
labial and lingual views (CY 62b), x1; M, lower(?) posterior, labial and lingual views (GPe), x1.

Plate 28. A-G, Carcharodon megalodon, all teeth x0.5, except G, x1; A, B, E, F, Grange Bum Fm.,
Hamilton, Victoria, Late Miocene-Early Pliocene: A, upper first anterior, labial and lingual views (NMV
P27421); B, upper lateral, labial view (NMV P 27422); C, upper posterior, labial and lingual views, (NMV
P13150), Black Rock Sandstone, Beaumaris, Victoria, Late Miocene-Early Pliocene; D, upper lateral, labial
views (WAM 68.9.3), Cape Range Gr., Shot Hole Canyon, Cape Range, Western Australia, Early-Middle
Miocene; E, upper second anterior, labial, mesial and lingual views (NMV P27419); F, upper lateral, labial
and lingual views (NMV P5469); G, upper posterior, labial, mesial and lingual views (NMV P27416),
Morgan Limestone, River Murray cliffs, 7 km down from Morgan, South Australia, early Middle Miocene.

Plate 29. A-D, Carcharodon megalodon, Grange Burn Fm., Hamilton, Victoria, Late Miocene-Early
Pliocene, x0.5: A, lower first anterior, labial and lingual views (NMV P27424); B, lower lateral, labial,
mesial and lingual views (NMV P27415); C, lower second anterior, labial and lingual views (NMV P27210),
D, lower lateral, labial, mesial and lingual views (NMV P 5514).

Plate 30. A-C, Carcharodon megalodon: A, lower lateral, labial and lingual views (NMV P27420), Grange
Bum Fm., Hamilton, Victoria, Late Miocene-Early Pliocene, x0.5. Note wrinkled base of distal margin,
which gives appearance of basal denticle; B, lower posterior, labial, mesial and lingual views (NMV
P27418), Gippsland Limestone, near Orbost, Victoria, Early-Middle Miocene, x0.5: C, upper(?) posterior,
labial and lingual views (BMcD), Cameron Inlet Fm, east coast Flinders Island, Late Pliocene, x1;

1

530 - N. KEMP

Carcharodon sp., lateral, labial and lingual views (WAM 68.9.2), Cape Range Gr., Cape Range, Western
Australia, Early-Middle Miocene, x1.

Plate 31. A-L, Carcharoides totuserratus, all x1.5: A, B, Point Addis Limestone Mbr, Jan Juc Fm., Airey's
Inlet, Victoria, Late Oligocene-Early Miocene; C, H, J-L, Waum Ponds Limestone Mbr., Jan Juc Fm., Waum
Ponds, Victoria, Late Oligocene-Early Miocene; D-F, Kawarren Limestone Mbr., Clifton Fm. Kawarren,
Victoria, Late Oligocene-Early Miocene: A, upper anterior, labial and lingual views (NMV P27213); B,
upper anterior, labial, mesial and lingual views (NMV P27212); C, lower anterior, labial, mesial and lingual
views (NMV P12636) (holotype of C. tenuidens); D, lower anterior, labial, mesial and lingual views (NMV
P27214); EE, upper lateral, labial and lingual views (NMV P27213); F, upper lateral, labial, mesial and
lingual views (NMV P13025); G, upper lateral, labial, mesial and lingual views (NMV P27211), Jan Juc Fm.,
Torquay, Victoria, Late Oligocene-Early Miocene; H, upper lateral, labial, mesial and lingual views (NMV
P27399); I, upper lateral, labial view, southem Victoria limestone (no other data available); J, upper lateral,
labial, mesial and lingual views (NMV P13036); K, lower lateral, labial and lingual views (TFF); L, lower
lateral, labial, mesial and lingual views (TFF).

Plate 32. A-O, Lamnidae, incertae sedis; all teeth except H, from a single block of Batesford Limestone,
Batesford, Victoria, Early Miocene; almost certainly representing one individual shark (block NMV P12984);
all x1 except E, x1.5: A, anterior, labial view; B, anterior, B1, labial view; B2, mesio-labial view; C,
anterior(?), distal view; D, lower(?), symphysial, labial, mesial and lingual views; E, upper(?), symphysial,
labial and mesio-incisal views; F, lower(?) lateral, labial, mesial and lingual views; G, upper(?) lateral,
labial, mesial and lingual views; H, upper(?) lateral, labial, mesial and lingual views, Fyansford Fm., Blue
clay, Balcombe Bay, Middle Miocene; I, anterior, lingual view; J, anterior(?), lingual view; K, lower(?),
lateral and labial views; L, lower (?) lateral and labial views; M, upper(?) lateral and lingual views; N,
lateral, lingual view; O, posterior, lingual view.

Plate 33. A, Carcharhinus plumbeus, teeth of 1.68 m specimen taken in 50 m off Eden, New South Wales,
left side of upper and lower jaws, labial view (MUGD F3906), x0.8; B, Carcharhinus brachyurus, teeth of
1.37 m specimen taken in 50 m off Eden, New South Wales, left side of upper and lower jaws, labial view
(MUGD F3904), x1.1. Note the dissimilarity of upper teeth and similarity of lower teeth between the two
species; C-O, Carcharhinus cf brachyurus; teeth of upper jaw from rows nearest symphysis progressing to
those nearest jaw articulation; C, E, N, Batesford Limestone, Batesford, Victoria, Early Miocene, D, F, H-M,
O, Black Rock Sandstone, Beaumaris, Victoria, Late Miocene-Early Pliocene: C, labial and lingual views
(MP 71); D, labial and lingual views (NMV P13396); E, labial and lingual views (MP 63); F, labial, mesial
and lingual views (NMV P13397); G, labial and lingual views (CY 15) Grange Burn Fm., McDonald's Bank,
Muddy Creek, Hamilton, Victoria, Late Miocene-Early Pliocene; H, labial and lingual views (NRK); I, labial
and lingual views (NMV P27444); J, labial and lingual views (NMV P27440); K, lower (?) lateral, labial
views (NRK); L, lower (?) lateral, labial views (NMV P13401); M, lower (?) lateral, labial views (NMV
P27441); N, labial and lingual views (MP 68); O, labial and lingual views (NRK).

Plate 34. A-CC, Carcharhinus sp.; A-Q, upper teeth; R-CC, lower teeth; all x2, except A, K, x1.5; A, K,
from unknown formation (?7Bookpumong Beds), Buckalow Bore No. 9730, 97.5 m, 80 km southsouthwest of
Broken Hill, New South Wales, Late Miocene(?); B, E, Q, R, S, V, X-CC, Black Rock Sandstone, Beaumaris,
Victoria, Late Miocene-Early Pliocene; C, D, F-J, L, N-P, Batesford Limestone, Batesford, Victoria, Early
Miocene; T, U, Jandakot Beds, Paulik's Bore, 34 m, Semple Road, Jandakot, Perth, Western Australia,
Pliocene-Pleistocene: A, labial and lingual views (MM F31323); B, labial and lingual views NMV P27422);
C, labial and lingual views (MP 75); D, labial and lingual views (MP 73); E, labial, mesial and lingual
views (NMV P27443); F, labial and lingual views (MP 74), G, labial and lingual views (MP 72); H, labial
and lingual views (MP 65); I, labial and lingual views (NRK); J, labial and lingual views (MP 64); K, labial
and lingual views (MM F31321); L, labial and lingual views (MP 66); M, labial and lingual views (TFF),
Waum Ponds Limestone Mbr.. Torquay Gr., Waurn Ponds, Victoria, Late Oligocene-Early Miocene; N, labial
and lingual views (MP 67); O, labial and lingual views (MP 70); P, labial and lingual views (MP 62); Q,
labial and lingual views (NMV P27445); R, labial and lingual views (NMV P27446); S, labial, mesial and
lingual views (NMV P27447); T, labial and lingual views (WAM 74.5.55.1); U, labial and lingual views
(WAM 74.5.55.2); V, labial and lingual views (NMV P13393); W, labial and lingual views (WAM
68.9.130), Cameron Inlet Fm,. east coast, Flinders Island, Tasmania, Late Pliocene; X, labial, mesial and
lingual views (NMV P13391); Y, labial and lingual views (NRK); Z, labial and lingual views (NRK); AA,
labial and lingual views (NMV P13392); BB, labial and lingual views (NRK); CC, labial, mesial and lingual
views (NMV P27448).

Plate 35. A-V, Galeocerdo aduncus, all x1.5;

A-D, F, H-J, O, P, T-V, Black Rock Sandstone, Beaumaris,
Victoria, Late Miocene-Early Pliocene; E, G, K-N, R, S, B

atesford Limestone Mbr., Torquay Gr., Batesford,

FOSSIL CHONDRICHTHYANS - 531

Victoria, Early Miocene: A-I, teeth of upper jaw from rows nearest symphysis progressing to those nearest
the jaw articulation; A, labial, mesial and lingual views (OPS 785); B, labial and lingual views (NMV
P26768); C, labial and lingual views (NMV P5480); D, labial, mesial and lingual views (NMV P26767); E,
labial and lingual views (MP 51); F, labial and lingual views (OPS 404); G, labial and lingual views (MP
54); H, labial and lingual views (NMV P26765); I, labial and lingual views (CMcC); J-V, teeth of lower jaw
from rows nearest symphysis progressing to those nearest the jaw articulation; J, labial, mesial and lingual
views (NMV P26763); K, labial and lingual views (MP 46); L, labial and lingual views (MP 53); M, labial
and lingual views (MP 49); N, labial and lingual views (MP 47); O, labial, mesial and lingual views (NMV
P26764); P, labial and lingual views (JP 29), Muddy Creek Marl Mbr., Port Campbell Limestone, Muddy
Creek, Hamilton, Victoria, Middle Miocene; Q, labial, mesial and lingual views (NMV P26770); R, labial
and lingual views (MP 55); S, labial and lingual views (MP 50); T, labial, mesial and lingual views (NMV
P26772); U, labial and lingual views (NMV P26773); V, labial and lingual views (NMV P26774).

Plate 36. Galeocerdo cuvier, teeth of 2.5 m specimen taken in 4 m off Southport, southern Queensland,
right side of upper and lower jaws, labial view (MUGD 3903), x0.5.

Plate 37. A, Sphyrna sp. 1, labial and lingual views (NRK), Cape Grim, Tasmania, Early Miocene, x3; B,
C, Sphyrna sp., Batesford Limestone, Batesford, Victoria, Early Miocene, x2; B, labial and lingual views
(PS); C, labial and lingual views (MP 61). D, E, Carcharhiniformes incertae sedis; D, labial and lingual
views (PS), Batesford Limestone, Batesford, Victoria, Early Miocene, x2; E, labial and lingual views (WAM
72.5.5), Ascot Beds, Lee's Bore, 21-23 m, Queen's Park, Perth, Western Australia, Pliocene, x5.

Plate 38. A, Dasyatis sp., labial and lingual views (NMV P12549), ?7Bookpumong Beds, Murray Gr.,
Mallee Bore No. 5, 49-52 m, Murrayville, northwestern Victoria, Late Miocene, x5; B-E, Myliobatis sp.; B,
D, E, Black Rock Sandstone, Beaumaris, Victoria, Late Miocene-Early Pliocene; B, occlusal and basal views
(NMV P161649a), x1.5; C, occlusal and basal views (CY 16), Grange Bum Fm., McDonald's Bank, Muddy
Creek, Hamilton, Victoria, Late Miocene-Early Pliocene, x1.5; D, occlusal and basal views (NMV
P161649b), x1.5; E, occlusal and basal views (NMV P161649c), x1.5; F, ?Myliobatis, part of caudal spine,
dorsal and ventral views (MM F31322), formation unknown (?7Bookpurnong Beds), Buckalow Bore No. 9370,
97.5 m, 80 km southsouthwest of Broken Hill, New South Wales, Late Miocene(?), x1.5.

Plate 39. A, Edaphodon mirabilis (holotype), right palatine, symphysial marginal and oral views (NMV
P13418); B, Edaphodon cf mirabilis , right palatine, oral view (NMV P161651B); C, Ischyodus mortoni
(holotype), left mandibular, oral and aboral views (NMV P9787), Fossil Bluff Sandstone, Fossil Bluff (Table
Cape), Tasmania, Early Miocene; D, Ischyodus cf dolloi, left palatine (posterior portion), oral, oral marginal
and aboral views (NMV P13416) (originally desczibed as vomerine of Edaphodon sweeti). All x1; A, B, D,
from Black Rock Sandstone, Beaumaris, Victoria, Late Miocene-Early Pliocene.

Plate 40. A-C, Edaphodon sweeti: A, right vomerine, aboral, oral marginal and oral views (NMV
P161651D); B, left palatine, oral, oral marginal and aboral views (NMV P161651A); C, holotype, right
mandibular, oral, oral marginal and aboral views, Grange Burn Fm., Grange Burn, Hamilton, Victoria, Late
Miocene-Early Pliocene (NMV P9111). D, E, Ischyodus cf dolloi: D, left palatine, oral, oral marginal, aboral
and symphysial marginal views (JP 42); E, left palatine, oral, oral marginal and aboral views (JP 43); all x1;
A, B, from Black Rock Sandstone, Beaumaris, Victoria, Late Miocene-Early Pliocene; D, E, from Muddy Creek
Marl, Hamilton, Victoria, Middle Miocene.

PLATE 1

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rrr ye YY
conan ea AGE

Ix—— po. —>|~ lat. ———>|«—. an. —>|S

PLATE 2 FOSSIL CHONDRICHTHYANS - 533

534 - N. KEMP PLATE 3

PLATE 4 FOSSIL CHONDRICHTHYANS - 535

536 - N. KEMP PLATE §

PLATE 6

FOSSIL CHONDRICHTHYANS - 537
ih dt i a dh A ed

meee anal: lt ole ae

538 - N. KEMP PLATE 7

PLATE 8 FOSSIL CHONDRICHTHYANS - 539

540 - N. KEMP PLATE 9

PLATE 10

FOSSIL CHONDRICHTHYANS - 541

542 - N. KEMP

PLATE

12

FOSSIL CHONDRICHTHYANS - 543

PLATE 13

PLATE 14

544 - N. KEMP

PLATE 15 FOSSIL CHONDRICHTHYANS - 545

546 - N. KEMP PLATE 16

PLATE 17 FOSSIL CHONDRICHTHYANS

PLATE 18

548 - N. KEMP

PLATE 19 FOSSIL CHONDRICHTHYANS - 549

B
(ie

=

550 - N. KEMP PLATE 20

PLATE 21

FOSSIL CHONDRICHTHYANS - 551

552 - N. KEMP PLATE 22

PLATE 23 FOSSIL CHONDRICHTHYANS - 553

554 - N. KEMP PLATE 24

PLATE 25 FOSSIL CHONDRICHTHYANS - 555

PLATE 26

556 - N. KEMP

PLATE 27 FOSSIL CHONDRICHTHYANS - 557

PLATE 28

558 - N. KEMP

PLATE 29 FOSSIL CHONDRICHTHYANS - 559

PLATE 30

560 - N. KEMP

PLATE 31

FOSSIL CHONDRICHTHYANS - 561

562 - N. KEMP PLATE 32

PLATE 33 FOSSIL CHONDRICHTHYANS - 563

SYN YOW WW

|

Sy ee ty pear re

564 - N. KEMP PLATE 34

Bu

PLATE 35 FOSSIL CHONDRICHTHYANS - 565

566 - N. KEMP PLATE 36

PLATE 37

PLATE 38

- 567

FOSSIL CHONDRICHTHYAN

PLATE 39

a
sy
&
cy
i
re
ba

PLATE 40

568 - N. KEMP

CHAPTER 16

AUSTRALIAN FOSSIL
AMPHIBIANS

1
Anne Warren

TMfOduUCHON Ms.. Seek ee ts he, eT LO
Lissamphibians ..............ccccsceeeeeeeeeeeeeeene eo QO
Labyrin thodomts.iceicsensstiecicces cetiew oa veeediacee’ 570

Australian Temnospondyl-Bearing Deposits....... 580
DEN ONIAIN ws Sek crete taco ucts ada enls dae sede datos ste 582
|e) 0410210 CR A en ne Ie 583
BSG io) (OR ae 583
JULASSIC S00). 352 Bsiiet osha eh hentia nae leeldalile aetite Wie 586
CHETACEOUSE 38 soko Ta ha Wojc wand od Ustelasde se a diabie ele siste 586

GOnNCIMSIONS coc i ad iaiils docah bees eet ace isle Le cen’ 586

REPETEN CES: Fi oct cain cach cee dean ecleens 4 Pe onG ale ceauassenae'e et 587

[sn nnn ene EEE Uy yISE yn

1 Zoology Department, LaTrobe University, Bundoora, Victoria 3083, Australia.

570 - WARREN

INTRODUCTION

The amphibians (Table 1) are a paraphyletic group of anamniotic tetrapods whose life
history usually includes a larval stage. While the three orders of extant Amphibia (Anura,
Urodela and Apoda) are largely defined on characters of their soft anatomy, the fossil
amphibians must be distinguished by osteological characters. The evidence that fossil
amphibians possessed an anamniotic egg is only implied, but larval stages are occasionally
preserved in the fossil record, and some fossil labyrinthodonts are perennibranchiate.

Aspects of their osteology that distinguish tetrapods from fish include a retroarticular
process on the lower jaw (not well developed in early labyrinthodonts), pentadactyl limb, the
lack of fin rays distal to the digits, loss of the bony connection between the skull and the
pectoral girdle, attachment of the dorsal blade of the ilium to the vertebral column by means of
one or more sacral ribs and the presence of a choana or internal nostril (present also in some
crossopterygian fish). Anamniote tetrapods (i.e. Amphibia) may be distinguished from the
amniotes by the usual possession of an otic notch in the posterior skull margin between the
squamosal and the tabular, the lack of a transverse flange on the pterygoid bone, the presence of
exposed lateral line sensory canals on the bones of the skull, and scales, where present, being
dermal rather than epidermal.

ee eee
es

Table 1. Systematic organisation of the Class Amphibia as it pertains to the fossil record of
Australia.

Class Amphibia

‘Labyrinthodonts'
Order Ichthyostegalia
Order Temnospondyli
Order Anthracosauria - not present in Australia

‘Lepospondyls' - not present in Australia

"Lissamphibians'
Order Anura - Frogs and toads
Order Urodela - Newts and salamanders - not present in Australia
Order Apoda - Blind worms - not present in Australia

oo SSsSsSsSs“_—v_O aon ={" ="

LISSAMPHIBIANS

Australia's few fossil lissamphibians, which are restricted to the Late Tertiary and
Quaternary and are all members of the Order Anura, are discussed in Chap. 17.

LABYRINTHODONTS

The labyrinthodonts can be divided into the Orders Ichthyostegalia (Devonian),
Anthracosauria (Carboniferous & Permian) and Temnospondyli (Table 2), which spanned the
Carboniferous, Permian and Triassic, and extended, in Australia, into the Cretaceous (Table 3).
All of Australia's labyrinthodonts were temnospondyls and most occurred in the Mesozoic.

In the field, labyrinthodont remains may be identified by a combination of characters (Fig.
1). Each vertebra is composed of four separate elements: a neural arch, two pleurocentra,
which may be entirely cartilaginous or, possibly, missing in some later temnospondyls, and an

AUSTRALIAN FOSSIL AMPHIBIANS - 571

intercentrum. The larger teeth are labyrinthine in cross-section, and the dermal bones of the
skull, lower jaw and dermal pectoral girdle are ornamented. The skull is traversed by sensory
canals that are sunken into the bone. These characters are also present in some bony fish and

A B C right

pulp cavity lateral = anterior posterior

ee) i oa

sensory canal

Figure 1. Some characteristic features of Triassic temnospondyls: A, posterior end of a left mandible
showing the postglenoid area (retroarticular process); also shown is ornament traversed by a sensory canal;
B, temnospondyl tooth in transverse section showing labyrinthine pattern; C, vertebrae of an Early Triassic
temnospondyl showing the arrangement of the neural arch (na), intercentrum (ic), and pleurocentra (pc).

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Table 2. Range of labyrinthodont families in time. Australian occurrences are Cross hatched.

572 - WARREN

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AUSTRALIAN FOSSIL AMPHIBIANS - 573

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574 - WARREN

Se Tre
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A, —

Figure 2. Family Brachyopidae: A, Xenobrachyops allos, Arcadia Formation, Early Triassic, Queensland
(after Howie 1972b); B, Blinasaurus townrowi, Knocklofty Sandstone, Early Triassic, Tasmania (after
Cosgriff 1974). This and the following reconstructions by David Keen show genera belonging to the
temnospondyl families that have been recorded from Australia's Early Triassic. For each case mirror imaging
of both halves of the holotype skull has been used to prepare as complete a reconstruction of the skull roof
as possible, and a complete restoration of the bony omament has been made for the right side of each skull.
In some cases extrapolation from other specimens or related species has been used to complete the
reconstructions, especially of the omament. Reference to the papers cited under each specimen will show the
actual extent of their preservation. Scale bar, 50 mm.

are thus primitive for amphibians. However, any one of them, together with the often
preserved postglenoid area of the lower jaw or the presence of well developed limb bones, may
be used to identify remains as labyrinthodont.

The earliest adequately known labyrinthodont is /chthyostega from the Late Devonian
(Famennian) of East Greenland. As all known Devonian and Carboniferous labyrinthodont
localities were in western Europe or North America, the prevailing idea in the '60s was that the
tetrapods must have originated in the Northern Hemisphere. In 1972 Warren and Wakefield
reported undoubted tetrapod trackways from the Late Devonian (Frasnian) Genoa River Beds of
eastern Victoria. Not only was this the earliest tetrapod trackway, but also the oldest record of
tetrapods, and its discovery forced a revision of the postulated Northern Hemisphere origin of
the group (Panchen 1977). The recent discovery of yet another probable tetrapod trackway in
the Grampians Ranges of western Victoria has extended the known record of tetrapods back at
least to the Early Devonian (Warren et al. 1986). It is assumed that both the Genoa River and
the Grampians trackways were made by labyrinthodonts, but it is not possible to determine
their ordinal position, as in neither can the number of digits on the manus be determined. The

Figure 3. Family Capitosauridae: A, Parotosuchus gunganj, Arcadia Formation, Early Triassic, Queensland
(after Warren 1980); B, Parotosuchus aliciae, Arcadia Formation, Early Triassic, Queensland (after Warren &
Hutchinson 1988); 1 - adult, 2 - post-metamorphic juvenile.; C, Paracyclotosaurus davidi Wianamatta Group,
Middle Triassic, New South Wales (after Watson 1958). Scale bar 50 mm (A, C); 10 mm (B1, B2).

AUSTRALIAN FOSSIL AMPHIBIANS - 575

576 - WARREN

forefoot of the ichthyostegids is unknown, but most temnospondyls have four manus digits
while anthracosaurs have five.

The lower jaw of Metaxygnathus denticulus, from the Cloghnan Shale (Frasnian -
Famennian) near Forbes, New South Wales, is the earliest, definitely identified labyrinthodont
body fossil. Described as a possible ichthyostegid by Campbell & Bell (1977), it may be close
to Doragnathus woodi, an Early Carboniferous temnospondyl from Scotland (Smithson 1980).
Clack (1988) has confirmed the probable tetrapod nature of this jaw. Such an affinity
reinforces the possible faunal link between Australia and the Arctic in the Devonian noted
earlier by Ritchie (1975) and Young (1974).

Figure 4. Family Trematosauridae. Erythrobatrachus noonkanbahensis, Blina Shale, Early Triassic, Wester
Australia. Ornament is not shown but centres of ossification are indicated (after Cosgriff & Garbutt 1972).
Scale bar, 50 mm.

Despite these finds in the Devonian, no labyrinthodont remains have been recovered in
Australia's Carboniferous, The rare finds of Carboniferous Amphibia elsewhere are usually
associated with an extensive fauna of acanthodian, palaeoniscid and occasional crossopterygian
fish (e.g. Smithson 1985). Similar fish assemblages are present at various Australian
Carboniferous localities (Long, this volume), so we may yet uncover Carboniferous amphibian
remains associated with them, A single Permian labyrinthodont, a brachyopid temnospondyl,
was found in oil shale within the Newcastle Coal Measures.

It is in the Triassic that Australia's labyrinthodont fauna is well known (Tables 2, 3, 4).
Early Triassic, probably Lystrosaurus Zone, temnospondyl faunas have been found in the Blina
and Kockatea shales of Western Australia, the Knocklofty Formation of Tasmania and the
Arcadia Formation of the Rewan Group in Queensland. The Sydney Basin has produced some
specimens from later in the Early Triassic and the only Middle Triassic fauna yet found in
Australia. No Late Triassic labyrinthodonts are known, although fish are present in the Late
Triassic Leigh Creek coal measures of South Australia (Wade 1953).

Of the eleven temnospondyl families recognised in the Early and Middle Triassic (Boy
1985, Warren & Black 1985, Carroll 1988) eight are present in Australia (Tables 2,4). These
eight are briefly discussed below.

The Brachyopidae is the most diverse labyrinthodont family in Australia, with five genera,
none of which is found elsewhere. Although brachyopids occur in all continents except South
America, no other continent has more than two genera. The Australian brachyopid skulls are
of two types; one with a preponderance of primitive characters, like Xenobrachyops allos
(Howie 1972b), with small orbits and occipital condyles almost level with the quadrate
condyles, and the other exhibiting more derived characters, like the three species of Blinasaurus,
with large orbits and occipital condyles placed well behind the quadrates (Cosgriff 1969, 1973,
1974) (Fig. 2).

Although species of the family Capitosauridae (Fig. 3) are restricted to two genera in
Australia, the species diversity within the genus Parotosuchus is greatest on this continent.

AUSTRALIAN FOSSIL AMPHIBIANS - 577

Figure 5. Family Micropholidae. Lapillopsis naas Arcadia Formation, Early Triassic, Queensland (after
Warren & Hutchinson in press). Scale bar, 10mm.

The three species of Parotosuchus from the Arcadia Formation (Warren 1980, Warren &
Hutchinson 1988) and the Parotosuchus sp. from Tasmania (Cosgriff & de Fauw 1987) are

578 - WARREN

more closely related to Parotosuchus from the Scythian Al (Cosgriff 1984) of South Africa and
Europe than to the Scythian A2 and Middle Triassic forms. However, P. brookvalensis and
Paracyclotosaurus davidi (Watson 1958) have the semiclosed otic notch typical of the later
forms. One of the Arcadia species, Parotosuchus aliciae, includes individuals which were
considered by Warren & Hutchinson (1988) to be recently metamorphosed. Capitosaurs are
absent from South America, but otherwise their distribution is cosmopolitan (Cosgriff 1984).

One well preserved member of the Trematosauridae, Erythrobatrachus noonkanbahensis
(Fig. 4), has been described from the Blina Shale (Cosgriff & Garbutt 1972), while fragments
of two trematosaurs occur in the Rewan Group (Warren 1985b). All three are long snouted
trematosaurs which are often found in near marine or deltaic environments, perhaps accounting
for their almost cosmopolitan distribution (Hammer 1987).

Figure 6. Family Lydekkerinidae. Chomatobatrachus halei Knocklofty Formation, Early Triassic, Tasmania
(after Cosgriff 1974). Scale bar, 50mm.

The remaining five families have a more restricted geographic range.

Members of the superfamily Dissorophoidea are relatively common in the Permo-
Carboniferous of Europe and North America, with a single species, Micropholis stowi (Boy
1985), surviving into the Early Triassic Lystrosaurus Zone of southern Africa, A second
Mesozoic dissorophoid, related to M. stowi and placed in the same family, the Micropholidae,
has recently been found in the Arcadia Formation. This specimen, Lapillopsis nana (Fig 5;
Warren & Hutchinson in press) serves to reinforce the close relationship between faunas of the
Arcadia Formation and the Lystrosaurus zone.

The family Lydekkerinidae contains three genera in South Africa and one in Antarctica, as
well as Chomatobatrachus halei (Fig. 6) from Tasmania (Cosgriff 1974) and one undescribed
species from Queensland.

AUSTRALIAN FOSSIL AMPHIBIANS - 579

SESS
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Y
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ie

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SY

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et
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Figure 7. Family Rhytidosteidae: A, Arcadia myriadens Arcadia Formation, Early Triassic, Queensland (after
Warren & Black 1985); B, Acerastea wadeae, Arcadia Formation, Early Triassic, Queensland (after Warren &
Hutchinson 1987; C, Deltasaurus kimberleyensis, Blina Shale, Early Triassic, Western Australia (after
Cosgriff 1965). Scale bar, 50 mm.

The family Indobrachyopidae was erected by Cosgriff & Zawiskie (1979) for members of the
Rhytidosteidae with rounded skulls. They united these two families in the superfamily
Rhytidosteoidea and suggested that Australia was the centre of radiation for the group. Warren
& Black (1985) could find no derived characters to unite the members of the Indobrachyopidae
and returned them to the family Rhytidosteidae. Australian rhytidosteids whose skulls have a
more rounded outline (formerly Indobrachyopidae) are Derwentia warreni from Tasmania

580 - WARREN

(Cosgriff 1974), Rewana quadricuneata (Howie 1972a), Arcadia myriadens (Warren & Black
1985) and Acerastea wadeae (Warren & Hutchinson 1987) from Queensland (Fig. 7). All
Queensland genera are remarkable for their vertebral columns: the three components of each
vertebra are subdivided into left and right parts, so that there are two neural arches, two
intercentra and two pleurocentra. This contrasts with the usual temnospondyl vertebra where
there is a single neural arch and intercentum. Two species of Deltasaurus, D. kimberleyensis
(Fig. 7) and D. pustulatus, are Australia's triangular skulled rhytidosteids. The former occurs
in great numbers in the Blina Shale and is present in the Knocklofty Formation, while the
latter, which has peculiar pustular ornament, is based on a single specimen from the Kockatea
Shale. Rhytidosteids have been found in South Africa, Antarctica, East Greenland, India and
Madagascar (Cosgriff 1984),

The family Plagiosauridae, a group of labyrinthodonts characterised by wide shallow skulls
with enormous orbits and (usually) pustular ornament was, until recently, known only from
the Middle to Late Triassic of Europe and Russia. Now fragments of a plagiosaur,
Plagiobatrachus australis (Warren 1985a) have been found in the Arcadia Formation, making
this the first member of the family from the Southern Hemisphere, and the oldest known
plagiosaurid. The family Plagiosauridae is often grouped with a second, older family, the
Peltobatrachidae, in the Superfamily Plagiosauroidea. The only member of the
Peltobatrachidae, Peltobatrachus pustulatus was found in the Late Permian of East Africa
(Panchen 1959), so the presence of the superfamily in Gondwana is not unprecedented,

The Family Chigutisauridae, which is united with the Family Brachyopidae in the
Superfamily Brachyopoidea (Warren 1981), is found only in Australia and South America. As
Australia is the only continent with both brachyopids and chigutisaurids and as our brachyopid
diversity is so great, it is probable that this superfamily also radiated from this region of
Gondwana. Keratobrachyops australis (Fig. 8) described by Warren (1981) from the Arcadia
Formation, is Australia's oldest chigutisaurid, and the first occurrence of the family outside
Argentina. It is known from three skulls and at least eight mandibles from a site near Bluff,
west of Rockhampton.

In 1977 Warren reported the discovery of a skeleton in the Early Jurassic Evergreen
Formation of Queensland, the first undisputed evidence of labyrinthodonts in the Jurassic. A
previous discovery in the Queensland Jurassic of a piece of mandible (Austropelor wadleyi) was
deemed insufficient evidence to extend the chronologic range of the labyrinthodonts (Colbert
1967), and was thought to have been reworked. Subsequently, a Middle Jurassic skull from
China (Dong, 1985), Late Jurassic labyrinthodonts from Soviet Middle Asia (Nessov 1988)
and a partial brachyopid jaw from the Early Cretaceous Strzelecki Group of southern Victoria
(Jupp & Warren 1986) have further extended their range.

The Australian Jurassic labyrinthodont, Siderops kehli (Warren & Hutchinson 1983) is a
well preserved specimen, lacking only the distal limb bones, part of the tail, and a few pieces
of skull (Fig. 8). It measured over two metres in total length being rivalled in size only by the
Late Triassic metoposaurs who were also closest to it in time. These late labyrinthodonts
represent an endpoint of a tendency to increase in size with time, seen in most Triassic
temnospondyl families, In Australia, the Brachyopoidea (families Brachyopidae and
Chigutisauridae) ranged from the Late Permian to the Early Cretaceous, an exceptionally long
period for a labyrinthodont superfamily.

AUSTRALIAN TEMNOSPONDYL-BEARING
DEPOSITS

Although Australian sediments in which temnospondy! labyrinthodonts have been found

AUSTRALIAN FOSSIL AMPHIBIANS - 581

were deposited by a wide range of environments, their included faunas were consistently
dominated by fish, or an amphibian - fish community. Only in the Jurassic and Cretaceous do

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= > Oa

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eu

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Figure 8. Family Chigutisauridae: A, Siderops kehli , Evergreen Formation, Early Jurassic,
Queensland (after Warren & Hutchinson 1983); scale bar, 250 mm; B, Keratobrachyops australis,
Arcadia Formation, Early Triassic, Queensland (after Warren 1981); scale bar, 50 mm.

reptiles predominate. The following is a list of these deposits with comments on their
depositional environment and included fauna (Fig. 9).

582 - WARREN

@ BLINA

ARCADIA
e

| | ARCADIA@
iin aT GLENIDAL

EVERGREEN
MARBURG

Brisbane

et

KOCKATEA

NEWCASTLE COAL

MEASURES NARRABEEN

|

~~. CLOGHNAN® ~~@ HAWKESBURY
| = C Sydney

\ Canberras’ wianaMATTA

ad Keo deg

Melbourne \
Yo} GENOA RIVER
BEDS

" STRZELECKI
evoonorrr| “Beau
Hobart

Figure 9. Distribution of labyrinthodont bearing localities in Australia.

DEVONIAN

GRAMPIANS GROUP. The Grampians Group as a whole is considered to be of fluvial
origin (Spencer-Jones 1965) with the sediments bearing the trackways being deposited in a
flooded backwater or overbank area, which also preserved as yet undescribed trace fossils of
invertebrate origin. Acanthodian spines and thelodont scales, perhaps referable to Austrolepis
seddoni occur in the Silverband Formation within the Grampians Group (Turner 1982a).

GENOA RIVER BEDS. The Genoa River trackway was preserved in a fine grained red
sandstone originally deposited under fluvial conditions. Bone beds containing disarticulated
plates of the crossopterygians, Bothriolepis, Remigolepis and Groenlandaspis are found
upstream and downstream of the footprint site (Young 1988).

CLOGHNAN SHALE. The fossiliferous part of the Cloghnan Shale consists of red to
brownish siltstone, shale and fine sandstone and is thought by Campbell & Bell (1977) to
represent overbank deposits. Its fish fauna includes the lungfish Soederberghia and the

placoderms Bothriolepis, Remigolepis and Phyllolepis.

AUSTRALIAN FOSSIL AMPHIBIANS - 583

PERMIAN

NEWCASTLE COAL MEASURES. A single poorly preserved but fairly complete
brachyopid, Bothriceps australis (Welles & Estes 1969) was recovered from lacustrine oil shale
near Airley, New South Wales. The palaeoniscid fish Elonichthyes and Urosthenes also occur
in these coal measures (Long 1982).

TRIASSIC

NARRABEEN GROUP. The three Narrabeen labyrinthodonts are thought to have come from
a fish bed in the Gosford Formation. This uppermost unit of the Narrabeen Series consists of
laminated mudstone and sandy shale suggesting a lacustrine environment. An extensive fish
fauna described by Woodward (1890), Wade (1940) and Ritchie (1981) includes a cestraciontid
shark, the dipnoan Gosfordia truncata, three palaeoniscids, three catopterids, four perleidids, a
single specimen of Belanorhynchus and two cleithrolepids. Recently a second locality about 8
kilometers away has yielded a similar fish fauna, but no amphibians (Ritchie 1987). At least
two reptilian trackways have been found in the Narrabeen Group.

HAWKESBURY SANDSTONE. In a quarry at Brookvale, north of Sydney, one
labyrinthodont consisting of the skull roof impression of a capitosaur, Parotosuchus
brookvalensis, was found associated with a collection of exquisitely preserved fish including a
lungfish, Ceratodus formosus and about twenty genera of actinopterygii, most of which are
palaeoniscids or catopterids, with a few holosteans (Wade 1935). These sediments are again
lacustrine shales occurring as lenses within crossbedded sandstones. Fragmentary plates of
labyrinthodont dermal bone have also been recorded from Hawkesbury Sandstone in the Sydney
area (e.g. Stephens 1886) as have temnospondy] footprints.

WIANAMATTA GROUP. Vertebrate fossils have been found in several quarries in the
Ashfield Shale near the base of the Wianamatta Group. Best known is the St Peters quarry, in
which the virtually complete capitosaur Paracyclotosaurus davidi was found inside a 2.8 metre
ironstone nodule (Watson 1958), Another nodule contained a fragmentary brachyopid.
Elsewhere from within the Ashfield Shale a small brachyopid amphibian Notobrachyops
picketti has been described (Cosgriff 1973), and a fragmentary labyrinthodont was reported in
the brickworks quarry at Bowral, while Pepperell & Grigg (1973) reported a temnospondy!
trackway west of Sydney. Woodward (1908) found two distinct fish bearing rock types at St
Peters. One, which was probably the same ironstone layer as produced P. davidi, contained a
xenacanthid shark similar to a specimen recently found by Ritchie (1987) in the Narrabeen
Group, as well as various actinopterygians and a lungfish, Sagenodus. Other actinopterygians
were found in a black shale. The fish of the Wianamatta Group are largely holostean, in
contrast to the chondrostean dominated faunas of the underlying Narrabeen and Hawkesbury

ARCADIA FORMATION. The Arcadia Formation is characterised by thick sequences of red
mudstone with thin green banding and was laid down by both meandering and anastamosing
stream systems (Jensen 1975). Its fauna contrasts with that of the Sydney Basin in that fish
are rare, and with that of typical Lystrosaurus Zone faunas in that synapsids are rare. Aside
from the eight families of labyrinthodonts, which constitute 90% of the Arcadia fauna
(Thulborn 1986) there are two lungfish (Turner 1982b), Saurichthyes (Turner 1982b) several
small lizard-like reptiles (Kadimakara, a prolacertid and Kudnu, a paliguanid (Bartholomai
1979)), an undescribed procolophonid, a thecodont, Kalisuchus (Thulborn 1979) and two
fragments of a dicynodont (Thulbom 1983).

584 - WARREN

Figure 10. Scene by David Keen depicting life beside a quiet pool in the Arcadia Formation of Queensland in
earliest Triassic times. The vertebrate fauna was dominated by a variety of temnospondyl amphibians.
1) Parotosuchus gunganj; 2) Parotosuchus aliciae - larger individual; 3) Parotosuchus

AUSTRALIAN FOSSIL AMPHIBIANS - 585

aliciae - small juveniles; 4) Xenobrachyops allos; 5) Arcadia myriadens; 6) Keratobrachyops australis; 7)
Ceratodus sp. cf C. phillipsi; 8) Saurichthys sp.; 9) Kudnu mackinlayi,; 10) ginkgo; 11) cycad; 12)ferns;
13) lycopod (Cylomeia); 14) giant horsetails; 15) charophyte algae; 16) dragonfly; scale bar, 1 m.

586 - WARREN

; GLENIDAL FORMATION. The environment of deposition of the Glenidal Formation is
similar to that of the Arcadia Formation, which it overlies. A trematosaurid amphibian is its
only recorded fossil.

KNOCKLOFTY FORMATION. While much of the Knocklofty Formation is composed of
floodplain deposits most fossil material was found in clay pebble conglomerate representing
stream or river channels in the sandstones of the flood plain. Fish associated with the
labyrinthodonts include the lungfish Ceratodus and the actinopterygians Acrolepis,
Saurichthyes and Cleithrolepis. A single proterosuchian, Tasmaniosaurus, is the only
known reptile (Camp & Banks 1978). Banks et al. (1978) considered that these fossils,
together with fragmentary arthropod and plant material, represent a natural stream-dwelling and
stream-margin community living under cool temperate conditions.

CLUAN FORMATION. A partial skull of Deltasaurus is the only known vertebrate fossil
from the Cluan Formation. It was found in a clay pebble conglomerate lens similar to those
in the Knocklofty Formation.

BLINA SHALE. The Blina Shale is a uniform sequence of buff-coloured sediments probably
deposited under estuarine or deltaic conditions (McKenzie 1961). In these deposits
labyrinthodonts again predominate. The few known fish include the lungfish Ceratodus, a
coclacanthid and the actinopterygian Saurichthyes.

KOCKATEA SHALE. This fine-grained, grey marine shale has produced a single rhytidosteid
amphibian, Deltasaurus pustulatus, associated with 'fish' and marine invertebrates (Dickens et
al, 1961).

JURASSIC

EVERGREEN FORMATION. Most of the sandstones of the Evergreen Formation were laid
down under continental conditions, but the vertebrate-bearing horizon is probably an extension
of the Westgrove Ironstone member. This consists of ferruginous sandstone, concretionary
ironstone, and an argillaceous, oolitic band which may represent a marine incursion. Two
plesiosaurian reptiles were found near the site of the labyrinthodont, Siderops kehli (Thulborn
& Warren 1980).

MARBURG SANDSTONE. The quartz-rich Marburg sandstones were deposited under fluvial
conditions. Apart from the jaw fragment of Austropelor wadleyi, the vertebrate fauna of the
Marburg is unknown.

CRETACEOUS

STRZELECKI GROUP. The rich fauna of the Strzelecki Group and its equivalent Otway
Group represents a stream deposited thanatacoenosis. Apart from the partial labyrinthodont
mandible, a fish, isolated bones of turtles, a pterosaur, at least two theropods and several
ornithopods have been recovered from a range of localities on the coastal rock platform. The
beds are sandstone, mudsione and shale with clay interclast conglomerates occasionally
predominant near the base (Flannery & Rich 1981).

CONCLUSIONS

In summary, Australia's amphibian record contains the earliest and the latest occurrences of
the labyrinthodonts, and shows the most diversity in the Early Triassic. Australia has no
endemic temnospondyl families, and no families with an otherwise worldwide distribution are
missing. No formation has produced as great a diversity of temnospondyls as has the Arcadia

AUSTRALIAN FOSSIL AMPHIBIANS - 587

Formation of Queensland. The area which is now Australia may have been the centre of
radiation within the Gondwanan block of Pangea for two groups of labyrinthodonts, the
thytidosteids and the brachyopoids. Australian Triassic vertebrate faunas are dominated by
temnospondy] labyrinthodonts or by fish, with reptiles a rare occurrence.

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COLBERT, E.H., 1967. A new interpretation of Austropelor, a supposed Jurassic labyrinthodont amphibian from
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COSGRIFF, J.W., 1973. Notobrachyops picketti, a brachyopid from the Ashfield Shale, Wianamatta Group, New
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COSGRIFF, J.W. & GARBUTT, N.K., 1972. Erythrobatrachus noonkanbahensis, a trematosaurid species from
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COSGRIFF, J.W. & ZAWISKIE, J.M., 1979. A new species of the Rhytidosteoidae from the Lystrosaurus zone
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DICKENS, J.M., MCTAVISH, R.A. & BALME, B.E., 1961. The Beagle Ridge Bore. Aust. Oil & Gas Journal 7:
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DONG, Z., 1985. A Middle Jurassic labyrinthodont (Sinobrachyops placenticephalus gen.et sp. nov) from
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FLANNERY, T.F. & RICH, T.H., 1981. Dinosaur digging in Victoria. Aust. Nat. Hist. 20: 195-198.

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HOWIE, A.A., 1972a. On a Queensland labyrinthodont. In Studies in Vertebrate Evolution, K.A. Joysey and T.S.
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HOWIE, A.A., 1972b. A brachyopid labyrinthodont from the Lower Trias of Queensland. Proc. Linn. Soc.
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JENSEN, A.R., 1975. Permo-Triassic stratigraphy and sedimentation in the Bowen Basin, Queensland. Bull.
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NESSOV, L.A., 1988. Late Mesozoic amphibians and lizards of Soviet Middle Asia. Acta Zool. Cracow 31:
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588 - WARREN

DP aed per 1988. Late Mesozoic amphibians and lizards of Soviet Middle Asia. Acta Zool. Cracow 31:

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WARREN, A.A. & HUTCHINSON, M.N., 1988. The morphology and relationships of a new capitosaurid
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AUSTRALIAN FOSSIL AMPHIBIANS - 589

WELLES, S.P. & ESTES, R., 1969. Hadrokkosaurus bradyi from the Upper Moenkopi Formation of Arizona
with a review of the brachyopid labyrinthodonts. Univ. Calif. Publs geol. Sci 84: 1-61.

WOODWARD, A.S., 1890. The fossil fishes of the Hawkesbury Series at Gosford. Mem. Geol. Surv. N.S.W.
(Palaeontology) 4: 1-55.

WOODWARD, A.S. 1908. The fossil fishes of the Hawkesbury Series at St. Peter's. Mem. Geol. Surv.
N.S.W. (Palaeontology) 10: 1-29. i .

YOUNG, G.C., 1974. Stratigraphic occurrence of some placoderm fishes in the Middle and Late Devonian.
Newsl. Stratigr.3: 243-261. ‘ .

YOUNG, G.C., 1988. Palaeontology of the Late Devonian and Early Carboniferous. In Geology of Victoria,
J.G. Douglas & J.A. Ferguson, eds., Victorian Division Geological Society of Australia, Melboume:
191-194.

590 - WARREN

Siderops kehli, a labyrinthodont amphibian from the Jurassic of Queensland. On the bank, from
a clump of horsetails and club-mosses, an individual of this species keeps tracks of a pterosaur.
The individual in the pond pursues some contemporary lungfish. (From Rich & van Tets 1985,
with permission of The Museum of Victoria).

CHAPTER 17

AUSTRALIAN FOSSIL
FROGS

Michael J. Tyler!

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1 Department of Zoology, University of Adelaide, Box 498, G.P.O., Adelaide, South Australia 5001,
Australia.

592 - TYLER

INTRODUCTION

At the latest assessment the world frog (anuran) fauna was estimated to include 3493
recognized species (Frost 1985) and, if the current rate of description is maintained, the actual
total will be at least 4000. Given that twenty years ago it was estimated at 2500 species, the
extent of taxonomic activity in recent years is evident.

Within Australia approximately 200 species are now recognized. This represents an
increase of more than 100% in twenty-five years. The greater part of that increase is a reflection
of improved knowledge of the distribution and diversity of species in northern Australia. For
example, in the same period the number of species known to occur in the Northern Territory
has changed from fourteen to forty-five: an increase of 214% (Tyler & Davies 1986).

The first fossil frog from Australia was reported from Lake Palankarinna in the Lake Eyre
Basin of South Australia (Tyler 1974). Subsequent studies have led to the location of twenty
species from eight sites throughout the continent (Tyler 1989a). The age of the material ranges
from the Late Oligocene through to the Late Pleistocene.

THE SKELETON

In comparison with other orders of animals, the skeleton in the Anura is remarkably
conservative. For example, the number of pre-sacral vertebrae in modern species ranges from
five to nine, and limb bones are commonly so similar that the long bones rarely are of any
value in familial or generic identifications. However, the vast amount of such material found
in Australia has not been subjected to a detailed analysis.

The extremes of osteological diversity that exist in Australian species is attributable in
some way to one of two processes. Firstly, there is failure of some bones to complete the
customary degree of ossification, so producing arrested development and hence paedomorphic
characters (for example, the exposure of a vast fronto-parietal foramen in the roof of the skull,
rather than complete median apposition or fusion of the frontoparietals). Loss of phalanges or
entire digits in many small burrowing species is probably attributable to a similar event. The
second process is basically just the opposite - namely the failure of bone deposition to cease at
its customary stage, so that an elaboration or exostosis of ridges or other sculpture is produced
on the outer surface of the skull. Exostosis is significant because it obscures and hence
reinforces sutures, conceivably reducing the likelihood of post-mortem disarticulation of bones
at those sites.

Accounts of frog osteology are widely scattered in the literature, but almost all species
descriptions of the Australian fauna have appeared within the last fifteen years, principally
through the work of M. Davies and her colleagues.

Essential reading preparatory to any investigation of frog osteology is the superb review by
Trueb (1973), who synthesises the literature available at that time, and provides a bone by bone
analysis of variation observed in anurans. Other publications particularly relevant to the
Australian fauna are those of Lynch (1971), which redefined most of the genera known at that
time, and Davies (1978), which presented an account of intraspecific variation in several
populations of the large tree frog (Litoria infrafrenata). Major contributions on broad aspects of
frog osteology or descriptions of cranial or post-cranial features are listed in Table 1.

AUSTRALIAN FOSSIL FROGS - 593

NOMENCLATURE AND PHYLOGENY

In an article in Rich & Thompson (1982) I described the nomenclature of Australian frogs
as being “in a state of flux" and expressed the hope that nomenclatural stability would be

reached soon.

a

Table 1: Sources of osteological data on Australian frogs.

Family
Hylidae

Leptodactylidae

Microhylidae

Genus
Cyclorana

Litoria

Nyctimystes

Adelotus
Arenophryne
Assa

Crinia

Geocrinia
Heleioporus
Kyarranus
Lechriodus
Limnodynastes
Megistolotis
Myobatrachus
Neobatrachus
Notaden
Philoria
Pseudophryne

Rheobatrachus

Taudactylus
Uperoleia

Cophixalus
Sphenophryne

Contributors

Stephenson (1965); Tyler (1976); Tyler, Davies & Martin
(1981a, 1982)

Briggs (1940); Buchanan (1921); Davies (1978); Davies,
Martin & Watson (1983); Davies & McDonald (1979);
Davies. McDonald & Adams (1986); Gilles & Peberdy
(1917); Tyler (1976). Tyler & Davies (1977, 1978a,
1978b, 1979, 1985); Tyler, Davies & Martin (1977,
1978)

Tyler & Davies (1979)

Lynch (1971)

Davies (1984)

Tyler (1976)

Davies (1984), du Toit (1934), Lynch (1971), Thompson
(1981), Tyler (1976)

Tyler (1976)

Lynch (1971); Tyler (1976); Tyler, Martin & Davies (1979)

Lynch (1971), Tyler (1976)

Lynch (1971), Tyler (1976)

Lynch (1971); Tyler (1978); Tyler, Martin & Davies (1979)

Tyler, Martin & Davies (1979)

Davies (1984), Lynch (1971), Tyler (1976)

Lynch (1971); Tyler, Davies & Walker (1985)

Lynch (1971), Shea & Johnston (1988)

Lynch (1971), Tyler (1976)

Davies (1984), Lynch (1971), Tyler (1976), Tyler & Davies
(1980)

Davies (1983); Davies & Burton (1982); Mahony, Tyler &
Davies (1984)

Czechura (1986), Liem & Hosmer (1973), Tyler (1976)

Davies & Littlejohn (1986); Davies, Mahony & Roberts
(1985); Davies & Martin (1988); Davies & McDonald
(1985). Davies, McDonald & Corben (1986); Davies,
Watson & Miller (1987); Stephenson (1965); Tyler &
Davies (1984); Tyler, Davies & Martin (1981a, 1981b.
1981c); Tyler, Davies & Watson (1987)

Zweifel (1985)
Fry (1912), Zweifel (1985)

594 - TYLER

That description is equally true today and, if anything, the state of nomenclature is even
more uncertain.

_Palaeontologists, like any group of investigators, seek stability of nomenclature to ensure
universality in the taxonomy of organisms. I think that we all recognize that changes are
inevitable with improved understanding of the nature and relationships of taxa.

Insofar as frog nomenclature is concerned, it seems to me that the instability, namely the
use of more than one name for a taxon, is attributable to two factors. Firstly, workers may
disagree on the weighting to attribute to the divergence that exists amongst organisms.

Whereas one person may choose to recognise two units amongst a group of, for example,
ten species, another, with perhaps a more conservative view, may recognise only one. This
sort of argument is healthy; concepts and criteria are challenged. If it results in change, then
the case is likely to be well argued and hence, hopefully, supported by all. Stability can be
expected to follow.

The second area producing instability of nomenclature is the existence of conflict between
associations of taxa derived from morphological and other techniques. As an example, Tyler &
Davies (1978a) produced a provisional subdivision of the Australopapuan hylid frogs in the
genus Litoria. On the basis of osteological, myological and biological features they recognised
37 infrageneric units, which they referred to as "species-groups". Mammalogists or
entomologists might have termed them "sub-genera”. But the term selected is unimportant. It
was one of convenience intended only to demonstrate that some species were more closely
related to one another than they were to other species within the genus.

King (1980) undertook a cytotaxonomic study of the Australian species of Litoria. He
postulated that given the chromosomal stability of many groups, the presence of shared, derived
states was of sufficient note to indicate the existence of genuine relationships. In general,
King's results suggested broadly similar groupings of species to those of Tyler & Davies.
Discordance, however, resulted in some of Tyler & Davies' groups appearing heterogeneous,
whereas others appeared unfounded.

More recently Hutchinson & Maxson (1986, 1987a) have employed micro-complement
fixation (M.C.-F.) techniques involving comparison of serum albumins to examine
phylogenetic relationships amongst some Australopapuan hylids. Their results have produced a
third perspective, agreeing in some areas with the previous contributors and differing in others.

Nomenclature has yet to come to grips with how to handle conclusions reached from totally
different perspectives. Morphology is not sacrosanct, but it is extremely disturbing to learn
that whereas morphological studies demonstrated conclusively that the species M. lignarius was
justifiably referred to a distinct genus (Megistolotis Tyler, Martin & Davies 1979), M.C.-F.
data indicated that it is as closely related to Limnodynastes convexiusculus as the remainder of
congeners are to each other (Hutchinson & Maxson 1987b).

Hillis (1987) has addressed the topic of conflicts derived from morphological and other
approaches but reaches no firm conclusion. It is a topic with which palaeontologists need to
become familiar.

I referred briefly above to the significance that different workers attribute to the degree of
morphological change between taxa. It is abundantly clear that fundamental differences exist in
concepts of genera and families in different classes. This may reflect the degree of lability of
morphological characters. In the case of frogs, problems are created by evolutionary
conservatism.

As an example of conflicting views of interpretation of significance, Tyler & Davies
(1978a) viewed Litoria caerulea and L. splendida, members of a separate species complex, the
L. caerulea group. Savage (1986), citing the diagnosis provided by these authors, considered
these frogs to be members of a distinct genus and so resurrected Pelodryas to accommodate

AUSTRALIAN FOSSIL FROGS - 595

them. It is worth noting that the names of all frog taxa included in Frost (1985) reflect the
most recent treatment; no assessment is made of the merit of the proposals.

Two alternative names are currently in use for Australian families: the Pelodryadidae for the
Hylidae, and the Myobatrachidae for the Leptodactylidae.

SPECIFICITY OF ANURAN BONES

All Australian material currently available comprises disarticulated and spatially separated
fragments. The association of isolated bones with individuals is currently not possible, and
there has been a need to select one bone upon which specific identification can be made in most
cases.

Without doubt, the ilium is the most satisfactory bone for purposes of identification of
extant species and for characterization of extinct forms. Certainly other bones have useful
diagnostic data, but amongst disarticulated bones the ilium is probably the best. The features
predisposing the ilium to this significance are as follows:

Dorsal crest

~~ Acet. fossa

llial shaft

V. acet.exp.
D. prom.
D. protub.
Se

sa

\
D. acet. exp.

pre-acet. zone

Figure 1. Lateral views of left ilia. Above possessing a dorsal crest; below, lacking crest. Key to
abbreviations: Acet fossa, acetabular fossa; D. acet. exp., dorsal acetabular expansion; D. prom., dorsal
prominence; D. protub., dorsal protuberance; pre-acet. zone, pre-acetabular zone; V. acet. exp., ventral
acetabular expansion.

596 - TYLER

(a) In assemblages of fossil bones from various vertebrates, the ilium is unlike any other
and hence is readily imprinted in the formation of a search-image. The elongate shaft and flat,
axe-like head bearing a crescentic portion of the acetabular fossa (Fig. 1) is rapidly
distinguished from long bones, maxillae, etc. Small rodent ribs are the nearest visual
candidates, and they can be distinguished macroscopically in the sorting process by rolling
them: the protrusion of the terminal articulation and curvature of the shaft of a rodent rib
prevent it from lying flat, whereas a frog ilium is always flat, and it cannot be rolled.

(b) Ilial shaft length is related to the total length of the head and the body (unlike limb
bones which can exhibit a 300% variation relative to total length). Hence, the size of the
donor animal can be calculated with reasonable precision.

c) The prominence of the rim surrounding the acetabular fossa and the width of the
acetabular bone indicates the surface area available for the attachment of muscle fibres and so
permits interpretation of body length and muscular development, giving an indication of the
habitus of the entire frog.

FEATURES OF THE ILIUM

Lateral views of two generalised ilia are depicted in Fig. 1. The position of the ilium in
the skeleton is illustrated in Fig. 2.

A comparative study of the ilia of Australian frog genera was undertaken by Tyler in 1976.
Each genus, except the more recently named Arenophryne and Megistolotis, is illustrated there;
a summary illustration from that paper is included in Figs 3-4. The Arenophryne ilium was
described by Davies (1984).

The ilium of frogs can be diagnosed with many features. The ilial shaft is straight or
slightly bowed in profile, and rounded or oval in section. Occasionally it bears a lateral groove
or a dorsal crest. On the dorsal surface near the origin of the shaft a dorsal prominence bearing
a small knob, the dorsal protuberance, is generally present. The dorsal prominence generally
lies on a level with the anterior extremity of the acetabular fossa, and it merges into the dorsal
acetabular expansion, which articulates with the ischium. Inferiorly, the shaft expands gently
or abruptly to form the pre-acetabular zone, which merges insensibly into the ventral acetabular
expansion that in turn articulates with the pubis. The portion of the acetabular fossa upon the
ilium varies in shape from a semi-circle or an ellipse to a roughly triangular form (Fig. 1).

THE TERTIARY FAUNA

It is only within the last two years that any major progress has been made in determining
the composition of the Australian Tertiary anuran fauna.

The first specimen obtained was reported by Tyler (1974) and subsequently described as the
new genus and species Australobatrachus ilius Tyler (1976). It was from the Ngapalkaldi
Fauna in the Etadunna Formation at Lake Palankarinna, South Australia. What set it apart
from extant species was the presence of a groove along the length of the ilium. Additional
specimens were reported from the same site (Tyler 1982, 1986) and also from the Namba
Formation at Lake Yanda to the south (Tyler 1986),

Additional elements in the Ngapalkaldi Fauna are a Litoria sp. cf. L. caerulea, two further
unnamed Liforia species (one reported by Estes 1984) and the extinct Limnodynastes archeri.

The age of the Ngapalkaldi Fauna initially was considered, based upon palynological
evidence, to be mid-Miocene (W.K. Harris, pers. comm.). However, Lindsay (1987) suggests
that it may be older, perhaps Late Oligocene - Miocene.

oN

Ly i

,

AUSTRALIAN FOSSIL FROGS -

598 - TYLER

The Riversleigh Station sites, northwest of Mt Isa, Queensland are extremely rich in frog
material. Since early 1988, 600 anuran ilia have been recovered. Dominant amongst them is
Lechriodus intergerivus Tyler (1989b). The remainder includes species of Litoria,
Limnodynastes, Crinia and Kyarranus. Although the material obtained to date is from strata
considered to be of Miocene and, particularly, mid-Miocene age, there are evident similarities to
the apparently older Ngapakaldi fauna.

As yet no fossils clearly dated as Pliocene have been reported. However, a single ilium of
the burrowing frog, Neobatrachus pictus, has been found at Curramulka, South Australia,
believed to be from sediments lying on the Pliocene/Pleistocene boundary (Tyler 1988).

THE QUATERNARY FAUNA

Most of our knowledge of the Quaternary fauna is derived from two pairs of juxtaposed
sites. The first to be reported were those from Victoria Cave, Naracoorte and Henschke's Cave
in the extreme southeast of South Australia (Tyler 1977). These localities yielded 166 ilia
representing five species common to the vicinity today: Litoria ewingi, Limnodynastes
tasmaniensis, Limnodynastes sp. cf. L. dumerilii, Crinia signifera and Geocrinia sp. cf. G.
laevis (Fig. 3). Uncertainty about the specific identity of the last two species is a consequence
of limited morphological divergence between them and cognate species that occur elsewhere in
southeastern Australia.

The second pair of significant sites are Skull Cave and Devil's Lair at Cape Naturaliste in
the extreme southwest of Western Australia, whose fauna was reported by Tyler (1985). There,
too, the species represented included only taxa living in that area today: Litoria adelaidensis,
Litoria sp. cf. L. cyclorhynchus and L. moorei, Crinia georgiana, Heleioporus/Neobatrachus
spp., Limnodynastes dorsalis and Pseudophryne guentheri (Figs 3, 4). A total of 409 ilia are
included in those samples.

To date all Quaternary fossil frogs can be associated with those that are extant. There is no
doubt a bias in favour of associating fossil forms with extant species, and it seems to be a
reasonable policy to assume close similarity rather than to split on the basis of minor points of
deviation. The material reported here is all considered to be of Late Pleistocene age and, in fact,
no more than 40,000 yBP. At each of the localities the fossils represent species that occur in
the same areas today (Tyler 1978; Tyler, Smith & Johnstone 1984).

DISCUSSION

The knowledge of the fossil fauna remains, like the material, very fragmentary. To date all
fossil species documented are members of the Hylidae and Leptodactylidae, which are considered
Gondwanan elements (Tyler 1979). The extant Ranidae and Microhylidae are represented
largely upon the tropical Cape York Peninsula of Queensland; one microhylid occupies the
northern periphery of Arnhem Land in the Northern Territory. Neither family is known from
the fossil record of Australia, and their origin is assumed to be from the Oriental Region, where
they are diverse and abundant, following the collision of the Australian and Oriental plates in
the Miocene.

It is worth noting that Savage (1973) postulated the existence of microhylids in Australia
for a longer period, with the group progressively retreating northwards associated with assumed
widespread aridity in the middle of the Cainozoic. I am uncertain of the evidence for that aridity

AUSTRALIAN FOSSIL FROGS - 599

but, if the presence of the family in Australia is of such antiquity, representatives could be

Figure 3. Pelvis or isolated ilium of hylid and leptodactylid frogs: A, Litoria caerulea, x2; B, L. lesueuri,
x2; C, L. eucnemis, x5; C, Nyctimystes zweifeli, x2; E, Adelotus brevis, x5; F, Assa darlingtoni, x5; G,
Crinia georgiana, x5; H., Cyclorana novaehollandiae, x2; I, Geocrinia laevis, x5;J, Uperoleia sp. x5; K,
Heleioporus albopunctatus, x2; L, Kyarranus kundagungan , x5; M, Lechroidus fletcheri, x5; N,
Limnodynastes peroni, x5; O, Mixophyes fasciolatus, x2.

600 - TYLER

Figure 4. Pelvis or isolated ilia of leptodactylid, microhylid and ranid frogs: A, Myobatrachus gouldii, x5;
B, Notaden melanoscaphus, x5; C, Neobatrachus centralis, x5; D, Philoria frosti, x5, E, Pseudophryne
bibroni, x5; F, Ranidella parinsignifera, x5; G, Rheobatrachus silus, x5; H, Taudactylus diurnus, x5; I,
Uperoleia sp., x5; J, Cophixalus ornatus, x5; K, Sphenophryne robusta, x12.5; Rana daemeli, x5.

AUSTRALIAN FOSSIL FROGS - 601

expected at sites such as Riversleigh. To date no microhylids have been detected there or
elsewhere.

In northern Australia it appears that the radiation of several frog species is a Holocene
phenomenon. Tyler (1972) examined the significance of Torres Strait as a barrier and a corridor
for dispersal of frogs from and to New Guinea. Where a species exhibited a widespread
distribution in northern Australia, but did not penetrate New Guinea, it was assumed that it did
not exist upon the Cape York Peninsula when fluctuating corridors permitted faunal
interchange.

Neither extinction nor ecological exclusion will explain the absence of Cyclorana species
from New Guinea. In Australia these burrowing frogs extend from the high rainfall periphery
of the northern wet-dry tropics, through to the central areas of unreliable rainfall and periodic,
prolonged drought. One species, Cyclorana australis, ranges across almost the entire northern
half of the continent. Yet this species, and congeners, failed to reach New Guinea.

Tyler, Davies & Watson (1986) expressed surprise at the absence of C. australis upon
Groote Eylandt in the Gulf of Carpentaria, whereas it occurs on the adjacent mainland, separated
by a marine barrier no more than 30 m deep. Given the dominance of the species in the frog
fauna of northern Australia, they concluded that it did not exist upon the adjacent mainland
when the opportunity for dispersal across land existed.

The significance of these observations to an understanding of the fossil record of Australian
frogs is profound, for it suggests that temporal changes of some magnitude in the distribution
of some species have been rapid. The finding of a Quaternary deposit in northern Australia
might permit clarification of the apparent change in at least a portion of the fauna.

Whereas it is customary to view the identification of the frog fauna solely in terms of the
existing native families, more primitive groups may once have occurred here. The most likely
is the Leiopelmatidae, now known from three species of Leiopelma in New Zealand. Worthy
(1987) has described and illustrated the extant species and described three new subfossil species.
I do not wish to try to make a compelling argument for the existence of leiopelmatids in
Australia at any time in the past. It is worth being familiar with their skeletal characteristics,
however, and to note that Worthy's data provide an excellent reference source.

ACKNOWLEDGEMENTS

I am indebted to Neville Pledge, Rod Wells and Mike Archer who collectively initiated and
maintained my forays into palaeontology. Veronica Ward has provided invaluable assistance in
my studies and Margaret Davies kindly provided Figs 1 and 2. Figs 3 and 4 are reproduced with
the permission of the Royal Society of South Australia. Financial support from the Australian
Research Grants Scheme is gratefully acknowledged.

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Western Australia. Rec. W. Aust. Mus. 13: 541-552.

TYLER, M.J., MARTIN, A.A. & DAVIES, M., 1979. Biology and systematics of a new limnodynastine
genus (Anura: Leptodactylidae) from north-western Australia. Aust. J. Zool. 27: 135-150.

604 - TYLER

TYLER, M.J., SMITH, L.A. & JOHNSTONE. R.E., 1984. Frogs of Western Australia. Western Australian

Museum, Perth.
WORTHY, T.H., 1987. Osteology of Leiopelma (Amphibia: Leiopelmatidae) and description of three new

subfossil Leiopelma species. J. R. Soc. N.Z.17: 201-251.
ZWEIFEL. R.G., 1985. Australian frogs of the family Microhylidae. Bull. Am. Mus. nat. hist. 82: 267-388.

CHAPTER 18

FOSSIL REPTILES IN
AUSTRALIA

Ralph E. Molnar!

FOO COM sits cca pabiccves'na¥ fase eevedtdiawadt 606
Mesozoic Record.....cccccceccesessscseeesereees 610
Procolophonians ............0..cecsceeeceeee 613
POSIT ER a rcs e ds eads dyyeasechataeoaacusvels 615
ANALOMY... oc. cececeeececcescesceeceecees 615
Evolution and Taxonomy........... 616
The Australian Record .............05 617
Palacozoogeography............00.005 619
PV APSIOS Lin lvceede ones vEveesTegadessodtes 620
Evolution and Taxonomy........... 620
The Australo-Antarctic Record..... 620
Palacobiology and
Palacozoogcography............. 621
Lepidosauls...........cccccesecseeceeeeeueeees 622
AMALOMY....cecccecsesececceeeceeecee sees 622
Evolution and Taxonomy........... 623
The Australian Record ............... 623
Palaeozoogeography..........0...006. 624
Ichthyoplerygians...........cceccceeeeeeeees 624
AMALOMY... ec eeceeccecceseeeeteeeesenees 624
Evolution and Taxonomy........... 625
Palacobiology..........ccceecsesceeeees 627
The Australian Record ............... 627
Palacozoogeography..........0..00088 628
SQULOPLCLYZIANS ...... cc cceceeceeeeeeeee eens 628
ANALOMNY....0i.cceccscscesscnserecnceaelss 628
Evolution and Taxonomy........... 629
Palacobiology..........ccesceseeeeenees 631
The Australian Record ..............: 631
Palacozoogeography..............eeeee 634
ProlacertifOrms ..........ccceseceeeeeeeeseeee 634
Thecodomls ........scsceceeeeceeeceeseeeeeeees 636

AMNalOMY 22.2... ceceeeeeececeeeeceeeeeees 636
Evolution and Taxonomy ........... 636
Palacobiology ...........:ccceceeeeecees 639

The Australian Record............... 639
Palacozoogcography ...............08. 641
Saurischians ..........ccccccecssceeceeeeceeees 642
AMNALOMY ..... cc ccececeeeceesecescncuenens 642
Evolution and Taxonomy ........... 643
Palacobiology ...........ccccceeeeeeeees 643

The Australian Forms................ 644
Palacozoogeography ..............060. 647
Ornithischians ...........0.0..ccceeeeeeeee ees 648
AMAtOMY ..... cece eecececeececeeaeeseaees 648
Evolution and Taxonomy ........... 648
Palacobiology ..........ccceeeeeeseeeees 649

The Australian Forms................ 650
Palacozoogeography .................. 657
PLCFOSAUTS ..... 2. ec ee cece ee cee cecceceeeuecn ene 657
ANalOMY ..... ce. cceececeececeseeseeaeeeee 657
Evolution and Taxonomy ........... 657
Palacobiology .........csecseeceeeeeeeee 658

The Australian Forms................ 658
Palacozoogeography .................. 658

Tracks and TrackwaYS........ccccsseee 659
SUN OYAR Ye at iret, daadiat edie yeeventiouss 662
Cainozoic Record .........cccececeeeceneseeeceuee 666
Crocodilians .............cccec cence wie be ndeles 671
SquamMatans ..........ccccceceeeeseececeeeseees 673
TEStudinesy sicscdiis cocipysese spinnedtabdestas 675
SPALPLIVI GEV, 5% sa) siisices len io tahelerspeeleeeans 676
FEBTOVETICES, 15.5 sos doesn sverstideswcnvteatalsteides 679
POORER: Vc onde:sn busine nde neaphevdanudsyentl 686
PEALE Ri ckeesehatetiaxstuitvolcieadlsalsteboese. 688

1 Queensland Museum, P.O. BOX 300, South Brisbane, Queensland 4101, Australia.

606 - MOLNAR

INTRODUCTION

From a rigorously cladist viewpoint, reptiles do not exist. Reptiles can only be defined
arbitrarily as those amniotes which are neither birds nor mammals. In a sense, this is not a
real group, for although its members all share derived features not inherited from their ancestors
(i.e. share synapomorphies), these synapomorphies are also shared by birds and mammals.
Reptiles are defined by subtraction, they are the amniotes that are not birds or mammals.
Although this rigorous cladist view is coming to be generally accepted, the more traditional
usage will be retained here.

Unlike amphibians, reptiles lay eggs (amniote eggs) with a suite of membranes - the
amnion, the allantois, the chorion, and the yolk-sac - which allow them to reproduce on land.
They need not lay eggs in water (or very moist environments) as do most amphibians. The
features distinguishing birds from reptiles are not generally agreed. One of the most obvious
characteristics of birds is their feathers. Although it has been suggested that some dinosaurs
may have had feathers, the possession of feathers is coming to be generally accepted as the
criterion of being a bird. For mammals the jaw structure is used. In mammals the jaw usually
consists of only a single element, the dentary (to be precise, other elements may be present as
long as the dentary participates in the jaw articulation). Another feature that distinguishes
reptiles (and birds) from mammals is cranial kinesis. This is not used in classification, but is
important for understanding reptilian biology. It is common among squamatans and dinosaurs.
"Kinesis", of course, refers to movement. Certain elements of the skull (and sometimes of the
jaw) can move on one another. Usually the mobile units are groups of bones, such as those
that make up the snout. In some cases kinesis allows reptiles to swallow larger prey than
otherwise could be accommodated (as pythons swallowing pigs). Another suggested function,
particularly for some dinosaurs where the amount of motion was small, is to reduce stresses
within the skull during feeding. Reptiles that lack kinesis, such as crocodiles, are said to be
akinetic. Further information on kinesis may be found in Hildebrand (1974).

Although now a broad range of features is used to define the various groups of reptiles (e.g.
Benton 1985), for most of this century the features used were based on the structure of the
skull. In order to understand most of the literature on the evolution of reptiles, especially
works that are old but still useful, it is necessary to understand these features. The openings in
the postorbital, or temporal region of the skull, the temporal fenestrae, were predominantly
important (Fig.1A). These fenestrae appear likely to be associated with the jaw musculature,
but the details of this association are unclear.

Temporal fenestrae come in four basic patterns. These fenestrae occur in different
relationship to two cranial bones, the postorbital and the squamosal (Fig.1). Each of the
patterns has been given a name, which designates both the taxonomic group having this
pattern, and the pattern alone.

It is important to remember the distinction between these two uses, for not all animals in a
given group show the fenestration pattern for which the group was named. For example, there
is a euryapsid member of the Diapsida (Fig. 1). In the anapsid condition there are no temporal
fenestrae. There often are, however, paired openings in the occipital face of the skull known as
posttemporal fenestrae (Fig. 2). In the synapsid condition the fenestra lies ventral to the
postorbital-squamosal junction, while in the euryapsid condition it lies dorsal to that junction.
A parapsid condition was also once defined, but was shown to be based on a misunderstanding
of the pattern of cranial bones. Ichthyosaurs, the forms thought to have hada parapsid
condition, in fact have the euryapsid condition. And finally, some forms have two temporal
fenestrae, one dorsal to the postorbital-squamosal junction and one ventral to it. These are
called diapsid. Synapsids and diapsids are now regarded as being taxonomic groups, while
euryapsids comprise three groups (Ichthyopterygia, Sauropterygia and Placodonta) that probably

FOSSIL REPTILES IN AUSTRALIA - 607

did not share a euryapsid common ancestor. The anapsid condition is plesiomorphic, inherited
from the amphibian ancestors of reptiles. Although the terminology directs attention to the
fenestrae, it 1s more helpful to consider the structure of the bones around the fenestrae, the
arches. This 1S to concentrate on the doughnut, not the hole. For example, in diapsids the
upper arch is formed by a postorbital and squamosal; both are triradiate and have only a small
mutual contact (Fig. 1C). In synapsids the lower arch is formed by the squamosal and jugal
(Fig. 1D) (in diapsids they are separated by the quadratojugal).

Figure 1. Patterns of temporal fenestration in reptile skulls. Skulls A-D show the four classical patterns,
while E & F show variations on the diapsid theme. A, anapsid pattem shown by Hylonomus, a paleothyrid;
D, euryapsid pattern shown by Araeoscelis, an eosuchian (probably); B, synapsid pattem shown by
Edaphosaurus, a pelycosaur; C, classical diapsid pattem shown by Youngina, a younginiform; E, squamatan
modification of diapsid condition, shown by Polyglyphanodon, a lizard - the lower bar of the lower temporal
fenestra has been lost; F, archosaur modification of diapsid condition, shown by Euparkeria , thecodont - in
addition to the two temporal fenestrae, a third opening (the anorbital fenestra) has developed on the snout
between the orbit and the naris. Abbreviations include j, jugal; p, postorbital; qj, uadratojugal; s, squamosal.
(Redrawn from Romer 1966, with Youngina from Carroll 1981).

Other features are also used in classification, such as the position of the functional hinge in
the ankle, or the structure of the pelvis. Brief descriptions of these features are given where
relevant. Basically the characters used are eclectic, as expected of a suite of adaptations to many
different environments.

Detailed information on the basic structure and taxonomy of reptiles may be found in
Romer (1956) and Carroll (1987), while the problems of the origin of reptiles have been
extensively and lucidly discussed by Carroll (1969, 1970; see also Carroll & Baird 1972).
Those who whole-heartedly adopt the cladistic viewpoint feel that attempting to identify
ancestors is in general a futile task: it is more practical to search for the closest relatives.
Nonetheless ancestors did exist, and are worth a brief discussion. Until cladistic influence was
widely felt in reptilian palaeoniology, during the last decade, the captorhinomorphs were
considered to be the group from which all other reptiles descended. Captorhinomorphs included
the protorothyrids, captorhinids and some of the “other anapsids" of Table 1. They were
(usually) small anapsid animals that probably looked like lizards. The group probably most
closely resembling the ancestral reptiles are the protorothyrids (Carroll 1987). In the

608 - MOLNAR

evolutionary sense these were successful animals, for not only do seemingly similar forms still
survive, but their descendants adopted life styles (such as flight) quite remote from those of the
first reptiles.

Figure 2. The posttemporal fenestra and its progressive enlargement in turtles. A, an anapsid reptile
(Procolophon), B, the Triassic turtle (Proganochelys); C, a modem turtle (Eremnochelys). The posttemporal
fenestrae (pf) on both sides and the foramen magnum (fm) are shaded. (After Romer 1956 and Gaffney 1979a).

A word on nomenclature is in order here. There is a trend, not yet widely accepted, to limit
the names of higher taxa to the descendents of the common ancestor of all modern species in
that group. Take for example, the birds (because they are more likely to be familiar than most
fossil reptiles). In this usage the well known ‘first bird' Archaeopteryx is not considered a bird.
In fact, the first bird would be one of the Late Cretaceous forms, such as Laornis. This usage
has the advantage of clearly defining what is meant by the term ‘bird’, which is otherwise
elastically expanded to include every new (relevant) fossil discovery. On the other hand, it de-
emphasizes the evolutionary continuity of the birds. We will not follow this usage here, but
anyone reading the references will encounter it.

Table 1 gives a classification of the major groups of reptiles. It is based largely on the
work of Carroll (1987), Benton (1985), Benton & Clark (1988), Gaffney (1980), Mazin (1981)
and Reisz (1980). This chapter discusses each of these groups of reptiles represented in the
Australo-Antarctic Mesozoic (Tables 2, 3). Groups represented by fragmentary or unstudied
specimens are treated only briefly. Each section gives some background information on the

FOSSIL REPTILES IN AUSTRALIA - 609

anatomy, evolution and palaeobiology of the groups to provide a perspective on the Australian
material. Further information on each of these groups can be found in Carroll (1987) and

Romer (1956), which, even taking into account subsequent discoveries, remains a useful
Starting point.

SS

Table 1. Classification of major groups of reptiles (alternative names of the groups are given
parenthetically).
Synapsids
other synapsids (‘pelycosaurs')
therapsids
Procolophonians
Mesosaurs
Other Early Reptiles
Anapsids
captorhinids
eunotosaurs
other anapsids
testudines (chelonomorphs)
Paleothyrids
Diapsids
araeoscelidians
claudiosaurs
mesenosaurs
sauropterygians
nothosaurs
plesiosauroids
pliosauroids
other sauropterygians
placodonts
ichthyopterygians
other diapsids
lepidosauromorphs
‘paliguanids'!
younginiforms (eosuchians)
saurosternids
lepidosaurs
sphenodontians
squamatans
lizards
amphisbaenians
snakes
archosauromorphs
rhynochosaurs
prolacertiforms (protorosaurs)
other archosauromorphs
archosaurs
proterosuchians
other archosaurs
crocodylotarsans
crocodilians
pseudosuchians
gracilisuchians
phytosaurs
ornithosuchians

610 - MOLNAR

omithosuchids

lagosuchians

pterosaurs

dinosaurs
herrerasaurs
saurischians
ornithischians

1 "Paliguanids” is a group of early lepidosaurs, until recently considered lizards, that are incompletely known.
They may not all be related, i.e., they may not be a real group in the cladistic sense. Information on the
“other” groups (e.g., “other anapsids", “other diapsids") may be found in Benton (1985) and Carroll (1987)
(see Appendix I-1).

MESOZOIC RECORD

During the Mesozoic, reptiles were the most diverse and numerous class of terrestrial
tctrapods, and were we able to visit a Mesozoic landscape they would be the most obvious.

The history of the reptiles in Australia may conveniently be divided into three parts
governed by the geographical history of Australia itself (Molnar 1985). During the first phase,
Australia was still geographically connected, via Antarctica, to the Gondwanan continents.
This lasted from Late Palaeozoic until Eocene time. Dispersal of terrestrial tetrapods into (or
out of) Australia was possible, in principle. Because dispersal to Antarctica was possible, the
few known Antarctic tetrapods will be included in this chapter, even with some discussion of
birds and mammals.

The second phase, commencing in the Eocene, is that of complete Australian isolation.
During this phase there were no immigrants into Australia, so all Australian reptiles of this
period were descended from Gondwanan ancestors.

Phase three probably commenced during the Pliocene. At this time Australia moved
sufficiently close to Indonesia for Asian reptiles to disperse into Australia. This phase
continues today.

The fossil record of reptiles in Australia is poor, with many gaps. One of these gaps
encompasses the transition from phase one to phase two. Thus, the fossil reptiles of the
Mesozoic (phase one) are of quite different aspect to those of the Cainozoic (phases two and
three). Even those lineages that were present during the Mesozoic and survived into the
Cainozoic show no continuity. The Mesozoic crocodilians, turtles and lizards seem not to be
ancestors of any known Cainozoic forms. Doubtless this is simply because the Mesozoic
forms are poorly known: the ancestors of the Australian Cainozoic reptiles almost certainly did
live in Australia. However, nothing is gained by discussing together the Mesozoic and
Cainozoic taxa, and a chronological treatment gives a better understanding of the Mesozoic and
Cainozoic times. So, this division will be reflected in the organization of this chapter into two
parts, the first dealing with the Mesozoic and the second with the Cainozoic reptiles.

FOSSIL REPTILES IN AUSTRALIA - 611

|

Table 2. Stratigraphic distribution of Australian Mesozoic Reptiles.
TRIASSIC
Western Australia
Blina Shale
?ichthyosaur
Tasmania
Knocklofty Formation
Tasmaniosaurus triassicus
Queensland
Arcadia Formation
dicynodont
procolophonid
Kadimakara australiensis
Kudnu mackinlayi
Kalisuchus rewanensis
Blackstone Formation (Striped Bacon Seam)
large tetrapod tracks
Blackstone Formation
Plectropterna sp. (tracks)
horizon? (N.E. Qld)
Agrosaurus macgillivrayi
unnamed Rhaetian beds
small theropod (?) tracks

JURASSIC
Queensland

Precipice Sandstone
omithopod tracks

Razorback beds
pliosaur
theropod tracks

Evergreen Formation
plesiosaur

Walloon Group (mostly Bruce & Wass Seams)
Changpeipus bartholomaii (tracks)
large theropod tracks
small theropod tracks
quadruped tracks

Hutton Sandstone
Rhoetosaurus brownei?

CRETACEOUS
Western Australia
Broome Sandstone
Megalosauropus broomensis (tracks)
Molecap Greensand
ichthyosaur
plesiosaur
mosasaur
Northern Territory
Bathurst Island Formation
Platypterygius sp.
elasmosaurid plesiosaur
South Australia
Maree Formation

612 - MOLNAR

plesiosaurs (possibly 3 spp.)
ichthyosaur
Kakuru kujani
2ornithopod
horizon? (Neales River)
?Woolungasaurus sp.
Victoria
Merino Group?
Chelycarapookus arcuatus
Otway Group
turtles
plesiosaur
Leaellynasaura amicagraphica
Atlascopcosaurus loadsi
Fulgurotherium australe
unnamed hypsilophodonts
theropods
pterosaur
Strezlecki Group
turtle
?lizard
Fulgurotherium australe
hypsilophodont
Allosaurus sp.
bird
New South Wales
Coreena Formation
ichthyosaur (AM F9924-5)
Cimoliasaurus maccoyi
? Trinacromeron leucoscopelus
theropod
Griman Creek Formation
turtle
plesiosaur
"Crocodilus" selaslophensis
Fulgurotherium australe
unnamed hypsilophodont
large ornithopod tracks
Muttaburrasaurus sp.
sauropod (AM F15555)
Rapator ornitholestoides
Queensland
Bungil Formation
plesiosaur
Minmi paravertebra
Griman Creek Formation
plesiosaur (QM L380)
?dinosaur (QM F11043)
Winton Formation
tortoise (QM F12413)
small ornithopod tracks (Wintonopus latomorum)
small theropod tracks (Skartopus australis)
large omithopod tracks
large theropod tracks (T'yrannosauropus sp.)
Austrosaurus
Mackunda Formation

FOSSIL REPTILES IN AUSTRALIA - 613

plesiosaur (QM F2637)
pliosaur cf. Trinacromerum (QM F3307)
Muttaburrasaurus langdoni
Toolebuc Formation
Notochelone costata
turtle
Platypterygius australis
Kronosaurus queenslandicus?
crocodile (QM F17070)
dinosaur
Muttaburrasaurus sp.
Minmi sp.
aff. Ornithocheirus sp.
Allaru Mudstone
Cratochelone berneyi?
Platypterygius australis
plesiosaur (QM F2464)
Muttaburrasaurus sp.
Austrosaurus mckillopi
Wallumbilla Formation
ichthyosaur (QM F551)
Kronosaurus sp.
Woolungasaurus glendowerensis
horizon? (Albian - Hughenden)
large sauropod
Normanton Formation
large reptile (QM F11042)

Those records not previously published include specimen number. A leading question mark indicates doubtful
taxonomic assignment; a trailing question mark doubtful stratigraphic assignment. Some stratigraphic
assignments for Mackunda, Allaru and Wallumbilla are based on geological mapping rather than field
observation.

PROCOLOPHONIANS: PRIMITIVE PLANT-EATING REPTILES

Procolophonians are small, superficially lizard-like reptiles (Fig. 3). The skull is anapsid
and quite broad at the back. The posterior margin of the orbit is usually extended posteriorly.
The bones of the limbs and vertebral column suggest that procolophonians moved much like
lizards, using lateral undulation of the vertebral column. They seem to have been the only
reptiles that had two coracoids on each side as adults, rather than one. Procolophonians are
thought to have been plant-eaters (Carroll 1987) and most probably looked like stout, short-
tailed lizards. Procolophonians were widely distributed throughout the Permo-Triassic world,
with a number of taxa in South Africa, and at least one each in Antarctica and South America.
The as yet unstudied Australian procolophonian (Bartholomai 1979) is very similar to those
from South Africa, South America and Antarctica. Thus, it supports a picture of free
migration during the Triassic to and from what is now Australia. .

Procolophonians seem related to paricasaurs and mesosaurs (Gauthier, Kiuge & Rowe
1988). Parieasaurs attained the size of an ox, but retained the sprawling posture typical of
smaller amniotes. These Permian animals lived in Northern Europe, South Africa and Brazil,
and may yet be discovered in Australia. Mesosaurs were small, graceful, marine Permo-
Carboniferous forms with long snouts and long teeth. They have been found only in Brazil and
South Africa, and their geographic distribution was used as an argument for continental drift
long before it was generally accepted. It has been suggested that parieasaurs, mesosaurs and
procolophonians all shared a common ancestor that was not on the line to modern reptiles.

614 - MOLNAR

Thus, some workers do not accept these animals as reptiles, but call them parareptiles
(Gauthier, Kluge & Rowe 1988). This is far from demonstrated, however, as even these
workers admit.

Table 3. Stratigraphic distribution of Antarctic Mesozoic and Cainozoic tetrapods. The data for this table
are derived from Case, Woodburne & Chaney (1987, 1988), Chatterjee & Small (1989), Colbert (1974, 1987),
Colbert & Cosgriff (1974), Colbert & Kitching (1975, 1977, 1981), Cosgriff (1983), Cosgriff & Hammer
(1984), Cosgriff, Hammer & Ryan (1982), Covacevich & Lamperein (1972), DeFauw (1988), Gasparini &
Goni (1983), Gasparini, Olivero, Scasso & Rinaldi (1987), Gasparini & del Valle (1984), Hammer (1988),
Hammer & Cosgriff (1981), Simpson (1946), Tonni (1982), Tonni & Tambussi (1983), Wiman (1905),
Woodbume & Zinsmeister (1982, 1984).

TRIASSIC

Transantarctic Mountains
Fremouw Formation

thecodont (proterosuchian or rauisuchian)
Prolacerta broomi
gomphodont
cynodont
Cynognathus sp.
Ericiolacerta parva
Kannemeyeria sp.
Kingoria sp.
Lystrosaurus curvatus
Lystrosaurus murrayi
Lystrosaurus mccaigi
Myosaurus gracilis
Padaeosaurus parvus
Rhigosaurus glacialis
Thrinaxodon liorhinus
Procolophon trigoniceps
Austrobrachyops jenseni
benthosuchid
capitosaurid
Cryobatrachus kitchingi
rhytidosteid
“very large” temnospondyl

CRETACEOUS

Vicecomodoro Marambio (Seymour) Island
Lopez de Bertodano Formation
elasmosaurid
Turnena seymoursensis
mosasaurid
diving bird
Vega and James Ross Islands
Snow Hill Group
plesiosaurs

James Ross Island
Santa Marta Formation
ankylosaurid

FOSSIL REPTILES IN AUSTRALIA - 615

EOCENE

Vicecomodoro Marambio Island
La Meseta Formation

turtle
phorhorhacoid?
penguin
Basilosaurus sp.
Antarctodolops dailyi
Eurydolops seymourensis

Formation not given
penguin
pseudodontomithid

OLIGOCENE-MIOCENE

King George Island
Fildes Peninsula Group
Antarctichnus fuenzalidae (bird tracks)
several unnamed bird tracks

MIOCENE

Vicecomodoro Marambio Island
Seymour Island beds

Anthropornis grandis
Anthropornis nordenskjoeldi
Archaeospheniscus wimani
Delphinornis larsenii
Icthyopteryx gracilis
Orthopteryx gigas
Palaeeudyptes gunnari
Wimanornis seymourensis

TESTUDINES: TURTLES AND TORTOISES

Turtles and tortoises (chelonians) are so familiar that we overlook just how unusual and
unique they are. Their possession of shells makes them unmistakable in the modern world,
although at the time that they originated, another group of reptiles, among the placodonts, also
possessed similar shells.

Anatomy

The basic features of the chelonian skeleton include a skull lacking lateral and dorsal
temporal fenestrae but with strong development of posttemporal fenestrae (Fig. 2), often deeply
emarginating the temporal region. The parietal foramen is absent, as are marginal teeth. The
middle ear region (otic capsule) is relatively larger than in most other reptiles. In all except the
very earliest forms, the skull is akinetic. The tail is often short, with the 18 presacral vertebrae
usually divided into ten dorsals and eight cervicals. The shell comprises a dorsal carapace and
ventral plastron. There is no sternum, and the pectoral and pelvic elements are rod-like (except
for the ischium). Chelonians are the only vertebrates with the pectoral and pelvic girdles

616 - MOLNAR

situated within the rib cage (because of the shell). The dorsal ribs are often fused to the
carapace. In swimming forms the limbs have become modified into paddles, while in terrestrial

Figure 3. The skeleton of Procolophon trigoniceps from the Triassic of South Africa. The Australian
Procolophon was similar. (From Watson 1914).

forms the proximal limb segments are held horizontally (again because of the shell) and give a
sprawling, lumbering gait.

Evolution and Taxonomy

The ancestry of turtles is unknown. The once presumed ancestor (the South African
Permian Eunotosaurus) is now recognized as quite unrelated (Cox 1969) (see Appendix I-2).
The reported Permian turtle, Archaeochelydium (Bergounioux 1938), was based on a

FOSSIL REPTILES IN AUSTRALIA - 617

pseudofossil (a septarian concretion). The oldest forms are known from the Late Triassic of
Germany, and although unlike modern forms in many characters, they are clearly turtles. In
1982, Triassic turtles were found in Thailand (de Broin 1984). Chelonians seemingly acquired
a shell and lost cranial kinesis during the Triassic. The limb and cervical adaptations then
followed (Gaf fney 1975). The specific adaptations for neck retraction developed even later,
although sull early in the evolution of turtles (possibly even in the Triassic). By the Late
Jurassic (or perhaps earlier) turtles were both common and cosmopolitan and have remained so.
Both freshwater and marine forms (thalassemyids) were present in the Late Jurassic. Modern
marine turtles (cheloniids) appeared in the Cretaceous, and tortoises (testudinids) in the early
Cainozoic.

Turtles as a whole come in two kinds (Gaffney 1975): the proganochelydians (the Triassic
forms) and the casichelydians (all the others). Presumably the latter are derived from the
former, although not from any presently studied taxa. The casichelydians are divided into the
pleurodires and the cryptodires, names reflecting, among other things, different mechanisms for
retracting the neck.

The Australian Record

Australian Mesozoic (and Cainozoic) chelonians have been surveyed by Gaffney (1981 and
this volume). Mesozoic forms all date from the Early Cretaceous and are mostly marine.
Most numerous of the Cretaceous forms is Notochelone costata (Owen 1882b) the fossils of
which are found along a crescent in north and west Queensland from Hughenden in the east
through Julia Creek to southwest beyond Boulia. They derive from the Toolebuc and Allaru
units. N. costata (Fig. 4) is a small form averaging less than a metre in overall length, and is
endemic to Australia. Almost all of the material is yet unstudied, hence its relationships are
uncertain: it was probably much like the modern Green Turtle (Chelonia mydas) in general
appearance and habits.

Like young green turtles (but unlike adults), Notochelone costata had a marked ridge along
the back of the carapace. Several shells and at least three well preserved skulls of N. costata
have been collected. In the western outcrops of the Toolebuc Formation, its fossils are more
abundant than those of other tetrapods.

Second and largest of the marine forms is Cratochelone berneyi found near Hughenden
(Longman 1915), probably in the Toolebuc. Cratochelone is represented by a single specimen,
consisting only of shoulder girdle and forelimb material plus some unidentified elements.
Comparison of the proximal part of the humerus with those of Chelonia mydas suggests that
Cratochelone was about 2.25 m long. The phyletic relationships of this form are unknown,
both because the specimen is incomplete and because there is little detailed knowledge of
shoulder-girdle and forelimb anatomy among chelonians.

A fluviatile turtle, with the delightful name of Chelycarapookus arcuatus (Fig. 5) was
collected in the early part of this century near Casterton, Victoria, probably from the Merino
Group (Warren 1969). Only the internal cast of the shell of a single specimen is known. The
shell is unusual in that an excavation, or fossa is present just anterior to the first rib on each
side, and the neural elements of the carapace become wider posteriorly. A feature related to the
broadened ncurals (the medial bones of the carapace), is that the necks of the posterior ribs are
lengthened. In Chelycarapookus these ribs, because of their elongate necks, fuse to the
carapace further from the vertebral column than in any other chelonians (Fig. 5).

In other fluviatile sediments of southern coastal Victoria (the Otway and Strzelecki groups),
limb bones, vertebrae, a lower jaw and numerous shell fragments of turtles have been found.
These are so primitive that they cannot be assigned to either the cryptodires or the pleurodires

(Rich & Rich 1989).

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Figure 4. Notochelone costata. A, type specimen; B, suggested restoration of carapace; and C, of plastron.
(From Owen 1882b and Gaffney 1981).

The impression of a turtle shell was found northwest of Winton, (PI.1.). The sediments
are mapped as Winton Formation, which is a terrestrial facies, but the flatness of the sheil
suggests that this turtle was a marine, or at least aquatic, form.

The carapace and vertebrae of a small marine turtle, apparently not Notochelone, have been
collected near Boulia in West Queensland. This form has yet to be studied. An incomplete
jaw and other bones of a small, probably freshwater turtle have been found in the Griman Creek
Formation at Lightning Ridge, New South Wales (Molnar 1980b).

FOSSIL REPTILES IN AUSTRALIA - 619

Figure 5. The Victorian turtle Chelycarapookus arcuatus. A, internal mold of the carapace in dorsal view; B,
plastron (as preserved); C, reconstruction of the dorsal vertebrae and ribs, for comparison with, D, those of
the extant chelid Emydura macquarii. The posterior ribs (pr) have much longer necks in Chelycarapookus than
in Emydura. f, fossa mentioned in text. (From Warren 1969 and Gaffney 1981)

Palaeozoogeography

Of the recent continental chelonians, the cryptodires live (or have recently become extinct)
on all continents but Antarctica. The pleurodires, once cosmopolitan, are now restricted to
Africa, South America and Australia. All modern Australian continental chelonians belong to
a single family, the chelids. Unfortunately, the known Australian Mesozoic turtles tell us

nothing about how this distribution came about.
Notochelone is a chelonioid (cryptodire). This taxon was seemingly widespread through

620 - MOLNAR

the Cretaceous oceans. Thus, they provide no evidence of unusual Australian taxa, such as will
be discussed later for other groups. But Chelycarapookus was decidedly unusual and unlike
other chelonians (in the fossa, broadened neurals and clongate necks of posterior ribs: Warren
1969), It was a continental river-dwelling animal. All other chelonian fossils are unstudied.

So, modern and Early Cretaceous Australian continental chclonians are unusual, but there is
no evidence that the one gave rise to the other,

SYNAPSIDS: ANCESTORS AND KIN OF MAMMALS

The synapsids, prominent in Permo-Triassic times, gave rise to the mammals. The cladist,
of course, recognizes that mammals are synapsids. All synapsids are characterized by the
possession Of a synapsid temporal fenestra (Fig. 1), hence their name.

Evolution and Taxonomy

Synapsids have been divided into two groups, primitive and advanced, that have been named
pelycosaurs and therapsids respectively. Pelycosaurs, like reptiles, are not a real group in the
cladistic sense (Reisz 1980). However, while realizing this, we will find it convenient to use
the term to designate primitive synapsids. Pelycosaurs appear in the fossil record very soon
alter the first reptiles, in the Early Carboniferous. They, together with the therapsids, form the
bulk of the reptilian faunas of the late Palaeozoic. They seem to have been mostly carnivores,
feeding, predominantly upon amphibious or aquatic animals, rather than terrestrial ones. After
all, most terrestrial animals presently known from this time were carnivores. A very good
reference on pelycosaurs and therapsids is Kemp (1982); it provides fascinating information on
the evolution, anatomy and life-styles of these beasts.

During the later part of the Early Permian therapsids evolved from the carnivorous
sphenacodont pelycosaurs. ‘There are several groups of therapsids, all characterized by the
structure of the angular bone in the mandible. The angular has a thin sheet of bone projecting
back from its lateral surface, called the reflected lamina. In therapsids the temporal fenestra has
become enlarged over those of pelycosaurs, and large single canine teeth developed in each jaw.
Pelycosaurs did show some dental differentiation, but had either no or multiple canines in each
jaw. ‘The story of how therapsids diversified and one group, the cynodonts, eventually evolved
mo mammals is beautifully detailed by Kemp.

The Australo-Antarctic Record

The major localities producing pelycosaurs are in the southwestern United States, especially
Texas, although significant finds have also been made in Nova Scotia (Canada) and Europe.
Most therapsids have been found in Russia and South Africa with increasingly important
discoveries in South America, China and even in Antarctica. With such a distribution, the
erstwhile absence of synapsids in Australia has been puzzling. Early Triassic faunas of similar
age to those of South Africa had been discovered in Queensland, Tasmania and Western
Australia, But there was no sign of therapsids, which are well represented in contemporancous
faunas elsewhere, ‘The importance of this absence was unclear, but disturbing. Some workers
felt the absence was real, ic. therapsids never existed in Australia, while others (myself
included) contended that therapsids had existed in Australia, but simply had not been found.

This incipient controversy was resolved in 1983 by R.A. Thulborn, who identified a
dicynodont, cither Kannemeyeria (Fig. 6) or a close relative, from Rewan, Queensland. The
identification was based on a large, but incomplete quadrate found by Thulborn during the
winter of 1982.

FOSSIL REPTILES IN AUSTRALIA - 621

Palaeobiology and Palaeozoogeography

Dicynodonts are unusual animals, unlike any now alive. They include the well-known
Lystrosaurus a strange, probably amphibious, dicynodont found through most of the
Gondwanan and some of the Laurasian continents. Dicynodonts tended to be massive, with
stout, sprawling limbs, short tails and relatively large heads (Fig. 6). They possessed a beak,
and sometimes rows of simple check teeth. Most had a large canine tusk in cach upper jaw,
but some late forms replaced these tusks with tusk-like processes of bone, a mystifying case of

Seuy Bees: The temporal fenestra was very large, accommodating greatly developed jaw
muscles.

Figure 6. Skeleton of Kannemeyeria, based on specimens from South Africa. The Queensland dicynodont
material likely came from an animal similar to this one (from Pearson 1924).

Plant food was apparently cut by the beak and then sliced into short sections by the action
of the jaws. A sharp vertical plate placed posteriorly on the mandible apparently sliced plant
forage against a flat horizontal plate on the palate, (Crompton & Hotton 1967) much as one
slices celery with a knife against a wooden cutting board. Oddly enough, this action was
duplicated in one group of theropod dinosaurs, the oviraptorosaurs. Dicynodonts had apparently
developed a greater stride of the forelimb than earlier therapsids, and some workers believe that
the hindlimb was held more or less erect. This view is not shared by all workers. Dicynodonts
may have been endothermic but there is no convincing evidence yet. Endocranial casts indicate
that the brain was developed to about the same degree as those of other therapsids.

Kannemeyerid dicynodonts have been found largely in Africa and South America, but also
in Europe, North America and China. Recently, Kannemeyeria has been reported in Antarctica
(Hammer 1988). Other dicynodonts inhabited Antarctica, including at least three species of
Lystrosaurus, sometimes considered a kanemeyeriid and sometimes put in its own family. It
seems to have been a common animal there.

Kingoria has also been reported (De Fauw 1988). This is unusual as the Antarctic Kingoria
was found in the Triassic Fremouw Formation. Outside Antarctica Kingoria is known from
southern and eastern Africa, but only from the Permian. Thus, if this report is correct, it was a
relict form in Antarctica, which lived on there after becoming extinct elsewhere. We shall see
this phenomenon again among Australian Cretaceous reptiles.

The finding of a dicynodont in Australia is welcome not only in that it furthers our
knowledge of Australia’s past, but also in lending confidence to theories of animal distributions

622 - MOLNAR

based on knowledge of plate tectonics.

Other therapsids have been discovered in the Antarctic, in fact they are the most common
reptiles known there. The cynodonts, Thrinaxodon and Cynognathus, have been found in the
Transantarctic Mountains (Colbert and Kitching 1977, Hammer 1988) They are well-known
from South Africa, and Cynognathus also lived in Argentina. Other advanced therapsids,
scaloposaurids, are known from Antarctica. (Colbert & Kitching, 1981). These are not as
closely related to mammals as cynodonts. The occurrence of such therapsids in Antarctica
suggests that they may have inhabited Australia as well. The Antarctic cynodonts are
noteworthy in this context, as Australia is the only continent host to all three major living
groups of mammals: monotremes, marsupials and placentals. Monotremes (so far) are known
nowhere else - perhaps they evolved from cynodonts in Australia.

LEPIDOSAURS: LIZARDS, SNAKES AND KIN

Lepidosaurs, thus far, are rare and few in the Australian Mesozoic, so only a short account
will be given here. For more complete information see Romer (1956), Carroll (1977, 1988)
and Benton (1985).

Anatomy

Lepidosaurs have a diapsid skull, often strongly modified postorbitally, especially in
squamatans (Fig. 1) The postparietal and tabular bones have been lost from the skull. The

Figure 7. The hooked v'h metatarsal in lepidosaurs. A, tarsal and metatarsal bones of a varanid in dorsal
view; B, the vith metatarsal of an iguanid in lateral view (not to scale). The vith metatarsal is gently curved
in lateral aspect, and bent at right angles ("hooked") in dorsal aspect. IV, IVth metatarsal. V, Vth metatarsal.
(From Robinson 1975).

FOSSIL REPTILES IN AUSTRALIA - 623

long bones have separate centres of ossification at their ends, epiphyses, that form the articular
surfaces. And, as in mammals, when these epiphyses fuse to the shaft, the animal's growth
ends. The feet are characteristic in three respects: the proximal tarsals (astragalus, calcaneum
and centrale) have become fused; distal tarsals 2 and 5 have been lost; and metatarsal V has
become "hooked". A bit of description of the "hooked" metatarsal V is in order, as this is often
not well understood. The fifth metatarsal of lepidosaurs has a shaft that is gently arched. In
addition, at its proximal end, the shaft is bent sharply medially, to project across the proximal
end of metatarsal IV (Fig. 7) (Robinson 1975). Lepidosaurs are also characterized by features
of the soft anatomy, given by Benton (1985) and Carroll (1988).

Evolution and Taxonomy

_ Lepidosaurs are the major group of the lepidosauromorphs, hence the name. They are
distinguished from archosauromorphs on essentially locomotory differences. Apparently
lepidosauromorphs initially adopted a mode of locomotion involving lateral flexing of the
vertebral column, whilst archosauromorphs (like mammals) adopted a mode that reduced lateral
motion of the vertebral column (Carroll 1988). Lepidosauromorphs, at least the land-dwelling
ones, hold the proximal limb elements, the humerus and femur, in a horizontal position,
whilst archosauromorphs, at least the advanced ones, hold the limbs directly beneath the body
(like mammals). Early lepidosauromorphs differ from other primitive diapsids in the
possession of a large sternum. This sternum is not homologous with the 'sternum' of
archosaurs, birds and mammals, but functions to increase the length of the stride in
lepidosauromorphs. Further details may be found in Jenkins & Goslow (1983).

The origins, early members and phyletic relationships of the lepidosaurs are still poorly
understood (Evans 1984, Carroll 1988). Certain long-established groups, such as the
Rhynchocephalia, are no longer recognized (Carroll 1977). Others, such as the eosuchians and
protorosaurs, have now been re-defined (Benton 1985, Carroll 1988) and no longer include
many forms that they once did. ‘ uch changes must be remembered when rzading older works,
and even in some modern works, for the names of some groups are not yet universally accepted
(e.g., Benton's Prolacertiformes is Carroll's Protorosauria).

Until recently it was thought that lizards originated late in the Palaeozoic and underwent an
adaptive radiation during the Triassic (Robinson 1967). Then a second radiation, later in the
Mesozoic, led to the modern groups. However, the Triassic forms, the paliguanids, are not
lizards. They are a group of primitive lizard-like lepidosaurs, that quite possibly are not real in
the cladistic sense. Several different groups of lizards were present in the Late Jurassic, all of
which looked like modern lizards. Modern squamatans consist of three groups: the lacertilians
(lizards and kin), serpents and amphisbaenians, a group of worm-like, limbless forms (Gans
1979). Early in the Cretaceous the snakes appeared (including some marine forms, the
symoliophids, which are the closest approach to sea serpents we know) and a group of marine
lizards, the mosasaurs. Mosasaurs developed highly kinetic skulls, altered their limbs into
paddles, and became quite large (for squamatans). They were apparently active predators,
feeding on ammonites (Kauffman & Kesling 1960) and likely fish, turtles and other large
marine reptiles (Russell 1967) as well. In Australia only one Mesozoic lepidosaur is clearly
represented: a mosasaur.

The Australian Record

A paliguanid has been reported from the Early Triassic at Rewan, Queensland (Bartholomai
1979). This animal, Kudnu mackinlayi, is represented by an incomplete snout (Fig.8). It was
originally thought a lizard, related to Paliguana from South Africa. However, in view of both
the incompleteness of the specimen and our poor understanding of the paliguanids, Kudnu can

624 - MOLNAR

be identified only as a small reptile. A second reported lepidosaur from this locality (Thulborn
1984), has not yet been studied.

Figure 8. Skull of Kudu mackinlayi, a Triassic reptile from Queensland. s, impressions of sclerotic plates.
(From Bartholomai 1979).

An incomplete humerus found near Cape Paterson, Victoria, originally thought dinosaurian,
most closely resembles that of a large lizard (Molnar 1980b). More informative lepidosaur
material has been found in the Late Cretaceous near Gingin, Western Australia. This is an
incomplete left forepaddle (Pl. 2) of a mosasaur (Lundelius & Warne 1960), apparently similar
to Platecarpus.

Palaeozoogeography

Both Triassic and Cretaceous lepidosaur fossils are too fragmentary for meaningful
comparison with overseas forms. The Western Australian mosasaur is one of the few yet
found in the Southern Hemisphere outside of New Zealand, where they were abundant.
Mosasaur vertebrae have also been found at Vicecomodoro Marambio Island (also known as
Seymour Island) off the Antarctic Peninsula. This indicates that mosasaurs were present in the
Australo-Antarctic region, and suggests that their absence from the Australian fossil record is
due to the paucity of Late Cretaceous marine deposits.

ICHTHYOPTERYGIAN: FISH-LIKE REPTILES OF THE SEA

Ichthyopterygians were a successful group of fish-like (or porpoise-like) marine reptiles.
The later, more widely known ichthyosaurs, had a familiar appearance of modern porpoises and
dolphins, with a long snout and prominent dorsal fin. Their sickle-shaped tail, however, was
vertical unlike the horizontal flukes of dolphins.

Anatomy

Until recently it had been supposed that ichthyopterygians showed a unique pattern of
temporal fenestration (called parapsid). However, upon detailed study it turned out that
ichthyopterygians, sauropterygians and placodonts exhibited the euryapsid pattern (Fig. 1)
(Romer 1968). Ichthyopterygian vertebrae have a characteristic discoid form and are strongly
biconcave. There is little regional differentiation of the vertebrae, and they decrease in size only
gradually in the tail. In post-Triassic forms the phalanges have become flattened, round or
polygonal elements, different from those of plesiosaurs (Fig. 9). The number of phalanges is
increased (hyperphalangy), and the number of digits is usually reduced (although it is increased
in the common Australian form).

FOSSIL REPTILES IN AUSTRALIA - 625

Q

oO Ps
accessory digit 006° accessory digit
fe)

a

primary digits primary digits

Figure 9. Ichthyosaur forepaddles showing the latipinnate condition (/chthyosaurus) (A) and the
longipinnate condition (Temnodontosaurus) (B). i, intermedium; R, radius; r, radiale; U, ulna; u, ulnare.
Scale bars represent 6 cm. (From McGowan 1972a).

Evolution and Taxonomy

Mazin (1981), studying the primitive Triassic ichthyopterygian Grippia (Fig. 10) found that
the fenestra was not exactly of the euryapsid pattern. In Grippia not only the postorbital and
Squamosal ventrally bound the fenestra, but also the postfrontal and quadratojugal. Since the
later ichthyosaurs clearly have euryapsid fenestrae, this suggests that the pattern is convergent.
Grippia resembles sauropterygians in having a distinct emargination along the bottom edge of
the skull behind the tooth row and in front of the jaw articulation. This suggests that, like
sauropterygians, they were derived from primitive diapsids.

Grippia was an early branch (the grippians) that diverged from the line leading to the
ichthyosaurs. More closely related to ichthyosaurs were the mixosaurs. Mixosaurs, although
basically porpoise-like in form, had a long, tapering tail with a low triangular tail fin dorsally.
None of these ichthyopterygians have yet been recognized in Australia, but there is a mixosaur
(Mixosaurus? timorensis) from Timor.

Ichthyopterygians are divided into two groups, the longipinnate and the latipinnate.
Longipinnate ichthyosaurs (Fig. 9) have three primary digits in the forepaddle, fewer and more
widely spaced phalanges in the forepaddle, and rarely or never have bifurcating fore-digits.
Primary digits are those that contact the distal margins of the distal carpals: accessory digits
may contact the carpals, but if so, only along their lateral (not distal) margins. The
latipinnates have four primary digits in the forepaddle, smaller, more numerous and more
closely spaced fore-phalanges and often have bifurcating fore-digits. Other differences and a
discussion of these two groups can be found in McGowan (1972a). Not all workers accept this
separation into two groups, however, pointing out that these two groups do not reflect
ichthyosaur phylogeny. Their ancestry is unknown: most groups of reptiles, and even the
labyrinthodont amphibians, have been considered likely ancestors. The best specimens of
ichthyosaurs come from the Northern Hemisphere, especially from the Jurassic limestones of
southern Germany (Hoffman 1958), but ichthyosaurs were basically cosmopolitan. Save for
the early genera, some of which had not developed the prominent dorsal and caudal fins,

626 - MOLNAR

A

Figure 10. The skull of the primitive ichthyosaur Grippia longirostris, from the Triassic of Spitsbergen, in
lateral (A) and dorsal (B) views - a restoration of Grippia is at the right. The lateral view shows the ventral
embayment of the skull just below and behind the orbit, which is taken to represent the remnant of a lateral

temporal fenestra. This suggests that ichthyosaurs are descendants of diapsid a ncestors. (From Mazin
1981).

ichthyosaurs all appeared much alike. Eurhinosaurus, with its long overshot snout and short
lower jaw (curiously paralleled in the Miocene dolphin, Euhinodelphis) is the only exception
(Abel 1927).

Corres eee ric
TH pry gpg pee eeeele gee eaenley
ean \
Ae
J -
?) ~~

Figure 11. The reconstructed skeleton of Platypterygius australis, in lateral view, with the paddles also
shown in plan view. (Drawn by L. Beirne).

FOSSIL REPTILES IN AUSTRALIA - 627

Palaeobiology

Their widespread distribution suggests that ichthyosaurs were quite capable of dispersal
across the high seas, while their abundance in the south German and the Australian inland seas
suggests that some forms at least enjoyed epeiric and near-shore seas. The Australian species
was limited to the great, apparently cool, (Frakes & Francis 1988) inland sea (Wade 1984).
Some ichthyosaurs fed on fish and cephalopods, as revealed by their fossilized stomach contents
(Pollard 1968, McGowan 1973). They were live-bearers, and some individuals seem to have
been cannibalistic (Ley 1966). The large orbits suggest acute eyesight, but the members of
this group are generally thought to have had poor hearing (McGowan 1973).

It has recently been suggested that ichthyosaurs may have used their forepaddles much as
penguins use their wings - to ‘fly' underwater (Riess 1986). This is not generally accepted, but

the suggestion that they had two modes of swimming, slow ‘flying’ and fast, using the tail, is
worth further thought

The Australian Record

Only one ichthyosaur, Platpterygius australis, (Fig. 11) about 5 m long, is well known
from Australia (McGowan 1972b, Wade 1984). This genus has also been found in central
North America, northern Europe and southern India, throughout most of the Early Cretaceous.
Doubts have been expressed that the Australian species is distinct, but the relationship of the
humerus to the more distal elements is unlike that in any other ichthyosaur. This indicates
that it is a valid species (Wade 1984).

The structure of the forepaddle of Platypterygius is distinctive in having two preaxial
(anterior) accessory digits, a condition apparently unique among ichthyosaurs (Fig. 12).
Platypterygius shows the same geographic range in Queensland as Notochelone and is found in
both the Allaru and Toolebuc formations. It lived in the large, shallow, food-rich inland sea.

Figure 12. Right forepaddle of Platypterygius australis, drawn from two specimens, one for the proximal
and the other for the distal part. This paddle shows the increased number of digits.

There are also fragmentary ichthyosaur remains known from South Australia (Molnar &
Pledge 1980), Western Australia (Teichert & Matheson 1944) and from near Darwin, Northern
Territory (Teichert & Matheson 1944). Those from South Australia and the Northern Territory
were contemporaneous with the Queensland ichthyosaurs, but those from Western Australia are
Late Cretaceous in age. Recent study of the ichthyosaur material from near Darwin (Murray
1985) suggests that these specimens had drifted dead for some time before falling to the sea-
floor to be preserved. Some of these specimens had lost their skulls, whilst in one from
Queensland the skull had apparently fallen to the seafloor, thus dragging the body with it, to be

628 - MOLNAR

buried with the snout penetrating vertically into the sediments. Cosgriff & Garbutt (1972)
suggested that there may be ichthyosaur material from the Blina Shale. If true, this would be
the oldest known ichthyosaur, as the Blina is Early Triassic in age. The Late Cretaceous
ichthyosaur from Western Australia may belong to a species other than P. australis and appears
to be the last known ichthyosaur.

Palaeozoogeography

Although the Australian ichthyosaur P. australis is endemic, it does not differ much from
those elsewhere. Ichthyosaurs demonstrate the basic similarity of marine reptiles in and out of
Australia. The Australian species may have arisen because the inland sea was not broadly
connected to the open ocean to the north. Hence, the Australian population was at least
somewhat isolated.

SAUROPTERYGIANS: MORE AQUATIC REPTILES

Another successful group of aquatic reptiles, both freshwater and marine, were the
sauropterygians. No indication has yet been found in Australia of the nothosaurs (probably
most diverse in China). The other major group of sauropterygians, the plesiosaurs, is usually
divided into two basic groups: the small-headed and long-necked plesiosauroids and the large-
headed and short-necked pliosauroids. These are informally known as plesiosaurs and pliosaurs,
although the former term is also often used to include both groups.

Anatomy

The basic skeletal features of plesiosauroids and pliosauroids include a low, akinetic skull,
with a euryapsid temporal fenestra (Fig. 1). The nares are placed high on the skull near the
orbits (as in ichthyosaurs), and there is a parietal foramen. The nasal bones are very small (or
are lost) and do not border the nares. The palate is well-developed, and the pterygoids extend
below the braincase. The vertebrae are quite distinctive and clearly unlike those of
ichthyosaurs. The centra are slightly biconcave, usually with characteristic paired ventral
foramina not found in other reptiles (Fig. 13C). At least some plesiosauroids or pliosauroids
had small, but well-developed chambers (centrocoels) situated centrally within the vertebral
centra (Fig. 13). The paired ventral foramina opened into these chambers. These foramina do
not always persist throughout the vertebral column, nor are they known in all plesiosaurs.
They are usually paired, but may range in number from one to four (Wiffen & Moisley 1986).

In the girdles, the scapulae and ilia are reduced and the coracoids, pubes and ischia developed
into great ventral plates. The limbs are modified into paddles, almost oars. They show
considerable hyperphalangy, but the individual phalanges are constricted to give a flattened,
hourglass-shape, quite unlike the phalanges of ichthyosaurs.

Plesiosauroids are distinguished by their usually small heads and long necks, although at
least one group, the cimoliasaurs, seem to have had rather large heads and short necks (Persson
1963). Unfortunately, the cimoliasaurs are poorly known. In plesiosauroids the pubis and
ischium are short, as are the femur and humerus. Often the humerus is longer than the femur.
By contrast, the femur of pliosauroids is usually longer than the humerus, and both elements
are relatively long and slender. Pliosauroids had non-uniform teeth and a short neck. The
cervical vertebrae have ventral keels, absent in plesiosauroids, and the pubis and ischium are
elongate.

FOSSIL REPTILES IN AUSTRALIA - 629

Figure 13. Plesiosaur vertebrae. Top: three views of a cervical vertebra of "Plesiosaurus” rostratus (an
English species): A, lateral view of vertebra; B, posterior view of centrum; C, ventral view of vertebra.
The paired ventral foramina characteristic of plesiosaurs may easily be seen in the ventral view. Bottom: the
chamber in the centrum of an Australian plesiosaur vertebra. D, in horizontal section; E, in frontal section.
Dotted lines indicate canals to foramina. (Top three illustrations from Owen 1884).

Evolution and Taxonomy

Sauropterygians may have originated near what is now Madagascar. From there has come a
reptile, Claudiosaurus, that may be close to the ancestry of the sauropterygians (Carroll 1981).
Benton (1985) concluded that while Claudiosaurus was a diapsid that shared some derived
features with nothosaurs, it was not clearly related to plesiosaurs. But in view of the six
derived features shared with nothosaurs, and the extension of the palate backwards to cover the
braincase as in nothosaurs and plesiosaurs, Claudiosaurus seems the best candidate so far for a
sister group, if not an ancestor, of the sauropterygians. Apparently the sauropterygian skull

630 - MOLNAR

Figure 14. The hypothetical sequence of changes in the diapsid pattern of temporal fenestrae during the
evolution of plesiosaurs: A, Youngina, a younginiform; B, Claudiosaurus, a primitive diapsid; C, Anarosaurus,
a nothosaur; D, Corosaurus; and E, Plesiosaurus, both plesiosaurs. The loss of the lower temporal bar and
subsequent reduction of the lateral temporal fenestra are shown. (From von Huene 1956 and Carroll 1981).

FOSSIL REPTILES IN AUSTRALIA - 631

form arose by the loss of the lower temporal arch (Fig. 14). In addition to the line leading to
plesiosaurs, there were several divergent branches of early sauropterygians (Sues 1987).
Among these were nothosaurs, moderately large reptiles ( a few metres long) with long necks
and long skulls, and pistosaurs. Pistosaurs were the sister group of plesiosaurs. These early
divergent branches show successive stages in adaptation to oceanic life (Sues 1987).
Plesiosaurs developed during the Triassic. Both pliosauroids and plesiosauroids survived until
the end of the Mesozoic, and members of both early invaded freshwater habitats, freshwater

pliosauroids being known from Queensland as early as the Early Jurassic. Both groups were
cosmopolitan in distribution.

Palaeobiology

The plesiosauroids, with their long flexible necks and flattened bodies, were apparently
fishing forms. The pliosauroids with their cylindrical, rather whale-like bodies, may well have
preyed upon other reptiles as well as large fish. Perhaps pliosauroids also attacked the giant
squid of those times. The Australian inland sea was home to both the large pliosauroid
Kronosaurus and a giant squid (not related to the modern species). Plesiosauroids, unlike
ichthyosaurs, used stomach stones for ballast to offset the buoyancy of their lungs, and may
have rested upon the bottom, to strike at prey with their long necks (Taylor 1981). Functional
studies of plesiosaur limbs by Jane Robinson (1977), have shown that the paddles may have
been hydrofoils generating thrust during the backstroke and also (from lift) during the recovery
stroke, much as do the flippers of sea turtles. This hypothesis has since undergone
modification, most recently by the suggestion that plesiosaurs "swam" much as do modern sea
lions (Taylor 1986). The broad ventral plates of the girdles of plesiosauroids and pliosauroids
anchored the locomotory musculature of the limbs. These girdles and the intervening gastralia
(or ventral ribs) were not rigidly attached to the vertebral column and rib cage, but were
suspended by muscles. This suggests that the vertebral column could be actively bowed, to
keep the lungs inflated, when plesiosauroids or pliosauroids ventured ashore. Thus, unlike
ichthyosaurs or modern cetaceans, plesiosauroids and pliosauroids might have come ashore to
give birth or lay eggs.

The Australian Record

Both plesiosauroid and pliosauroid material is rather common in Australia, and good
specimens are known from Queensland, South Australia and New South Wales. Two
plesiosauroids are reasonably well known in Australia, the cimoliasaur Cimoliasaurus maccoyi
and the elasmosaurid Woolungasaurus glendowerensis. Other named forms ("Plesiosaurus"
sutherlandi and "Plesiosaurus" macrospondylus) are too incomplete for comparison.

Cimoliasaurus is a poorly known cosmopolitan form, reported from western Europe, the
east coast of North America and South Island, New Zealand, as well as Australia. The
Australian material is Early Cretaceous in age, and the genus is reported from the Late
Cretaceous in New Zealand. Cimoliasaurus seems to have had a relatively short neck, and high
slender, gracefully recurved teeth for holding fish. The Australian species, C. maccoyi, differed
from the others in lacking lateral ridges on the cervical centra (Persson 1960). C. maccoyi
comes from White Cliffs, New South Wales, and seems to have been only about 3 or 4
metres long. Persson suggested that C. maccoyi may be the same as Cimoliasaurus planus
known from England and France. co

Having written this, it must be cautioned that the type species of Cimoliasaurus, from New
Jersey, is among the most incomplete plesiosaur specimen known from North America. In
fact, its describer, Joseph Leidy, referred only other species to it: both from North America.
European workers referred eleven species to this genus, however, and, thus, it came to be

632 - MOLNAR

reported from several continents. Some modern workers feel that only the New Jersey species
is valid, and the others (including the Australian) are unrelated.

Woolungasaurus glendowerensis (Pl. 3) had the small head and a very long neck
characteristic of elasmosaurids. All of its cervical centra were longer than high (Persson
1960). Only in regard to its girdle anatomy is Woolungasaurus unusual, as its girdles are
much like those of the more primitive plesiosaurids while its limbs and vertebrae show
advanced, typically elasmosaurid features. Woolungasaurus is known from north Queensland,
from Richmond to Prairie, with a likely specimen from the Neales River, South Australia.

Australian pliosauroids include Trinacromerum leucoscopelus, Kronosaurus queenslandicus
(Longman, 1930) and an early Leptocleidus- or Bishanopliosaurus-like form. Trinacromerum
is known from White Cliffs, with a possible second specimen from near Richmond, north
Queensland. Persson (1960) thought the material so far collected was too incomplete for
specific identification and hence referred to it as Dolichorhynchops? sp. (now replaced by the
name Trinacromerum). I have retained the specific name (leucoscopelus) here for convenience,
not because I am convinced it is valid. The cervical vertebrae are deeply concave, as is
characteristic of polycotylid pliosauroids. The White Cliffs material is similar to that of
Trinacromerum osborni from the midwestern U.S.A. particularly in vertebral and tooth form.

Figure 15. Incomplete skull of Kronosaurus (QM F2446). The skull is slightly distorted by shear, which is
corrected in the outlines of the missing portions. Most of the bone surface has been weathered away, but it
appears that the frontal region was depressed below the level of the snout, and bore three longitudinal
grooves. The teeth are reconstructed from the left side, and may be drawn too small. Apparently, the lower
teeth projected lateral to the snout, while the upper projected medial to the jaw. Diagonal lines designate
regions covered by matrix, while vertical lines designate broken bone. Scale bar represents 25 cm.

FOSSIL REPTILES IN AUSTRALIA - 633

T. osborni_ has a long, ichthyosaur-like rostrum, and a high, strong sagittal crest, implying the

existence of a strong jaw musculature. Trinacromerum was interesting in another way, but that
is better discussed later.

Kronosaurus queenslandicus is a large pliosaur, which is familiar from a reasonably
complete specimen mounted at Harvard University. The Harvard specimen, however, scems not
to be K. queenslandicus, but possibly a second species. K. queenslandicus is based on very
incomplete material, a poorly preserved segment of the jaw symphysis with the teeth broken.
More complete specimens of probable K. queenslandicus (no attempt at adequate comparison
with the type material has yet been carried out) have recently been collected by the Queensland
Museum. One of the reasons for doubt about the identity of the Harvard kronosaur is that it
was collected near Richmond, from the Wallumbilla Formation, while the type material (and a
second partial skull) was recovered from near Hughenden where the Wallumbilla is replaced by
the younger Toolebuc Formation. Also the second partial skull (QM F24446) differs in form
from that of the Harvard specimen. The second skull is very broad, low, and flat (at least 87
cm across by only 13 cm high), with large upwardly directed orbits (Fig. 15). There is no
indication of crushing, although the Harvard skull (MCZ 1285) seems deeper.

The Harvard skeleton is less complete than appears from the mount. Nothing of the
forelimbs was preserved, for example, and much of the skull and vertebral column was
incomplete (Romer & Lewis 1960). Nonetheless, enough material is preserved to give a
reasonable idea of its general form. The skull is just over 25% of the entire length, and the
dorsal column is also elongate. The neck is very short. The general build is massive and
robust, and the ventral, plate-like regions of the girdles elongate. Zygapophyses are lacking
from both the posterior dorsals and the caudals, which suggests marked flexibility in that
region. The Harvard skeleton is just under 13 m in length (12.8 m to be exact), but may be
smaller than the recently collected material in the Queensland Museum, The British
pliosauroid Stretosaurus was larger.

A freshwater pliosauroid has been found in the Early J urassic near Mt Morgan, Queensland.
The bone itself has been lost, but the matrix has retained good impressions of the vertebrae,
teeth and some of the limb elements (Fig. 16). This specimen was initially described by
Bartholomai (1966b) as Early Cretaceous (or perhaps Late Jurassic) on the basis of comparison
with the Wealden freshwater pliosauroid, Leptocleidus superstes. I had suggested (1980b) that
this constituted evidence for free migration into and out of Australia during the Early
Cretaceous. However, it is now known that these beds are Early Jurassic, and hence this
“evidence” is invalid. A pliosaur similar to that from Mt Morgan has been described (Dong
1980) from the Chinese Jurassic (Bishanopliosaurus youngi), but detailed comparison with the
Mt Morgan form has not been carried out.

The Mt Morgan pliosauroid together with two unstudied forms, possibly plesiosauroids,
from near Springsure, represents the earliest known Australian freshwater sauropterygian.
Apparently freshwater pliosauroids retained primitive features (as do the freshwater platanistid
dolphins), particularly in the structure of the girdles. The marine Trinacromerum retains these
states as well, so that Trinacromerum may have been a secondarily marine form derived from
freshwater ancestors (Andrews 1922).

Other more fragmentary plesiosaur material, has been found elsewhere in Australia: from
the Walsh River, Cape York Peninsula; Lightning Ridge, New South Wales; Andamooka
(probably three kinds, Molnar & Pledge 1980) and Coober Pedy, South Australia; and the
Otway Ranges, Victoria. Remains have also been found in the Late Cretaceous at Dandaragan,
Western Australia (Teichert & Matheson 1944) , and, as mentioned previously, in the Early
Jurassic Evergreen Formation near Springsure, southeastern Queensland (Thulborn & Warren
1980). One of the better preserved South Australian plesiosaurs, appears to be unique,
reportedly with the hind paddles inclined backwards on the femur, but it has yet to be studied.

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Figure 16. Vertebrae (A, C), ischium (B) and neural arch (D) of the fresh-water pliosaur from near Mt
Morgan, Queensland. The vertebrae are characterized by the deep transverse processes, similar to those found
in Leptocleidus and related forms. These elements are each drawn from impressions left in the matrix, with no
bone actually preserved. Scale bar represents 5 cm.

Palaeozoogeography

As with the ichthyosaurs, there are no very distinctive endemic forms in Australia.
Cimoliasaurus appears to have been cosmopolitan. But in view of the doubt that it is valid, no
conclusions may be drawn from the reported Australian occurrence (other than that the
specimens are scrappy). Woolungasaurus is an endemic genus, but not greatly different from
elasmosaurids found elsewhere. Kronosaurus has been reported, at least tentatively. from
Colombia, in South America (Acosta-A. et al. 1979). Two large complete and articulated
specimens are known. Hopefully these will soon be studied and their relationships to the
Australian Kronosaurus revealed. The remaining Australian material has either yet to be
studied or is too incomplete for zoogeographical analysis. Like ichthyosaurs, the plesiosaurs
suggest the continuity of marine reptiles around the world.

Elasmosaurids and cryptocleidids have been found in the Late Cretaceous of the Antarctic
Peninsula (Gasparini & Goni 1985, Chatterjee & Small 1988). This, together with their
occurrence in New Zealand and Australia, indicates that they lived in polar latitudes.

PROLACERTIFORMS: LIZARD-LIKE ANCESTORS OF ARCHOSAURS

The prolacertiforms (see Appendix I-4,5) are a group of small to moderately small
Permo-Triassic reptiles. They have long been of obscure relationships, so obscure that their

FOSSIL REPTILES IN AUSTRALIA - 635

former name, protorosaurs, has been discarded by some as being imprecise. Prolacertiforms
share with archosaurs certain features, such as posteriorly curved (recurved) teeth, set in distinct
sockets (thecodont implantation), an ear region with a high backwardly concave quadrate
contacting the paroccipital process, reduction or loss of the posttemporal fenestrae, and an ankle
in which the astragalus and calcaneum contact each other on both sides of a perforating
foramen. Other similarities are given by Gow (1975) and Benton (1985).

The prolacertiforms are named for the prolacertids. Long considered ancestral to, or at least
related to, lizards the prolacertids were restudied by Gow (1975), who concluded they were
related to thecodonts, perhaps being ancestral to that group. A prolacertid has been reported in
the Early Triassic of Rewan Station, southeastern Queensland. It is represented by two pieces
of the skull (Bartholomai 1979). This animal, Kadimakara australiensis (Fig. 17) was
interpreted as closely resembling Prolacerta. Benton (1985) redefinded the prolacertids as
possessing a tetraradiate squamosal; long, narrow palatal bones with long choanae; and a gap
between the pterygoids and posterior vomers to accommodate part (the cultriform process) of
the parasphenoid. Since Benton's work followed Bartholomai's, I re-examined the material to
find out if Kadimakara was related to Prolacerta. A gap beween the pterygoids to accommodate
the parasphenoid is present (Bartholomai 1979), whilst close examination of the squamosal
shows that it is tetraradiate. Bartholomai was in doubt about this and presented two alternative
interpretations of this region: his second seems to be correct (Fig. 17). So, Kadimakara is a
prolacertid (Fig.18). When alive it was probably an inscct eater.

= ae,

Figure 17. A, Prolacerta, skull and lower jaws (Gow 1975); Kadimakara australiensis, a Triassic
prolacertiform from Queensland. B, C, reconstruction of skull and lower jaws; D, back of skull in dorsal view
(tetraradiate squamosal (sq) at right) (From Bartholomai 1979).

Prolacertans are found in South Africa and Antarctica, as well as Australia, but some forms
(e.g. Boreopricea) lived in the Northern Hemisphere. The same species, Prolacerta broomi, is
known from both South Africa and Antarctica, where it is very abundant (Colbert 1987),
highlighting the faunal continuity of these lands during the Early Triassic. This species may

636 - MOLNAR

yet be found in Australia.

THECODONTS : PRIMITIVE ARCHOSAURS

Thecodonts are the earliest members of the archosaurs, and evolved into dinosaurs,
crocodilians and birds. Thecodonts include all forms that are clearly or reasonably considered
archosaurs, but are more primitive than members of the other archosaur taxa. As such, this
group includes disparate forms, that are difficult to relate to each other. For this reason
thecodonts need further study. In the cladistic view thecodonts, like reptiles, are an invalid
taxon.

Figure 18. The skeleton of Prolacerta. Presumably Kadimakara also looked very much like this. (From
Gow 1975).

Anatomy

Thecodont characters are basically archosaur characters, including a diapsid skull with an
antorbital fenestra (Fig. 1) and usually a mandibular fenestra, in the posterior portion of the
mandible. Thecodonts typically have an imperforate acetabulum, and broad, plate-like pubis
and ischium, a primitive pelvic structure. Unfortunately, these characters are typical, but not
diagnostic. This is not to say that the individual groups of thecodonts cannot be well defined,
for they can. As expected for a group that includes the early members of several divergent
lineages, thecodonts show a variety of forms. Some were quadrupedal and others bipedal, some
were herbivores and others carnivores, several groups were armoured, some were land-dwellers
and others were amphibious. Further comments on the thecodont anatomy will be given in the
section on evolution and taxonomy. The defining features (autapomorphies) of the Archosauria
are given by Benton and Clark (1988).

Evolution and Taxonomy

As the group responsible for the archosaur radiation, and hence the bird radiation, the origin
of the thecodonts is of considerable interest. Thecodonts were usually thought to have derived
from the eosuchian (now the younginiform) lepidosaurs, and Gow (1975) nominated the
prolacertiforms (which he called the parathecodonts) as ancestral to, or close to the ancestry of,
the thecodonts. This view is now generally accepted. Other views had been aired as well.
Cruickshank (1970) has suggested that archosaurs share ancestry with sphenodonts. Reig
(1970) made an impressive, but apparently not convincing, case for the origin of archosaurs
from pelycosaurs. Many of Reig's similarities were plesiomorphies, and this seriously
weakened his case in spite of a few shared, derived features that he found.

The problem of thecodont classification is the determination of the relationships of these

FOSSIL REPTILES IN AUSTRALIA - 637

groups to one another and to their descendants. Certain individual thecodonts do resemble later
archosaurs, e.g. Lagosuchus, the dinosaurs, and Scleromochlus, the pterosaurs (Padian 1984).
None of the major groups of thecodonts, however, show any clear-cut relationships to the later
forms, although such relationships have been proposed (Walker 1964).

There has long been no generally accepted classification for thecodonts. Recently Gauthier
(1986) and Benton and Clark (1988) have looked at the interrelationships of archosaurs in terms
of their relationships to the two surviving groups, crocodilians and birds. For convenience, we
may divide the archosaurs into three groups: those which diverged from the lineage leading to
the latest common ancestor of crocodilians and birds; those that derive from the common
ancestor of crocodilians and birds but are more closely related to crocodilians than to birds
(crocodylotarsans), and; those that derive from the common ancestor and are more closely
related to birds than to crocodilians (ornithosuchians). Those taxa of the second and third
groups, all descendants of the common ancestor of crocodilians and birds, are known as crown-
ue archosaurs. Thecodonts include all of the first group, and the primitive members of the
other two.

The classification here used follows that of Benton and Clark (1988). Four groups of
archosaurs diverged before the common ancestor of crocodilians and birds:

Proterosuchia. The oldest forms, mostly sprawling crocodile-like predators. Found in
the Russia, China, India, Australia and South Africa.

Erythrosuchia. Also quadrupedal predators, but with a more upright posture and broader,
deeper skull. Found in South Africa, Russia and China.

Proterochampsidae and Doswellia. The proterochampsids were crocodile-like forms,
with a broad, low, flat skull with slit-like nares. Found in Brazil and Argentina. Doswellia
has a similar skull but its body is sheathed in armour. Found in Virginia (U.S.A.).

Euparkeriidae. Small, bipedal predators. Found in South Africa and China.

In addition to the Crocodylia, the Crocodylotarsi includes four groups of thecodonts:

Poposauria. Large bipedal carnivores, superficially resembling dinosaurs such as
Tyrannosaurus. Found in the U.S.A., Germany and the British Isles.

Pseudosuchia. Also includes a group of large, bipedal carnivores (Rauisuchia), as well
as quadrupedal, armoured herbivores (Aetosauria). Rauisuchia found in Argentina, Brazil, India,
Tanzania, Morocco and Switzerland. Aetosauria found in Scotland, Germany, China, Chile,
Argentina and the U.S.A. (It is important to realize that the usage of the term 'Pseudosuchia’'
by Benton and Clark, while close to the original usage of the last century, is different from that
used for much of this century.)

Gracilisuchus. A small bipedal carnivore. Found in Argentina.

Phytosauria. Quadrupedal, amphibious predators, very crocodile-like in appearance.
Found in Morocco, China, India, Madagascar, Turkey, England, Germany, and the U.S.A.

Ornithosuchians include the dinosaurs and two groups of thecodonts.

Ornithosuchidae. Large, bipedal predators. Found in Scotland and Argentina.

Lagosuchidae. Small, bipedal or quadrupedal predators. Found in Argentina.

This scheme leaves some groups unclassified. Most of these are either incompletely
preserved or not easily accessible for study. While the bulk of thecodonts are accounted for,
those left out are among the most bizarre and interesting, and so will be bricfly mentioned:

Lotosauria. Large beaked quadrupeds, with a low sail along the back. Found in China,
Tanzania, Germany, Brazil and the U.S.A.

Longisquama. A small, apparently quadrupedal form with greatly elongate scales along
the back, perhaps used for gliding (Haubold & Buffetaut 1987). Found in Soviet Central Asia.

Megalancosaurus. A small, perhaps arboreal animal, with a number of bird-like
features in the skull and pelvis and a very large ‘hand’. Found in Italy.

Some workers include the lotosaurs in the rauisuchians. The term "pseudosuchian" was
until recently widely used for rauisuchians, actosaurs and protodinosaurs. Originally it meant

638 - MOLNAR

groups.

The oldest thecodont, a proterosuchian, comes from the Permian of Russia. What may be
the most primitive of thecodonts, Cosesaurus aviceps, is known from a single specimen from
Europe. It is a small, gracile, contentious form. Some workers believe that it shares no
derived features with archosaurs and regard it as a prolacertiform. Other views are possible,
however, as the specimen seems to show an antorbital fenestra. If this is correct, this animal
may link archosaurs with prolacertiforms. Another proposed primitive archosaur is
Mesenosaurus romeri, which also reportedly has an antorbital fenestra, albeit a very small one
(Ivakhnenko & Kurzanov, 1978). Mesenosaurus, from the Permian of northern Russia was
previously considered a pelycosaur.

Figure 19. Basic pattems of archosaurian tarsal structure. The left ankle of three genera is shown, with
the elements pictured above and the pattern diagrammed below. A, D, the advanced archosaurian condition
(Thescelosaurus), with the hinge passing between the proximal and distal tarsals; B, E, the crocodilian
condition (Crocodylus), with the hinge joint passing between the proximal tarsals, and with a peg (p) on the
astragalus (a) fitting into a socket on the calcaneum (c); C, F, the crocodilian-reversed condition
(Riojasuchus), with a peg on the calcaneum fitting into a socket of the astragalus: this condition is not
widespread. Other abbreviations: i, first metatarsal; f, fibula; t, tibia; v, fifth metatarsal. (From Bonaparte

1971 and Thulborm 1980).

The relationship of the thecodonts to dinosaurs has occasioned much interest over the past
ten years. The studies appear, however, to have produced more enthusiasm than
enlightenment. Basically they have centred on the structure of the ankle. Archosaurs exhibit
two types of ankle joint (Fig. 19) (in fact there is a third variant among one group of
thecodonts). One is a simple mesotarsal joint, in which the "hinge" about which the foot
rotates lies between the distal row of tarsals and the two proximal tarsals (astragalus and

FOSSIL REPTILES IN AUSTRALIA - 639

calcaneum). This a feature of the ornithosuchians. In the other, the crocodiloid joint, the
hinge passes between the two proximal tarsals. The astragalus is fixed to the calf, while the
calcaneum is fixed to the foot. The astragalus articulates with the calcaneum via a peg-and-
socket joint, the astragalus bearing the peg and the calcaneum the socket. This is found in the
crocodylotarsans. Further details of the anatomy may be found in Thulborn (1980). A third
form, known as the crocodile-reversed joint, is essentially like the crocodiloid joint, except that
the astragalus bears the socket and the calcaneum the peg. Difficulties lay in understanding, or
reconstructing, how one of these forms might have evolved into another, as it appears that
some lines of archosaurs would have had to reverse the trend of ankle joint evolution. A
further difficulty is that various dinosaurs (such as Allosaurus) have various combinations of
pegs and sockets at the astragalus-calcancum junction. This has been compounded by
misidentifications of the type of joint present in certain archosaurs. The whole situation has
not been wholly resolved to general satisfaction, although Bonaparte (1984) has presented a
significant analysis of locomotion in thecodonts, and Thulborn (1980) has suggested, sensibly,
that both archosaur ankle joints may have evolved from a more primitive, intermediate
condition.
Crocodilians are less modified from their ancestors than birds and dinosaurs. Crocodilians
should be sufficiently familiar to need no detailed introduction, although an excellent summary
can be found in Buffetaut (1979).

Palaeobiology

In keeping with the variety of different forms among the thecodonts, there was a similar
variety of life-styles. Such diversity is often found in the radiation of successful evolutionary
lines. Also, like other successful lines, the thecodonts were first and primarily carnivores. The
armoured aetosaurs were probably plant-eaters. The aetosaur skull had an extended tip to the
snout, so maybe they rooted for roots and tubers like modern pigs. The lotosaurs, with their
parrot-like beaks and dorsal sails, have so far defeated attempts to understand their mode of life.

Most thecodonts, including those found in Australia (proterosuchians), were carnivores.
Little work has been done on the mode of life of the proterosuchians. As mentioned above,
they probably lived much like crocodiles, as sprawling, amphibious predators.

Crocodilians need little description, as modern ones are quite familiar. Information on the
Australian species is provided by Webb & Maniolis (1988a, 1988b). However one important
point must be made. Modern crocodilians, formidable as they are, give little indication of the
diversity found among the Mesozoic crocodilians. There were marine crocodilians
(thalattosuchians), with long snouts, paddles instead of feet and dorsal tail fins. There were
large forms (stomatosuchians, and the Cainozoic nettosuchians) with broad flat skulls and
minute teeth. These may have lived in ponds, and scooped up to eat whatever was drifting in
the water (Langston 1965). The uruguaysuchians were short-snouted crocodilians with tecth
like those of herbivorous lizards and dinosaurs, and hence may have been plant-eating
crocodiles. Others, the ziphodont crocodiles (which include members from several unrelated
lineages), had laterally compressed shearing teeth like those of the theropod dinosaurs. These
became more prominent during the early Cainozoic, and seem to have been land-dwellers rather
than amphibious animals. This great diversity of forms and habits is now sadly reduced.

The Australian Record

Only proterosuchian thecodonts are known so far in Australia. Two genera,
Tasmaniosaurus and Kalisuchus (Fig. 20), are from Tasmania and Queensland respectively.
Tasmaniosaurus triassicus has been found in flood-plain pond deposits of the Early Triassic
Knocklofty Formation (Camp & Banks 1978). This environment was shared with a diverse

640 - MOLNAR

Knocklofty Formation (Camp & Banks 1978). This environment was shared with a diverse
fauna of temnospondyls upon which Tasmaniosaurus may have fed (Thulborn 1986a). The
skull of this thecodont was moderately low and long, as in other proterosuchians. Its reported
relationship to phytosaurs is in error (Thulborn 1986a). Although the postcranium is
preserved, the quality of the preservation is poor. The vertebrae appear typical of
proterosuchians, and there is a well-developed interclavicle, a primitive character not found in

most other proterosuchians. Tasmaniosaurus is a proterosuchian resembling the well-known
Chasmatosaurus.

i <— ,

Figure 20. Reconstructed skulls of the two known Australian thecodonts. A, Tasmaniosaurus triassicus, B,
Kalisuchus rewanensus. (From Thulbom 198€a).

Kalisuchus rewanensis, from the Early Triassic of southeastern Queensland (Thulborn
1979), was also found with a diverse fauna of temnospondyl amphibians. The material is
isolated fragments, but shows the characteristic proterosuchian features: subthecodont teeth and
triple-headed ribs. In addition, there are numerous detailed resemblances to Chasmatosaurus.
Differences from Chasmatosaurus include a crocodiloid calcaneum, strongly developed

FOSSIL REPTILES IN AUSTRALIA - 641

maxillary shelf. This latter suggests that Kalisuchus had a broad snout. The slender limb
bones, expanded snout, calcaneum and the long (for a proterosuchian) neck and tail suggest an
approach to the dinosauran condition. Nevertheless, when alive, Kalisuchus (and
Tasmaniosaurus) probably looked much like crocodiles.

Cosgriff (1983) reported two thecodont fossils from the Lower Triassic Fremouw
Formation of Antarctica. Two bones, both incomplete, were found: a vertebra and a humerus.
Cosgriff was uncertain as to what kind of thecodont they belonged. The vertebra resembled
those of ornithosuchids and proterosuchians, and the humerus was like those of the
rauisuchians. The evidence at hand is not adequate to resolve this uncertainty, but the forms of
the two bones do suggest a more derived creature (or creatures) than those known from
Australia. A wide variety of thecodonts lived in South Africa, South America and probably
India, so the occurrence of other kinds of thecodonts in Antarctica is also likely. In turn, this
suggests that other thecodonts may eventually be found in Australia.

Isolated crocodilian material has been found in the Early Cretaceous Griman Creck
Formation at Lightning Ridge (Molnar 1980c). This material, consisting of portions of a jaw
(Pl. 4) and skull, some limb bones and some vertebrae (PI. 5), is interesting, because it
includes procoelous cervical vertebrae. Most Mesozoic crocodilians (protosuchians and
mesosuchians) had amphicoclous vertebrae, that is the centra were excavated at both ends. The
modem crocodilians are characterized by procoelous centra, which are concave on the anterior
face and convex on the posterior. This pattern is thought to optimize mobility and strength of
the vertebral column, although further work is needed to clarify the function of procoelous
vertebrae. Until recently, it was thought that the earliest procoelous crocodilians dated from
the latest Jurassic of western Europe (//ylaeochampsa), and that they did not become
“widespread” until the Late Cretaceous. Thus, the Lightning Ridge crocodile ("Crocodilus"
selaslophensis) seemed to be an early example of a procoelous crocodile rather far from their
presumed area of origin. It now appears, due to the work of J.M. Clark at Chicago, that
procoelous crocodilians were already well-developed in the Late Jurassic of North America.
Thus, they are older than previously believed, and their appearance in the Early Cretaceous of
Australia is less surprising.

Procoelous vertebrae are generally taken to characterize eusuchian crocodiles, although
Benton and Clark (1988) have recently suggested that procoely developed prior to the diagnostic
eusuchian feature, choanae entirely surrounded by the pterygoids. So, while the Lighting Ridge
crocodile is a procoelous crocodile, it is not known to be a eusuchian.

Palaeozoogeography

Both Australian proterosuchians belong to endemic genera, but both show detailed
similarities to taxa known overseas. Tasmaniosaurus is unusual in its possession of a well-
developed interclavicle. The similarities of both forms to overseas taxa are consistent with the
palaeogeographic interpretation of Triassic Australia as part of a single land mass, Pangaea.

The Lightning Ridge crocodilian is quite unlike later Australian (or overseas) forms. The
mandibular teeth are all of the same size and set in adjacent sockets. Later, and most
Cretaceous, crocodiles have mandibular teeth that vary markedly in size, by a factor of about
two. Furthermore the teeth of "C." selaslophensis are set in sockets in a groove in the dentary.
This is rare in crocodiles, being found only in two other, also poorly known, Mesozoic
species. However, such a groove is also found in the jaw of the Eocene Argentinian Eocaiman,
which is similar to the jaw fragment from Lighting Ridge in other ways. A fragment of the
skull or jaw of a large crocodilian was recently found in the Toolebuc Formation, near
Hughenden.

642 - MOLNAR

SAURISCHIANS: MEAT-EATERS AND DINOSAUR GIANTS

Saurischian dinosaurs are well known, for they include such perennial favourites as
Apatosaurus (Brontosaurus) and Tyrannosaurus. They had long been recognized by the pelvic
structure, in which the pubis and ischium diverged (in lateral view) at an angle, often about 30
degrees (Fig. 21B). This was distinguished from the ornithischian condition in which part of
the pubis was inclined posteriorly parallel to the ischium (Fig. 21A). However, in the late
1970s saurischians were discovered in Mongolia that had the pubis and ischium parallel and
posteriorly directed. And this form of pelvis (opisthopubic) was found in two only distantly
related saurischians. One group was the segnosaurs (Barsbold & Perle 1980), which resemble
prosauropods (Fig. 21C). The other was the genus Adasaurus, a small dromaeosaurid theropod
(Fig. 21D), closely related to theropods thought to be well known from North America. These
discoveries led to revision of the standard definition of saurischians. Gauthier (1986) proposed
several new defining features: among them having the jaw musculature attach (in part) to the
dorsal face of the frontals, long posterior cervicals, a distinctly asymmetrical forefoot, and an

enlarged claw on the pollex (thumb).

Figure 21. Dinosaur pelves. The top row represents the two classical pelvic types. A, omithischian, as
shown by /guanodon; B, saurischian, as shown by Allosaurus. The bottom row shows two of the newly
discovered Mongolian saurischians with pelves that approach the omithischian in form. C, a Segnosaurus,
D, a small theropod, Adasaurus.

eR

Many workers now accept the contention of Bakker & Galton (1974) that the dinosaurs are
a real (i.e. monophyletic) group. Dinosaurs, like Gaul are divided into three parts,
saurischians, ornithischians and herrerasaurs (previously considered saurischians). Saurischians
come in two varieties, theropods (the carnivores, by and large) and sauropodomorphs (the
giants, by and large).

Anatomy

Saurischian dinosaurs show two basic body forms or bauplans, that of the theropods and

FOSSIL REPTILES IN AUSTRALIA - 643

that of the sauropodomorphs. Sauropodomorphs all have relatively small skulls and relatively
long necks. They have four or five digits in the hindfoot and leaf-shaped or peg-like teeth.
Theropods usually have large skulls on short necks (but may have small skulls on long necks).
They have three-toed hindfeet and either flattened, knife-blade-like teeth or no teeth at all. Most
of the features that characterize theropods, and distinguish them from sauropodomorphs, are
obscure. However, two more obvious features are the greater development of the anterior
moeity of the ilium (Fig. 21B) and the absence of neural spines and transverse processes from
the distal portion of the tail in theropods.

Evolution and Taxonomy

Another "feature" shared by theropods and sauropodomorphs (see Appendix I-6) is that their
taxonomy and evolution are poorly understood. Both increased in size during their evolution,
although sauropodomorphs reached their maximum size early in the Cretaceous, while
theropods were largest late in the Cretaceous. The sauropod form was established early in the
Jurassic and remained until their extinction. The theropods, on the other hand, continued to
develop new forms right up until their extinction. Oddly enough, the most derived sauropods
seem not to have been the latest. In fact, the most derived sauropods seem to have been those
of the Late Jurassic. They were replaced by seemingly more plesiomorphic sauropods (the
titanosaurids) in the Cretaceous, especially in Gondwanaland. Further details on theropods and
sauropodomorphs may be found in Norman (1985).

The earliest dinosaurs are the herrerasaurs, from the Triassic of Argentina and Brazil.
Unfortunately, little has yet been published on these animals, although it has been suggested
(Bakker & Galton 1974) that they are ancestral not only to later saurischians, but to
ornithischians as well. Their occurrence in South America suggests that they may also have
occurred in Australia, and more importantly that dinosaurs mzy have originated in
Gondwanaland.

Palaeobiology

Three major groups of saurischians are known, prosauropods, sauropods (both
sauropodomorphs) and theropods. The prosauropods were small to moderately large herbivores,
probably chiefly quadrupedal, of the Triassic. They survived into the earliest Jurassic, but soon
thereafter became extinct. They have been found on all continents except Antarctica. The
segnosaurs are thought to be relatives of prosauropods that survived into the Late Cretaceous
in east Asia and North America.

Sauropods, also found everywhere except Antarctica, first developed during the Early
Jurassic and survived until the end of the Cretaceous. They seem most diverse, or at least best
known, from the Late Jurassic. They are often considered to be amphibious, although there are
no convincing reasons for this opinion. Trackways show they were capable of travelling on
dry land, but others also show that they spent at least some time in the water, and could swim.
Stomach contents of one individual (Stokes 1964) indicate an eclectic diet of plant material, and
likely dead animal material as well. Early sauropods may have been carnivores (Raath 1972)
although the evidence for this is less than convincing. Development of large cavities in the
vertebrae (pleurocoels) seems to have been progressive, later forms having such large
pleurocoels as to convert the vertebral column functionally into an I-beam. Earlier ones had
only slightly developed pleurocoels. Sauropods are among the largest of the dinosaurs, and
the largest individuals were considerably larger than the largest whales, reaching maybe 45 min
length (Dutuit & Quazzou 1980, Anderson 1987). More information on sauropod life styles

may be found in Coombs (1975).

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Figure 22. Tibia (A-D, F) and claw (E) of Agrosaurus macgillivrayi, presumably from Queensland. A, lateral
view; B, dorsal view; C, cross section of shaft; D, distal view; F, anterior view. (From Seeley 1891).

Theropods appeared in the Triassic and survived until the end of the Cretaceous, and again
have been found on all continents except Antarctica, even occurring in New Zealand. They
developed too great a variety of forms for any easy, brief description. Some had sabre-like
shearing teeth, others no teeth at all; some were ponderous and large, others small and graceful;
some had sails along their backs, or horns or crests of various forms on their heads, and others
did not. They were bipedal (so far as we know) and probably mostly carnivorous. It was
usually assumed without evidence that the large carnivores did not venture into water, but it is
now accepted that they probably could swim well (Coombs 1980).

The Australian Forms

Prosauropods, sauropods and theropods all have been found in Australia. A single
fragmentary specimen of a prosauropod, Agrosaurus macgillivrayi (Fig. 22), was collected
during the voyage of the H1MS Fly (Sceley 1891). It is presumably from Triassic beds

FOSSIL REPTILES IN AUSTRALIA - 645

somewhere on the northeastern coast of Australia, probably at the very tip of the Cape York
Peninsula. Little is known of this beast, but the tibia suggests a small form much like

Thecodontosaurus" minor (probably Massospondylus) of South Africa. It indicates the
presence in Australia of prosauropeds like those found elsewhere.

Both Jurassic and Cretaceous sauropods have turned up in Australia. The Middle Jurassic
Rhoetosaurus brownei (Fig. 23), from southeastern Queensland, is one of the earliest sauropods
(Longman 1926, 1927). It is known largely from the hindquarters and tail and is currently
under study by Dr R.A. Thulborn. It was a very peculiar beast, about 12 m long.

Figure 23. The right hind foot (A) and a dorsal vertebra (B) of Rhoetosaurus brownei. The presumed extent
of the claws is shown by the dashed lines. (Courtesy of L. Beirne & M. Wade).

A second sauropod, Austrosaurus mckillopi, is represented by dorsals and ribs from the
Allaru Mudstone near Maxwellton in northern Queensland (Longman 1933). Although from
the late Early Cretaceous, the dorsals are primitive and resemble those of Middle Jurassic
forms from overseas. Pleurocoels are very rudimentary, and the centra are composed of spongy
bone. Further specimens, probably Austrosaurus sp., come from the slightly younger Winton
Formation, near Winton, central Queensland (Coombs & Molnar 1981). This material is
somewhat more complete and shows an elongate metacarpus, which in turn suggests a
relatively long forelimb, and simple caudals. Again the resemblance is to Middle and Upper
Jurassic sauropods from overseas. The posterior portion of the cervical of the large sauropod
was found near Hughenden (Fig. 24). This closely resembles the corresponding bone of the
gigantic Brachiosaurus brancai of east Africa and suggests a 20 metre long animal. However,
as little of the neck of Austrosaurus is known, this cervical may come from Austrosaurus,

rather than from a brachiosaur.

646 - MOLNAR

Figure 24. Incomplete cervical vertebra of the large "Hughenden sauropod" (QM F6142) in lateral (A) and
posterior (B) views. Diagonal lines indicate broken surfaces, and patterning indicates matrix still present on
the specimen.

Theropod material from Australia is generally scrappy, much less complete than the
sauropod material: usually only single bones. One of these from the Early Cretaceous
Strzelecki Group near Wonthaggi, Victoria, by a happy coincidence tured out to be an
astragalus. Now while it is often believed (by non-palacontologists) that a palaeontologist can
identify, and provide sundry other information about a fossil animal from a single bone, this is
very rarely the case. In this instance, however, it was true, due to a comprehensive study by
Welles & Long (1974) of the astragali of theropods. Astragali turned out to be diagnostic
elements of theropod skeletons, and hence the Victorian astragalus could confidently be
identified as Allosaurus sp. (Molnar et al. 1981). Allosaurus is a well known theropod from
North America and east Africa. The Victorian Allosaurus represents either a juvenile or small
individual, no more than about 6 m long. Several other small theropod bones are now known,
but unstudied, from the Early Cretaceous Otway and Strzelecki groups of southern Victoria
(Rich & Rich, 1989).

Theropod material has also turned up at Lightning Ridge, New South Wales, particularly
vertebral centra. One of these has been named Walgettosuchus woodwardi (v. Huene 1932), but
is unfortunately so incomplete that it could belong to any of three theropod families (including
the Allosauridae). A single, unusual metacarpal from Lightning Ridge forms the type material
of Rapator ornitholestoides (v. Huene 1932). This metacarpal, about the size of that of a small
adult Allosaurus possesses an elongate posteromedial process not found elsewhere (Fig. 25).
A much smaller process is found in the smaller North American Ornitholestes hence the
specific name "ornitholestoides" . Thus, Rapator seems to be a distinct and valid, though not
well known, form.

A third fragmentary theropod is Kakuru kujani from the Maree Formation at Andamooka,
South Australia. It is known from a single, almost complete, but crushed, tibia (Molnar &
Pledge 1980). Kakuru was a relatively small theropod with very slender proportions (to judge

FOSSIL REPTILES IN AUSTRALIA - 647

from the tibia) matching those of herons and cranes, and recently discovered very bird-like
dinosaurs from Mongolia (Avimimus and Borogovia). The distal end of the tibia retains the
facet where the astragalus attached so that the form of the astragalus, usually well-reflected by
this facet, is known. Again, thanks to Welles & Long (1974), it was possible to determine
that this astragalus differed significantly from that of any other theropod (but one, and that yet
to be studied) in that it was very high but did not extend across the entire width of the tibia.
Thus, Kakuru seems not closely related to any known forms from overseas.

Figure 25. First metacarpals of theropod dinosaurs, shown in dorsal and ventral views. A, B, Rapator
ornitholestoides (New South Wales); C, D, Struthiomimus altus (Alberta, Canada), E, F, Deinocheirus
mirificus (Mongolia): G, H, Allosaurus fragilis (western U. S. A.). I, Therizinosaurus cheloniformis, dorsal
view only (Mongolia). The metacarpal of Raptor differs from all of these by having the strong posteromedial

process (bar).

Finally, the cervical vertebra (Pl. 6) of what seems to be a very small theropod has been
found in the Early Cretaceous Toolebuc Formation of western Queensland.

Palaeozoogeography

The fragmentary remains of Agrosaurus provide little information, but suggest that this

648 - MOLNAR

Australian prosauropod resembled those from other lands. Rhoetosaurus is still to be fully
studied, but Austrosaurus seems to resemble forms (cetiosaurs) from Europe, Africa and South
America that are much older than it. Australian theropod remains show that both large
(Allosaurus and Rapator) and small (Kakuru) animals were present. The latter two seem to
represent endemic forms of dubious relationship to those known elsewhere, while Allosaurus is
quite similar to overseas forms, but survived longer in Australia. This suggests some isolation
of Australian terrestrial tetrapods in the Cretaceous. Of the four Cretaceous saurischian genera
known in Australia, two seem endemic and two survived later than their known overseas
relatives.

ORNITHISCHIANS: PLANT-EATING DINOSAURS

Ornithischians include the remainder of the dinosaurs, the horned, armoured, plated and
duck-billed dinosaurs, among others. All were herbivores of small to moderately large size.
All ornithischians have a single, median bone at the front of the lower jaws, anterior to the
dentaries. This is the predentary, and ornithischians were once known as predentates. Among
the other features characterizing ornithischians are: a jaw joint depressed below the level of the
toothrow, at least one ossified cartilage (palpebral) in the eyelid, reduced antorbital fenestra
(even, in some cases, almost lost) and the fifth digit of the foot reduced to a metatarsal only
(Gauthier 1986).

Anatomy

Ornithischian dinosaurs come in a wide variety of forms, and any attempt to describe their
anatomy must be either short and abstract or long and detailed. We shall follow the first course
here, and those interested in detail should consult Norman (1985). Ornithischians have an
anterior projection of the pubis, known as the prepubis (Fig. 21A). This is often well-
developed, but may be short as in ankylosaurs. The crowns of the cheek teeth of ornithischians
are usually low, with the edges lined by denticles (Pl. 9I-L). These teeth often have a bulbous
base (cingulum) and are basically triangular in profile. In other features ornithischians, like
other successful vertebrate groups, vary widely. Some were bipedal, others quadrupedal, some
were armoured, others were not, some had shearing teeth, others grinding teeth.

Evolution and Taxonomy

As with saurischians, the evolution of ornithischians is poorly understood. Four to seven
(depending upon the authority) major groups of ornithischian dinosaurs are recognized. Only
those found (ornithopods and ankylosaurs) or likely to have occurred (fabrosaurs,
heterodontosaurs, pachycephalosaurs and stegosaurs) in Australia will be discussed. The others
(ceratopsians and scelidosaurs) seem restricted to Laurasia.

Ornithischians appear late in the Triassic. One Early Jurassic ornithischian, the South
African Geranosaurus, has been known for over sixty years (Broom 1911). However, it was
overlooked until the discovery of new material in the 1960s. This was both because of its
peculiar and unexpected dentition and the lack of any further specimens. Predictably, once new
material was recognized from South Africa, more types promptly turned up elsewhere (Canada,
China and Argentina). It also developed that other Early Jurassic ornithischians had been found
in the interim, and not recognized.

Much of the difficulty arose from the dentition. Instead of uniform teeth, as in later
ornithischians, many of the Early Jurassic species had a mammal-like dentition. This included
incisor-like, canine-like and molar-like teeth. This explains how one Early Jurassic
ornithischian was mistaken for a therapsid.

FOSSIL REPTILES IN AUSTRALIA - 649

The Early Jurassic ornithischians are often considered to belong to two groups, fabrosaurs
and heterodontosaurs, although neither may be a real group in the cladistic sense.
Heterodontosaurs had the mammal-like dentition, and developed cheeks. Fabrosaurs had a
simpler dentition, and little indication of cheeks. While most experts consider both of these
groups to be ornithopods, the matter is not settled to everyone's satisfaction, and the question
will be avoided here.

Ornithopods appeared in the Middle Jurassic and, in addition to probably giving rise to most
of the other ornithischian groups, developed in two main directions: towards small gracile
running herbivores (hypsilophodonts) or large to very large, bulky quadrupeds (iguanodonts
and hadrosaurs) with a recurrent tendency for cranial elaboration (horns, crests, etc.). The
hypsilophodonts appear in mid-Jurassic and last through the Cretaceous. Similarly, the
iguanodonts also appear in the Jurassic and last out the Cretaceous, evolving into hadrosaurs,
with impressive grinding batteries of teeth, late in the Early Cretaceous, at about the time that
angiosperms began their successful radiation. Hadrosaurs, often called duck-billed dinosaurs
from the shape of their bills, are usually considered amphibious, although there is good
evidence that some were not (Galton 1970). There is also evidence that some of the large
species (and they reportedly reached lengths in excess of 20 metres - Morris 1970) were indeed
aquatic (Morris 1981).

Pachycephalosaurs seem to have evolved from hypsilophodonts. They developed a great
thickening of the skull roof presumably used in clashes between males, as with modern sheep
and goats. In the Late Cretaceous they spread through Asia and North America. A single
isolated specimen from Madagascar suggests that they were more widespread than has been
realized (Sues & Taquet 1979) and may have occurred in other parts of Gondwana.

Until recently, ankylosaurs were considered the major Cretaceous armoured dinosaurs, while
the stegosaurs and scelidosaurs, were the armoured dinosaurs of the Jurassic. However,
ankylosaurs could not have appeared fully developed precisely at the end of the Jurassic.
Recent discoveries of Jurassic ankylosaurs (Galton 1980) have shown that they were present
through most of the Jurassic, but were simply not prominent in the known faunas. The first
dinosaur found in the Antarctic was an ankylosaur.

Stegosaurs have also been confirmed at a time when they were thought not to have existed.
Presumably having evolved from a heterodontosaur early in the Jurassic, they reached their
greatest diversity in the Late Jurassic and survived into the Early Cretaceous in Europe, Africa
and China. In India (which during the Cretaceous made its way across the Indian Ocean isolated
from other lands) small (almost pygmy) stegosaurs reportedly survived until the end of the
Cretaceous (Yadagiri & Ayyasami 1979), long after they had become extinct elsewhere.
Appropriately for such storybook-dragon-like creatures, stegosaurs were most abundant and
diverse in China. Their occurrence in Africa and India suggests that they may have lived
elsewhere in Gondwanaland.

Palaeobiology

Fabrosaurs and heterodontosaurs were small, bipedal ornithischians. The primitive
fabrosaurs did not possess cheeks and hence probably fed much like modern herbivorous lizards.
The heterodontosaurs, however, did have cheeks, which allowed them to masticate their food
much more efficiently, as checks and tongue could be used to hold the food between the
toothrows of both jaws. This allowed more effective grinding. Heterodontosaurs also had
canine-like tusks, apparently present only in the males, and hence presumably used in courtship
combat or display.

Omithopods were herbivores, the smaller ones bipedal and the larger at least facultatively
bipedal. The later groups (iguanodonts and hadrosaurs) developed great batteries of grinding
teeth, composed of several adjacent rows in hadrosaurs. In later ornithopods, the maxilla was

650 - MOLNAR

passively rotated about a longitudinal axis at its dorsal margin during chewing. This system,
termed pleurokinesis, increased the efficiency of mastication and may have been responsible for
the great proliferation of ornithopods (Weishampel 1984).

Pachycephalosaurs, except for their thickened skulls, seem much like the more primitive of
the ornithopods.

Stegosaurs also retain more primitive jaw and tooth structure. They specialized in the
development of bony plates and spikes held erect along the back. Although the spikes bore at
the distal end of the tail may well have been defensive, it seems likely that the plates along the
back were not. Farlow et al. (1977) have suggested that the plates functioned in absorbing and
dissipating heat.

Ankylosaurs were also armoured dinosaurs, but had scutes within the skin much like those
of crocodiles, rather than erect plates like stegosaurs. Rather little is known of ankylosaur
palaeobiology, probably because although their scutes are fairly common, well-preserved and
reasonably complete skeletons are not.

The Australian Forms

Few ornithischians are known from Australia, but one of them (an iguanodont) is the most
completely known Australian dinosaur. So far, only ornithopods and ankylosaurs are known,
but the distribution of the others (with the exception of ceratopsians and scelidosaurs) is such
that any of them may be expected to be found here. Hypsilophodontid material is known from
New South Wales (Molnar 1980b) and Victoria (Flannery & Rich 1981, Rich & Rich, 1989)
(Pl. 7-10). So far, only isolated elements (often femora) and two partial skeletons are known
but they indicate forms generally similar to those from overseas. A single incomplete femur
from Lightning Ridge, the type of Fulgurotherium australe, was originally interpreted as a
theropod (v. Huene 1932). Six more femora referred to Fulgurotherium (Molnar & Galton
1986) are similar to those of Hypsilophodon and Othnielia (Fig. 26). A seventh incomplete
femur more closely resembles that of Hypsilophodon and represents a distinct form. A single
tooth from the Ridge likely pertains to the animal from which most of these femora derive (PI.
11) and other material, including parts of skulls, one associated with a partial skeleton, has
been found in the Early Cretaceous rocks along the south coast of Victoria.

This material represents at least two genera, Atlascopcosaurus and Leaellynasaura, in
addition to Fulgurotherium. A juvenile specimen of Leaellynasaura includes both the skull and
the cast of the brain (Rich & Rich 1989). It was a large-eyed, large-brained animal at least
when compared to other ornithopods. Although the relatively large size of the brain may
reflect its youth, the large eyes suggest accommodation to the conditions of winter darkness of
the south polar regions. Victoria, after all, was within the Antarctic Circle of the Early
Cretaceous. The femur of Leaellynasaura shows some primitive features, recalling those of the
fabrosaurids (Rich & Rich 1989). Atlascopcosaurus is known from less complete material, and
differs from leaellynasaurs in having more ridges on the cheek teeth. Femora from Victoria are
known from Leaellynasaura and Fulgurotherium as well as two other forms, one of which may
be Atlascopcosaurus.

The iguanodont, Muttaburrasaurus langdoni, comes from the marine Early Cretaceous
Mackunda Formation of central Queensland (Bartholomai & Molnar 1981). It is represented by
a reasonably complete skeleton lacking much of the tail (Figs. 27-32). Much like Iguanodon,
except for the skull, and it is a massive form about 8m long. Muttaburrasaurus seems most
like the robust iguanodonts in its proportions, and has similarities to Camptosaurus in cranial
proportions. Probably quadrupedal, it is characterized by a broad, low skull with a remarkable
expanded hollow chamber on the snout. Unlike other ornithopods, Muttaburrasaurus had a
shearing, not grinding dentition, roughly like that of ceratopsians. The function of the snout
chamber, associated with the nares, is not known, but either resonance, for calls or enhance-

FOSSIL REPTILES IN AUSTRALIA - 651

Figure 26. The femur of Fulgurotherium australe from Lightning Ridge, New South Wales. This
reconstruction is based on two specimens. (From Molnar & Galton 1986).

ment of smell is likely. Like the European Iguanodon, Muttaburrasaurus seemingly developed
a thumb spike, but in the Queensland form the spike is flattened (as in only some European
species) and rather large. Like Camptosaurus, Muttaburrasaurus retained four large metatarsals
in the foot. Muttaburrasaurus has been found from Hughenden, Queensland, in the north to
Lightning Ridge, New South Wales, in the south.

In spite of an early report of armoured dinosaurs from near Darwin, Northern Territory
(which turned out to be fossilized bivalve shells), the only ankylosaur remains are from a small
animal found near Roma, Queensland. This individual of Minmi paravertebra is known from
part of the rib cage and vertebral column, a foot and armour from the belly (Figs 33, 34)
(Molnar 1980a). It represents a small ankylosaur, probably no more than 4 m long, but likely
as much as 0.75 m broad. The material is just sufficient to show, in addition to a diagnostic
ankylosaurian character, the existence of unusual little bones (paravertebrae) that lay alongside
the neural spines of the dorsal vertebrae. These bones resemble, both in form and in their
relationship to the ribs and vertebral column, tendons and tendinous sheets (aponeuroses) found
in crocodiles (Fig.35) (Molnar & Frey 1987). In crocodiles these tendons serve to

652 - MOLNAR

Figure 27. Reconstruction of skull of Muttaburrasaurus langdoni. A, lateral view; B, posterior view; C,
cross-section through snout bulla; D, dorsal view. (From Bartholomai & Molnar 1981).

Figure 28. Composite of two dorsal vertebrae of Muttaburrasaurus. Centrum and prezygapophysis of
anterior dorsal with spine and postzygapophysis of posterior dorsal. A, anterior; and B, lateral, views; C,
cross-section through spine at level of bar. (From Bartholomai & Molnar 1981).

FOSSIL REPTILES IN AUSTRALIA - 653

Figure 29. Left humerus of Muttaburrasaurus. A, anterior, B, medial; and C, proximal views. (From
Bartholomai & Molnar 1981).

Figure 30. Right pelvis of Muttaburrasaurus. A, lateral view of elements as preserved; B, same view of
pelvis reconstructed; and C, medial view of ilium. (From Bartholomai & Molnar 1981).

654 - MOLNAR

Figure 31. Right femur of Muttaburrasaurus. A, femur as preserved, B to D reconstructed. A and B, medial;
C, posterior; D, distal views. (From Bartholomai & Molnar 1981).

AS CA

Figure 32. Tibia, fibula, astragalus and calcaneum of Muttaburrasaurus. A, as preserved; B to D
reconstructed. A, B, posterior views; C, lateral view; D, distal view. (From Bartholomai & Molnar, 1981).

FOSSIL REPTILES IN AUSTRALIA - 655

strengthen and support the back during the high walk, providing additional area for attachment
of the back muscles. In other archosaurs these attach to the neural spines. While most
quadrupedal archosaurs have elongate neural spines, those of crocodiles and ankylosaurs are
rather short, seemingly correlated with the presence of dorsal armour. The tendons and
aponeuroses in crocodiles attach to the dorsal armour, and are not ossified. In Minmi there is
no convincing evidence for dorsal armour, although it did have belly armour, and this may be
the reason for the ossification of the tendons and aponeuroses into paravertebrae.

Figure 33. Those portions of the skeleton represented in the type specimen of Minmi paravertebra
(shaded). The skeleton outline is based on Sauropelta . (From Molnar & Frey 1987).

B

Figure 34. Dorsal vertebra of Minmi paravertebrata in lateral (A) and posterior (B) views.

Minmi exemplifies the unusual character of Australian dinosaurs. Although clearly related
to other nodosaurid ankylosaurs found overseas, it is also unique in its possession of
paravertebrae. This both permits insights into the palaeobiology of these animals not available
from overseas specimens (which apparently never ossified these back tendons) and also (in the
same feature) shows the unique adaptations of some Australian forms.

The paravertebrae suggest that Minmi may have relied more on running to escape predators
than on its armour (Molnar & Frey 1987). Interestingly Minmi seems to have lived on a large
island, as what is now eastern Queensland was at that time completely separated by the inland
sea from the rest of Gondwanaland.

656 - MOLNAR

Figure 35. The paravertebrae of Minmi compared with the tendons of the back musculature of crocodiles.
A, C, Minmi dorsals with one paravertebra shown on the right; B, D, crocodilian dorsals (and scutes at left)
with a tendon of the dorsal trunk musculature (in black) on the right. The general similarity in form may be
seen. A, B, dorsal views; C, D, lateral views. (From Molnar & Frey 1987).

Another possible armoured dinosaur has been collected in Queensland near Hughenden.
Oddly, it seems represented by the same portions of the skeleton as that of Minmi. Another
Minmi specimen (PI. 19) has also just been found, which has the dorsal armour present, but

reduced.

FOSSIL REPTILES IN AUSTRALIA - 657
Palaeozoogeography

_ The hypsilophodont material from Australia shows no evidence (perhaps because of its
incompleteness) of any unusual Australian taxa, although there are three endemic genera. On
the other hand, Muttaburrasaurus is unusual and endemic. It most closely resembles
Camptosaurus of North America and Europe, but shows development of the snout into a
hollow chamber. Other iguanodonts, and their descendants, the hadrosaurs, show a variety of
cranial omament” but nothing like this. Muttaburrasaurus also has a shearing dentition,
unlike anything seen in other iguanodonts and hadrosaurs. This suggests that it was an
unusual, endemic form. Minmi is unlike overseas relatives in the possession of the
paravertebrae. Again, this is an unusual, endemic form.

PTEROSAURS: ARCHOSAURS OF THE AIR

Although pterosaurs are among the longest-known fossil vertebrates they also seem to
have provided the most surprises and been the least well understood.

Anatomy

Being flying forms, the characteristic features of pterosaurs are those concerned with flight.
The pterosaur wing is supported by a greatly elongate digit IV, and is thus different in structure
from the bird wing, which is supported only relatively little by the skeleton of the hand, and
from the bat wing which is supported by digits II, III, IV and V. The scapula and coracoid of
pterosaurs are fused and rod-like in form, somewhat like those of birds. The scapula articulates
with a fused set of dorsal vertebrae (the notarium), a condition found in no other tetrapod. This
presumably helped support the body weight by passive, rather than muscular, means.
Pterosaur bones are very highly pneumatic, the cavities probably having been occupied by air
sacs, as in birds, and helping to reduce the body weight of even the largest forms to a
minimum. A massive flight musculature, probably similar at least in function to that of birds,
is indicated by well-developed sternal keels and even better developed deltopectoral crests on the
humeri. The skull is very lightly built and usually has a long, slender snout.

Evolution and Taxonomy

Pterosaurs are classed into two groups, the rhamphorhynchoids and the pterodactyloids
(pterodactyls). This appears to be a division into primitive (rhamphorhynchoids) and advanced
forms (pterodactyloids). Rhamphorhynchoids tend to have teeth and long tails (and hence be
stable fliers), while the pterodactyls tend to be larger and lose their teeth. Pterodactyls have no
tails and were, thus, unstable, and hence more highly manoeuvrable, fliers (Smith 1952).
Rhamphorhynchoids are characterized by the absence of the advanced features of the
pterodactyls. The earliest pterosaurs are known from the Late Triassic of Italy (Wild 1978) and
were derived from unknown ancestors. Although Wild suspects that these ancestors were not
archosaurs, this has not gained general acceptance. Instead, it has been suggested that
pterosaurs are closely related to dinosaurs (Padian 1984). By the Jurassic, pterosaurs had
become widespread and diverse. Their diversity is hard to gauge, as most fossils come from
marine deposits and, thus, preferentially represent marine forms. Consider what would be
known of modern birds if only those dying at sea were available. We do know from some few
scattered remains that inland forms existed, such as the gigantic Titanopteryx and
Queizalcoatlus, the latter reaching 13 to 14 m in wing span.

658 - MOLNAR

Palaeobiology

Recent work on pterosaurs has done much to increase our understanding of their life and
habits (Wellnhofer 1978, and his earlier papers cited therein, Langston 1981). Computer
simulations of their flying and gliding abilitics (Bramwell & Whitford 1974, Stein 1975) have
suggested that various forms were either efficient flyers or very good gliders (comparable to
modern sailplanes). Working models have shown that some of the small forms may have been
very good fliers (v. Holst 1957, D. Attenborough documentary 1989, British Broadcasting
Corporation). Recent studies of the wing have shown that it was probably much more efficient
aerodynamically than was previously believed (Frey & Riess 1981, Padian 1979). The wing
membrane included parallel fibres (actinofibrills; Wellnhofer 1987), which would have given
the wing different physical properties from those of any living animals. The implications of
the fibrous wing with anisotropic properties (properties different in different directions) are not
understood. On the whole, there seems no good reason to think that pterosaurs were not
excellent in the air. Evidence for webbed feet in some forms, suggests that they could swim on
the surface like ducks or gulls (Wellnhofer 1978). It has been argued that pterosaurs, being
active fliers, probably generated sufficient body heat to maintain a constant body temperature
(Nopsca 1916, Desmond 1975), and this argument is supported by the existence of a coat of fur
in at least some primitive forms.

Most marine pterosaurs were long-beaked fish eaters (Pterodactylus and Rhamphorhynchus),
while others had a deep snout (Anurognathus) like those of puffins and auklets. Others
paralleled the great whales rather than birds, and these forms (Pterodaustro) had multitudes of
thin, baleen-like teeth and seemingly fed upon surface plankton. Still others (Gnathosaurus)
had long thin teeth that anteriorly formed a "spoon" reminiscent of that of the spoonbill. But
whatever their specialities, they all disappeared with the coming of the Tertiary.

The Australian Forms

In Australia, pterosaurs have been found only recently (1979) from the Early Cretaceous
Toolebuc Formation, near Boulia, west Queensland. This material represents a moderately
small (wingspan of about 2 to 4 m) marine fish-eater, much like Pteranodon and
Ornithocheirus (Molnar & Thulborn 1980). So far, little of the skeleton is known, but what
has been studied is almost unique in that the material is uncrushed and undistorted. (Fig. 36,
Pl. 12). Pterosaur material is almost always completely flattened, so that one has virtually
only a two-dimensional projection of the bones. The Queensland material shows that not only
the limb elements but also the vertebrae are hollow, virtually only a surface layer being
present, internally supported by struts much like those of bird and aircraft construction. The
acetabulum of the pelvis is directed laterally and not at all downward.

Isolated pterosaur elements have also been found in the Early Cretaceous Otway Group of
Victoria (Rich & Rich 1989).

Palaeozoogeography

As far as can be told from its scattered remains, the Queensland form is like those known
from overseas. This is just what would be expected as pterosaurs were likely excellent fliers,
and some taxa (Pteranodon) are reported to have had wide distributions (North America,
England, Russia and Japan). }

Pterosaurs have recently been found in New Zealand, which was virtually part of Antarctica
in the Late Cretaceous. This indicates that even shortly before they became extinct, pterosaurs
were capable of surviving in cool polar regions.

FOSSIL REPTILES IN AUSTRALIA - 659

Figure 36. _ The pelvic girdle of a pterosaur from the Early Cretaceous of western Queensland, in lateral
view. Specimen at left, diagrammatic reconstruction at right. Abbreviations: a, acetabulum; il, ilium: is,
ischium; p, pubis. (From Molnar 1987).

TRACKS AND TRACKWAYS

While fossil skeletal remains of Mesozoic and older reptiles have been quite rare in
Australia, tracks and trackways have been more common. Most trackways derive from
dinosaurs, although there are unstudied reptilian trackways from the Triassic of Berowra Creck,
New South Wales (Fletcher 1948) that appear not to be dinosaurian. The best known and most
informative are those that apparently represent a dinosaur stampede.

There is quite a literature on reptilian (and other) tracks. Much of it is restricted to
morphology (Lull 1953, Kuhn 1958, Haubold 1971), although some embodies a more
profound approach (Abel 1935, Casamiquela 1964). While individual tracks often serve only
to confirm what one would have guessed anyway, valuable information may be derived from
the study of trackways. Trackways can provide information both on the dimensions and speed
of their makers, and the speed can in turn be used to investigate the metabolic rates of the
trackmakers. Trackways can also reveal the agility of the tackmakers. This information can
be used to provide insight into the habits and physiology of extinct animals, a topic difficult to
investigate in other ways.

Tracks are known from each of the three periods of the Mesozoic in Australia, but those of
the Triassic are limited to southeastern Queensland, with the exception of those from New
South Wales mentioned previously. A large theropod, much like Eubrontes of North America,
left its tracks at Rhondda Colliery, near Dinmore (Fig. 37P) (Bartholomai 1966a). The tracks
are about 46 cm in length with a stride of about 2 m, and suggest an animal about 6 m in
length. A second type of track, very similar to those of Plectropterna from North America, has
been found near Goodna (Fig. 370). It is not clear what kind of animal made these tracks, but
it may have been a rauisuchian. The tracks are 19 cm long and hence represent a smaller beast
than that recorded near Dinmore.

Recently tridactyl tracks have been discovered at two localities in the Callide Basin of
southeastern Queensland. These date from the end of the Triassic (Fig. 37Q). Some of the
tracks are striking in that the track impressions are dark brown on an almost white background.

More tracks are known from the Jurassic. The earliest are from the Lower Jurassic
Precipice Sandstone, of Carnarvon Gorge, Queensland. These are being studied by Tony

660 - MOLNAR

Thulborn and John Draper, who have identified them as ornithopod tracks. They were made by
small beasts, less than two metres long, and represent the oldest ornithischian dinosaurs known
from Australia. Most of the Jurassic dinosaur tracks, however, come from the coal measures of
southeastern Queensland (Fig. 37C-E, H-N). Many of these represent moderate to large
theropods, and one of them (Changpeipus bartholomaii) is similar to a contemporaneous track
from China (Fig. 37C) (Haubold 1971). They indicate the presence of large carnivorous
dinosaurs in Australia at this time. One colliery, however, Balgowan on the Darling Downs,
has yielded tracks of several different animals (Fig. 37D, E-F). In addition to the large
theropod, there was a small theropod (tracks about 12 cm long) and probably a four-toed
quadruped (Hill et al. 1966). It has been suggested this latter track is from a stegosaur, but any
other contemporaneous quadrupedal dinosaur (ankylosaur, sauropod, scelidosaur) could equally
well have left it.

The large theropod track from Balgowan (Fig. 37F) suggests an animal 10 m or more in
length, which ranks close to the largest known theropods. The abundance of theropod tracks in
what are coal swamp deposits seems somewhat unusual. There are more herbivores than
carnivores in any terrestrial food web, and the prey of these theropods must have been fairly
large. The only obvious suggestion is that the feet of the prey being broader than those of the
theropods, and possibly less loaded (the prey walking on four rather than two of them), made
less clear impressions. These might then have been easily overlooked (except at Balgowan).
The only Jurassic tracks to occur outside of the Darling Downs-Brisbane area have been found
at Mt Morgan, Queensland (Fig. 37H, I). These also are of a theropod, and are interesting in
that both fore and hind prints are preserved (Bartholomai 1966a). These tracks were originally
reported as Early Cretaceous in age, from which I (1980b) had concluded that this theropod was
an archaic one, as it reportedly retained five digits on the manus (most Late Jurassic forms have
only three). As it turned out, the theropod was not archaic, but the dating was incorrect and the
tracks are Early Jurassic in age.

Cretaceous tracks and trackways are rather more widespread, occurring in Western Australia
and Victoria as well as Queensland. The Western Australian trackways, Megalosauropus
broomensis (Fig. 37A), from a large theropod, come from the Early Cretaceous Broome
Sandstone of Gantheaume Point, near Broome (Colbert & Merrilees 1967). This form is quite
similar to tracks from Germany, Portugal and Texas, and demonstrates that not all Australian
saurischians were endemic or relict. The tracks suggest an animal about 6 to 7 m long. The
Victorian track (Fig. 37B), from near Knowledge Creek in the western Otway Ranges also
represents a theropod, but a smaller one (Flannery & Rich 1981). This track is only about 10
cm long, and somewhat like the tracks from the Callide Basin, the track impression is dark on
a light grey matrix.

Figure 37. Tracks of Australian archosaurs. A, Megalosauropus broomensis, Early Cretaceous, Ganthaume
Pt, Western Australia; B, small theropod, Early Cretaceous, Knowledge Creek, Victoria; C, Changpeipus
bartholomaii, Middle Jurassic, Westvale No. 5 Colliery, Queensland (QM F5702); D, small theropod, Middle
Jurassic, Balgowan Colliery, Queensland (QM F3278); EE, quadrupedal dinosaur, same source as D (QM
F5701); F, large theropod, same source as D (QM F3278); G, Changpeipus bartholomaii, same source as Cc
(QM F5700). C, G, come from the same trackway; H, I, large theropod, Early Jurassic, Mt. Morgan,
Queensland; J, theropod, Middle Jurassic, Lanefield Extended Colliery; K, L, M and N, theropods, Middle
Jurassic, Lanefield Colliery; M, seemingly consists of two superimposed tracks; O, Plectropterna, Late
Triassic, Goodna, Queensland; P, large theropod, Late Triassic, Rhondda Colliery, Queensland; Q, theropod,
Latest Triassic, Callide Basin (this drawing is based on a photo taken at a slight angle); R, Tyrannosauropus
sp., Upper Cretaceous, Lark Quarry (Winton); S, Wintonopus latomorum, Upper Cretaceous, Lark Quarry; T,
Skartopus australis, Upper Cretaceous, Lark Quarry; the two small images below R, represent the tracks of
Skartopus and Wintonopus to scale with that of Tyrannosauropus. Other images not to scale. (A from Haubold
1971; B from Flannery & Rich 1981; H, I from Staines 1954; J-N from Ball 1934a, 1934b, 1946; R, T from
Thulbom & Wade 1984).

FOSSIL REPTILES IN AUSTRALIA - 661

“l yee ee

We yy WM

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The most impressive, spectacular, abundant and significant tracks, however, are those
discovered to the south of Winton, Queensland, by some of the local residents. These tracks
(Fig. 37R-T) were first brought to scientific attention in 1971, later excavated under the
direction of Mary Wade, and studied by R.A. Thulborn and Mary Wade. The results (Thulborn
& Wade 1979, 1984) suggest that most of the tracks (which number into the thousands of
individual prints) were made during a stampede of small ornithopods and theropods, frightened
by a large theropod, whose tracks (Fig. 37R) are overlain by those of the stampede. These
tracks are from the Winton Formation and, oddly, represent only forms not represented by

662 - MOLNAR

skeletal remains (yet), while there are no tracks (yet) from any of the Winton Formation
sauropods, which are represented by skeletal material. The Winton Formation seems to be
earliest Late Cretaceous (Cenomanian) in age, and the footprint locality appears to have been
deposited over a point bar, and hence relatively near a river or waterhole. The tracks have been
impressed into a thin layer of mud overlaying sand. The scenario envisioned by Thulborn &
Wade (1984) is that a solitary large theropod approached across the mudflats from the northeast,
leaving a line of prints. Large numbers of small theropods and ornithopods had congregated in
the vicinity of the river or waterhole somewhere to the south of the locality. Whether these or
the water itself attracted the large beast is not known. This beast for some reason made an
abrupt turn to the right. Very shortly thereafter a large number of small dinosaurs (at least 150
individuals) dashed across the mud and left their tracks over those of the large theropod.
There is no evidence that any of these animals had crossed the mudflat before the coming of
the large theropod. This suggests that there had been another route to the area of their
congregation now blocked, presumably by the large theropod.

This scenario was reconstructed from the evidence afforded by the trackways. These
indicated that the large theropod's tracks were overlain by the others and all of the smaller
animals’ trackways headed in a single direction (about 55° east of north). There is no
indication of any prolonged period of track impression. The mud presumably dried fairly
rapidly, and all of the tracks were impressed during the same state of the mud. The trackways
of the small dinosaurs were all straight, or slightly zig-zag where the beasts apparently careened
together or swerved to avoid one another. There are no tail drags and no impressions of hands,
and most (99%) of the foot impressions have the imprints of the toes only.

In addition to this interpretation of the dinosaurian behaviour resulting in the Winton
trackways, other less spectacular but more general and hence significant results were reached.
The first concerns estimating the size of a dinosaur from that of its footprint - in the fashion of
Sherlock Holmes. The size of an animal may be most reliably estimated not from the length,
or any other single measurement of the footprint, but from the "size index". The size index is
the square root of the ratio of the width of the print divided by its length.

The nature of some of the footprints also allowed estimation of the height of the hip above
the ground. Together with the "stride length", the distance between successive tracks of the
same foot, this can be used to estimate speed (details given in Thulborn & Wade 1984). These
give speeds of 16 km/h for the omithopods and 12 km/h for the small theropods (the large
theropod was walking at only 7 km/h). Both ornithopods and small theropods were moving at
what is termed a "physiologically similar" speed, presumably as fast as they could go across
soft mud!

This work exemplifies how trackways can be used to indicate physiological properties and
behaviour of extinct animals.

The most recently discovered (1988) dinosaur tracks in Australia (Fig. 38) were found in the
ceiling of an opal mine at Lightning Ridge. At least two kinds of animals seem to be
represented : from an examination of snapshots Tony Thulborn feels that at least one of these
was an ornithopod.

SUMMARY

The record of Mesozoic reptiles in Australia is obviously poor. The Mesozoic lasted for
180 million years. Australian fossil reptiles are known from only about 30 of these. And that
is a very liberal estimate of the period of time for which we have some representation of
terrestrial Mesozoic reptiles. Even looking over the whole world, great gaps in the record of
terrestrial reptiles remain. In the Jurassic there is a gap of some 30 million years (almost half
the length of the Cainozoic) from which only a few bones of terrestrial animals are known,
while similar, but shorter, gaps exist in the Triassic and Cretaceous. It is important, especially

FOSSIL REPTILES IN AUSTRALIA - 663

in Australia where gaps are so prominent and extensive, to reiterate their existence.

a
=

Figure 38. A large omithopod(?) track from Lightning Ridge, New South Wales. These tracks are in the
ceiling of a mine, and this track is drawn obliquely from below.

The Triassic reptiles known from Australia derive from the Scythian stage at the very
beginning of the Triassic. No other Triassic forms are confidently known from skeletal
remains, although Agrosaurus presumably derives from late in the Triassic. All of the
Scythian reptiles known - prolacertiforms, procolophonids, thecodonts and dicynodonts - are
closely related to those known from overseas, especially from South America, South Africa and
Antarctica. Thus, in the Early Triassic there is no reason to suggest any significant variation
in terrestrial reptile faunas. However, looking only at the reptilian faunas can be misleading.
Thulbomn (1986b) has considered the entire tetrapod faunas from Australia and found that they
do indeed differ from those elsewhere in the world. They are about 90% temnospondyl
amphibians, which make up less than 20% of the non-Australian Scythian faunas. This, he
suggests, might be due to some isolation of what is now Australia, perhaps due to barriers to
migration in the Antarctic region, or to some consequence of the peninsula effect.

Rhoetosaurus, from the early Middle Jurassic, is poorly known and under study. It does not
seem different from sauropods known from overseas. The J urassic plesiosaurs have yet to be
studied.

The marine Cretaceous forms, from the Aptian and (mostly) Albian stages, are similar to
those from overseas. There are endemic genera (Notochelone and Woolungasaurus) that are
reasonably well known, but they differ little from similar forms overseas. Terrestrial
Cretaceous reptiles are also generally similar to those from elsewhere. There is as yet little
evidence for endemic families. Some endemic genera (Minmi, Muttaburrasaurus) show unique
structures not found in their relatives known from overseas, while others (Allosaurus,
Austrosaurus) are seemingly archaic compared to contemporaneous forms from overseas. The
greatest similarity seems to be to those from South America. Thus, by the Cretaceous,
terrestrial reptiles suggest a distinct Australian fauna, related to, and, possibly derived from,
that of South America. ,

The unusual nature of the Cretaceous terrestrial tetrapods of Australia may be related to
Australia's geographic location. During the Early Cretaceous Australia lay astride the Antarctic
Circle. The Victorian dinosaur localities were well south of the Circle, and Lightning Ridge
was on the Circle. Even the Queensland localities were further south than almost all other
Cretaceous tetrapod sites, except for the southernmost in South America. Thus, the difference

664 - MOLNAR

of the Australian dinosaur fauna from others of the same time may reflect its near-polar
location. Similarly, during the Late Cretaceous, New Zealand was south of the Antarctic
Circle, so that its vertebrates of this age also reflect near-polar faunas. Elsewhere such faunas
are known only from Alaska and northern Canada (Davies 1987, Brouwers et al. 1987) and
from some tracks and trackways in Spitsbergen (de Lapparent 1962, Edwards et al. 1978). The
Alaskan material is Late Cretaceous and the Spitsbergen tracks Early Cretaceous.

Polar regions have been suggested as source areas of evolutionary innovations in both
marine (Zinsmeister & Feldmann 1984) and terrestrial animals (Hickey et al. 1983). The latter
suggestion is based on observations of the first appearance of tortoises and certain mammals
(e.g. perissodactyls) in Arctic North America (Hickey e¢ al. 1983). In North America these
forms later migrated south and established themselves over the continent. In Australia there
was nowhere further from the pole to which to migrate. Thus, if we had only the early Arctic
North American forms to compare with those of other continents, as we have only the near-
polar Australian Cretaceous tetrapods, the Arctic North American forms would likely appear
just as unusual.

However, Australian land-dwelling tetrapods also include relicts, not noted among the near-
polar North American beasts. A speculative explanation of this second faunal component
involves Vermeij's (1987) proposal that polar regions constitute "safe places" from predation.
This is not to argue that there are no polar predators, but to suggest that the most intense
predation occurs outside of the polar regions. This being so, the chance of any given species
being driven to extinction by intense predation is reduced in polar regions. Vermeij's work
involves only marine organisms, but the intensity of predation may have been reduced among
the south polar Cretaceous tetrapods as well, and this may account for the Australian relicts.
The absence of evidence for substantial armour in the dinosaur Minmi, whose overseas relatives
are all well-armoured, is consistent with this proposal - but other explanations are possible.
Specimens of large theropods are rare in Australia, which is also consistent with this proposal.
But again other explanations are possible - the Australian fauna is poorly known, so this could
be simply a sampling effect. Although other explanations for these observations are possible,
the observations are consistent with Vermcij's proposal, and so far no contradictory data are
available.

Very little is known of near-polar dinosaurs in general, and those from Victoria provide
important information. Most important is the conclusion that near-polar dinosaurs probably
existed throughout the Cretaceous. The most common dinosaurs of southem Victoria are small
hypsilophodontians, which are also represented in Late Cretaceous New Zealand. Interestingly,
the New Zealand form appears to be a relict dryosaurid, a family otherwise known from the
Late Jurassic and Early Cretaceous.

The Victorian and New Zealand hypsilophodontians, and even those from Lightning Ridge,
are small forms. The largest dinosaur in Victoria is the Allosaurus , and that was only about 6
m long, while the largest North American specimens were about 13 m long. At least two
other, smaller theropods were present also. Hypsilophodontians, the smallest of the
ornithopods, are the most common dinosaurs in Victoria and at Lightning Ridge. No very
large dinosaurs, such as sauropods, have been found in Victoria, and, while seemingly present,
they appear to have been rare at Lightning Ridge. Small size is unexpected in near-polar
tetrapods - most modern vertebrates are larger in colder climates than their relatives in warm
climates (Bergmann's Rule). In hypsilophodontians the reverse is true, with the larger forms,
such as Parksosaurus and Thescelosaurus, found in areas closer to the equator. Even among
the other dinosaurs represented, the near-polar ones seem to be smaller than their more tropical
relatives, at least those in the Southern Hemisphere.

The existence of a fauna of near-polar dinosaurs raises several interesting questions. What
was the climate? If cold, at least seasonally, were the dinosaurs endothermic? Did they spend
their entire lives at this location, which experienced at least three months of total darkness

FOSSIL REPTILES IN AUSTRALIA - 665

yearly? Many of these questions are treated by Rich & Rich (1989) and Rich et al. (1988) and
by Paul (1988), for northern dinosaurs. The question of climate is unresolved. There is much
evidence that it was at least cool, although probably not glacial at the poles as it is now.
Evidence for coolness includes oxygen isotope ratios, which suggest mean annual temperatures
between 8° C. and -4° C. (Rich et al. 1988), consistent with at least some of the Northern
Hemisphere data (Paul 1988). The geographic position of the poles implies that they were the
coolest Places on earth (at sea level). There is some evidence for ice-rafting of stones in the
Australian region (Frakes & Francis 1988).

On the other hand, there are complicating factors: the dinosaurs, for one. Ceratodontid
lungfish have been found in these deposits in Victoria: today they cannot breed in water colder

than 10° C. (Kemp 1981). Forests were present in the Antarctic Peninsula (e.g. Jefferson
1982): these could not have grown under present-day polar conditions. There is no evidence of
glaciation or cold mean annual temperatures in Antarctica (at sea level) until the Eocene
(Birkenmajer & Zastawniak 1989), although there has been unpublished speculation that
transient high-altitude glaciation did occur in Antarctica.

None of this evidence is definitive. Oxygen isotope temperatures may be "reset" by later
geological conditions, and even when not, might reflect groundwater temperatures possibly of
runoff from local high-altitude areas. The geographic position implies that the poles were the
coolest places, but doesn't say how cool. Evidence for transport of stones in masses of ice may
be "counterfeited" by rafting of stones by masses of floating vegetation. Similarly, earlier
claims for varves (Waldman 1971), that in turn indicated winter freezing, could be based on
similarly laminated sediments that may be deposited under much milder climates.

On the other hand, that modern ceratodont lungfish cannot survive near-freezing
temperatures does not rule out such tolerances in their ancestors. Most workers on dinosaurs
do not accept that they were endothermic (myself included), but everyone has been wrong one
time or another. Paul (1988) cites evidence for regular or periodic frosts and possibly even
riverine or coastal sea ice in the Cretaceous Arctic. He uses this as evidence that at least some
(Arctic) dinosaurs were endothermic. But, again, other explanations are possible. However, a
wide variety of marine reptiles are known from the Antarctic seas during the Late Cretaceous.
These include plesiosaurs, mosasaurs and even turtles (Wiffen 1981). No one has suggested
that these were endothermic animals, yet this would be consistent with arguments of the type
used by Paul. Perhaps the Antarctic seas were substantially warmer than the Antarctic land
(and note that here we refer to the Late Cretaceous, not the Early Cretaceous of the Victorian
dinosaurs and oxygen isotope temperatures). Or perhaps the Antarctic was just warmer than
some evidence suggests. This issue is clearly still open.

The question of dinosaur endothermy ("warm-blooded dinosaurs") is best treated by Thomas
& Olson (1980). Most workers regard dinosaur endothermy as unlikely, except perhaps in the
line leading to birds. Paul (1988) cites suggestive, but not conclusive, evidence that
hypsilophodontians may have had some form of insulation, like hair or feathers. However, the
evidence for dinosaur endothermy is inconclusive and remains as mere speculation.

Modern animals (especially birds and caribou) migrate north to the shores of the Arctic
Ocean to exploit rich seasonal food resources. Such food allows them to raise more young
than if they remained in their "winter quarters." The abundance of juvenile hypsilophodontian
material in Victoria has suggested to Rich et al. (1988) that the area was a “dinosaur nursery."
Perhaps dinosaurs also made seasonal migrations to exploit seasonal food resources? Perhaps,
but as Rich et al. (1988) point out even evergreen plants may have grown in this region. They
may not have grown well during the total darkness of mid-winter, but they would have made up
for this during the 24-hour-daylight of mid-summer. Paul (1988) suggests that although
endothermic dinosaurs may have been capable of such migrations, there are no compelling
reasons to posit these migrations. Rich et al. (1988) agree. The nearest regions north of
Victoria without 24-hour winter darkness were about 1000 km away, not impossible according

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to Paul's calculations. But, unless the juveniles grew very rapidly, this would be a strenuous
trek for them. Like the other questions discussed here, migration is speculative, and while
there is no compelling evidence against it, there is no conclusive evidence in its favour.

The dinosaurs of Victoria, New South Wales and New Zealand stimulate interesting
questions about the nature of the Cretaceous world, and the nature of dinosaurs themselves.
Further work on these animals, and further discoveries in these regions, promise to provide
important insights into the history of life on our planet.

Two conclusions may be drawn from the record of Mesozoic terrestrial vertebrates in
Australia. First, it is very incomplete. Of the entire Mesozoic, a period of time almost three
times as long as the Cainozoic (approx. 180 as opposed to approx. 65 million years), less than
one-seventh is represented in Australia. Hence, any conclusions derived from this record must
be very tentative.

Second, the Australian fauna appears different from those known elsewhere even when it is
first known (in the early Triassic). There is little indication of this difference from the
individual elements (taxa) of the fauna, but their proportions in the fauna are different
(Thulborn 1986b). Later (Cretaceous) forms show some differences - unique morphologies -
from their non-Australian relatives, but these relatives are still easily recognized. Unlike the
Cainozoic, when isolation was complete, the individual elements are obviously related to
overseas forms, but differences and distinctions occur in these elements and in their
proportional representation in the fauna.

CAINOZOIC RECORD

Although reptilian fossils date back to the Eocene in southeast Queensland, most Australian
Cainozoic reptilian material is Pleistocene in age. With few exceptions, this material more or
less certainly pertains to modern genera. Thus, there is little to be gained by discussing this
material that is not readily apparent from the stratigraphic table (Table 4). Instead I will
concentrate on the extinct forms.

Table 4: Stratigraphic Distribution of Australian Cainozoic Reptiles and Amphibians.

The data for this table are derived from Archer et al. (1989), Bartholomai (1977), Estes (1984),
Gaffney (1981), Gaffney & Bartholomai (1979), Gorter & Nicoll (1978), Hecht (1975), Hecht &
Archer (1977), Molnar (1982a), Pledge (1984), Smith (1976), Smith & Plane (1985), Tyler (1979),
and Woodburme (1967). Additional data are from the other contributions to the first edition of this
volume, especially those of Tyler (1982) and of Rich et al. (1982) and Rich et al., this volume. The
term 'crocodilian’ here excludes ziphodont crocodilians.

SOUTH AUSTRALIA
Miocene-Pliocene
Corra-Lynn Cave sediments (Curramulka Local Fauna)
Neobatrachus pictus
turtles
Wonambi sp.
elapid
scincid
Tiliqua sp.
Oligo-Miocene
Etadunna Formation (Ngapakaldi Local Fauna)
Australobatrachus ilius
Litoria sp. cf. L. caerulea

FOSSIL REPTILES IN AUSTRALIA - 667

Limnodynastes archeri
Emydura sp.
Meiolaniidae
Egernia sp.
Varanus sp.
Ophidia
crocodilian

Etadunna Formation (Ngama Local Fauna)
cf. Emydura sp.
Egernia sp.
crocodilian

Namba Formation (Tarkarooloo Local Fauna)
Emydura sp.
Meiolania sp.
cf. Egernia
crocodilian

Namba Formation(Y anda Local Fauna)
crocodilian

Namba Formation (Ericmas Local Fauna)
chelid
crocodilian

Namba Formation (Pinpa Local Fauna)
cf. Emydura sp.
meiolanid
crocodilian

Wipajiri Formation (Kutjamarpu Local Fauna)
Emydura sp.
Meiolania sp.
Egernia sp.
Tiliqua sp.
agamid
crocodilian

Pliocene

Mampuwordu Sands (Palankarinna Local Fauna)
Crocodylus
?sebecosuchian or pristichampsine

Pleistocene

unnamed beds at Lake Kanunka
Megalania prisca

Katapiri Sands (Kanunka Local Fauna)
?chelid
Megalania prisca
crocodilian

Naracoorte Cave deposits
Geocrinia sp. cf. G. laevis
Limnodynastes tasmaniensis
Limnodynastes sp. cf. L. dumerili
Litoria ewingi
Ranidella signifera
Amphibolurus sp. cf. A. barbatus
Egernia sp. cf. E. whitet
Notechis sp. cf. N. scutatus
Pseudechis sp. cf. P. porphyriacus
Pseudonaja sp. cf. P. nuchalis
?2Sphenomorphus sp.
Tiliqua nigrolutea

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Trachydosaurus rugosus
Varanus gouldii
V. varius
Wonambi naracoortensis
unnamed beds of the Warburton River
Megalania prisca
unnamed beds at Burra
frog

WESTERN AUSTRALIA
Pleistocene
cave deposits at Windjama Gorge
?Carettochelys sp.
crocodilian
Cave deposits at Devil's Lair and Skull Cave
Crinia georgiana
Heleoporus sp.
Platyplectron dorsalis
Litoria adelaidensis
Litoria sp.
Neobatrachus sp.
Ranidella sp.
TASMANIA
Oligocene or Miocene
unnamed formation at Taroona
Emydura macquarii

NORTHERN TERRITORY
Oligo-Miocene
Camfield Beds (Bullock Creek Local Fauna)
chelonian
crocodilian
Morelia antiquua
unnamed beds at Kangaroo Well (Kangaroo Well Local Fauna)
chelonian
crocodilian
Miocene-Pliocene
Waite Formation (Alcoota Local Fauna)
chelonian
"Crocodylus" sp.
cf. Pallimnarchus sp.

NEW SOUTH WALES
Pliocene
Gulgong Deep Lead
Emydura sp.
Meiolania sp.
unnamed beds at Bow (Bow Local Fauna)
chelonian
lizard
unnamed beds (Krui River Local Fauna)
crocodilian
Pliocene-Pleistocene
Blanchetown Clay (Bone Gulch Local Fauna)
chelid
Mooma Formation (Fisherman's Cliff Local Fauna)

FOSSIL REPTILES IN AUSTRALIA - 669

Emydura sp. cf. E. macquarii

unnamed beds at Oaky Creek
Meiolania sp.

unnamed beds at Cuddie Springs
Megalania prisca

Pleistocene

Wellington Cave deposits
Trachydosaurus rugosus
Varanus sp. cf. V. giganteus

beach deposits, Lord Howe Island
Meiolania platyceps

QUEENSLAND
Eocene? (Early Tertiary)
unnamed beds at Boat Mountain (Boat Mountain Local Fauna)
crocodilians
lizards
snakes
Redbank Plains Series
chelonians
crocodilian
Oxley Group
chelonian
crocodilian
Rundle Oil Shale
chelonian (chelid?)
crocodilian
Oligo-Miocene
Carl Creek Limestone (Riversleigh Local Fauna)
Crinia sp.
Kyarranus spp.
Lechriodus intergerivus
Limnodynastes spp.
Litoria spp.
chelids (Emydura/Elseya)
crocodilians
ziphodont crocodilians
Meiolania sp.
scincids
Physignathus sp.
varanids
gekkonids
Montypythonoides riversleighensis
?Rhamphotyphlops sp.
elapids
madtsoids
Pliocene
Chinchilla Sands
Emydura sp.
trionychid
meiolanid
Megalania sp.
Pallimnarchus pollens
?Quinkana sp.
Allingham Formation (Bluff Downs Local Fauna)
chelonian

670 - MOLNAR

Crocodylus porosus
agamid
Varanus sp.
?acrochordid
?Morelia sp.
elapid
unnamed beds at Tara Creek
Chelodina sp.
Crocodylus porosus (=C. nathani)
cave deposit (Rackham's Roost Local Fauna)
frogs
squamates
Pliocene-Pleistocene
unnamed beds at Floraville (Floraville Local Fauna)
chelid
Varanus sp.
snake
unnamed beds at Floraville Crossing (Floraville Local fauna)
trionychid
Pallimnarchus pollens
Dry River
Meiolania sp. cf. M. platyceps
Megalania sp.
?Crocodylus sp.
Pleistocene
Eastern Downs soil horizon
Meiclania owenii
chelonian
Chlamydosaurus kingi (= C. bennetti)
Megalania prisca
Varanus emeritus (? = V. salvadori)
Texas Cave breccia
frog
Quinkana sp.
?agamid
geckonid
?Amphibolurus
snake
Tea Tree Cave breccia
Quinkana sp.
Tiliqua sp.
Varanus sp.
snake
"Glen Garland" swamp deposits
chelonian
Crocodylus sp.
Quinkana sp.
Megalania sp.
Gore cave and fissure deposits
Tiliqua scincoides
"Alehvale" channel deposits
Crocodylus sp.
Quinkana sp.
unnamed beds at "Rosella Plains"
ziphodont crocodilian
unnamed beds at Emerald

FOSSIL REPTILES IN AUSTRALIA - 671

unnamed beds at Emerald
trichonychid

unnamed beds at Cape River
Megalania sp.

unnamed beds at Springsure
Megalania sp.

unnamed beds at Tambo
Pallimnarchus sp.
ziphodont crocodilian

unnamed river gravels at Riversleigh
chelids (Emydura/Elseya)

WESTERN AUSTRALIA
Late Cainozoic
unnamed clays at Jubilee Dam (Quanbun Local Fauna)
crocodylid

CROCODILIANS

Much Cainozoic crocodilian material is either fragmentary or unstudied. Since crocodiles
have been reported from the Eocene Redbank Plains Series, near Brisbane (Reik 1952) and
from the Early Cretaceous Griman Creek Formation, near Lightning Ridge (Molnar 1980c), it
is likely that the crocodilians have been in Australia through the Cainozoic. Although the
reported Eocene Redbank Plains material has never again been located, there is an Eocene
partial crocodile jaw from Eight-Mile Plains (a suburb of Brisbane). Only the symphysial
region is preserved, which suggests a narrow snout. An almost complete crocodilian jaw
(Pl. 13) from the Rundle Oil Shale deposits, is also probably Eocene. Oligo-Miocene material
from Lake Palankarinna, South Australia and Riversleigh Station, Queensland, is now under
study. Jaw fragments from Murgon, southeastern Queensland, represent crocodilians similar to
those from Lake Palankarinna. The material from Riversleigh indicates at least three, and
maybe as many as eight species - most of which were quite unlike that from Lake
Palankarinna. Of the two modern species the Fresh Water Crocodile, Crocodylus johnstoni,
was present in the Pleistocene (Willis & Archer 1990), while the Salt Water Crocodile (PI.
14), Crocodylus porosus, dates well back into the Pliocene (Molnar 1979).

Two extinct crocodiles, neither well represented, differ from the living crocodiles.
Pallimnarchus pollens (de Vis 1886) was a very large animal, about the size of an adult
Crocodylus porosus. It is represented largely by fragmentary material, although there is an
almost complete skull in the Mirani Shire Council Museum, near Townsville in Queensland.
Pallimnarchus is now being studied by P.M.A. Willis and myself. Its skull had a broad snout,
broader even than C.porosus, and was very flat with upwardly directed orbits. It resembles
skulls of the nettosuchian crocodiles or some of the larger temnospondyls. Preliminary results
indicate that a crocodile snout (Fig. 39) from Lansdowne, thought to be C. porosus (Molnar
1982b), is from Pallimnarchus.

Pallimnarchus inhabited the inland waterways of east Queensland during the Pliocene and
the Pleistocene, but only at one locality is there any suggestion that both Pallimnarchus and
Crocodylus may have lived together, and even here the evidence is not clear.

The second extinct crocodilian is a different kind of beast. It was a ziphodont crocodilian,
reaching a length of perhaps 3 m. Ziphodont crocodilians all share a suite of features of the
jaws and teeth. Originally they were thought to be a single, closely related group, although it
is now generally recognized that at least five separate lineages are involved. They all share

672 - MOLNAR

deep, usually laterally compressed snouts, armed with laterally compressed, serrate shearing
teeth. These teeth are much like those of carnivorous dinosaurs, and in some cases were
mistaken for dinosaur teeth.

Figure 39. The tips of the snouts of Crocodylus porosus (right) and Pallimnarchus pollens (left). The snout
of Pallimnarchus is the broader. Upper, dentaries; lower, upper jaws.

The Australian ziphodont, Quinkana fortirostrum (P|. 15), has a relatively broad, although
still deep, snout (Molnar 1981). It is known largely from cranial fragments ranging from an
almost complete snout to small pieces with a few teeth still in place. Isolated teeth have also
been found. The beast apparently ranged throughout eastern Queensland. Although the
maxillary teeth have long roots, accommodated in part in an alveolar process projecting below
the palate, the snout is more than deep enough to house them. This suggests that the depth of
the snout is related to strengthening the snout, rather than simply housing the teeth. The
orbits face laterally, not dorsolaterally as in modern crocodiles, and are placed high on the side
of the skull. Quinkana is not related to the well known South American ziphodont sebecoid
crocodiles (Colbert 1946) nor, apparently, to any of the other ziphodonts known overseas. A
fragment of maxilla with two teeth still in place, from the Pliocene Mampuwordu Sands of
South Australia described as an Australian sebecoid (Hecht & Archer 1977), may represent a
similar beast.

Quinkana was unusual in that it survived some ten million years later than overseas
ziphodonts. In the Northern Hemisphere ziphodonts became extinct by the end of the Miocene.
Only in Australia did they survive as late as the Pleistocene.

FOSSIL REPTILES IN AUSTRALIA - 673

Figure 40. Maxilla (A) and dentary (B) of the lizard Egernia sp., from the Etadunna Formation, South
Australia. This animal is very similar to (from the same genus as) lizards still living. Note that the maxilla
is seen in lateral view, but the dentary in medial view. (From Estes 1984).

SQUAMATANS

Although there is evidence of Mesozoic squamatans in Australia, little can be said of most
fossil forms older than Pleistocene. Most Pleistocene forms belong to genera still alive, such
as Amphibolurus (the dragons), Chlamydosaurus (the frilled lizard), Varanus (the goannas),
Trachydosaurus (the shingle-back), Notechis (the tiger snake) and Pseudonaja (the brown
snakes), among others (Archer 1978, Archer & Wade 1976, Smith 1976). Our knowledge of
Cainozoic squamatans in Australia can best be described as almost non-existent. With the
exception of the extinct forms shortly to be discussed, all we know is that the various living
forms must have existed in Australia for variable periods into the past. Agamids, varanids,
Egernia (Fig. 40), Physignathus and Tiliqua all are at least as old as the Miocene in Australia.

Only three extinct squamatan genera are known from the Cainozoic of Australia. Megalania
prisca was a large goanna, possibly as much as five metres long (Hecht 1975, Rich & Hall
1979). It inhabited at least the eastern half of the continent (although unknown in Tasmania),
and is represented mostly by vertebrae (Fig. 41), with a few parts of limbs and girdles and
skull. Isolated teeth have also been found. Recently more cranial bones have been found,
some representing previously unknown parts of the skull. When studied, these should allow a
new and more accurate reconstruction of the skull, They confirm that the cranial kinesis
characteristic of varanids was suppressed in Megalania prisca (Hecht 1975). The vertebrae of
Megalania are similar to those of living varanids, strongly concave anteriorly and convex
posteriorly with both articular surfaces inclined to the long axis of the centrum - but much
more massive. Hecht has suggested that both neck and tail were shortened, but in the absence
of any complete vertebral columns, this must remain but a suggestion. The humerus is quite
stout, much more so than in other varanids, even Varanus komodoensis. The pelvis was like
those of modern varanids. The phalanges of the feet are basically bird-like but may be

674 - MOLNAR

distinguished from those of birds in having a prominent rectangular flexor knob on the ventral
surface. The hindfoot (pes) was rather different from those of other varanids, especially in the
great expansion of the distal end of the fifth metatarsal. Unfortunately, no known fifth
metatarsal is complete, so that the extent and significance of this difference is unclear.

Megalania teeth are generally similar to the ziphodont teeth of Quinkana and like forms.
But whereas the ziphodont teeth are sharply edged and serrate both anteriorly and posteriorly,
Megalania teeth have a sharp, serrate edge posteriorly but a rounded edge anteriorly with
serrations only near the tip. The enamel is wrinkled into longitudinal grooves around the neck
of the tooth in Megalania.

A

Figure 41. A dorsal vertebra of the giant varanid Megalania prisca from eastern Australia, in anterior (A)
and lateral (B) views. (From Owen 1884).

A few vertebrae from Chinchilla, southeastern Queensland, indicate a second, earlier
(Pliocene) form of Megalania (Hecht 1975).

Not only were there giant lizards in Australia during the Pleistocene, but large snakes as
well, although probably no larger than the living Python amethystina (the Amythestine
Python). A large constrictor, Wonambi naracoortensis, known from vertebrae (Fig. 42) and a
portion of the maxilla, was probably about 5 m in length (Smith 1976). Its vertebrae are
similar to those of the living pythons but differ in having backwardly sloping neural spines,
paracotylar foramina and no accessory processes. In general, Wonambi shows resemblances to
a much older constrictor from South America and Africa, Madtsoia (Smith & Plane 1985).

Two further fossil snakes have been recently described from the Miocene, one from the
Northern Territory, the other from Riversleigh Station, Queensland. Morelia antiquua, from
the Territory, is known from a dentary about 4.5 cm long. The presence of weak cutting ridges
on the lingual and labial sides of the teeth link this species to the living Australasian pythons
of the genus Morelia (Smith & Plane 1985).

FOSSIL REPTILES IN AUSTRALIA - 675

Figure 42. A vertebra from the snake Wonambi naracoortensis, anterior (A), lateral (B), and posterior (C)
views. For comparison, a vertebra of Madtsoia bai in lateral (D) and posterior (E) views. Not to scale. (From
Simpson 1933 and Smith 1976).

The Riversleigh form, Montypythonoides riversleighensis, is represented by a maxilla and
several vertebrae (Smith & Plane 1985). While clearly a boid of the subfamily Pythoninae,
Montypythonoides differs from other pythons by the absence of a lingual cutting ridge on its
teeth (maxillary, at least) and by the presence of a distinct ridge laterally along the hinder
portion of the maxilla. Although both Morelia antiquus and Montypythonoides
riversleighensis represent large snakes (of about the same size) neither are larger than the large
living Australian constrictors.

TESTUDINES

With two exceptions, Australian Cainozoic chelonians are just what would be expected
from a knowledge of the modern forms. Turtle shell fragments date back to Eocene deposits
from Brisbane (Eight-Mile Plains). Some come from shells 1 cm in thickness indicating
that large turtles were present at that time. Others (from Redbank Plains) have arched shells,
like modern Galapagos tortoises. There is no indication of what kind of turtles they were.

Fragments of chelids, indistinguishable from corresponding parts of the modern Emydura
macquarii, are known from the Oligocene or Miocene of Tasmania. Chelids of the
Elseyal/Emydura group are common at Riversleigh. '

The most spectacular extinct tortoise was the horned tortoise Meiolania oweni (Owen 1881,
1882b, Woodward 1888). Horned tortoises, which all appear to be related, are known also from
Lord Howe Island (Pl. 16), New Caledonia and Argentina. M . oweni had a large, broad skull
with long, stout horns above and behind the eyes. When first describing the beast, Owen
suggested that there might well have been other horns, one on the snout behind the nares, and

676 - MOLNAR

another on the skull roof behind the orbits (Pl. 17). The shell of the Australian species is
poorly known, as are the limbs, although it is known that the end of the tail was encased in
bony rings (PI. 18). Each of these rings bore a pair of stout spikes on either side. Meiolania
had free cervical ribs, unusual among chelonians, and characteristic caudals with broadened
transverse processes and chevrons to support the armour rings of the tail. One cannot help but
wonder from what all this armour protected Meiolania.

The first reliable records of Meiolania are shell fragments from the Miocene, and it survived
through the Pleistocene. Material of Meiolania has been found in Pleistocene deposits of Tiga
Island and Walpole Island (both off New Caledonia), New Caledonia itself and Lord Howe
Island (see Balouet, this volume). Although some workers used to be insistent about the
aquatic adaptations of Meiolania, it, in fact, has none. Its distribution, too, on these islands is
still mysterious. Its distribution on Australia is less so. All three horned tortoises, the South
American Crossochelys and Niolamia, as well as Meiolania, are related members of a single
family. Presumably, this family dates back to the Cretaceous when Australia was connected to
South America via Antarctica, and they presumably ranged over all three "continents." The
islands' populations thus predate the breakup of that part of Gondwanaland. If so, older
remains of meiolaniids should be uncovered in Australia, and in Antarctica.

Three cow-like horn cores of large Meiolania have recently been found near Townsville,
Queensland (Gaffney & MacNamara 1990). This animal was apparently not quite as large as
M. oweni, but more similar in skull form to M. platyceps from Lord Howe Island. This
discovery indicates that platyceps-like tortoises were found in Australia as well as on the
southwest Pacific Islands. And the north Queensland animals were as large as M. oweni,
having the bulk, if not the height, of a cow. So, surprisingly, two rather different forms of
horned turtles dwelt in Australia. It seems that M. oweni followed, but probably did not
evolve from, the platyceps-like form. The oldest Meiolania remains are Miocene, but the
skull form is unknown. Since M. oweni seems not derived from the platyceps-like Meiolania,
presumably either both forms lived in Australia, or the platyceps-like tortoise somehow arrived
from the Pacific Islands.

As mentioned above, meiolanid tortoises have been found on Tiga Island, in the Loyalty
Group, and on New Caledonia itself (Gaffney, Balouet & de Broin 1984)(see also Balouet, this
volume). These finds represent more than a simple extension of geographic range, for the Tiga
Island specimen is from an uplifted reef dated as no more than 120,000 years old. This is quite
informative in the absence of any comparably precise dating of the Australian and Lord Howe
remains, but the really surprising date is from the New Caledonian material. This specimen
was associated with charcoal which was dated at only 1700 years old. Thus meiolanid tortoises
were living in New Caledonia in historical times, while the Romans were in England. Since
New Caledonia was populated by at least about 1000 B.C. the extinction of these tortoises may
well have been due to humans.

The remaining form to be mentioned is a trionychid (soft-shelled turtle). These had been
reported in Australia by de Vis (1894), but because of the absence of living specimens, and the
fragmentary nature of his material, the reports were generally not accepted. Recent restudy of
de Vis' material (Gaffney & Bartholomai 1979) indicates that de Vis was correct and such forms
did inhabit Australia during the past. Trionychids today live in North America, Africa and Asia
(into Indonesia), and fossil specimens have turned up in South America as well, so their
occurrence in Australia is not too surprising. They are an old group, dating back to the
Jurassic.

SUMMARY

In the absence of much of a fossil record of Cainozoic reptiles in Australia, many workers
have tried to deduce the histories of various groups based upon their patterns of speciation and

FOSSIL REPTILES IN AUSTRALIA - 677

have tried to deduce the histories of various groups based upon their patterns of speciation and
present distribution. Where these could be checked in the fossil record, as with varanids and
Crocodylus porosus, they appear earlier in the record than had been deduced from modern
information (Molnar 1985). The Tertiary record of reptiles in Australia documents the
existence of yawning chasms in the fossil record and the hazards of deducing history from
modern data alone.

The Pleistocene reptile faunas are better known and essentially composed of two
components. The first is those forms that survive today or are obviously similar to surviving
forms. The second component is the extinct giant forms, such as Megalania, Meiolania and
Wonambi. So far, only Quinkana does not fit comfortably into one of these. Otherwise
there seem to be no "small” reptiles unrelated to those now living. This suggests that the
Pleistocene reptile fauna was basically a modern fauna (or vice versa) with the extinct giants
added. Of the land-dwelling reptiles, the small ones have preferentially survived, but the large
ones did not.

Note that this division involves two different criteria, which happen to produce the same
groups. The first is whether or not the Pleistocene taxa have left descendants. The second is
the size of the animals.

Two of the extinct giants, Meiolania and Wonambi, seem to be archaic or relict forms, but
the status of the small, still-living forms is unknown. Most of these have no fossil record
prior to the Pleistocene and none of them a record prior to the Miocene, so it cannot be
determined which are relicts. We might be able to infer this from their relationships. We
divided the Australian Pleistocene reptilian fauna into two components based on their size and
whether they became extinct. The modern Australian reptilian fauna may also be divided into
constituent parts, but this division is based on the time of their (evolutionary) origin.

The modern Australian reptile fauna is expected to have three components (not to be
confused with the two of the Pleistocene fauna) based on time of evolutionary origin, rather
than relationship to surviving forms. The first of these components includes those reptiles that
migrated to, or originated in Australia before the separation of Australia and Antarctica from
South America. The second includes those forms that evolved in Australia after its separation
from Antarctica, and the third includes those whose ancestors arrived recently as immigrants
from Asia (especially Indonesia), The first component would be most closely related to the old
Gondwanan (South American or African) forms, while the second would also be related but
have undergone evolutionary change. Thus, the first component would consist of relict forms
that had relatively low rates of evolution, at least as expressed in their morphology. The
second component would have had higher rates of evolutionary change. Thus, those reptiles
related to South American or African forms would belong to the first two components, while
those related to Asiatic forms would make up the third. Confirmation of this scheme, and
determination whether those of the Gondwanaland component were relict or had undergone
considerable evolution since the isolation of Australia, would come from the fossil record,
when it becomes better known.

In comparing the three proposed components of the modern Australian reptilian fauna to the
two of the Pleistocene fauna, nothing can yet be said of the component that left modern
descendants. However, the extinct giants Meiolania and Wonambi belong to the first, relict
component; none of these taxa still survive. All modern Australian reptiles seem to belong to
the second or third components: derived (as opposed to relict) Gondwanan descendants or Asiatic
descendants. . . . .

Another interesting feature of the Australian Cainozoic faunas is that the large predators
appear to have been all reptiles (Fig. 43). Among these large reptiles were Megalania,
Pallimnarchus, Wonambi and Crocodylus porosus. The largest predatory mammal,
Thylacoleo, is rather smaller than those of other continents, such as bears, hyaenas, lions and
tigers - it is comparable in size to a leopard. Australia is also where ziphodont crocodiles

678 - MOLNAR

Viverid - Viverra civetta Felid - Panthera leo

Sarcophilus harrisii ~ . wi Aro
Dasyurus BURNS See

Thylacinus oh Canid - /
capris a Canis lupus Mustelid -
ees Gulo gulo
Y Hyaenid -
Ursid - Ursus arctos Crocuta crocuta

Varanus giganteus
Ct Se 7 ea

Megalania prisca

— Pallimnarchus

RFE pollens

fd ra

Qinkana fortirostrum

4,

Crocodylus porosus

Figure 43. The large reptilian predators of the Australian Pleistocene compared with contemporary
mammalian predators and with mammalian predators now living on other continents. In Australia only the
reptiles provided large predatory land vertebrates. (Drawn by L. Beime).

FOSSIL REPTILES IN AUSTRALIA - 679

survived the longest. This suggests the unusual situation in which the major tetrapod predators
are members ofa different class from the major herbivores. This may also have been the case
for early Cainozoic South America, in which large flightless birds were prominent predators,
although carnivorous marsupials and ziphodont crocodilians were also present.

Such unusual features of the Australian fossil record, particularly that of reptiles, shows

that there is much of importance still to be learned from vertebrate palaeontology and
palacoecology in Australia.

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686 - MOLNAR

APPENDIX I

Cladogram 1. Relationships of the major groups of reptiles and their sister-group, the anthracosauroids. Based on the
work of Gauthier, Kluge & Rowe (1988). These workers define reptiles as the descendants of the common ancestor of living
reptiles (see text for further discussion of this convention). Thus, they consider that synapsids, pareiasaurs, procolophonians
and mesosaurs are not reptiles. Not all workers agree that anapsids are more closely related to diapsids than are synapsids.

Cladogram 2. Relationships of turtles, based on Gaffney & Meylan (1988). Groups with Australian members are
emphasized. The term “other groups” on this and the following diagrams indicates one or more that may not be related.

Cladogram 3. Relationships of ichthyopterygians and sauropterygians, based on Mazin (1981) (ichthyoptergians) and Sues
(1987) (sauropterygians). These relationships are obviously unclear, but, even so, not all workers agree with this.

Cladogram 4. Relationships of archosauromorphs and lepidosauromorphs, based on Benton (1985).

Cladogram 5. Relationships of archosaurs and their sister-group, the prolacertiforms. The mixed character of the
thecodonts, including forms not closely related, may be seen. Based on Benton & Clark (1988).

Cladogram 6. Relationships of dinosaurs and pterosaurs and their sister-group, the ornithosuchids, based on (Gauthier
1986).

FOSSIL REPTILES IN AUSTRALIA - 687

Anthracosauroids Captorhinids

Synapsids Proganochelydians

Pareiasaurs Pleurodires

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688 - MOLNAR

PLATES

Plate 1. Internal mould, in dorsal view, of a chelonian carapace from the Winton Formation of central
Queensland (QM F12413).

Plate 2. The only known Australian mosasaur bones from the Late Cretaceous of Western Australia. An
ulna (A) and unidentified bone (B), possibly a limb bone. (Courtesy of the Department of Geology,
University of Western Australia and J. Wiffen).

Plate 3. Left femur from the type specimen of Woolungasaurus glendowerensis.

Plate 4. The type jaw of "Crocodylus" selaslophensis from the Cretaceous of Lightning Ridge, New South
Wales. The mass of opal projecting into the foreground from the lower jaw seems not to represent a part of
the fossil. (Courtesy of the University of New South Wales and Teaching Hospitals and The Australian
Museum).

Plate 5. Cretaceous crocodilian material from Lightning Ridge, New South Wales. Left is a procoelous
cervical centrum in dorsal (A) and lateral (B) views; C, tibia. (Courtesy of the University of New South Wales
and Teaching Hospitals and The Australian Museum).

Plate 6. Dorsal vertebra probably from a very small theropod dinosaur. This was found in the Lower
Cretaceous rocks near the Hamilton River, western Queensland. The centrum is about 3 mm long.

Plate 7. Leaellynasaura amicagraphia, holotype (NMV P185990). A, C, E ventral view of rear of frontals
plus the parietal in normal and reverse; B, D, dorsal view of the counterpart to A,C,E showing endocast of
brain. Skull is 67 mm in length. (From Rich & Rich 1989).

Plate 8. Right femur of Leaellynsaura amicagraphica (from the Early Cretaceous near Cape Otway, Victoria).
A, C, anterior view, stereo pair; B, D, posterior view, stereo pair; E, G, proximal view, stereo pair; F, H,
distal view, stereo pair (length of femur 67 mm). (After Rich & Rich 1989).

Piate 9, Atlascopcosaurus loadsi from the Early Cretaceous near Cape Otway, Victoria. A-D, maxilla with
upper cheek teeth; A, C, lateral view, stereo pair, B, D, medial view, stereo pair, all x1. E-H, mandible with
two unermupted cheek teeth; E, G, lateral view, stereo pair, x0.66; F, H, medial view, stereo pair, x1.33; I-L,
lower cheek teeth; I, K, (same individual as in E-H) lateral view, stereo pair, J, L, lateral view, stereo pair,
all x2. (After Rich & Rich 1989).

Plate 10. Right femur of Fulgurotherium australe, from the Early Cretaceous near Cape Otway, Victoria. A,
C, anterior view, stereo pair; B, D, posterior view, stereo pair, E, G, medial view, stereo pair; F, H, lateral
view, stereo pair; I, J, proximal view stereo pair (all x0.5). (After Rich & Rich 1989).

Plate 11. A hypsilophodontid tooth, possibly from Fulgurotherium, from Lightning Ridge, New South
Wales; x3.

Plate 12. The shoulder girdle of a pterosaur from the Early Cretaceous of western Queensland.
Plate 13. Jaw of a crocodile from the probably Eocene oil shales at Rundle, eastern Queensland.

Plate 14. The snout of a young Crocodylus porosus from the Pliocene deposits at Allingham Creek,
northern Queensland. This is the oldest specimen of the Salt-Water Crocodile in Australia. (From Molnar
1979).

Plate 15. The type skull (i.e. snout) of the Australian ziphodont crocodilian, Quinkana fortirostrum, A,
dorsal view; B, ventral view. (From Molnar 1981).

Plate 16. The skull of Meiolania platyceps from Lord Howe Island, New South Wales. Two different skulls
are shown in posterior view (A, C) to demonstrate the variation in development of the hom cores. B, dorsal;
D, lateral views. (From Gaffney 1983).

FOSSIL REPTILES IN AUSTRALIA - 689

3 . . a
abe 17. ea skull of Meiolania oweni from the Darling Downs, Queensland. Continuous lines indicate
reconstructed portions of the skull, while dashed lines indicate Owen's locations for homs. (From Owen 1884).

Fo The tail armour of Meiolania oweni, viewed from above (A), laterally (B) and below (C). (From Owen

Plate 19. Ankylosaur skeleton (Minmi sp.) collected in January 1990, from near Richmond, north Queensland. The
triangular skull is visible at the top, and the scutes of the dorsal armour may be seen on the neck (just below the skull)
and between the ribs of the back. The small ossicles of the armour are visible in the intervening areas (and between the
ribs) of the back. The left ilium is visible at the bottom left: that of the right is still covered by rock. The skeleton is
viewed from above, and the neck was somewhat longer than as laid out here.

ADDENDUM

; Since this chapter was completed there have been several significant, relevant discoveries. New work on the
primitive African procolophonian Owenetta has shown ten synapomorphies with early chelonians, leading to the
conclusion that procolophonians are the sister-group of turtles (Reisz & Laurin 1991). The diversity of Mesozoic
crocodilians has been enhanced by discovery, in Lower Cretaceous rocks of Malawi (central Africa), of crocodilians
with a differentiated dentition, similar to that of mammals (Clark ef al. 1989).

New material has been found in Australia as well. Footprints of several kinds of dinosaurs have been found near
Broome (W.A.), including those of sauropods, omithopods and possibly stegosaurs (Long 1990). The first Australian
oo reptile, apparently a Quetzelcoatlus-like pterosaur, has also been recognised in Westem Australia (Long

In assessing the problem of south polar temperatures and dinosaurian physiology (pp.664-665), I pointed out that
marine reptiles are known from the Late Cretaceous of both New Zealand and Antarctica. Further, I commented that no
one had suggested that they maintained an elevated body temperature. But the recent work of Paladino et al. (1990)
shows that leatherbacks (Dermochelys) can maintain a body temperature of 25°C. in a water temperature of Jess than
8°C. These workers further argue that larger extinct reptiles could have held their body temperatures at Icast 30°C.
above that of their surroundings - without needing metabolic rates comparable to those of mammals. So, it seems
possible that large marine (and terrestrial) reptiles could have survived temperatures like those postulated for southem
Victoria during the Early Cretaceous.

A new ziphodont crocodilian, Baru darrowi, has been described from Riversleigh (Qld.) and Bullock Creek (N.T.)
(Willis et al, 1990). Baru is similar to Quinkana, Pallimnarchus and undescribed crocodiles from South Australia. It
suggests that there was an indigenous radiation of Australian Cainozoic crocodilians. The Riversleigh agamids
(Covacevich et al, 1990) include Physignathus (Water Dragon), indicating a Late Miocene time for the coming of
Asiatic lizards into Australia, for Physignathus seems to have originated in Asia. Although most of the Miocene
lepidosaurs still survive, demonstrating the conservatism of the Australian small lepidosaur fauna, two genera,
Montypythonoides and Sulcatidens, have not (Covacevich et al 1990). So, it is not just the "giants" that became
extinct. The discovery of Meiolania in the Miocene Camfield beds (N.T.) (Merigan 1989) supports the suggestion that
it was a relict form that lived in Australia throughout the Cainozoic. And finally, the newly studied skull roof elements
of Megalania (Molnar 1990) support Hecht's contention that the skull was akinetic, and show that Megalania sported a
low medial crest on lits head (see reconstruction including this feature by P. Trusler in Vickers-Rich & Rich, 1991).

CLARK, J.N., JACOBS, L.L. & DOWNS, W.R., 1989. Mammal-like dentition in a Mesozoic crocodylian. Science 244: 1064-1066.

COVACEVICH, J., COUPER, P., MOLNAR, R.E., WITTEN, G. & YOUNG, W., 1990. Miocene dragons from Riversleigh: new data on
the history of the family Agamidae (Reptilia: Squamata) in Australia, Mem. Qd. Mus. 29: 339-360.

LONG, J.A., 1990. Dinosaurs of Australia. Reed Books, Balgowlah (Sydney).

MEGIRIAN, D., 1989. A description of homed-turtle remains (Testudines: Meiolaniidae) from the mid-Miocene Camfield beds of
northem Australia, The Beagle 6: 105-1 13.

MOLNAR, R.E., 1990. New cranial elements of a giant varanid from Queensland. Mem. Qd. Mus. 29: 437-444.

PALADINO, F.V., O'CONNOR, M.P. & SPOTILA, J.R., 1990. Metabolism of leatherback turtles, gigantothermy, and thermoregulation
of dinosaurs. Nature 344: 858-860.

REISZ, R.A. & LAURIN, M., 1991. Owenetta and the origin of turtles. Nature 349: 324-326,

VICKERS-RICH, P & RICH, T-H., 1991. Wildlife of Gondwana. Reed Books, Balgowlah (Sydney).

WILLIS, P., MURRAY, P. & MEGIRIAN, D., 1990. Baru darrowi gen. et sp. nov., a large, broad-snouted crocodyline (Eusuchia:
Crocodylidae) from mid-Tertiary freshwater limestones in northem Australia. Mem. Qd. Mus. 29: 521-540.

PLATE 1
PLATE 2

690 - MOLNAR

PLATE 3 FOSSIL REPTILES IN AUSTRALIA - 691

692 - MOLNAR PLATE 4

PLATE 5

PLATE 6 FOSSIL REPTILES IN AUSTRALIA - 693

PLATE 7

fe
Cerebral
hemisphere

F- |

oe)

Parietal

body
Cerebellum

PLATE 8

694 - MOLNAR

PLATE 9 FOSSIL REPTILES IN AUSTRALIA - 695

PLATE 10

696 - MOLNAR

P
mae er et FOSSIL REPTILES IN AUSTRALIA - 697

PLATE 12

PLATE 13

pr Of BE fe Of GE WE GE ZENE OF HZ OL Oe Ge me BE EE Ie OF GH wm

Pid ititiiiiiid

698 - MOLNAR PLATE 14

A

PLATE 15 FOSSIL REPTILES IN AUSTRALIA - 699

700 - MOLNAR

PLATE 16

PLATE 17

FOSSIL REPTILES IN AUSTRALIA - 701

PLATE 18

PLATE 19

702 - MOLNAR

CHAPTER 19

THE FOSSIL TURTLES OF
AUSTRALIA

Eugene S. Gaffney!
MifOMUE HOR 517 he. Ae Pin CR ER eet Ds, 704
Family Qhelidaes. 3..)0), ee OE Seen 704
Diagnosis 8. Lath. eee See ee hs 704
References sie tc7. seed | ee eas 705
Australian Records of the Chelidae .............. 705
Family Meidlaniidae. 2.4.08. pe ee 708
DEARNOSIS2 et OR SR cer ee 708
Reféronces 4 ty 9. he So Rn eae Ue: 709
Australian Records of the Meiolaniidae......... 709
Meiolania platyCeps .......ccccccecccceecenecees 709
Meiolanid OWENI....cr.cccccesecececesecseceeeecs 710
Meiolania cf. platyceps ..........cccc ccc eeceues 710
Undetermined Meiolaniids from the
Tertiary of Mainland Australia........... 710
Meiolaniids from New Caledonia........... 713
Family Desmatochelyidae, D10,
Unnamed Taxon of Gaffney
& Meylan (1988)........ccceeeeceeeseees 713
AA ENDSES 1.) ches Mele Lee eh eee 714
IRELCIENCES SEA Tees ek ee ee 714
Family Trionychidaes: 7.0! che ee 714
Diagnosise: 00s: ee ee ee 714
References 204.0000 ae ee 714
Australian Taxa of Trionychids................... 714
Mesozoic Turtles of Undetermined
Relationships iy..0. +. ...c03) ccasseecets Sentient 714
REfEreNnCES sr mi Goes ee es dete tah eet es 715
Plates f. 035th s, Te Soece cls ee ante eee te eo 716

a ns

1 Department of Vertebrate Paleontology, The American Museum of Natural History, Central Park W. at 79th
St., New York, New York 10024, U.S. A.

704 - GAFFNEY

INTRODUCTION

Turtles are often common elements of both living and fossil faunas and have the potential
to provide significant perspectives on biogeography, habitat and evolutionary relationships.
The Australian turtle fauna is less diverse than that of other continents (except Antarctica) in
terms of species and higher taxa, but the taxa that are present are very interesting and
significant from an evolutionary point of view.

There are some general works on turtles that the interested reader should be aware of. A
compilation of the living species of turtles with figures can be found in Wermuth & Mertens
(1961, 1977) and Pritchard (1979), while a compilation of fossil turtles is in Mlynarski (1976)
and Kuhn (1964). The higher level systematics and relationships of living and extinct turtles
to genus is in Gaffney & Meylan (1988) (see also Fig. 1): Cranial morphology of turtles is
presented in Gaffney (1979a), and the best description of turtle postcranial morphology is
Bojanus (1819, reprinted 1970). A history of chelonian classifications and phylogenies is in
Gaffney (1984). The living turtles of Australia are reviewed in Cogger (1975), and the fossil
turtles of Australia are reviewed in Gaffney (1981, in press).

The vernacular use of the word "turtle" to refer to marine forms and "tortoise" to refer to
non-marine forms is essentially a local usage peculiar to British-influenced regions. The more
widespread usage is "turtle" meaning all chelonians and "tortoise" restricted to the family
Testudinidae, a family of dryland turtles not found in Australia. As in other scientific works on
Australian chelonians, I use "turtle" to mean all chelonians.

FAMILY CHELIDAE

The most common non-marine chelonians in the recent and fossil biota of Australia are the
chelids, a freshwater aquatic to semi-aquatic family of pleurodires or side-necked turtles. The
recent and fossil members of this family are restricted to South America (about 12 living
species) and Australia (about 15 living species). The fossil record of chelids is not extensive,
although fragmentary fossils of chelids are known from most Tertiary and Quaternary vertebrate
sites in Australia (Figs 1-3). Species not assignable to the recent fauna have not yet been
objectively diagnosed.

DIAGNOSIS

Lateral cheek emargination of the skull bones unusually well-developed; quadratojugal
absent; mesoplastra absent; neural bones extensively reduced or absent; cervical vertebral
formula is (2((3((4((5))6))7((8).

The above features are synapomorphies for Chelidae, but the group has other characters that
differentiate it from most or all of the remaining Australian turtles. Because chelids are such a
prominent Australian group, it is worth listing plesiomorphic characters found in chelids.

1. Fore and hindlimbs of generalized chelonian pattern, with distinct joints, and
clawed, webbed digits not developed into flippers or paddles.

2. Nasal bones present.

3. Prefrontals do not meet in midline.

FOSSIL TURTLES OF AUSTRALIA - 705
REFERENCES

‘ The systematics of the entire family down to genus is dealt with by Gaffney (1977).
pecies level taxa of recent Australian chelids are in Cogger (1975), Goode (1967), Cann
(1978), Leagler & Cann (1980), and Burbidge, Kirsch & Main (1974).

AUSTRALIAN RECORDS OF THE CHELIDAE

The three recent generic level taxa of chelids as recognized by Gaffney (1977, in press) are

Pseudemydura, Chelodina and the Emydura Group (Emydura, Elseya, and Rheodytes). These
three taxa are also present in the Australian fossil record (Fig. 2).
_ _Pseudemydura is known from the recent fauna as one rare species, on the brink of extinction
in Western Australia. The only fossil record of this genus is a fragmentary specimen from the
Riversleigh localities of Queensland (Gaffney, Archer & White 1989). This Miocene record
provides evidence that Pseudemydura was more widespread in the Tertiary and has a history of
some antiquity.

Pseudemydura is diagnosable by many synapomorphies (Gaffney 1977, in press) and has
been hypothesized (Gaffney 1977) as the sister group of all other chelids.

The Emydura Group contains nine (or more, depending on the author) Australian and New
Guinean species that are very similar in osteology and general habitus. Most of the characters
diagnosing the Emydura Group (Gaffney 1977, Goode 1967, Cogger 1975) are probably
primitive for chelids. The only likely synapomorphy for the group is the relatively heavy
lower jaws with a wide triturating surface (Gaffney 1977). The possibility is very good that
dividing the Emydura Group (that is, the old genus Emydura) into three genera, Emydura,
Elseya, and Rheodytes, is not based on a systematic hypothesis that there are three
monophyletic groups defined by synapomorphies. Rather, it reflects the recognition of a few
species that are more readily distinguishable on the basis of unanalyzed characters (Gaffney, in
press),

The current state of species level systematics in the Emydura Group does not allow an
objective basis for the naming of extinct species. Few of the recent taxa are described
osteologically, and the absence of a phylogenetic analysis based on synapomorphies prevents
the identification of these synapomorphies among fossil taxa. However, the Emydura Group
has been widely identified in the Tertiary and Pleistocene of Australia (Figs 2-5, Pls 1-2).
Most of these records are related in Gaffney (1981). Descriptions of the best known Emydura
material, including skulls are in Gaffney (1979b) with well preserved shells described by Burke,
Gaffney & Rich (1983) and Warren (1969a). More recent records and discussions of problems
concerming the Emydura Group are in Gaffney (in press) and Gaffney, Archer & White (1989).

Again, it should be expected that any primitive chelid will look like Emydura, and the
identification of a fossil as Emydura is often the same as saying that it is a primitive chelid.

Chelodina is a long-necked chelid (in contrast to the short-necked Pseudemydura and
Emydura) that is diagnosed by a series of synapomorphies (Gaffney 1977, in press). Six or
seven species are present in the recent Australasian fauna, but the species level systematics of
Chelodina are in just as primitive a state as in the Emydura Group. In contrast to the
widespread occurrence of Emydura Group fossils in Australia (Figs 2-3), Chelodina is known
from only two localities: the Tara Creek Pliocene of eastern Queensland (Gaffney 1981) and
the Riversleigh Miocene of western Queensland (Gaffney, Archer & White 1989).

In addition to the better known material mentioned above, there are chelid fossils that extend
the family into the Eocene of Australia. The Rundle Formation near Gladstone, Queensland,
and the Cape Hillsborough Formation of Cape Hillsborough, Queensland, have yielded
diagnosable shell parts that are chelid (Gaffney, in press).

706 - GAFFNEY

and the Cape Hillsborough Formation of Cape Hillsborough, Queensland, have yielded

diagnosable shell parts that are chelid (Gaffney,

in press).

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FOSSIL TURTLES OF AUSTRALIA - 707

CHELONIIDAE
TRIONYCHIDAE
EMYDIDAE
“BATAGURIDAE”
TESTUDINIDAE

DERMOCHELYIDAE
CARETTOCHELYIDAE
DERMATEMYDIDAE
KINOSTERNIDAE

OSTEOPYGIDAE

PROTOSTEGIDAE
ADOCIDAE

MILLIONS OF YEARS | 300

Figure 1. The relationships of the main groups of turtles. This is the cladogram presented in Gaffney &
Meylan (1988), placed on a time scale showing the ranges of the groups. Recent groups are shown by a
living turtle, while the extinct groups are represented by a skeleton. Groups found in Australia are shown

with an asterix (*).

708 - GAFFNEY

FAMILY MEIOLANIIDAE

The extinct horned turtles of Australia and South America are probably the most fascinating
and bizarre of chelonians. Their relationships have been controversial for more than a century,
but recent work (Gaffney 1983) argues that they are primitive eucryptodires, the sister group of
the living cryptodires. The meiolanids are known only from South America and Australasia
(mainland Australia plus Lord Howe Island, Walpole Island and New Caledonia). The South
American forms are the oldest, the Eocene record being the first well-documented meiolaniid,
The Australian forms are known from the Miocene to the Late Pleistocene (Figs 2-3); the
island species are Late Pleistocene.

CHELIDAE CHELONIOIDEA TRIONYCHIDAE CARRETOCHELYIDAE
REGENT MEIOLANIIDAE ge

PLEISTOCENE 9

PLIOCENE

Bey
i.
ano

MIOCENE

OLIGOCENE

TERTIARY

EOCENE

PALEOCENE

UPPER

CRETACEOUS

Figure 2. Temporal ranges of turtle families in Australia.

DIAGNOSIS

Squamosal and supraoccipital produced into large posteriorly and posterolaterally directed
processes that extend clear of the skull; medial plate of pterygoid separated ventrally from
basisphenoid to form the intrapterygoid slit; broad squamosal - quadratojugal contact ventral to
quadrate; heavily ossified tail club; ossified tail rings; cervical ribs unusually large; pes and
manus digits and with two or fewer phalanges.

FOSSIL TURTLES OF AUSTRALIA - 709

REFERENCES

Skull morphology can be found in Gaffney (1983) and Owen (1881), vertebral morphology

in Gaffney (1985) and aspects of the entire skeleton in Anderson (1925, 1930). A comparison
of known records is in Gaffney (1981).

AUSTRALIAN RECORDS OF THE MEIOLANIIDAE

Meiolania platyceps

The abundant and well-preserved material from Lord Howe Island (Figs 3, 6, Pl. 3 ) has
made this species the best known member of the family. The specimens all occur on a small
volcanic island in calcarenites tentatively dated at 100,000-120,000yBP (see Gaffney 1983, for
geology and discussion). Although the Lord Howe specimens show a great deal of variation,

the species is characterized by large, recurved horns, no occipital frill and a total size of about
two metres,

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710 - GAFFNEY

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Figure 4. Drawings of shell in Pl. 1A, B, with bones and scales identified.
Meiolania oweni

This species, from the Darling Downs Pleistocene of Queensland, is known only from the
skull and tail. It was described by Owen (1881, 1882) and named by Woodward (1888, 1901).
Meiolania oweni differs from M. platyceps in lacking recurved horns, in having a large
occipital frill and in being one and one half to two times larger.

Meiolania cf. platyceps

Three horn cores and a caudal from Late Pleistocene (30,000-200,000 yBP) are known from
Wyandotte Station, northern Queensland (Gaffney & McNamara 1990). The horn cores are
very similar to the Lord Howe species, except that they are more than twice the size. It is
likely that the Pleistocene of Queensland had two, contemporary species of giant horned turtle
wandering about, wreaking havoc.

Undetermined Meiolaniids from the Tertiary of Mainland

Australia

Etheridge (1889) and Gaffney (1981) describe meiolaniid fragments from Gulgong, New
South Wales, that may be Miocene in age. This species seems to be about 20-30% smaller
than Meiolania platyceps.

The Miocene of Central Australia has yielded meiolaniid fragments (Gaffney 1981) similar
in size and morphology to the Gulgong material. The Riversleigh Miocene of western
Queensland also has meiolaniid fragments, but all of these specimens are too fragmentary to
make definitive comparison as yet.

FOSSIL TURTLES OF AUSTRALIA - 711

Figure 5. Skull of Emydura macquarii, Chelidae, Recent, South Australia. (From Gaffney 1977).

712 - GAFFNEY

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Figure 6. Skull of Meiolania platyceps, Meiolaniidae, Pleistocene, Lord Howe Island, Australian Mus.
57984. (From Gaffney 1983).

FOSSIL TURTLES OF AUSTRALIA - 713

Meiolaniids from New Caledonia

Anderson (1925) described fragments of a small meiolaniid from Walpole Island as
Meiolania mackayi, and some meiolaniid cervicals were reported from the main island of New
ceca (Gaffney, Balouet & de Broin 1984). These all appear to be Late Pleistocene or

r.

It should be emphasized that the meiolaniids were not aquatic turtles, but had limb
morphology similar to the tortoises (Testudinidae) of today. They are unlikely to have been
transported from the Australian mainland by swimming, but may represent relict faunal
elements surviving on repeatedly emergent volcanoes.

Family Desmatochelyidae, D10, Unnamed Taxon of Gaffney
& Meylan (1988)

In the Early Cretaceous Toolebuc Limestone of Queensland, there are two sea turtles,
Cratochelone and Notochelone. They have never been studied in detail, but Notochelone (Fig.
7) appears to be similar to the North American Desmatochelys and the European Allopleuron.
All three of these latter genera have been united with the protostegids and dermochelyids in a
monophyletic group, "D10", by Gaffney & Meylan (1988). Cratochelone is based on very
fragmentary material, but appears to be a protostegid and would, therefore, belong in this
taxon. bes

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Figure 7. Shell of Notochelone costata, Chelonioidea, Cretaceous, Toolebuc Limestone, Queensland.
Restoration based on Queensland Mus. F2249. (From Gaffney 1981).

714 - GAFFNEY

DIAGNOSIS

Plastron with very large central and peripheroplastral fontanelles, reducing the hyoplastral-
hyoplastral contact to a narrow projection; medial process of jugal absent; scapular angle wide,
in contrast to other chelonioids.

REFERENCES

The Australian material is not well described; the best figures are in Gaffney (1981)

FAMILY TRIONYCHIDAE

The trionychids or soft-shell turtles, are cryptodires that occur today in Africa, Asia and
North America, but not in South America or Australia. Fossils of the group, however, are
found in South America and Australia, leaving Antarctica as the only continent lacking a record
of the family.

DIAGNOSIS

Peripheral bones absent (except in Lissemys); pygal and suprapygal bones absent;
boomerang-shaped entoplastron; scales of shell absent; centrum of last cervical not articulating
with centrum of first thoracic; jaws covered by fleshy lips.

REFERENCES

Meylan (1987) is the best reference for recent trionychids.

AUSTRALIAN TAXA OF THE TRIONYCHIDAE

De Vis (1897) identified some shell fragments from Darling Downs as trionychid. Gaffney
& Bartholomai (1979) re-affirmed the accuracy of de Vis’ identifications and provided more
material from other localities. None of the specimens are diagnosable beyond family. All of
the descrobed material is from Queensland and is at least Pliocene in age, but newly discovered
specimens from Mrgon may be Palaeogene.

MESOZOIC TURTLES OF UNDETERMINED RELATIONSHIPS

The only Mesozoic turtles from Australia that are known well enough to determine
relationships are the chelonioids. There are, however, fragments of other turtles from the
Cretaceous that suggest the presence of more primitive turtles, probably below the level of
Centrocryptodira as diagnosed by Gaffney & Meylan (1988).

The best of these specimens is the steinkern, Chelycarapookus (Warren 1969b) (Fig. 8),
which is probably a primitive cryptodire (Gaffney, in press). It is probably from the Early
Cretaceous Merino Formation of Victoria. Molnar (1980) described some fragments from the
Early Cretaceous Griman Creek Formation of New South Wales, that are probably from
cryptodires below the level of Centrocryptodira. T. and P. Rich and associates have discovered
fragments from the Early Cretaceous Otway and Strzelecki groups of southern Victoria that

FOSSIL TURTLES OF AUSTRALIA - 715

pia also be referred to non-centrocryptodiran cryptodires. Even if taken together, this material
ie not allow definitive identification of particular cryptodire groups, but it suggests that
ustralia, like the other continents, had representatives of the primitive cryptodires.

Figure 8. Intemal mould of carapace of Chelycarapookus arcuatus, Cryptodira indeterminant, ?Cretaceous, ?
Merino Group, ?Victoria. (From Gaffney 1981).

REFERENCES

ANDERSON, C., 1925. Notes on the extinct chelonian Meiolania, with a record of a new occurrence. Rec.
Aust. Mus. 14: 223-242.

ANDERSON, C., 1930. Paleontological notes no. Il. Metolania platyceps Owen and Varanus (Negakabua)
priscus (Owen). Rec. Aust. Mus.17: 309-316.

BOJANUS, L.H., 1819. Anatome Testudinis europaeae. Soc. Study Amph. & Rept. facsimile reprints in
herpetology (reprinted 1970), 26: i-vi, 1-178.

BOULENGER, G.A., 1889. Catalogue of the chelonians, rhynchocephalians and crocodiles in the British
Museum (Natural History). British Museum, London.

BURBIDGE, A.A., KIRSCH, J.A.W. & MAIN, A.R., 1974. Relationships within the Chelidae (Testudines:
Pleurodira) of Australia and New Guinea. Copeia2: 392-409.

BURKE, A.C., GAFFNEY, E.S. & RICH, T.H., 1983. Miocene turtles from Lake Tarkarooloo, South Australia.
Alcheringa 7: 151-154.

CANN, J., 1978. Tortoises of Australia. Angus & Robertson, Sydney.

COGGER, H., 1975. The Reptiles and Amphibians of Australia. Reed, Sydney.

DE VIS, C.W., 1897. The extinct fresh-water turtles of Queensland. Ann. Qd. Mus.3: 1-7.

ETHERIDGE, R., 1889. On the occurrence of the genus Meiolania in the Pliocene Deep Lead at Canadian,
near Gulgong. Rec. Geol Surv. N.S.W.1: 149-152.

GAFFNEY, E.S., 1977. The side-necked turtle family Chelidae: a theory of relationships using shared derived

characters. Am. Mus. Novit.2620: 1-28.
GAFFNEY, E.S., 1979a. Comparative cranial morphology of recent and fossil turtles. Bull. Am. Mus. nat.

hist. 164: 65-375. ;
GAFFNEY, E.S., 1979b. Fossil chelid turtles of Australia. Am. Mus. Novit. 2681: 1-23.
GAFFNEY, E.S., 1981. A review of the fossil turtles of Australia. Am. Mus. Novit.2720: 1-38.

716 - GAFFNEY

GAFFNEY, E.S., 1983. Cranial morphology of the extinct horned turtle, Meiolania platyceps, from the
Pleistocene of Lord Howe Island. Bull. Am. Mus. nat. hist. 175: 361-480.

GAFFNEY, E.S., 1984. Historical analysis of theories of chelonian relationships. Syst. Zool. 33(3): 283-
301.

GAFFNEY, E.S., 1985. The cervical and caudal vertebrae of the cryptodiran turtle, Meiolania platyceps, from
the Pleistocene of Lord Howe Island. Am. Mus. Novit.2805: 1-29.

GAFFNEY, E.S., in press. An introduction to turtles, with a review of the turtles of Australia.

GAFFNEY, E.S. & BARTHOLOMAI, A., 1979. Fossil trionychids of Australia. J. Paleont. 5§3(6): 1354-
1360.

GAFFNEY, E.S., BALOUET, J.C. & DE BROIN, F., 1984. New occurrences of extinct meiolaniid turtles in
New Caledonia. Am. Mus. Novit. 2800: 1-6.

GAFFNEY, E.S. & MEYLAN, P.A., 1988. A phylogeny of turtles. In The Phylogeny and Classification of
Tetrapods. M. J. Benton, ed., Clarendon Press, Oxford: 157-219.

GAFFNEY, E.S. & MCNAMARA, in press. A meiolaniid turtle from the Pleistocene of northem Queensland.
In Vertebrate Zoogeography & Evolution in Australasia, M. Archer & G. Clayton, eds., Hesperian Press,
Carlisle.

GAFFNEY, E.S., ARCHER, M. & WHITE, M., /989 Chelid turtles from the Miocene freshwater limestones
of Riversleigh Station, northwestem Queensland, Australia. Am. Mus. Novit.2959: 1-10.

GOODE, J., 1967. Freshwater tortoises of Australia and New Guinea (in the Family Chelidae). Lansdowne
Press, Melboume.

KUHN, O., 1964. Fossilium Catalogus. 1: Animalia, Pars 107, Testudines. Junk, The Hague.

LEGLER, J. & CANN, J., 1980. A new genus and species of chelid turtle from Queensland, Australia. Contr.
Sci. Nat. Hist. Mus. Los Angeles Co. 34: 1-18.

MEYLAN, P.A., 1987. The phylogenetic relationships of soft-shelled turtles (Family Trionychidae). Bull.
Am. Mus. nat. hist.186: 1-101.

MLYNARSKI, M., 1976. Testudines. In Encyclopedia of Paleoherpetology, Part 7, O. Kuhn, ed.: 1-130.

MOLNAR, R., 1980. Australian late Mesozoic terrestrial tetrapods: some implications. Mém. Soc. géol. Fr.,
n.s. 139: 131-143.

OWEN, R., 1881. Description of some remains of the gigantic land-lizard (Megalania prisca, Owen) from
Australia. Part Il. Phil. Trans. R. Soc. (1880) 171: 1037-1950.

OWEN, R., 1882. Description of some remains of the gigantic land-lizard (Megalania prisca Owen) from
Australia. Pam Il. Phil. Trans. R. Soc. (1881) 172: 547-556.

PRITCHARD, P.C.H., 1979. Encyclopedia of Turtles. T. F. H. Publications, Neptune.

WARREN, J.W., 1969a. Chelid turtles from the mid-Tertiary of Tasmania. J. Paleont. 43(1): 179-182.

WARREN, J.W., 1969b. A fossil chelonian of probable lower Cretaceous age from Victoria, Australia. Mem.
natin. Mus. Vict.29: 23-28.

WERMUTH, J.W. & MERTENS, R., 1961. Schildkréten, Krokodile, Briickenechsen. Gustav Fisher Verlag,
Jena.

WERMUTH, H. & MERTENS, R., 1977. Liste der rezenten Amphibien und Reptilien, Testudines, Crocodylia,
Rhynchocephalia. In Das Tierreich. H. Wermuth, ed., Walter de Gruyter, Berlin, New York.

WOODWARD, A.S., 1888. Note on the extinct reptilian genera Megalania Owen and Meiolania Owen. Ann.
Mag. nat. Hist. 1(6): 85-89.

WOODWARD, A.S., 1901. On some extinct reptiles from Patagonia, of the genera Miolania, Dinilysia, and
Genyodectes. Proc. zool. Soc. Lond. 1901: 169-184.

PLATES

Plate 1. Shell of Emydura sp., Chelidae, Wipajiri Formation, Miocene, South Australia. (From Gaffney
1979a). Plastron (A) and carapace (B) (Univ. California, Berkeley, specimen UCMP 77348). See Fig. 4 for
labelled drawing.

Plate 2. Left lateral views of Recent and fossil Emydura skulls, Chelidae. A, Emydura australis, Amer. Mus.
Nat. Hist. 108857, Recent; B, Emydura sp., Univ. California (Berkeley) UCMP 57253, Etadunna Formation,
South Australia. (From Gaffney 1979a).

Plate 3. Skeleton of Meiolania platyceps, Meiolaniidae, Pleistocene, Lord Howe Island, restored cast based
on Australian Mus. 57984. (From Gaffney 1983).

PLATE 1A FOSSIL TURTLES OF AUSTRALIA - 717

718 - GAFFNEY PLATE 1B

PLATE 2

FOSSIL TURTLES OF AUSTRALIA - 719

720 - GAFFNEY PLATE 3

CHAPTER 20

THE MESOZOIC AND
TERTIARY HISTORY OF
BIRDS ON THE
AUSTRALIAN PLATE

Patricia Vickers-Rich!

TIMER CTIOIN ior hcl Paddys died wah cocencatty adaiiee!
Historical Perspective ........cccccccceeeececees
Temporal Distribution of Avian Fossils
in Australia and New Guineaa.......
Geographic Distribution of Avian Fossils
in Australia and New Guinea.......
Biases of the Mesozoic and Tertiary
Avian Record in Australia
and New Guinea
The Avian Fossil Record (Australia-
New Guinea): Mesozoic............
The Avian Fossil Record (Australia-
New Guinea): Palacogene..........
The Avian Fossil Record (Australia-
New Guinea): Neogene
Dromormithidae..., c..ct An tesneedare
Casuariidae
POUICIPEMIN ACs hs sa.0k Heres cidoeden's Seseen oe
Spheniscidae
WiGOMed HAAS s bs. hss css ser cewcich oan han ce

Ardeidae

eee eee eee eee eee ee ee ee ee ee)

—_—$—$—$—$_$—_———————————————————— ———————————————————————————————————————

726

729

Threskiornithidae.........cee cece eeeeeeeeeee 752

PROCMICOPIETOAC 0. eda cciccsiceces bee sii 153;

Palaclodidae..........ccccccccceeeeeeeeecenees 756

ASTIALIGAGH Make Een e MN. Ae te 757

Accipitridae and Falconidae............... 757

Megapodiidae.............cccsccceseccseceeees 758

UEMICIMAG SS 5; SEH oe a Boal seus ee 758

Gruidag fed. to RN Re es 758

Rallidaey. 9.5 fect al Wt A Mavenee cle be 760

Onitdan a tee, SES A 760

Charadritformes ai ¥s 4) Stele 760

Tytonidae and Strigidae.....0000......... 761

Podargidae and Aegothelidae ............. 761

APOdIGAGS 5. i eee ancien 762

Columbidae Wee eee hittin 762

Psittaciformesss, wheat Maevaee 762

Rasseniformes.t 48.0 See tine 762
Origin of the Australian Avifauna:

Dispersal, Vicariance or Both? .......... 763
Summary and Conclusions.....0...........0.. 768
IRELEVENCEST A Sete Ph kc eel oe Sieh ae 769
lates ee ad eevee eels oo 781

1 Earth Sciences and Ecology/Evolutionary Biology Departments, Monash University, Clayton, Victoria 3168

Australia.

722 - RICH

INTRODUCTION

Bones of extinct birds were amongst the first fossil vertebrates recovered from Australia. In
1831 George Ranken, a local grazier helping Sir Thomas Mitchell (Mitchell 1838) explore the
Wellington caves of New South Wales (see Rich & Archbold, this volume), deftly attached his
rope to "what seemed a projecting portion of rock" in order to support the next stage of his
descent. Unfortunately for Mr. Ranken, but quite fortunately for the science of
palaeornithology, the projecting portion of rock turned out to be the femur of a large ground
bird, now known to belong in the family Dromornithidae (Owen in Mitchell 1838). The aim
of this article is to detail the avian fossil record for Australia and New Guinea. This includes
an historical account of the discovery of the fossil birds as well as a critical look at how the
record is biased and how it can be used as a basis for outlining the development of the
Australasias' unique avifauna.

HISTORICAL PERSPECTIVE

Mr. Ranken's discovery of a fossil bird bone in the Wellington Valley was the beginning of
palaeornithology on the Australian continent in 1831. Over the next half century bird bones
were occasionally discovered by state surveyors, such as H. Y. L Brown (1894) in South
Australia and W. B. Clarke (1869, 1877) in New South Wales. They also were tumed up in
Aboriginal and European wells and were reported on by Owen (1874, 1879a,b), Etheridge
(1889, 1894) and Lydekker (1891, 1892), amongst others. The Reverend J. E. T. Woods
(1862, 1866, 1882) both collected and reported on bones of the large dromornithids from the
southeastern part of South Australia, particularly around Penola.

It was in the latter decade of the 19th century when a great deal of fossil avian material was
found and early part of the 20th century when it was described, mainly through the efforts of
Charles de Vis (1885-1911) while working in the service of the Queensland Museum. He
worked with a small modern comparative collection and, evidently, a paradigm that most fossil
material must represent extinct avian taxa. His material came mainly from the Darling Downs
of Queensland and Cooper Creck in South Australia, the latter collected either by H. Y. L
Brown or J. W. Gregory (1906) and his group of Melbourne University students in the early
20th century. Many of these "new" taxa have been synonomized with extant forms (Rich &
van Tets 1981, 1982, Rich, van Tets & McEvey 1982, Rich et al. 1986, van Tets & Rich
1987), but it was his work that painted the first pictures of the fossil birds from central
Australia, most of which were of Quaternary age.

In the latter part of the 19th century, the spectacular discovery (Newton 1893) of fossil
vertebrates at Lake Callabonna in eastern South Australia certainly led to increased research on
avian fossils. The first articulated skeletons of any fossi} Australian birds, in this case the
dromomithids, were found bogged in the Pleistocene clays uf the lake. A series of well written
and superbly illustrated papers by E. C. Stirling and A. H. C. Zietz (1896-1913) clearly
outlined this endemic Australian group that is unrepresented in the modern fauna.

After such a burst of activity, little transpired in the field of Australian palaeornithology
until the mid-20th century, except for a few papers on Quaternary fossils, such as those from
the Bass Strait islands or Tasmania (Milne-Edwards & Oustalet 1899, Giglioli 1907, Legge
1907, Spencer & Kershaw 1910, Anderson 1914, Dove 1926, Howchin 1926, Morgan &
Sutton 1928, Jouanin 1959) or such prolific localities as the Cuddie Springs bone bed in New
South Wales (Anderson 1889), which produced a variety of giant marsupials as well as
dromornithids, all disarticulated. Chapman (1910) mentioned a fossil feather from Western
Victoria, which may only be of recent derivation. Finlayson (1938) reported on a penguin

MESOZOIC & TERTIARY BIRDS OF AUSTRALIA - 723

from the Miocene of South Australia. Many of these early discoveries were reported in
Lambrecht's survey of palaeornithology in 1933.

In the early 1950s, R. A. Stirton, R. H. Tedford, Paul Lawson and their colleagues mainly
from the University of California (Berkeley) and the South Australian Museum, discovered
Tertiary vertebrates in northern South Australia (Stirton 1955, Stirton, Tedford & Woodburne
1967). They also visited some of the original J. W. Gregory sites along Cooper Creek and
discovered several more. Their collections added great time depth to the history of birds, as
well as for other vertebrates, on the Australian continent. It also served as a tremendous boost
for research in vertebrate palaeontology on this continent.

Alden Miller was one of the members of the mid-twentieth century Stirton expeditions, and
he energetically began describing the new material deriving from a series of University of
California-South Australian Museum expeditions (Miller 1962-1966). His untimely death cut
his research short, but a student of his, Patricia Rich, continued the work he had begun (1975-
1982). She was later joined by A. R. McEvey (Rich & McEvey 1977, 1980), G. F. van Tets
(Rich & van Tets 1980-1985, Rich, van Tets & McEvey 1982, van Tets & Rich 1987), R.
F. Baird (Rich & Baird 1986, in press, Baird & Rich, in press), C. Patterson (Patterson 1983,
Patterson & Rich 1987), T. H. Rich (Rich e¢ al. 1982) and W. Boles (in press) in describing
the bulk of the Stirton material.

In the 1950s and 1960s concurrent with the work by Stirton, Edmund Gill (National
Museum of Victoria, now the Museum of Victoria, Melbourne) described fossil penguin
material from Victoria (1959a,b) as did G. G. Simpson (American Muscum of Natural History,
New York) (1957-1970). In 1969 Ron Scarlett (Canterbury Museum, Christchurch) reviewed
an Australian "moa" described by de Vis, finding it to be conspecific with a New Zealand
form, and, thus, suggested that it had not really been found in Australia. Stirton's colleagues
continued to discover more Tertiary bird-bearing sites, such as those at Alcoota (Woodburne
1967) in the Northern Territory (worked by M. O Woodburne and J. Mawby in the 1960's),
Bullock Creek, also in the Northern Territory (Plane & Gatehouse 1968) and Riversleigh in
northwestern Queensland (originally reported on by Tedford in 1968 but currently being
excavated and studied under the direction of Dr Michael Archer, Univ. of New South Wales).

During the 1970s and 1980s field recovery and research on fossil birds expanded greatly.
Rich & van Tets (see van Tets & Rich 1987 summary), as well as Storrs Olson (National
Museum of Natural History, Smithsonian Institution, Washington) (1975-1977), have
reviewed de Vis's fossil birds. Other workers active during this period include Allan McEvey
(Museum of Victoria; now retired), who has studied Cainozoic fossils and the morphology of
extant taxa, Edmund Gill (deceased, formerly of the Museum of Victoria), who worked on
fossil footprints (Rich & Gill 1976), and Dominic Williams (deceased, formerly of Australian
National University and Flinders University), fossil eggs (this volume, Williams & Rich in
press). Williams (1981) was the first to report on dromornithid egg shells from the Quaternary
of South Australia. Robert Baird (Earth Sciences Department, Monash University,
Melbourne), Charles Meredith, Greg McNamara (James Cook University, Townsville) have all
been students of P. V. Rich. Baird's work (Baird 1986) on the cave faunas (with emphasis on
the passeriforms) of the southeastern part of Australia has been an important contribution
demonstrating the usefulness of fossil avifaunas in charting changing climatic regimes through
time and understanding the factors that bias avian cave assemblages. Charles Meredith (1985)
analyzed the Holocene fossil avifauna of Norfolk Island emphasizing seabird evolution in the
Pacific. Van Tets has also been active in description of fossil seabirds from Australasia as well
as cave and archaeological faunas (e.g. van Tets & Smith 1974, van Tets & O'Connor 1983).
Greg McNamara studied the Quaternary avifauna of McEachern's Cave (McNamara 1981) in
Victoria, and Chris Patterson (1983) the fossil history of the emus on the Australian continent.
Shane Parker (South Australian Museum) has discussed the dwarf emus of Kangaroo and King
islands south of Australia, as well as the dromaiines of Tasmania.

724 - RICH

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17 (Ngama Local Fauna) M.Mio 15 & ~~ R
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18 (Yanda Local Fauna) M.Mio | 45 ail R
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19 (Ericmas Local Fauna) 15 | - LR
Lake Pinpa, Billeroo Ck., S.A. 11-
20 (Pinpa Local Fauna) 15 7~ M
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21 (Ngapakaldi Local Fauna) 15 & Lon! A
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26 Koonwarra, Vic. E.Apt 1/110 & He M
27 Hamilton Hotel, Qld Albian hi0 & R
Tabie 1. Distribution of birds in pre-Quaternary sediments of Australia. Method of Dating: 4 :
palynology; om , vertebrate fauna, mainly diprotodontid marsupials; @ , microinvertebrates and/or
macroinvertebrates; y*, , radiometric dating; [+] , fission track dating. Depositional Environment: F,
fluviatile; L, lacustrine; P, paludal. Listed in order of abundance of sedimentary types. Quality of

Preservation: E, exceptional; G, good detail preserved but bones often fractured and not complete; P, poor.

MESOZOIC & TERTIARY BIRDS OF AUSTRALIA - 725

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large (Emu-sized or larger, O , average; e, small (songbird-sized). Abundance of Fossil Material:
A, abundant (more than 50 bones known; M, moderate (10-49); R, rare (less than 10 bones); S, single
bone only. Number of Taxa: Number indicates number of taxa (species, etc.) recognized currently.

726 - RICH

Walter Boles (Australian Museum, Sydney) is currently studying the avian material from
the Tertiary-aged Riversleigh deposits (Boles, in press) in northwestern Queensland as well as
others from central Australia. Natalie Schroeder (Monash University) and Michael Plane
(Bureau of Mineral Resources, retired) have both carried out research on New Guinea fossil
avifaunas (Rich, Plane & Schroeder 1988). Peter Murray, Michael Plane and P. V. Rich have
all been involved in describing new Miocene material from the Bullock Creek site in the
Northern Terrtitory.

Palaeormithology in Australia is currently the most vibrant that it has ever been. Now,
instead of one or two active workers, there are many. Now, instead of simply describing the
fossils to widen the data base (although careful descriptions are certainly still vitally essential),
a great deal of interpretive analysis is underway, ranging from bioclimatic analyses (Baird 1986)
to functional morphological studies (Rich, McEvey & Baird 1985) and the biogeographic
implications of the total continental avifauna. Together with the greater number of workers
involved in palaeornithological studies in Australia and the interpretive techniques that are
being applied there has also been a great expansion in field operations over the last decade. The
next decade will certainly see a marked increase in the knowledge of the fossil birds on the
Australian continent.

TEMPORAL DISTRIBUTION OF AVIAN FOSSILS IN
AUSTRALIA AND NEW GUINEA

Unlike the rich tapestry of birds recorded in the Quaternary caves of Australia's south and
east, the pre-Quaternary record is poor (Rich & Baird 1986). Table I summarizes the sum total
of late Mesozoic and Tertiary fossil bird-producing sites. When compared with the records of
North America and Europe, fossil bird producing localities in Australia, and especially in New
Guinea, are few and far between and most localities have produced quite small avifaunas. There
are exceptions to this rule, however, such as the Miocene-aged sites at Lake Palankarinna in
northern South Australia and Riversleigh in northwestem Queensland.

One other aspect of the fossil bird record from Australia is that it is quite lengthy. The
oldest fossil birds on the continent are of Early Cretaceous age, probably Aptian (app. 120
myBP) (Dettman 1986). The Strzelecki Group lacustrine sediments of the Koonwarra locality
in southeastern Australia have produced a number of feathers (PI. 1) (Talent, Duncan &
Handley 1966, Waldman 1970) associated with a rich fauna and flora. Even at such an early
date in avian history, then, birds were already quite widespread, far from the location of the
oldest bird-like vertebrates reported from the Late Jurassic lagoonal sediments of southern
Germany, and they were inhabiting polar latitudes that may have been as far south as 85° with
near freezing temperatures at times (Rich & Rich 1989).

Lengthy though the record is, however, it is characterized by great gaps. No fossil birds are
known from the Albian until the Eocene, and then only a few isolated bones of penguins have
been recovered in both Eocene and Oligocene deposits. The first diverse arrays of forms occur
in the Late Oligocene to Early Miocene, and then the record is mainly from two geographic
locations, central and northern Australia.

Several localities in northeastern South Australia (Lake Kanunka, Lake Ngapakaldi, Lake
Pitikanta, and Lake Palankarinna in the Lake Eyre Subbasin; Lake Pinpa and Lake Yanda in
the Tarkarooloo Subbasin) (Fig. 1) have produced fossil avifaunas that range in age from
Oligo-Miocene to Pleistocene, dates based primarily on palynology (Rich & Baird 1986).
Most of this material occurs in fluviatile deposits (Table 1) and is disarticulated and
disassociated due to the fact that the bones have experienced transport of varying distances after
wie death of the birds.

MESOZOIC & TERTIARY BIRDS OF AUSTRALIA - 727

| ALICE
SPRINGS

ADELAIDE

MELBOURNE *~_
{ .

11
oe

Figure 1. Localities producing Mesozoic and Tertiary fossil birds in Australia. Black circles, Mesozoic;
white squares, Eocene; white circles, Oligocene; black squares, Miocene (or Oligo-Miocene); black triangles,
Pliocene. Numbers refer io those used in Table 1.

A second set of localities are known from northern Australia, Riversleigh in northwestern
Queensland and Bullock Creek in the northern part of the Northern Territory (Fig. 1).
Preservation at these sites occurs in sediments deposited by lime-rich streams that coursed
across a limestone plateau in the mid- to late Tertiary. This resulted in the bones being
cemented into solid limestone, evidently only a short time after burial. Although most of the
material, like that from South Australia, has been transported after the death of the birds, the
preserving medium is such that little post-depositional alteration has occurred. Vertebrate

728 - RICH

material is removed from these limestones by soaking in a dilute solution of acetic acid
(Whitelaw & Kool, this volume); what remains after preparation is uncrushed material with
exceptional detail preserved, Bullock Creek, for example, has yielded the only known good
cranial material of the dromornithid birds. The Riversleigh localities, likewise, are producing
an array of avian materials with exquisite preservation. Over the next decade or two with the
further collection and processing of material from these sorts of sites, our knowledge of the
mid-Tertiary radiation of birds on the Australian continent is bound to dramatically increase.

Alcoota, another locality in northern Australia has produced literally tons of bones, a large
portion of which are dromornithid birds (up to 60-70% of the total excavated biomass, which is
a similar situation to that at Bullock Creek). Alcoota differs from Bullock Creek, however, in
the quality of preservation; the material is preserved in the fluviatile and lacustrine sands and
clays of the Waite Formation. These are not lime rich. The bones, although abundant, are
quite often crushed and fragmented, and very difficult to excavate and prepare, and even more
difficult to restore to their original three-dimensional form.

A real difficulty with all of the localities discussed above of Tertiary age is that the dating
is only approximate, not absolute. Ages for such sites have been assigned by using contained
vertebrate remains, mainly diprotodontid marsupials (see Rich, T. H. et al., this volume). The
stage of evolution or position in a phylogenetic sequence has been used to produce a relative
date of one site to another. Some sites, such as Beaumaris in coastal Victoria, contain both
diprotodontids and marine invertebrates which in turn can be related to invertebrate faunas in
rock sequences elsewhere in the world that have associated absolute radiometric ages.
Beaumaris is considered to be about 5 million years old, near the Mio-Pliocene border, based on
its contained marine invertebrate fauna which includes both macro- and microfauna. The
diprotodontids that are known from Beaumaris seem to be more advanced than those found in
the centra! Australian sites previously discussed, which could mean that those Iccalities were
older than 5 million years. But what of relictual distributions, which we know do occur today?
It could be that pockets of more primitive diprotodonts survived in some areas (such as central
Australia?) longer than they did elsewhere. We cannot rule out that possibility at present, until
we have a much better record. However, as a first approximation, as a working hypothesis,
most terrestrial biostratigraphers make the assumption, until they have evidence to the
contrary, that such evolutionary change in the diprotodontids was instantaneously recorded
continent-wide.

A second avenue for dating the central Australian sites, particularly those in northeastern
South Australia, has been the used of pollen and spores. These microfossils sometimes occur
associated with the vertebrate faunas themselves, such as at Lake Palankarinna, or in well cores
that sample facies lateral to the vertebrate-bearing sequences. Similar microfossils occur in
marine sequences in southern Australia, which in turn contain marine microfossils that can be
used to tie the Australian sequences with those elsewhere in the world, namely Europe. Most
of the palynological correlation has been carried out by W. K. Harris, and is reported on in
unpublished South Australian, Department of Minerals and Energy reports. Harris concluded
that the age for many of the Lake Palankarinna sites was Miocene, which gives some support
to the evidence based on diprotodontid marsupials. T. H. Rich er al., in this volume, details
the support for such dating.

A third method just recently being applied is an attempt to separate minute particles of
volcanic ash or other volcanic airfall particles from the lime-rich sediments of Bullock Creek
(T. H. Rich, pers. comm.). If this can be done, an absolute date on sediments containing
vertebrate fossils may not be impossible.

Most of the localities that have produced birds in Australia are isolated spots. They do not
occur in a sedimentary succession where one physically occurs above or below another. Lake
Palankarinna in northern South Australia is one of those rare areas where superposition of
terrestrial vertebrate faunas can be demonstrated. Here at least one can be sure of the sequence

MESOZOIC & TERTIARY BIRDS OF AUSTRALIA - 729

of occurence of avian palaeofauna, keeping in mind the complex cut and fill stratigraphy that
characterizes this fluvio-lacustrine system.

Absolute dating yet remains a significant problem to determining Australia-wide avifaunal
evolution during the Mesozoic and Tertiary. Only three localities have reliable absolute dates
assigned to them: Koonwarra, Bugaldi and Boat Mountain. Koonwarra, the oldest record of
birds on the Australian continent, has been dated by using fission track techniques to between
120-110 million. This date has been refined palynologically to about 119 myBP (Dettman
1986). A second locale, Boat Mountain, containing some bird material has been dated as 54
myBP using potassium-argon dating. A third locality, dated as medial Miocene, yielded a
skeleton of a primitive aegothelid, Quipollornis (Rich & McEvey 1977). Volcanic lavas
above the fossiliferous diatomites were dated using the K-Ar technique as between 13.5 and 17
myBP, giving a minimum age.

A number of localities producing birds in Australia have been called Pliocene. Only one of
these has an absolute date associated with it, the Bluff Downs site (Archer & Wade 1976) near
Townsville in eastern Queensland. Dates of between 4-4.5 myBP were determined for a basalt
flow thought to be the same as that overlying the fossiliferous Allingham Formation, which
contains the bird bones. Further dates need to be determined on rocks that can be shown to
directly overlie the fossiliferous beds before this date can be accepted. One further locality,
Awe, in Papua~-New Guinea, has an absolute date associated with it from which cassowary
fossils have been reported. Page & McDougall (1972) gave a minimum of 3.1-3.8 myBP for
this locality based on igneous intrusions that affected the fossiliferous sediments. Dating based
on the included marsupial fossils does not contradict this.

GEOGRAPHIC DISTRIBUTION OF AVIAN FOSSILS IN
AUSTRALIA AND NEW GUINEA

Localities producing fossil birds in Australia and New Guinea are not evenly spread; there
are concentrations of sites in northern South Australia and in the northern part of Australia;
there are ghastly blanks in other areas, such as Western Australia. Most sites occur on the
eastern half of the continent (Fig. 1, Table 1). The Bullock Creek site in the Northern
Territory is the most northerly at 17° S$. The most southerly site has produced only footprints
of Oligocene age, the Endurance tin mine near South Mt Cameron in northeastern Tasmania at
41° S.

One locality, Koonwarra, has produced most of Australia's Mesozoic birds. It occurs in
southeastern Victoria, near the present coast. The extent of outcrop of rocks of similar age
(Rich et al. 1988, Rich & Rich 1989) in this area of Australia may eventually yicld more avian
material, and it is certainly the most probable spot to expect further material in the future.
Molnar (1986) has recently reported a second Mesozoic locality at Slashers Creek, near
Hamilton Hotel, western Queensland.

Eocene and Oligocene localities are mostly restricted to the southeast of Australia in
southeastern South Australia, southern Victoria and northeastern Tasmania. Most of the fossil
remains are penguins or other seabirds except for the Tasmanian occurrence of dromornithids.
One new locality of probable Eocene age is that of Redbank Plains near Brisbane, Queensland.

Miocene localities have the widest geographic spread and have produced the most diverse
pre-Quaternary avifauna in Australia. Sites of this age (Fig. 1, Table 1) occur over the eastern
half of the continent from southern Victoria to the northern part of the Northern Territory and
Queensland. The furthest east is the Bugaldi locality in nortneastern New South Wales.

730 - RICH

Pliocene localities are less numerous and do not have quite the geographic spread of the
Miocene site. Pliocene locales often occur associated with Miocene sites, such as at Lake
Palankarinna and Lake Kanunka in northern South Australia and at Riversleigh in northwestern
Queensland. In the case of the South Australian sites, direct superposition of sites can be
demonstrated. Several isolated sites thought to be of Pliocene age also occur in eastern
Australia: Allingham Creek, Peak Downs and Chinchilla in Queensland, Bow in eastern New
South Wales and Fisherman's Cliff in western New South Wales, and Morwell in southern
Victoria.

The only pre-Quaternary avian fossil locality in New Guinea is Awe in Papua-New Guinea.
This site has produced only a few fragments of cassowary material. The record from this
extremely interesting area is a blank slate, however.

BIASES OF THE MESOZOIC AND TERTIARY AVIAN RECORD
IN AUSTRALIA AND NEW GUINEA

The fossil bird record of Australia and New Guinea pales before that of North America and
Europe. There are a variety of reasons for this, some of which may eventually be diminished,
but others will always remain. These factors that bias the Australian avian record include:
degree of exposure of rocks of appropriate age, degree of post-depositional alteration, type of
original depositional environment, accessibility of rock outcrop, number of trained personnel
to prospect, collect and study material, availability of funds to support palaeornithological
exploration and research, and types of techniques applied in collecting and preparing material.

One factor that will always bias the Australian fossil record is that Australia lacks much
topographic relief. This is not a problem in New Guinea, but a severe one in Australia. The
tallest mountain in Australia is only 2228 m in height. This means that because of the lack of
uplifted rock sequences, there is little exposure of rocks that could, through continued erosion,
bear fossils. Compare this to western North America or Europe, or even Asia and South
America, where extensive chains of mountains and plateaus expose thousands of metres of
rocks. Australia has no chance of competition with such land masses. Further inhibiting the
exhumation of potentially fossil bearing rocks is that many of those areas where topographic
relief exists in Australia, also experiences high rainfall. This results in dense vegetation cover,
further inhibiting the exposure of bare hillsides. Only in areas such as central and western
Australia is there a combination of some topographic relief and aridity, which results in rock
and sediment exposure. In both of these cases, yet another problem rears its head: the rocks
are of the wrong age to contain fossil birds. In both areas the rock sequences exposed are
primarily Archean, Proterozoic, or Palaeozoic. Birds did not appear until the mid- to late
Mesozoic. Only in northern South Australia and parts of the Northern Territory and western
Queensland are the conditions right for some outcrop of the right-aged sediments, but in all
these cases, topographic relief is limited to at best a few hundred metres and often much less,
frequently only a few tens of metres along the western sides of playa lakes.

Chemical weathering of Mesozoic and Tertiary sequences has been extensive and disastrous
from a palaeontological point of view. During much of Australia's history since the
Carboniferous, the continent has remained far south attached to Antarctica (see Frakes & Rich,
this volume). This has positioned the continent in a high rainfall regime for much of its pre-
Pliocene history. High rainfall results in high watertable and thus passage of ground water
through near surface sedimentary sequences. Bones and shells are often dissolved as a result.
Thus, although bones may have originally been preserved, they are often postdepositionally
destroyed geochemically. What remains behind are the red iron oxides and silica that so
characterize much of Australia. This factor, like that of low topographic relief, will not go

MESOZOIC & TERTIARY BIRDS OF AUSTRALIA - 731

Carpal phalanges Cranium
RG Carpometacarpus

eed
SAN

i
tS Quadrate
* Mandible
Ulna %,
~ Humerus us Cervical
vertebrae
Caudal \ ENE Digs 73

vertebrae

Pygostyle Coracoid
% R MN “lan >, RY : Y %
Wee S y
sous Yr Clavicles
Ys
V/ Y
ternum
Tibiotarsus
ff Tarsometatarsus
LM SEN
SS iy PN ea

=

Skeleton of a bird indicating the susceptibility of bones to wear in stream environments. a
nape ie humerus and tarsometatarsus, are the most resistant to wear; parallel-lined bones are oO
hite bones are very fragile and tend to erode very quickly, thus not often being

Fig
bones, such as the n
intermediate durability; w
preserved as fossils if transported any distance.

732 - RICH

away even with the concerted effort of the palaeontologist, and thus will always bedevil the
quality and quantity of the Australian fossil record.

Another biasing factor is that of the enclosing sediments. Bones are better preserved in
some environments than in others. Ideal are lacustrine deposits which preserve material in situ
without moving fossils from point of death. Material preserved under such conditions is often
whole or nearly so, such as the aegothelid skeleton from the Miocene Bugaldi site in New
South Wales or the Genyornis skeletons preserved just as they were bogged on the margins of
Pleistocene Lake Callabonna in South Australia. Some associated material of flamingoes and
palaelodids has been recovered from Miocene lacustrine sediments at Lake Palankarinna.
Fluviatile sediments, however, seem to be the most frequent preservers of fossil bird bones in
the Australian Tertiary. Streams tend to concentrate bones in various ways (Voorhies 1969,
Wolff 1973, 1975, Behrensmeyer 1975, Baird, this volume) but in doing so both disarticulate
and erode the transported bones. This must be taken into account when comparing such
occurrences with others, both in Australia and elsewhere, and it also explains to some extent
why the Australian record is so limited. Obviously, those bones which can withstand wear
best (Rich 1980a, Napawongse 1981, Rich & Baird 1986), such as humeri, tibiotarsi,
tarsometatarsi and coracoids of medium-sized (Fig. 2), but not small or very large, birds are
most frequently preserved. Others, such as cranial remains, delicate bones or those with a large
surface to volume ratio, such as wing phalanges, scapulae, radii, fibulae, and pelves, are rare.
Some localities, such as the Miocene Tom O's Quarry in the Tarkarooloo Subbasin and the
Pliocene-aged Hamilton locality of Victoria, contain no bird bones at all, and the bones of
other vertebrates clearly show that lengthy transport is involved. The fragile bird bones that
most likely were present in the stream load were totally destroyed during transport, whereas
mammalian teeth have remained intact, but extremely eroded. Tumbler experiments designed to
simulate a variety of stream environments like those in the mid-Tertiary of central Australia
show the same representation and erosion patterns (Figs 3-6) as occurs in the fossil
assemblages from those areas (R, Berra, P. Napawongse & P. Rich, unpublished data,
Napawongse 1981).

Such biasing, as mentioned above, also determines the size of species best preserved. There
is not only a selection against small, delicate bones that are part of a larger bird, but also
against entire skeletons of small birds. So, the record is automatically biased toward medium-
sized birds (e.g. ducks, flamingoes, burhinids) or larger birds (emus) and excludes smaller ones.
Such an influence also means that relationships of isolated bones are not always understood, so
proportions of limb elements within one individual skeleton cannot be determined with
certainty. Sometimes different disassociated elements can be assigned to a single taxon because
of the common occurrence at a single locality, common identity to a single family, and
similarity of size. Usually, however, one cannot be certain of the association to one
individual, so mean measurements must be utilized.

Fluviatile fracturing of bones also means that very often proximal and distal ends are not
associated. Trying to associate fragments can be difficult. It is only because of such localities
such as Lake Palankarinna, Bullock Creek and Alcoota, where associations of avian material
can be demonstrated that association of the isolated, out-of-context fragments from other
localities can be made.

The kinds of depositional environments also determine the kinds of animals that will be
preserved - namely those that lived close to the environments where deposition was occurring.
In the Tertiary sites of central Australia, for example, waterbirds are by far the most common
(see Table 1). Ducks, flamingoes, rails and charadriiforms are far more common than are
pigeons and parrots. This probably bears no relationship to the actual proportions of species
that inhabited and died in central Australia 10 to 20 million years ago. It is, however, related
to the probability that birds living, feeding, and nesting near or in rivers or lakes are more
likely to die there and to be preserved than those living elsewhere. Because of such biasing

MESOZOIC & TERTIARY BIRDS OF AUSTRALIA - 733

ELEMENTS

FEMUR

DEEP

ORIENTATION OF LONG LIMB BONES IN FLUME EXP.
SHOWING EFFECT OF DEPTH OF WATER

POLARITY
(Parallel to
current)

TIBIOTARSUS

TARSOMETATARSUS

HUMERUS

ORIENTATION:

Transverse

Parallel

MOVEMENTS:

=a :
-——J Rolling

Continuous

| Intermitent

RADIUS

SKULL

PELVIS

INCREASING VELOCITY

F Finite

0.016

STERNUM

DIGITS

FURCULA

CORACOID

SCAPULA

HUMERUS

ULNA

RADIUS

CARPOMETA —
CARPUS

FEMUR

TIBIOTARSUS

TARSOMETA —
TARSUS

Figure 3. Effect of moving water at different speeds and at different depths on the orientation of bones

(After Napawongse 1981).

734 - RICH

factors, one should always be cautious of interpreting low representation or absences. For
example, both the parrots and the pigeons have a very dismal record in the Tertiary of
Australia. In this case, the fossil record should not be used as evidence of a fairly recent
radiation or invasion of the Australian continent by either of these groups. Rather, there is the
real possibility that low numbers are directly related to the non-fluviatile and non-lacustrine
habitats that these birds prefer.

100 90 8c 70 60 50 40 30 20 190 9
= = AQACE—RK>—>pz:< qv AK Femur
| S —— Radius
Pachyptila \ \\ Una — WINGS
belcheri | \ feline
NN Digit
=z << Synsacrum PELVIS
ra CDWyYy NE Si P. GIRDLE
& \ Gecees — STERNUM
= \\) Mandible SKULL
=
Zz
z Fg <Ce me
oO ibiotarsus athe
od RM Q uu uuwWwDW \\ Tarsometatarsus —
ag KX GG \l|§ L LA NY Humerus —
o cea ale : — cee WINGS
arpometacarpus
: \ Syreacrom ——_—PELMS
& \Seraears P. GIRDLE
WW SJ Furcula — ;
a
L BRA i HIND
——— Tareomstataraig Tae
eC Humerus
Coturnix WY Ves | wines
japonica \N ire
— Synsacrum PELVIS
QM Gh’ Coracoid P. GIRDLE
QQ Anterior end ——— STERNUM
IMDM UNNNKQAAY Mandible ————— SKULL

Figure 4. Erosional effect of moving water (simulated fluviatile environment) on bones of two species of
birds, a seabird (Pachyptila belcheri) and a terrestrial bird (Coturnix japonica). The horizontal scale represents
the hours bones were tumbled in a water-moderatedly sorted gravel mixture (mean grainsize -3.18 phi). Bones
could be recognized for the period indicated by the hatched column. (After Napawongse 1981).

Yet another of the many biases is the accessibility of the rocks that might actually contain
fossils. Many of the most productive of the avian fossil sites are in very remote areas. Most
of these sites were discovered only in the 1950s or more recently. There are still many parts of
central Australia and Western Australia that need either initial prospecting or further detailed
prospecting to capitalize on initial discoveries. One should realise that Australian
palaeornithology is still in its infancy. Even though its record will most likely never match

MESOZOIC & TERTIARY BIRDS OF AUSTRALIA - 735

that of North America and Europe, it still has vast areas in remote places that are in great need
of exploration and development.

SKULL STERNUM PECTORAL VERTEBRAE RIBS PELVIS WINGS HIND
GIRDLE LIMBS

100%
, a
belcheri
(Procellariiformes,
small sea bird)
ie)

TIME ———»

T
Coturnix
| japonica
{ (Galliformes,
| | a small land bird)
fe] Le L aa]

TIME

Figure 5. Graph showing how quickly bones differentially wear in a fluviatile environment. In this
tumbler experiment where the matrix was gravel (mean grainsize -3.18 phi) birds, one a seabird with densely
ossified bone is compared with a less densely ossified terrestrial bird. Percentage of original material is
calculated at the end of each of 10 runs (approx. 150 hr./run, for a total of 1650 hours). (After Napawongse
1981).

The degree to which a continent or an area is developed is related to many factors. In
Australia, besides the remoteness of many of the possible areas for fossil vertebrate localities,
there is a shortage of trained vertebrate palaeontologists. The population size of Australia
places a limit on the number of positions that can be occupied by professional
palaeontologists. This is to a large extent controlled by availability of funding, which, of
course, is directly related to the population base providing the funding. It is also related to the
philosophies that the population and its representatives in government have towards funding of
the sciences. Australia has in the past and even today underfunded scientific research when
compared to countries like the United States. Traditionally, many new discoveries on this
continent have been made by people not professionally engaged in palaeontological research.
Graziers, tourists, mapping geologists are only a few of the many sources of new vertebrate
palaeontological finds. Likewise, both funding and professionals from overseas have been
critical to new discoveries on this continent, the most spectacular example being R. A. Stirton
from the University of California (Berkeley), who discovered the first diverse assemblages of
pre-Quaternary marsupials and birds in central Australia in the 1950s (see R. H. Tedford, this
volume). He also trained many students and rekindled an interest in vertebrate palaeontology
on this continent, which still has a strong influence on the field.

Fortunately, the bias provided by low levels of funding has been subdued during the past
decade. The Australian Research Grants Scheme, now under the umbrella of the Australian
Research Council, has provided some sizeable grants in support of vertebrate palacontology,
especially benefiting palaeornithology. Universities and museums have given support, as have
a number of organizations associated with such institutions (e.g. Friends of the Museum of
Victoria, The Australian Museum Society, etc.). Private industry and friendly societies have
likewise come to the aid of palaeontological research, examples including the National
Geographic Society, Alas Copco (a Swedish mining equipment company), Esso, Kelloggs,
the Ingram Trust, the Ian Potter Foundation, Mobil Oil, Dick Smith, Shell, Safeway, Coles
New World, CRA, ICI, Western Mining, Ingersoll-Rand, Utah Mining, Danks Trust, the
Myer Foundation, Qantas, Ansett, Australian Airlines, David Holdings, McDonalds, to

736 - RICH

mention just a few. With continued support at current levels, the future seems bright for
advances in the next decade.

SKELETAL ELEMENTS GROUPED ACCORDING TO THEIR CHARACTERISTIC
SUSCEPTIBILITY TO TRANSPORT

GROUP |: IMMEDIATELY GROUP II: ReEMoveED GROUP Ill: LAG DEPosit

REMOVED, TRANSPORTED BY GRADUALLY, TRANSPORTED BY

SALTATION OR FLOATATION TRACTION

RIB PELVIS
VERTEBRA STERNUM
MANDIBLE
CORACOID
SCAPULA
HUMERUS

ULNA

RADIUS

CARPOMETA-
TARSUS

FEMUR
TIBIOTARSUS

TARSOMETA-
TARSUS

MICHEAL VOORHIES'’ DISPERSAL GROUPS

RIB FEMUR SKULL
VERTEBRA TIBIA MANDIBLE

SACRUM HUMERUS
STERNUM METAPODIAL
PELVIS

RADIUS

Figure 6. Classification of avian skeletal elements according to their susceptibility to transport in a
fluviatile environment. Results summarized in this figure are based on simulated fluviatile environments in
tumbler experiments. (After Napawongse 1981).

A final set of factors which can bias the fossil record relates to how material is processed
once it is actually located. Collecting itself can bias samples even though fossil material may
be abundant in the sediments. In soft sediments, such as sands, silts, and clays, the only way
to sample completely is to first surface collect, and then follow this up by careful, controlled
(often mapped) excavation, which may be carried out in the first instance with small tools. All
the matrix (sediment) that results from such an excavation needs then to be wet-screen washed
(McKenna 1962, Whitelaw& Kool, this volume). If wetscreening, as opposed to dryscreening,
is not carried out, then a significant part of the smaller vertebrates could be lost. If the matrix
surrounding the bones is hard and calcareous, as in the case at Riversleigh and Bullock Creek,
ihen mechanical preparation with dental drills and related tools is not sufficient. Acid etching

MESOZOIC & TERTIARY BIRDS OF AUSTRALIA - 737

(Whitelaw & Kool, this volume) is the only method that will result in the recovery of the total
death assemblage (thanatocoenose). This technique removes all of the adhering matrix, leaving
mainly bones, both large and small. Both Bullock Creek and Riversleigh are excellent
examples of how a technique can influence the faunal lists. Both sites have been known for
more that two decades, but until the early 1980s most material from these sites represented
medium-sized or large animals. With the use of acid etching since that time, the faunal
diversity from both sites has exploded, especially for small vertebrates. It is important, then,
to understand the methodology used in any study that leads to the conclusions presented. How
an area was prospected, excavated, and then what methods were applied to prepare, sort, identify
and, in the end, study the material can have a marked impact on the final decisions about the
identity and diversity of the fauna. Time, technique and philosophy can all play a part.

THE AVIAN FOSSIL RECORD (AUSTRALIA-NEW GUINEA):
MESOZOIC

Two localities have produced the sum total of Australia's Mesozoic birds, one in Victoria
and a second in Queensland. Koonwarra in southeastern Victoria has yielded a variety of
feathers in the last two decades (PI. 1) (Talent, Duncan & Handley 1966, Waldman 1970), but
unfortunately no bony remains. Although clearly feathers, detail on these specimens is not
well enough preserved to allow taxonomic assignment to any particular avian group, despite a
recent attempt to do this (Rogers 1987). The Koonwarra locality is important in establishing
that birds had reached Australia by Early Cretaceous times (?Barremian to Aptian times based
on contained pollen; Dettman 1986). This occurrence is important in a second way, in that the
locality lay at very high latitudes (Rich, et al. 1988), about 75-85° S. This means that birds
were capable of utilizing such high latitudes, whether by migration or permanent residency.
They were not restricted to the lower, perhaps more benign areas.

At least 5 feathers, some with counterpart impressions, have been recovered from the
Koonwarra locality, one of which appears to have been a down feather, and the others contour
or wing feathers (Rogers 1987). Despite a number of researchers examining the specimens,
insufficient detail has been preserved to allow identification beyond Aves. The carbon film that
represents the feathers is patchy and irregular in its distribution, and the specimens are
significantly compressed (Rogers 1987). This does not, therefore, preserve the fine detail
necessary for recognition of taxonomic subdivisions with the class Aves (Chandler 1916,
Rogers 1987). After detailed comparison across the Class Aves, Rogers (1987) concluded that
one feather studied in detail (Mus. Vict. P26059) could be distinguished from all modern orders
of birds. Thus, unless better preserved specimens or actual bony remains are recovered, it
appears that little else can be said about the Early Cretaceous birds from southern Australia.
All of the bird feathers were recovered from a quarry in lacustrine sediments of the Strzelecki
Group that has produced a great number of fish skeletons. It is not unlikely that in the future,
especially since a significant amount of field work is underway in the Strzelecki sediments and
the related Otway Group in the south central part of Victoria, that avian skeletal material will
be forthcoming (Rich ef al. 1988, Rich & Rich 1989).

The only other avian remains from the Mesozoic of Australia is a single tibiotarsus
recovered from the Early Cretaceous (Albian) Toolebuc Formation from Warra Station, on the
east side of the Hamilton River near Hamilton Hotel, west Queensland (Molnar 1986). The
enclosing limestone is of marine origin, and recovery of the single avian fossil was by use of
acetic acid digestion. Although only a single bone, the Queensland fossil Nanantius eos
appears to be related to the enantiornithines, known also from Mongolia, Mexico, and
Argentina. Occurrences outside of Australia are all from the Late Cretaceous (Campanian to

738 - RICH

Maastrichtian), so the Australian fossil is the oldest member of this subclass of birds (Molnar
1986). Nanantius was a small bird, about the size of the European blackbird (Turdus merula),
and the size is consistent with the size of birds indicated by the Koonwarra feathers, which
range up to 2-3 cm in length. It is impossible to decide at this point if the feathers and the
tibiotarsus of the Queensland enantiornithine originate from the same kind of bird.

The enantiornithine tibiotarsus is quite unique amongst vertebrates in having a distal end
with a very narrow intercondylar sulcus and the unequal development of the distal condyles, the
medial being elongate and the lateral narrow and about half the length of the medial. This
combination distinguishes the enantiornithines from pterosaurs, which have a more pulley-like
condylar arrangement where the condyles are more equal in development, from ornithischian
dinosaurs that lack a tibiotarsus, from theropod dinosaurs and non-enantiornithine birds where
the condyles are more equally developed (Molnar 1986).

THE AVIAN FOSSIL RECORD (AUSTRALIA-NEW GUINEA):
PALAEOGENE

The Palaeogene record of birds in Australia and New Guinea consists of dromornithid
trackways in the Late Oligocene of Tasmania and penguin bones from a few localities of Late
Eocene and Oligocene age.

Dromornithid tracks and trackways have been known for some time (Rich & Green 1974)
from the Endurance Tin Mine near South Mt Cameron in northeastern Tasmania (Fig. 2, Pls
2,3). The prints occur as isolated, single specimens as well as in trackways, varying in total
length from 147-240 mm in total length (Fig. 8). Ali prints show the impression of three toes
only with the middle (III) digit longest and the inner and outer digits of nearly equal length.
The terminal phalanges on the digits were apparently broad and gently rounded distally.
Originally the age of the clays present at South Mt Cameron were thought to be Early Miocene
in age, but recent palynological investigations suggest an Oligocene age (Hill & Macphail
1983). The vegetation near the site where the prints were made was dominated by species
typical of a closed, temperate rainforest dominated by Nothofagus johnstonii. Rich & Green
(1974) concluded because of the age and the size of the prin‘s that their identity was most likely
dromornithid, not dinosaur or dromaiine.

The remaining Palaeogene records of birds in Australia are of penguins (Finlayson 1938,
Glaessner 1955, Jenkins 1974, Jenkins et al. 1982, Rich 1975a,b, Rich & van Tets 1982)
from southeastern South Australia. Beginning in the early 1930s fossil penguin bones were
found in the Late Eocene Blanche Point Formation at Blanche Point southwest of Adelaide
(Jenkins et al. 1982). From the same formation at Witton Bluff, near Port Noarlunga, more
material was found (Jenkins in Rich & van Tets 1985). Some of this material (Fig. 12) came
from an unusually large penguin, a giant, that was originally described by Jenkins (1974) as a
new species, Pachydyptes simpsoni. Further research by Jenkins (1985 in Rich & van Tets)
indicated that, instead of a new form, Pachydyptes simpsoni was actually conspecific with
Anthropornis nordenskjoeldi, a form already known from the Late Eocene and Early Oligocene
sediments of Seymour Island in the Antarctic Peninsula (Marples 1953, Simpson 1971,
Zinsmeister 1982).

In addition to Anthropornis two other smaller penguins are known from Australia's
Palaeogene marine sediments. One form reported originally by Finlayson (1938) apparently is
closely related, if not congeneric, with Palaeeudyptes, a form known from Seymour Island as
well as New Zealand. A third, smaller form is now known from both Blanche Point as well as
pea cee Eocene rocks of Browns Creek, Otway Ranges, Victoria (Jenkins in Rich & van

‘ets 1985).

MESOZOIC & TERTIARY BIRDS OF AUSTRALIA - 739

By the end of the Oligocene large penguins were evidently no longer present in southern
Australia. Jenkins (in Rich & van Tets 1985) has suggested that the changes occurring in the
oceanic regimes of the Late Eocene and Early Oligocene brought about by marked climatic
deterioration and the increasing separation of Australia and Antarctica had a decided effect on the
penguin populations of the time (see also Fordyce, this volume).

THE AVIAN FOSSIL RECORD (AUSTRALIA-NEW GUINEA):

NEOGENE

With the beginning of the Miocene the avian fossil record improves markedly. Instead of
an isolated bone, feather, or footprint, or a partially articulated limb, varied assemblages are
known as are occasional nearly complete, articulated skeletons. Most of the Neogene material
has only recently been found, since 1952, and there is much room for discovery if some of the
known localities are worked again over a period of time.

The Ngapakaldi Local Fauna (see Rich, T. H. et al., this volume) is recorded from a
number of localities in northern South Australia, probably best known at Lake Palankarinna in
the Lake Eyre Subbasin. It contains the first diverse avifaunas (Table 1) which consist
predominately of waterbirds, e.g. flamingoes (Phoenicopteridae), the flamingo-like palaelodids
(Palaclodidae), ducks (Anatidae) and related anseriforms, and a variety of charadriiforms, best
represented by the thick-knees (Burhinidae). Represented, but rare, are pelicans (Pelicanidae),
hawks, eagles and kin (Falconiformes), rails and related forms (Gruiformes), pigeons
(Columbidae), and songbirds (Passeriformes). Several other local faunas are known from the
Tarkarooloo Subbasin (Pinpa, Ericmas, Yanda) and the Ngama Local Fauna from Lake
Palankarinna (summarized in Table 1) are of nearly equivalent age or slightly younger.
Together they add a number of avian families to the mid-Tertiary record including grebes
(Podicipedidae), cormorants (Phalacrocoracidae) and songbirds (Passeriformes). As Table 1
clearly illustrates the remaining Neogene record is characterized by the gradual occurrence
upsection of more and more families, but it is not until the Early Pleistocene that there is a
dramatic increase in diversity of forms represented. Such a burst of new appearances is directly
related to the quality and type of fossil record available. R. F. Baird, in two chapters of this
volume, discusses in detail the Quaternary record and the reasons for its richness in comparison
to the more ancient record of birds on this continent.

Of the fossil birds known in Australia today, only a few groups have both a good
representation through time and have been studied sufficiently to allow reconstruction of
evolutionary and biogeographic patterns over the past 20 or so million years. These include
the Dromornithidae, Casuariidae, Phoenicopteridae, Palaelodidae, Aegothelidae, and to a lesser
extent the Pelecanidae. As work progresses in the future, other groups, such as the
Phalacrocoracidae, Burhinidae, Anatidae and Rallidae, may provide useful phylogenetic
information.

The main problems in construction of such lineages have been, in part, mentioned above:
the material is often fragmentary, quite incomplete, and rare. Often taxa being compared are
not represented by comparable elements. There are also a great many hiatuses in the rock
sequences from site to site, and thus more or less continuous sequences, where gradual or
punctuate changes might be observable, are rare or non-existant.

Despite these problems, there are some intriguing absences of groups in certain geographic
areas that may be of some significance. For example, emus are absent in the oldest
fossiliferous sediments in both the Lake Eyre and Tarkarooloo subbasins. In fact, what is
present in the later Miocene in both the Kutjamarpu Local Fauna at Lake Ngapakaldi in the
Lake Eyre Subbasin and at Riversleigh in western Queensland is a casuariid that appears
intermediate between emus and cassowaries, lacking the specializations of the open country

740 - RICH

dromaiines (Boles, in press). The lack of cursorial adaptation may indicate that conditions
favouring such specializations were not yet dominant in Central Australia, thus differing from
today (Rich, Rich & Flannery, ms.). Neogene flamingoes and palaelodids are absent in most
of Australia, occurring only in Central Australia. They, like flamingoes of today, appear to
have been tied to the shallow, permanent alkaline lakes such as those that occur in the East
African Rift Valley today. In areas where such lakes did not occur, flamingoes, likewise, did
not occur. Similarly, with the disappearance of these permanent water bodies sometime in the
Pleistocene, the flamingoes and palaelodids became extinct in Australia. Other birds, such as
the Pink-Eared Duck (Malacorhynchus membranaceus) and the Freckled Duck (Stictonetta
naevosa), whose feeding habits in some ways parallel those of flamingoes, seem to have coped
with the disappearance of permanent shallow lakes by utilizing alternative sites in times of
drought and moving back into the interior lakes in times of wetter conditions. Flamingoes,
whose reproductive cycles depended on the constant presence of these sheltered lakes, failed to
survive. Another survivor that utilizes the salt lakes for breeding and feeding, the Banded Stilt
(Cladorhynchus leucocephalus), is able to delay breeding until the salt lakes are filled. These
birds appear to be more adaptable than the Australian flamingoes, and the palaelodids as well.

Absence and presence, as has been discussed in detail above, can very much be controlled by
the depositional environment and the habits of the birds. Parrots (Psittaciformes) and pigeons
(Columbidae) are both extremely diverse in Australia today, and yet their pre-Quaternary record
is almost non-existant. Because neither group is composed primarily of species that seek out
riverine or lacustrine conditions, their chances of being preserved in these types of
environments are slight. In fact, the record of these two groups on a world scale is poor.
Because of this bias, we know little of the evolutionary patterns of these two groups in
Australia, the area where they are most diverse on a world scale. It would be quite premature,
however, to conclude that because the record is poor, that the parrots and pigeons are recent
arrivals on this continent that have explosively radiated. In fact, we can conclude very little
about the evolutionary history of this group on a world scale. Quite the opposite is the record
of ducks. Their chances of preservation are excellent since they often live in close proximity to
aqueous environments, and their record is quite rich. This may have little to do with their
original diversity relative to such groups as parrots and pigeons, but a great deal to do with
their habitat preferences!

As a summary of the information available on fossil birds from the Neogene of Australia
and New Guinea, each family will be discussed in turn. Emphasis will be placed on the
biostratigraphic and palaeoenvironmental utility of each group. Appendix I presents
illustrations of the living relatives of most non-passeriform (and a representative passeriform)
avian families known from Australia. This appendix is for use in identification of fossil and
recent material.

THE DROMORNITHIDAE - MIHIRUNGS (Figs 7-8, Pls 2-19)

The dromornithids, the mihirungs, were a group of birds endemic to Australia that are now
extinct. Dromornithids first appeared in the late Palacogene (see above) and survived until at
least 26,000 years ago. These were large, ground-dwelling birds whose relationships are still
not well understood. They have previously been allied with ratites and galliform birds based on
both postcranial and quadrate morphology. The quadrate is certainly very un-ratite like (Olson
1986 and pers. obs.), Recent preparation of four fragmentary skulls from the Bullock Creek
site in the Northern Territory indicates that the basicranial structure is not typically ratite or
galliform. In fact, the skull structure is so highly derived, it is difficult to associate with any
known avian group. The Bullock Creek skulls show some similarity to living parrots (Pls

MESOZOIC & TERTIARY BIRDS OF AUSTRALIA - 741

OV WUT
“acyiss yy .

aU ee ;
wi ul eee £
*, y ra

i

Figure 7. Reconstruction (by P. Trusler) of the dromomithid Genyc.nis newtoni being attacked by the giant
goanna, Megalania prisca, both known from the Pleistocene of Australia.

742 - RICH

5,6), but this most likely is a reflection of functional similarities and not phylogenetic
affinity. Both the cranial material (Miocene and Pleistocene in age) and the lower jaws (from
the Pleistocene of Lake Callabonna, South Australia) (Pl. 4) are robust in the same fashion as
the Psittaciformes and Alcedinidae, but in detail are unique to the Dromornithidae.
Dromornithids appear to have been herbivorous because they lack specialized structures such as
a hooked beak or hooked ungual phalanges characteristic of such terrestrial carnivores as the
New World phororhacoids. They also often form the bulk of the biomass (up to 60-70%) of
the fauna at such sites as Bullock Creek and Alcoota (both Late Miocene in age) together with
the herbivorous diprotodontid marsupials. In such sites, vertebrates known to be carnivorous,
such as the marsupial lions (thylacoleonids) and varanid lizards, are quite rare, indicating that
the proportions of carnivores to herbivores approaches what is expected in natural living
populations.

Dromornithids are known primarily from bony remains, but they are also represented by
tracks and trackways (Rich & Green 1974, Rich & Gill 1976), gizzard stones (Stirling & Zietz
1900, pers. obs. at Bullock Creek and Alcoota sites), and eggshells (Williams 1981, Williams
& Rich, in press, Williams & Rich, this volume). One eggshell fragment associated with this
group (cf. Dromornis) of Miocene age from near Marree, South Australia, has a thickness of at
least 4 mm. The shell has a unique morphology, a rugose outer surface made up of subparallel
ridgelets and slitlike pore openings aligned along the longitudinal axis of the egg (Williams &
Rich in press, Williams & Rich, this volume). Both eggshell fragments and gizzard stones can
be abundant at certain localities. Perhaps the best representation of gizzard stones definitely
associated with skeletons of dromornithids are those known from Lake Callabonna in South
Australia. There, entire masses of smooth stones (jasper, siliceous sandstone, claystone and
quartz) were recorded associated with complete skeletons of Genyornis newtoni , one such
weighing 0.4 kg (Stirling & Zietz 1900).

Width at base of Mid - Toe (G)

160 180 200 220 240 260 280 300
Width of Foot (D)

Figure 8. Measurements of the foot of living emus (Dromaius novaehollandiae), black circles, and a
possible dromomithid, black triangles, from the mid-Tertiary of northeastern Tasmania (Rich & Green 1974).

Five genera and eight species of dromornithids are currently known. The greatest diversity
of the group occurs early in its history, during the Miocene, suggesting that the family had a
much longer history stretching well back into the Palaeogene. Most of the detailed
osteological information on the dromornithids derives frora the hind limb, although rare cranial
remains have been found (including the posterior half of three crania and a partial juvenile
cranium from the Miocene, a poorly preserved skull from the Pleistocene of Lake Callabonna,

MESOZOIC & TERTIARY BIRDS OF AUSTRALIA - 743

lower jaws and quadrates from both the Miocene and Pleistocene; see Pls 5,6). Hind limb
morphology indicates that both graviportal (Dromornis) and cursorial (IIbandornis lawsoni)
forms developed in this group, sometimes coexisting such as at the Late Miocene Alcoota
localities in the Northern Territory. This suggests that both forested and Open country
environments coexisted in the Late Miocene in Central Australia (Rich & Baird 1986, Rich e¢
al. ms.). It seems, however, that the dromornithids, although experimenting with different
lifestyles, were not particularly successful at invading the open grasslands, They may have lost
out to emus and kangaroos that were radiating rapidly in this developing environment as aridity
gripped the continent during the latest part of the Cainozoic (Hope 1980, Rich & Baird 1986).
Dromornithid species do have restricted time ranges and so can be used biostratigraphically.
Genyornis is restricted to the Pleistocene. Dromornis extends from the Miocene into the
Pliocene. Ilbandornis and Bullockornis are known only in the Late Oligocene to Miocene,
whereas Barawertornis may be restricted to the Early Miocene. Further work on the
superposition of the many localities in the Riversleigh complex in northwestern Queensland
may extend the ranges of the latter two genera.
Evolutionary patterns within the group include: (1) reduction in the size of digit II, (2)
increase in the size of digit IV until it is nearly coequal in size with digit III, (3) increased
mediolateral compression of the cnemial crests on the tibiotarsus and (4) general increase in
size, from earlier to later forms. These trends may simply reflect the development of a more
cursorial lifestyle, with I/bandornis and Genyornis representing the more cursorially adapted
members of the family. Only three fragmentary bones of Barawertornis (femur,
larsometatarsus, vertebra) are reported in the literature and thus offer little phyletic information,
but this may soon be augmented by new material being recovered from Riversleigh (W. Boles,
pers. comm.). Ilbandornis and Dromornis best known from the Late Miocene Alcoota sites,
and Genyornis form a related group within the dromornithids defined by quite uniform
morphology, whereas Bullockornis is related but distinct from all other dromornithids. Further
study of the dromornithid material from Alcoota, Bullock Creek and Riversleigh will be of
great value in resolving the relationship of these unique birds both within the family and with
other avian groups.

THE CASUARIIDAE - CASSOWARIES AND EMUS (Figs 9-11, Pls 20-22)

The fossil record of emus and cassowaries begins in the Miocene, most likely because of
the bias of the record. Like the record of the dromornithids, it is quite likely that forms related
to the casuariids were present in Australia during the Palaeogene.

The oldest member of this group is a mosaic that lies intermediate between the two living
families. Originally described as a primitive emu, Dromaius gidju (Patterson & Rich 1987)
(Pl. 20), the casuariid material from the Late Oligocene to Middle Miocene of northern (Archer
et al. 1986, 1988) and central Australia apppears to represent a form intermediate between
cassowaries and emus (Boles, in press). The more distal limb elements (tibiotarsi and
tarsometatarsi) are more emu-like, whereas the femur more closely resembles that of the
cassowaries. Based on the discovery of new material subsequent to Patterson & Rich (1987),
Boles (in press) proposed a new generic name for D. gidju to emphasize the intermediate nature
of this early casuariid. Thus far the material is limited to hind limb and foot, scapulocoracoid
and an upper mandible, which bears the longitudinal grooves indicating that the overlying
rhamphotheca characteristic of the ratites and tinamous was divided into distinct segments.
Increased cursoriality of D. gidju when compared to cassowaries is apparent when the ratios of
the length of the femur to those of the tibiotarsus and tarsometatarsus are calculated (Howell
1944, Boles, in press). Reduction in digits II and IV of D. gidju may, likewise, be related to
increased cursoriality. Boles (in press) suggests that such increased cursoriality might be cause

744 - RICH

for allying D. gidju slightly closer to the emus than to the cassowaries, but it is clearly a form
that lies close to the ancestry of both groups in the early Neogene or late Palaeogene.

A small casuariid is known from the Late Miocene or Early Pliocene Alcoota and Bullock
Creek localities in the Northern Territory that is of similar size to. Dromaius gidju, but due to
the small number of specimens and the poor preservation, the affinities of this form are
unresolveable at present.

Figure 9. Right tarsometatarsi of emus and cassowaries, posterior view: A,B, Dromaius novaehollandiae;

C, Dromaius ocypus type: and D, Casuarius unappendiculatus. All to the same scale, A is about 430 mm in
length. (After Miller 1963a).

Dromaius ocypus (Miller 1963a,b, Patterson & Rich 1987) (Fig. 9) is another casuariid
that has been designated as an emu. It is based ona partial hind limb, including a complete

MESOZOIC & TERTIARY BIRDS OF AUSTRALIA - 745

larsometatarsus and fragments of the femur and the tibiotarsus. The bones may not belong to a
single individual, but they all come from the same locality (Lawson Quarry, Palankarinna
Local Fauna, Middle to Late Pliocene) at Lake Palankarinna in northern South Australia. This
species is smaller than the living Emu, but significantly the tarsometatarsus of D. ocypus is
markedly shorter relative to width than is the same element in the living D. novaehollandiae.
The proportions of D. ocypus lie intermediate between those of the living cassowaries and
Emu suggesting a less cursorial lifestyle for the fossil relative to the living Emu.

All other definite emu (dromaiine) fossils are of Pleistocene age and have been assigned to
the living species, Dromaius novaehollandiae, including the de Vis taxa D. patricius and D.
gracilipes, or to one of two dwarf species, now extinct, that occurred on King and Kangaroo
islands (D. ater and D. baudinianus respectively (Parker 1984)). Yet another species of small
emu may have inhabited Tasmania during the Quaternary, but the validity of this species has
not been firmly established (Parker 1984).

ed
: 7
g Miocene DARWIN
a Pliocene ;
@ Pleistocene or ae Riversleigh
Quaternary Aas Bullock
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TROPIC OF. CAPRICORN |___.. 12s, PONT Sas pe nraen, aN. «1d or Cooper Creek localities
seepeeeteneee ALICE SPRINGS | 2 a Lake Callabonna
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Moore | @2 e Wellington Caves
PERTH ADELAIDE ° Wombeyan Caves
Lake Kanunka Aaurs SYDNEY
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Lake Palankarinna : CANBERRA
Brothers Island eee e
Bone: Gave Kangaroo I'sland = take: Menindee
Burra Baldina Ck. ELBOLAM \> Lake, tandou

Naracoote Area Buchan District Caves

McEachern’s Cave P '
Fisherman's Clift
Scotchtown Cave = Frenchman's Creek
owbray p Lancefield
trishtown HOBART King Island

Mole Creek

Figure 10. Localities producing fossil emus (Dromaiinae).

Pre-Quaternary cassowary (Casuariinae) fossils are limited to one Pliocene locality near
Bulolo, Papua New Guinea. Plane (1967) reported toe bones and assigned them to Casuarius.
The material is insufficient to allow any phylogenetic conclusions to be drawn. Pygmy
cassowary material has recently been described (Rich et al.1988) from Pureni in the highlands
of Papua New Guinea, a late Quaternary site. This material, which includes a number of
postcranial elements, is indistinguishable from Casuarius lydekkeri, a form defined originally
on a single distal tibiotarsus, whose provenance is uncertain.

746 - RICH

iii

i

CAPE

< / york : @ PURENI FAUNAL
AUSTRALIA o SITE

/
= AWE FAUNAL
Casuarius unappendiculatus Bees PENINSULA =} @ SITE

Casuarius bennetti

Casuarius Casuarius M

Figure 11, Localities producing fossil cassowaries (Casuarinae) plotted on a map with the distribution of
living cassowaries. (After Rich, Plane & Schroeder 1988).

THE PODICIPEDIDAE - GREBES

Although never an abundant element in fossil avifaunas, grebes are nonetheless present in
sediments as old as Late Oligocene to Miocene, the oldest record recovered from the Namba
Formation of Lake Pinpa in the Tarkarooloo Subbasin of South Australia. Other specimens
have been recovered from Pleistocene sediments along Cooper Creek in South Australia.
Because of the rarity of this group in the record, however, it will probably never be very useful
biostratigraphically.

THE SPHENISCIDAE - PENGUINS (Figs 12-13)

Penguins are first known from Australia in the Palaeogene, from the southeast of the
continent in South Australia. At least three different penguins are known from the Late Eocene
Blanche Point Formation (Jenkins et al. 1982, Jenkins 1985) near Blanche Point and Witton
Bluff about 40 km southsouthwest of Adelaide. One form is thought to be closely related to
the genus Palaeeudyptes also known from fossils in New Zealand and Seymour Island in West
Antarctica. A second, somewhat larger form is known from both the Tuketja Member of the
Blanche Point Formation and from Late Eocene rocks of Browns Creek, Otway Ranges,
Victoria Jenkins 1985). A more spectacular, giant penguin, Anthropornis nordenskjoeldi has

also been recovered from
1955, Jenkins 1974, 198
(Jenkins 1974) (Fig. 12),
best known from Seymo na ;

height and 90 kg in weight, significantly larger than the largest living penguin, paar
forsteri, which stands about 90 cm tall and is restricted to the Antarctic today. oe
nordenskjoeldi, additionally possessed an unusually long neck for a penguin (Jenkins in Ric
van Tets 1985).

| |
10cm

F gure omparison 0} e wing 4 i j t the Late Eocene Blanche

12 ¢ i f th i of Anth. opornis nordensk oeldi (A) from ;

Hei fasils near Adelaide South usiatin with that of the largest living penguin, Aptenodytes forsteri.
i ’

i related to a mid-Oligocene form from the South Island

Seah 197 eae by a single bone that bears grooves apparently

oe tan, f mohair (Jenkins 1985). This form from rocks of similar age to the New

esa ae s recovered from near Mt Gambier, South Australia (Glaessner 1955,

zone 1957) Fis, 13). An extraordinary feature of this Australasian species was its long
ee ie buen was nearly twice the length of the skull Jenkins 1985).

MESOZOIC & TERTIARY BIRDS OF AUSTRALIA - 747

the Late Eocene rocks at Blanche Point (Finlayson 1938, Glaessner
5). Originally described as a new genus Pachydyptes simpson
it was later found to be a synonym of Anthropornis nordenskjoeldi
ur Island in Antarctica. This giant form may have reached 135 cm in

748 - RICH

The end of the Eocene and earliest part of the Oligocene was characterized by marked
climatic deterioration (see Fordyce, Chap. 26, this volume). This was linked to the progressive
separation of Australia and Antarctica and the strengthening of the Circum-Antarctic Current.
This was also the time when the previously successful giant penguins became extinct, and thus
some relationship between these events seems likely.

1 Pachydyptes
2 Anthropornis

3 Palaeeudyptes

4 Wimanornis

5 Archaeospheniscus
6 Delphinornis

7 Indeterminate

vee Warm currents
Cool currents

Figure 13. Distribution of fossil penguins from Late Eocene localities, plotted on a palaeogeographic map

appropriate for this time period. Arrows indicate hypothetical pattern of oceanic surface circulation. (After
Jenkins 1974).

Three Miocene penguins are known in Australia, two from the Cheltenhamian rocks of
Victoria (Pseudaptenodytes macraei from Beaumaris and Spring Creek; P. minor from
Beaumaris) and another form from the Balcombian deposits north of Dartmoor on the Glenelg
River (Anthropodytes gilli). Pseudaptenodytes is unique in having an unusually shaped
tricipital fossa on the humerus, which allows no easy linkage to other groups of penguins.
Simpson (1970) observed that in the Miocene most larger species of penguins have simple
fossae, and most smaller species have bipartite fossae, as do the living penguins.
Pseudaptenodytes has the double fossa but is not clearly related to any living genus of penguin.

In the Pleistocene, penguin bones are common in dune and midden deposits along the coasts
of southern Australia and Tasmania, as well as in older sand deposits of Lord Howe Island

MESOZOIC & TERTIARY BIRDS OF AUSTRALIA - 749

(Rich & van Tets 1982). Most of the bones are indistinguishable from those of the Little
Penguin (Eudyptula minor), which today is known as far north as Australia and New Zealand.
Van Tets & O'Connor (1984) have recognized a new form Tasidyptes hunteri recovered from
800 year old midden deposits on Hunter Island, Tasmania. It is similar in size to the living

Rockhopper Penguin (Eudyptes chrysocome) and most similar to Eudyptes and Megadyptes in
overall morphology (see Baird, Chap. 21, this volume).

THE DIOMEDEIDAE - ALBATROSSES

The oldest record of this group in Australia is from the Cheltenhamian-aged (Late Miocene)
marine sediments of Beaumaris in the suburbs of Melbourne, Victoria. Diomedea thyridata
(Wilkinson 1969) is based on only a smali bill fragment, and little can be said about its
relationships. All other records of diomedeids are of Holocene age, mainly from sand dune and
midden deposits along coastal Australia (Rich & van Tets 1982).

| MapA

L “apB a 13305) a 7 & a7
| Se. id SX
eae eee : SS Ra \ \
Se ee ey 0 ee eS 5
| vis Y \—¥9
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a i$ 5 \ \ ka OPE
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| i A > S
rT 3 Se Ge 4 @6
AP Nicos
Lake Eyre YY Y >
{ ‘Qe 10 &
Xe 13
F 1412
3 q

— ae
eA:

s/

f ‘
Y ah ea

Ve , |
| re ie) 100 Km
| res ae
Figure 14. Localities producing pelican fossils in Australasia: 1, Pelecanus proavus, holotype; 2-5, P.

novaezealandiae; 6-7, P. tirarensis; 8, 10, 13, P. cadimurka; and 8-9, 11-14, P. conspicillatus. (After Rich &
van Tets 1981).

o

750 - RICH
THE PELECANIDAE - PELICANS (Figs 14-16, Pls 23-24)

Pelicans have a lengthy history in the fresh-water/terrestrial record, but not in marine
deposits of Australia. The oldest pelican fossils occur in Miocene sediments of the Lake Eyre
and Tarkarooloo subbasins of northern and northeastern South Australia (Fig. 14). The
Ngapakaldi, Pinpa and Kutjamarpu local faunas all contain pelican fossils, albeit not large
numbers of specimens. Pelecanus tirarensis (Fig. 15) is the single species recorded from these
three faunas, based on five specimens. These are fragmentary specimens, but they demonstrate
that in the morphology of the trochlea II of the tarsometatarsus, the oldest known species
is distinct from all later forms (Miller 1966b; Rich & van Tets 1981). Two other species,
both from the Pleistocene, are known, restricted to the northern South Australian basins.
These include a small form, Peiecanus cadimurka, as well as a form indistinguishable from the
living Australian Pelican, P. conspicillatus. Pelecanus proavus, first described by de Vis
(1892), from Quaternary sediments in the Darling Downs of Queensland has been lost. It is
impossible to determine from the illustrations if this specimen represents a unique species
name. Pelecanus validipes (de Vis 1894 in Brown) originally part of the Archaeocycnus
lacustris material reported on by de Vis (1905) and P. grandiceps (de Vis 1905) seem to be
synonyms of the living Australian pelican, P. conspicillatus (Rich & van Tets 1981, 1982)
(Fig. 16).

4 \
aye
Gi

LS

Figure 15. Tarsometatarsi of pelicans. Pelecanus tirarensis (type) from the Miocene of central Australia:
A, medial; C, posterior; E, anterior views. P. conspicillatus, the living Australian Pelican: B, medial; D,
posterior, and F, anterior views. Width of the distal end of D, 24 mm. (From Miller 1966b).

Evolutionary trends evident in the Australian pelicans can only be observed on the distal end
of the tarsometatarsus because of the material available in the fossil record. These include (1)
increases in trochlea II width along its posterior border from the Miocene to the Pleistocene, (2)

minor changes in the details of the shaft and trochlear morphology (Rich & van Tets 1981,
1982).

MESOZOIC & TERTIARY BIRDS OF AUSTRALIA - 751

P. novaezealandiae 4 onocrotalus
a *P. validipes’
a ire
Ae.

P. crispus we,

10 P. philippensis os Ns

— P. grandiceps'

e
; Me,
a == eo
=| ¢ a ok
Bg M
= P. tirarensis . * bg
2 o——___ P. erythrorhynchos
= e
g 8 e = ——P. conspicillatus
=
& * o- P. rufescens
6 Ww
7 ¢
P. occidentalis
| S
®
6 w— P. cadimurka

8 9 10 aa 12 3 #14 #+15 ‘6 17 #18 «19 20
External Depth of Trochlea II (mm)

Figure 16. Comparison of width and depth measurements on the tarsometatarsus of living pelicans and
Australasian fossil pelicans. (From Rich & van Tets 1981).

THE PHALACROCORACIDAE AND ANHINGIDAE - CORMORANTS,
SHAGS AND DARTERS

Cormorants are first recorded in the Miocene lacustrine and fluviatile sediments of northern
South Australia, but it is not until the Quaternary that their bones become a common element
in the palaeofaunas of Australia. Most, if not all, of the Quaternary forms appear to belong
within modern species known from Australia. One form, the Black-faced Shag (Leucocarbo),
now restricted to marine waters, may have previously occurred in Quaternary central Australia
(van Tets, pers. comm.), but this is yet to be documented in the literature. The fossil
cormorant material collected on many expeditions to central Australia since the 1950's is
currently under study by G. F. van Tets at C.S.I.R.O. Wildlife and Ecology in Canberra.

There may be an anhingid from Lake Kanunka, of Pliocene age, but this needs to be
reexamined.

THE ARDEIDAE - HERONS

Heron bones are rare in the Australian fossil record, and they are known primarily in
Quaternary deposits, mainly from northern and northeastern South Australia. One occurrence at
Lake Kanunka may be a Late Pliocene occurrence. The group has not been sufficiently studied
to make any further comment on it, however.

752 - RICH

THE CICONIIDAE - STORKS

Today there is only one living stork species in Australia, the Jabiru or Black-necked Stork,
Ephippiorhynchus asiaticus, also known in southern Asia. In Pleistocene deposits of northern
South Australia and in the Darling Downs of Queensland, bones similar to this living form
have been found. A smaller form is also known that has been named Ciconia nana. This
material needs further study before any decisions can be made regarding its affinities.

The oldest stork material from Australia, still undescribed, comes from Riversleigh Station
in northwestern Queensland (Miocene) (W. Boles, pers. comm.) and Allingham Creek,
Queensland, a Pliocene site that is radiometrically dated at 4.0-4.5 myBP (Archer & Wade
1976). The Allingham stork appears to be closely related to the living Xenorhynchus (G. F.
van Tets, pers. comm.). Unfortunately, it is represented by a single specimen, so more
material is required before much inference can be made about relationships. There is a
questionable occurrence at the Late Pliocene Chincilla site, but this needs further study to
confirm its identity,

THE THRESKIORNITHIDAE - IBISES AND SPOONBILLS

Specimens of ibises and spoonbills are exceedingly rare in Australia. Records are restricted
to the Late Pliocene and Quaternary, except for a possible occurrence in Miocene sediments of
Riversleigh Station in northwestern Queensland. Platalea subtenuis of de Vis from the
Darling Downs area of Queensland is in need of review, but may represent Gallinula (G. F. van
Tets, pers. comm.).

Figure 17. Palmar and distal views of the distal end of right humeri of: A, Black-necked Stork,
Xenorhynchus asiaticus, B, cf. Xenorhynchopsis tibialis (UCMP 56324); C, Greater Flamingo, Phoenicopterus
ruber, D, Straw-necked Ibis, Threskiornis spinicollis; E, Lesser Flamingo, Phoeniconaias minor; F, Bush

Stone-Curlew, Burhinus grallaria; G, Banded Stilt, Cladorhynchus leucocephalus. Scale bar, 20 mm. (From

Rich et al. 1987).

MESOZOIC & TERTIARY BIRDS OF AUSTRALIA - 753

THE PHOENICOPTERIDAE - FLAMINGOES (Figs 17-21)

_ Flamingo diversity was high throughout the Neogene of Australia. This group first appears
in the Late Oligocene-Miocene lacustrine deposits of the Lake Eyre and Tarkarooloo subbasins.
Miller (1963b) described the first of this material establishing several new taxa:

Phoenicopterus novaehollandiae and Phoeniconotius eyrensis from the Etadunna Formation in
the Lake Eyre Subbasin.

Phoeniconotius eyrensis, based on part of a tarsometatarsus and a few phalanges (Fig. 20),
is of similar size to Phoenicopterus ruber, the living Greater Flamingo, larger than a
contemporary Miocene form, Phoenicopterus novaehollandiae (Fig. 20). Phoeniconotius
differed, however, from both in being decidedly more robust, and, according to Miller (1963b),
possessed a much stronger hind toe (digit I) than either of these species. Phoenicopterus
novachollandiae (based on a nearly complete tarsometatarsus) similarly differed from all
flamingoes except Phoeniconotius in having a well-developed scar for the metatarsal I,
suggesting a robust first digit, and perhaps meaning that both of these fossil Australian forms
were more terrestrially adapted than any other species in this family. Thus, during the Late
Oligocene-Miocene, at least two genera and two species of flamingoes inhabited the Lake Eyre
Subbasin, both included in the Ngapakaldi Local Fauna. If habits of flamingoes alive in the
Oligo-Miocene were within the range of tolerance of species alive today, then at minimum
there must have been permanent lakes in the Lake Palankarinna area of South Australia, lakes
that were reliable enough on a year to year basis to accommodate feeding and breeding of these
specialized colonial nesters.

Desiccation of central Australia during the late Cainozoic led to the demise of such
dependable conditions, and birds such as the flamingoes, which were unable to cope with long
droughts and ephemeral lacustrine conditions, were destined for extinction. They did survive for
quite some time, however. Pliocene sediments at Lake Kanunka in the Lake Eyre Subbasin
contain three species of flamingoes that very likely coexisted: Ocyplanus proeses (Figs 18-19,
21), a form smaller than any known flamingo except for the African Miocene Leakeyornis
aethiopicus, Xenorhynchopsis minor, similar in size to the living Phoeniconaias minor; and
Phoenicopterus ruber, a form with living representatives and the largest of the three fossils
(Rich et al. 1987). As Rich et al. (1987) point out, however, this material is not easily
evaluated, as it is based on very few specimens. Hopefully, future excavations, especially at
lakes Kanunka and Palankarinna will allow the species diversity hypothesis to be tested as
more significant samples come to light.

Flamingoes are also known in Pleistocene sediments (see R. F. Baird, Chap. 21, this
volume), but dating of these specimens is difficult, if not impossible at present.
Xenorhynchopsis tibialis, larger than any living flamingo and on par with the Miocene
Phoeniconotius, occurs only in Pleistocene sediments (from along Cooper Creek in South
Australia) together with Phoenicopterus ruber and Xenorhynchopsis minor. Only Ocyplanus
proeses may be restricted to the Pliocene. It has never been found anywhere together with X.
tibialis, and is known from only one locale along the Cooper, which may be a remainé fossil.

The disappearance of flamingoes sometime in the last million years most surely
corresponds to the disappearance of permanent lakes in the Great Artesian Basin and the
establishment of the current environmental conditions which include only ephemeral lakes. If
the flamingo material could be dated, then it would be critical in determining just when this
crucial climatic alteration took place.

Some confusion exists as to the names of Australian flamingoes, due, in part, to the lack of
comparative material available to Charles de Vis, and due to the lack of availability of the de
Vis material to Alden Miller when he originally studied the flamingo fossils from Australia.
De Vis was unaware that some of the bones he named were those of flamingoes and identified

754 - RICH

C
D
F
F
G

Figure 18. Proximal, anterior and posterior view of left femora of: A, Black-necked Stork, Xenorhynchus
asiaticus; B, Greater Flamingo, Phoenicopterus ruber; C, Lesser Flamingo, Phoeniconaias minor, D, Straw-
necked Ibis, Threskiornis spinicollis; E, cf. Ocyplanus proeses (=Ibis conditus, QM F5519); F, Bush Stone-
Curlew, Burhinus magnirostris; G, Banded Stilt, Cladorhynchus leucocephalus. Scale bar, 50 mm. (After Rich
et al. 1987).

Figure 19. Anterior and distal views of the distal end of right tibiotarsi of: A, Black-necked Stork,
Xenorhynchus asiaticus; B, cf. Xenorhynchopsis tibialis (QM F5515); C, Greater Flamingo, Phoenicopterus
ruber; D, cf. Xenorhynchopsis minor (QM F5517); E, Straw-necked Ibis, Threskiornis spinicollis; F, Lesser
Flamingo, Phoeniconaias minor; G, cf. Ocyplanus proeses (UCMP 56887); H, Bush Stone-Curlew, Burhinus
grallaria; 1, Banded Stilt, Cladorhynchus leucocephalus. Scale bar, 20 mm. (From Rich et al. 1987).

MESOZOIC & TERTIARY BIRDS OF AUSTRALIA - 755

\ } AN \
4% ANS We X
\l

yi
we

wo ij,
YY, ‘uly ‘willl
OMIT, . Tm,

ies Pre

Figure 20. Fossil and living flamingoes. A-C, Phoeniconotius eyrensis, type: (UCMP 13649) A, medial
view; B, plantar view; C, anterior view; length of digit I, 45.5 mm. D, F, Ocyplanus proeses
(=Phoeniconaias gracilis), UCMP 13650: D, medial view; F, plantar view; width across trochleae, 13 mm.
E, G, Lesser Flamingo, Phoeniconaias minor: E, medial; G, plantar views. H, I, anterior views of the
tarsometatarsi of Phoenicopterus novaehollandiae (H) (width across trochleae, 17.6 mm) and Phoenicopterus

ruber roseus (1). (From Miller 1963b).

G H
sp cP

Figure 21. Posterior and distal views of the distal end of left tarsometatarsi of: A, Black-necked Stork,
Xenorhynchus asiaticus; B, Greater Flamingo, Phoenicopterus ruber, C, Straw-necked Ibis, Threskiornis
spinicollis; D, Lesser Flamingo, Phoeniconaias minor; E, cf. Ocyplanus proeses (=Phoeniconaias gracilis)
(SAM P13650); F, Bush Stone-Curlew, Burhinus magnirostris; G, Banded Stilt, Cladorhynchus leucocephalus.
Scale bar, 20 mm. (From Rich et al. 1987).

them as follows: Xenorhynchopsis tibialis and S. minor as Ciconiidae, storks; Ocyplanus
proeses as a Charadrii, wader; and Jbis (?) conditus as a Threskiomithidae, ibis. Miller was
unaware that de Vis had already assigned names to existing material and thus set up
Phoeniconaias gracilis on the basis of new material discovered by R. A. Stirton in the middle
of the 20th century. Phoeniconaias gracilis is, in fact, the same species as Ocyplanus proeses,
which holds priority (Rich et al. 1987). To further complicate the issue, Lambrecht (1933)
reassigned Ocyplanus proeses to the Laridae and Condon (1975) assigned it to the Rallidae,
both of which were incorrect.

Determining the phylogenetic relationships of Australian flamingo taxa is not possible at
present due to the fragmentary nature of the record. It is clear, however, that the taxa are unique
to the Australian continent in most cases. Sample sizes of both the fossil and recent taxa of
flamingoes need to be increased, and the completeness of individual species needs to be
improved. Two sites in the Oligo-Miocene of Lake Palankarinna offer marked potential for
recovery of partial skeletons as well as cranial material, and thus further field work could
markedly increase our knowledge of this highly endemic group in Australia.

Because flamingoes occur in most of the central Australian Miocene sites, from Lake
Palankarinna and Lake Pinpa in the south to Alcoota and Bullock Creek in the Northern
Territory, and they range into the Pliocene and Pleistocene as well, they show promise for
defining relative time, once the samples improve. Previous field work at several of these
locales indicates that additional excavation at selected sites would likely be quite fruitful.

THE PALAELODIDAE - PALAELODIDS (Pls 25-26)

Palaelodids have been known in Australia since 1982, having previously been reported only
from Europe and North America. Two new Australian species have been described: Palaelodus
sp. A, from the Ditjimanka Local Fauna of Middle Miocene age at Lake Palankarinna, South
Australia and Palaclodus sp.B with a long time range beginning with the Middle Miocene

MESOZOIC & TERTIARY BIRDS OF AUSTRALIA - 757

locales of Lake Palankarinna (Diljimanka Local Fauna), Lake Yanda (Yanda Local Fauna or
Ericmas Local Fauna) and Lake Pinpa (Pinpa Local Fauna) and extending to the Middle
Pleistocene of Kudnampirra Waterhole along Cooper Creek (Kutjitara Formation), thought to
be about 200,000 years of age (Baird & Rich, in press).

Palaelodus sp. Bis as large or larger than the largest European species in the genus,
based on the libiolarsus. Besides the large size, P. sp. B differs from the European forms in
having proportionally broader tibiotarsii, both distally and proximally. The shaft of the
libiotarsus in P. sp.B is swollen laterally and has a proportionally narrow dorsal surface
(Baird & Rich, in press).

Palaelodus sp Ais smaller than P. gracilipes, the previously smallest species in the
eee based on the tarsometatarsus. Shaft depth is less than that of all European species in

€ genus.

Prior to the Australian record of palaclodids, this group was known to occur from the Early
Miocene to the Middle Miocene of Europe and the Early Miocene to the Early Pliocene of
North America. The group appears to have survived longer in Australia than anywhere else
(Baird & Rich, in press). Finally, the group succumbed in Australia at some time during the
Late Pleistocene or Holocene as aridity expanded and the interior lakes dried out.

ANATIDAE - DUCKS, GEESE AND SWANS

Australasia has relatively few species of Anatidae when compared to the rest of the world
but does have several endemic genera, including many that are extant. Although the record of
the anatids extends far back in the Australian record, only the Pleistocene and Holocene forms
have been adequately studied. Despite the diversity of extinct forms reported by de Vis in
several papers, Olson's restudy (1977) demonstrated clearly that all nine species described by de
Vis were assignable to extant forms. Nyroca effodiata was shown to be a junior synonym of
Leucosarcia proevisa and, in fact, not a duck at all (van Tets & Rich 1980).

The oldest known duck remains in Australia come from the mid-Tertairy deposits of Central
Australia, for example from Lake Palankarinna and Lake Ngapakaldi in the Lake Eyre Subbasin
and from Lake Pinpa in the Tarkarooloo Subbasin. Younger forms are known from Alcoota,
Allingham Creek and Lake Kanunka in central and northern Australia. As many as 6 different
taxa may have been present at the Late Pliocene Chinchilla sites in Queensland. The Tertiary
fossils are largely unstudied but are under examination by W. Boles (Australian Museum).

ACCIPITRIDAE AND FALCONIDAE - HAWKS, EAGLES AND FALCONS

Australia appears to have had more kinds of large accipitrids during the Pleistocene than
now, including one form from the southern part of the continent that exceeded the size of the
living Wedge-tailed Eagle (Aquila audax) significantly (Rich & van Tets, pers. comm.).

The oldest records of this family come from both central and northern Australia, from Lake
Palankarinna (in both the Ngapakaldi and Ngama local faunas) and Riversleigh (Archer et al.
1989). A large eagle, about the size of the living Wedge-tailed Eagle is known from a single
tarsometatarsus from Alcoota of Late Miocene age. Both Chinchilla and Lake Kanunka have
also yielded accipitrids.

The record of falcons, thus far, is restricted to the Pleistocene and is limited to extant
species. Of those Pleistocene forms, even Asturaetus furcillatus described by de Vis (1905) and
renamed Plioaetus by Richmond (1909) from Cooper Creek, is indistinguishable from the
living Brown Falcon (Rich, van Tets & McEvey 1982).

758 - RICH

MEGAPODIIDAE - MOUNDBUILDERS (Fig. 22)

All extant genera of moundbuilders occur in Australasia, excluding New Zealand, but
including Sulawesi. Only one genus of living megapode, Megapodius, has a distribution that
extends westwards to a few coastal islands in southeastern Asia, and eastwards to islands in the
southwestern Pacific Ocean. This spread may be in part explained by prehistoric transport by
humans. In addition, there are likely two extinct megapodes that occur in the southwest Pacific
basin, one in the Fiji islands (yet undescribed) and possibly a second from New Caledonia
(Sylviornis ; see Balouet, this volume).

The extant genera of megapodes are exceeded in size by Pleistocene relatives, now extinct,
Progura (Fig. 22) in Australia and possibly Sylviornis in New Caledonia serving as good
examples. Progura may have reached or exceeded 7 kg in weight (van Tets 1974). Several
fossil forms from the Pleistocene, assigned to a variety of avian families have been found to be
megapodes. Progura was originally thought to be a giant pigeon (de Vis 1888b),
Palaeopelargus nobilis a stork (de Vis 1891a). One form described as a bustard (de Vis 1888b)
was, likewise, found to be a megapode. De Vis was correct, however, in his assignment of
Chosornis praeteritus to the Megapodiidae.

The only pre-Pleistocene record of megapodes in Australia is a single form from Chinchilla.

Interestingly, Mourer-Chauvire (1982) reported the occurrence of small megapodes in the
Late Eocene deposits of Quercy in France. Perhaps the present distribution of this group in
Australasia is relictual. Another possibility is that megapodes dispersed northwards from
Australia into Europe in the early Tertiary. Until more is known about the early Tertiary
record of Australasia, it is difficult to favour either hypothesis.

TURNICIDAE - BUTTON-QUAIL

Bones similar to those of extant species of Turnix are, like those of Coturnix, common in
Quaternary cave and midden deposits in Australia. Older records of this family from the
Tertiary of northern Australia seem to be rails, rather than turnicids (W. Boles & R. Baird,
pers. comm.)

GRUIDAE - CRANES

The limited amount of Pleistocene and Holocene material of cranes found in Australia
appears to be indistinguishable from that of the Brolga, Grus rubricundus, which is now
limited in its distribution to Australia and New Guinea. There are rare bones in the Early
Miocene and Late Pliocene (Lake Palankarinna, Lake Ngapakaldi and Lake Kanunka) of central
Australia that may represent this group, but these need further study.

Figure 22. Fossil and recent megapodes: A, right tarsometatarsus, Progura naracoortensis; B, right
tarsometatarsus, Megapodius reinwardt; C, left reversed tarsometatarsus, Progura; D, E, right tarsometatarsi,
Progura; F, right femur, Progura naracoortensis; G, right femur, M. reinwardt; H, right tibiotarsus, M.
reinwardt; I, right tibiotarsus, P. naracoortensis; J, synsacrum, P. naracoortensis; K, synsacrum, M. reinwardt.

MESOZOIC & TERTIARY BIRDS OF AUSTRALIA - 759

760 - RICH
RALLIDAE - RAILS

Rails occur in some of the oldest Cainozoic bird-bearing deposits in Australia, probably
Late Oligocene-Early Miocene in age, at Lake Palankarinna (both the Ngapakaldi and Ngama
local faunas). At least two types of rails have also been recognized in the Early Miocene or
Late Oligocene deposits of Riversleigh, one of which had reduced power of flight or was
flightless (W. Boles, pers. comm.). Two different forms are known from the Chinchilla
deposits. The group is never particularly diverse either in the Tertiary or Quatemmary deposits.

The Pleistocene forms appear to be indistinguishable from extant forms (see Olson's review
of de Vis' work, 1975). Of greatest interest is the Tasmanian Native Hen (Gallinula (Tribonyx)
mortierii) whose remains have been found in Quaternary deposits of southeastern South
Australia, Victoria, New South Wales and the Darling Downs, Queensland. Historically, the
Tasmanian Native Hen has been restricted in its distribution to Tasmania, and, thus, its
Quaternary range was decidedly broader than that of today. Olson (1975) has suggested that this
species probably evolved on the mainland, spread to Tasmania, was isolated there after the
flooding of Bass Strait, leaving the mainland population to become extinct and the Tasmanian
population to remain as a relict where it evolved slightly larger size. This is certainly one
possibility, but areas of origin are difficult to assess with confidence without a good fossil
record.

OTIDIDAE - BUSTARDS

The record of bustards in Australia is very limited. The oldest known occurrence is from
very Late Pliocene deposits at Lake Kanunka. All of the Quaternary material, also very
limited, appears to be indistinguishable from the living Australian Bustard (Ardeotis australis),
now restricted in its distribution to Australia and New Guinea.

CHARADRIIFORMES - WADING BIRDS (Pls 27-28)

The Charadriidae, Scolopacidae and cf. Laridae material known from Pleistocene and
Holocene deposits in Australia appear to be indistinguishable from that of extant species. A
variety of charadriiforms also are known in older sediments back as far as the Miocene, but they
are yet to be analyzed. Incomplete comparative collections in this diverse group have still
hampered work on these taxa.

Of the limited amount of Quaternary material of stone curlews (Burhinidae) that has been
found in Australia, all appears to be indistinguishable from the living Bush Stone-Curlew
(Burhinus grallarius), which is now restricted to Australia and New Guinea. The pre-
Quaternary record, especially the Oligo-Miocene, has an abundant record, however. One large
collection from the Namba Formation at Lake Pinpa, Tarkarooloo Subbasin contains more
than 100 bones, and this record rivals the oldest record of the group from North America.
Other material is known from similar-aged sediments in the Lake Eyre Subbasin, the Etadunna
Formation at Lake Palankarinna. The material from Lake Pinpa is significant in that although
disarticulated, most elements of the skeleton are represented, including a partial skull. It is
clearly a form distinct from the Australian forms of today, at least rating a new specific
recognition. It appears most closely related to Esacus magnirostris (Perry 1983), the living
Beach Stone-Curlew of northern Australia.

A partial skeleton similar to that of the extant Plains-Wanderer (Pedionomus torquatus) (P|s
27, 28) has been recovered in Pliocene or Pleistocene fire-hole deposits in the Morewell coal

MESOZOIC & TERTIARY BIRDS OF AUSTRALIA - 761

seams of Victoria (Rich & McEvey 1980). There is not general agreement on the age of the

Five ee deposits, but this occurrence is the only possibly pre-Quaternary record of this

pale All other fossil pedionomids are from Pleistocene or younger deposits within
ia.

Olson & Steadman (1981) have presented a strong case for allying this group to the
charadriiform Thinocoridae of South America. ‘ te

TYTONIDAE AND STRIGIDAE - BARN OWLS AND TYPICAL OWLS

Most fossil owl material in Australia has been recovered from Quaternary caves (see Baird,
this vol.), One bone tentatively assigned to Tyto novaehollandiae (Rich, McEvey & Walkley
1978) was recovered from fluviatile or lacustrine sediments along Cooper Creek in central
Australia, of probable Quaternary age, although dating of many of the bones in this area,
unless well documented, is not totally reliable. The remains of Quaternary owls in Australia
seem indistinguishable from extant species. To date, there are no pre-Quaternary fossil owls
from this continent, even though the record of this group elsewhere in the world span back to
the beginning of the Tertiary.

PODARGIDAE AND AEGOTHELIDAE - FROGMOUTHS AND OWLET
NIGHTJARS (Pls 29-30)

The fossil record of frogmouths is restricted, in Australia, to the Quaternary, which is of
some interest, since it has a much older record in Europe. Mourer-Chauvire (1982) reports
fossils of this group in the Late Eocene of France that show affinities with the living
Podargus. The current distribution of the podargids in Australia and southeastern Asia may, in
fact, be of a relictual nature.

Aegothelids, by contrast, have a much longer fossil record in Australia, A nearly complete
skeleton of a primitive member of this family, Quipollornis koniberi, is known from mid-
Tertiary deposits in the Warrumbungle Mountains near Bugaldi in New South Wales (Rich &
McEvey 1977) (Pls 27, 28). This fossil was preserved in volcanic caldera lake deposits. The
volcanics forming the caldera were dated as between 13.5 and 17 million years old.
Quipollornis is without doubt an owlet-nightjar, but it possesses a number of character-states
that demonstrate that it is not as specialized as living members of this group, a phenomenon
typical of birds of this age from around the world. The main differences lie in the limb
proportions: Quipollornis, for example, has longer wings relative to hind limbs, which is
more characteristic of the non-aegothelid caprimulgiforms, especially the Caprimulgidae. Such
proportions suggest that emphasis on lengthening the hind limb and a terrestrial lifestyle had
not begun at this time, and this mid-Tertiary aegothelid was more of an aerial, rather than a
terrestrial, feeder. Further study of this fossil form is warranted, however, in that Rich &
McEvey limited their study to only some living Aegotheles. Further comparisons with other
members of this genus would be worth carrying out to see if proportions for A. cristatus are
typical for all species in this genus, in light of a study by Olson, Balouet & Fisher (1987).

Fossils of aegothelids are also known from Australia, as well as New Caledonia and New
Zealand, in the latter case where they do not occur today. Although the record in Australia
dates from the Miocene, it may not be the oldest record of the family, as Mourer-Chauvire
(1982) has reported the fragment of a sternum from the Late Eocene of France that may belong
in this family. At present the European record is tentative due to the fragmentary nature of this
material, and, thus, it is not possible to comment further on the possible dispersal pattern of

this family relative to Australia.

762 - RICH

APODIDAE - SWIFTS

The record of swifts in Australia begins in the Early Miocene or Late Oligocene, based on
material from Riversteigh, Queensland (W. Boles, pers. comm.). This record rivals or perhaps
predates the previously oldest record from the Early Miocene (Aquitanian) of France (Olson
1986), although a new report of Eocene material from Asia may precede this (Boles, pers.
comm.) The only other record of this family in Australia comes from Quaternary deposits.

COLUMBIDAE - PIGEONS

Pigeons and doves have a long record in Australia, dating back to the Early Miocene in
central Australia. A few bones are known from both Lake Palankarinna (Etadunna Formation)
and Lake Pinpa (Namba Formation), in the Lake Eyre and Tarkarooloo subbasins respectively.
Once again, this record rivals any previously known records for this group, the oldest being
from the Early Miocene (Aquitanian) of France (Olson 1986). At least two different sizes of
columbids are represented, one dove-sized (from Palankarinna) and one with medium-sized
pigeon measurements.

Similar to the record of this group elsewhere in the world, fossils are rare. Quaternary
material is known from several localities in Australia and on Lord Howe and Norfolk islands,
most or all of which is conspecific with living forms.

PSITTACIFORMES - PARROTS

Parrots are first known in Australia from Early Miocene or Late Oligocene deposits at
Riversleigh, remains assigned to the Cacatuidae (W. Boles, pers. comm.). This is not the
oldest record for the family on a world scale, it being from the Eocene of the Old World, either
from the Early Eocene (Palaeopsittacus georgei) of the London Clay or from the Late Eocene of
France (Olson 1986). All other parrot fossils from Australia found thus far are from the

Quaternary.

PASSERIFORMES - SONGBIRDS

The oldest records of passeriforms in Australia come from the Late Oligocene to Middle
Miocene sediments of Riversleigh in Queensland (W. Boles, pers. comm.) where at least one
family that now occurs in Australia has been recognized, the Orthonychidae (W. Boles & R. F.
Baird, pers. comm.). A bird similar to the lyrebirds (Menuridae), has been tentatively identified,
as well (W. Boles, pers. comm.). Other yet unidentified passeriform material is known from
Riversleigh as well as from central Australia from Lake Palankarinna and Lake Pinpa in South
Australia, and from Bullock Creek in the Northern Territory. These records are as old as any
records known from elsewhere in the world, the oldest being material from the Early Miocene
(Aquitanian) of Langy, France (Olson 1986), and probably older from the Late Oligocene of
France (Mourer-Chauvire et al. 1989.). Certainly by this time songbirds were widespread. It is
interesting that even though the European and North American records are rich in fossil birds,
there are no passeriforms present before this time, and it makes one wonder if, in fact, this
group might have invaded the Northern Hemisphere from the south. Only a better fossil
record, which is almost non-existent for birds in Australia prior to the Miocene, will resolve
this problem.

MESOZOIC & TERTIARY BIRDS OF AUSTRALIA - 763

ihe rhe record of passcriforms, excepting these mid-Tertiary occurrences, is entirely restricted to
- Quaternary, where it is quite diverse (see Baird, this volume).

ORIGIN OF THE AUSTRALIAN AVIFAUNA: DISPERSAL,
VICARIANCE OR BOTH?

Since P. L. Sclater divided up terrestrial biotas of the world into a series of biogeographic
regions, there have been many discussions and no lack of debate about how such units came
into being. As knowledge of the fossil record of many of the living organisms became better
documented, it became clear that today's biogeographic regions did not apply to all times in the
past. Even with this realization, however, for the first few decades of the 20th century, most
biogeographers were content to move their animals and plants about on a static geography, one
Just like that of today. It wasn't, in fact, until the early 1960s that biogeographers began to
entertain seriously the idea that continents had moved with respect to one another during the
history of life on earth. Once this was accepted as a viable hypothesis, Pandora's box was
indeed opened. Where once there had been a single explanation for the distribution of
organisms, many hypotheses were possible, unless a good fossil record was available. On the
other hand, many insoluble problems had instant solutions.

« Lydekker's Line
Muller's Line
Murray's Line
Sclater’s Line

o Wallace's Line (original)
Wallace's Line (of Huxley)
Weber's Line (faunal balance)
Limit of Marsupials
200 metres depth

_, Limit of Native Placental Mammal!s other
than bats, Muridae, Sus & Cervus

AUSTRALIA
JS 14

Figure 23. The Malay Archipelago, a mixing area for the Oriental and Australasian biotas, showing several
different lines proposed to delimit the extent of each biogeographic region. (Modified after Simpson 1977).

Most work prior to the 1960s that dealt with the origin of the Australasian avifauna,
suggested that birds entered Australia from the north across the Malay Archipelago (Fig. 23), in
a series of waves, a valuable concept put forward by Ernst Mayr in his classic paper on the
birds of Timor and Sumba (1944b). Mayr was assuming a static, not mobile, earth geography,
and his five waves of immigrants were arranged, oldest to youngest, according to the level of

764 - RICH

endemism (specific, generic, familial, etc.) these groups had within Australasia. Mayr
characterized these waves as:

1. Dromaiidae, Casuariidae, Megapodiidae, Loriinae, Cacatuinae,
Platycercinae, Podargidae, Menuridae, Atrichornithidae,
Grallinidae, Artamidae, Neosittidae, Meliphagidae, Struthideinae,
Ptilonorhynchidae and Cracticidae (old endemic families, subfamilies whose
nearest relatives are uncertain).

2. Pedionomidae, Ptilinopinae, Pachycephalinae, Sphecotheridae,
Cinclosomatinae, Acanthizinae, Pardalotinae and Paradisaeidae
(families and subfamilies that are clearly related to Old World
families).

3. Numerous genera which are endemic in "Australo-Papua”
but which are clearly related to Asiatic genera (e.g. Synoicus,
Geopelia, etc.).

4. Numerous species that are clearly related to Old World species
(e.g. Coturnix pectoralis, Elanus axillaris, etc.).

5. Numerous subspecies that are in the same species as old world
forms.

—H
TickeTS PLEASE Y /
L. = |

a

Figure 24. Cartoon depicting biogeographic theory on the origin of Australia’s avifauna primarily from
dispersal of northem forms southward, and idea championed by Mayr (1944). (Drawn by R. Plant).

MESOZOIC & TERTIARY BIRDS OF AUSTRALIA - 765

Mayr Suggested that Group 1 represented the oldest colonists, which arrived in the early
Tertiary (or possibly earlier), Group 2 arrived in the early or middle Tertiary, Group 3 in the
Miocene or Pliocene, Group 4 in the Pliocene or Pleistocene, and Group 5 very recently. At
the time that Mayr wrote his paper there was little palaeontological data to test this hypothesis
against, and still there is little fossil information older than Miocene in Australia that bears on
his ideas. One set of fossils that have been found, however, since Mayr's study, are the Early
Cretaceous feathers in the Strzelecki Formation near Koonwarra in south-central Victoria.
These document that birds were in Australia early in the history of this group and suggest that
it had a very long history on this continent, not necessarily dominated by outside recruitment.

In the early 1960s biogeographers were faced with a new paradigm, a new set of
possibilities that must be considered in any study of palaeobiogeography dealing with Australia
as well as all other continents of the globe. Seismologists, structural geologists, marine
geologists and geophysicists threw their data and support behind the concept of continental and
ocean basin mobility, and such theories as continental drift, sea floor spreading and plate
tectonics emerged (Tarling & Tarling 1975). To the palaeobiogeographer, this meant
consideration of at least two possibilities to explain past distribution patterns of their
organisms: dispersal (Fig. 24) of organisms from one place to another using their own
devices about crossing barrier that might vary from time to time (due to variable sea levels,
advancing and retreating glaciers, development and erosion of mountain ranges, etc.): or
vicariance (Fig. 25), which implied that the distribution of an organism was determined by

hp, frm t
amt Hy Gah
i di | Tl

R TR fo) De, it rea: I)
| i iE SE-ASIA Pius “em ji ke haa

Ll

(taken Yo} ee)
| Seth ?? hy
\ forth a Sth ea

A

" ADIACENT ISLANDS

1
2
Yaa

Figure 25. Cartoon depicting the influence of moving continents on the origin of Australia's avifauna.
(Drawn by R. Plant).

766 - RICH

the movement of continents - when continents were adjacent, biotas could move between them,
and when continents separated, biotic interchange stopped. McKenna (1972, 1974) introduced
several new biogeographic concepts that flowed directly from the addition of mobile continents.
Noah's Arks, for example, were continental masses that broke away from another mass and
carried with them a biota, which could later be juxtaposed to another biota if the "Ark"
continent docked next to a new continental mass. Viking Funeral Ships were those same,
moving continental masses, which carried not only a living biota but fossils of animals and
plants that had lived, died and left their fossil remains on a continent that moved away from the
place where those fossils had lived. Once the continental "Ark" docked somewhere else in the
world, those fossils might be far from the place they had lived and died, and would be exotics in
the new location juxtaposed by the continental perigrinator.

As a result of the proposal of these several theories of past continental mobility, several
papers were produced in quick succession, many of a summary nature, that dealt directly with
the origin of the Australia and Australasian avifauna (Keast 1971, 1972, 1981, 1982, Serventy
1972, 1973, Cracraft 1972, 1973, 1980, Rich, 1973, 1975a, 1975b, 1976, 1981, 1982,
Schodde 1980, Schodde & Calaby 1972), The general outcome of these has been that most
suggest mixed origins for Australia's birds, some having a Gondwanan origin, while others are
northern invaders, some very recent. Some groups appear to have moved from south to north.

Rich (1975a) divided Australia's non-passeriform birds into two groups, one which probably
utilized the Indomalaysian route to move between Australia and the rest of the world, and a
second whose route was uncertain:

Indomalaysian Route (primarily southward movement):
Podicipedidae, Anhingidae, Ardeidae, Ciconiidae, Anatidae
in part (Cygnini, Tadomini, Anatini, Aythyini), Accipitridae
in part (Elaninae, Circinae, Accipitrinae), Pandonidae (?N or
?S movement), Falconidae, Phasianidae, Gruidae, Rallidae,
Otididae, Jacanidae, Charadriidae, Arenariidae, Phalaropididae,
Scolopacidae, Glareolidae, Laridae in part (Larinae), Cuculidae,
Strigidae, Tytonidae, Caprimulgidae, Hemiprocnidae (?N or ?S
movement), Apodidae, Alcedinidae, Meropidae, Coraciidae,
Bucerotidae.

Route Uncertain (Antarctic, Indomalaysian and in some cases
oceanic dispersal possible): Casuariidae (including the
Dromaiinae and the Casuariinae), Dromomithidae, Pelecanidae,
Phalacrocoracidae, Anatidae in part (Anseranatinae, Cereopsini,
Dendrocygnini, Oxyurini), Accipitridae (Milvinae, Perninae),
Megapodiidae, Turnicidae, Haematopodidae, Recurvirostridae,
Burhinidae, Laridae in part (Sterninae), Rostratulidae,
Pedionomidae, Columbidae, Psittaciformes, Podargidae,
Aegothelidae.

Cracraft (1972) had, previous to Rich's work, suggested that many of the groups in her
“Route Uncertain", such as the Dromornithidae, Casuariidac, Megapodiidae, as well as some
suboscine passeriforms and penguins, had utilized the Gondwana configuration to aid in
dispersal to or from Australia. These groups either traversed the landmass itself, or dispersed
along its margins in the continental shelf seas.

Certainly these ideas have appeal, especially when near relatives can be found on various
fragments of Gondwana. Olson & Steadman (1981) have pointed out the close relationships of
the Pedionomidae of Australia and the Thinocoridae (seedsnipes) of South America. One

MESOZOIC & TERTIARY BIRDS OF AUSTRALIA - 767

interpretation of this data could be that their present distribution reflects a vicariance event,
their common ancestor lived on Gondwana (or some part of it) and with the break-up of this
Supercontinent and the loss of communication between South America and Australia sometime
in the Eocene, the two populations drifted apart evolutionarily, leaving the current distribution
of related families. Another interpretation, which is also possible, but perhaps not as
appealing, is that these two families are relicts of a group that was once much more widely
distributed and has been replaced by other taxa in the Northern Hemisphere. Because the fossil
record of both of these groups is known only into the Quaternary, perhaps the Late Pliocene in
the case of the Pedionomidae, the dilemma yet remains.

Muluple hypotheses on area of origin and dispersal history have been presented for another
Australasian group, the Megapodiidae (the Moundbuilders). Cracraft (1972) supported a
Gondwana dispersal, but both Olson (1980) and Rich (1975a) were unwilling to make a final
decision on the basis of current evidence. Olson favoured an Indomalaysian origin, and pointed
out that within the galliforms the Megapodiidae and the Phasianidae are ecological competitors,
and they have a mutually exclusive distribution. He further pointed out that the phasianids are
primarily restricted to continental Asia and the larger islands in the Indomalaysian archipelago
and practically absent east of Wallace's Line. He suggested that the non-migrating phasianids
were unable to reach Australia and the megapodes, which once may have had a much larger
distribution, remained as relicts in Australia, to which the phasianids were unable to disperse.
In 1982 Mourer-Chauvire reported the occurrence of megapode bones in the Late Eocene of
France (Quercy), and suggested that this evidence supported Olson's hypothesis. Until older
material of birds is found in Australia, at least as old as the French record, it is difficult to
make a convincing argument as to which direction the megapodes moved. They could have
moved northwards from Australia into the Asian region via island-hopping, even at a time
before phasianids had a foothold there, or perhaps even before phasianids had developed (the
oldest known phasianid occurs in the late Oligocene of Europe (Palaeortyx, Olson 1986) and
not until the Miocene of Asia (Linquornis, Rich et al. 1986).

Schodde (1980) and others have suggested that perhaps further groups of birds placed in the
uncertain category by Rich (1975a) might have utilized the Gondwana route: the
Caprimulgiformes, the vanelline plovers, the cuculine cuckoos, tytonid owls and perhaps even
the Australian warblers, wrens, babblers, tree-creepers, robins, flycatchers and butcherbirds. To
this list should be added many of the highly endemic Australian songbirds such as the
Menuridae, Atrichornithidae and the Grallinidae and their near relatives. Here again, however,
the basis for denoting such groups as of possible Gondwanan origin is (1) their endemic nature
and (2) the lack of near relatives in the Old World. Unfortunately, so far, the fossil record of
birds in Australia, Asia, Antarctica and South America is not sufficiently long enough and/or
diverse enough to properly test these biogeographic hypothesis. The record in North America
and Europe is reasonably good, but to adequately test any biogeographic hypothesis regarding
Gondwanan vs. Northern Hemisphere origin, a good record on the southern continents is also
necessary, even despite the interesting findings of such groups as the Podargidae and
Aegothelidae in the Eocene of France (Mourer-Chauvire (1982). Even at this stage there may
have been possibilities of northward (or southward) dispersal of birds along an island chain
from Asia to Australia. ;

We are lefi, then, with at least two possibilities (both of which are in certain cases correct,
but both of which are very dependent upon the group being considered) to explain where many
of Australia's birds came from. One hypothesis is that some terrestrial birds moved back and
forth across a Gondwana landmass that existed in parts up until the Eocene. Highly endemic
forms, those that are familially distinct from birds elsewhere, are the most likely to fit this
hypothesis. A second hypothesis is that birds utilized an Indomalaysian dispersal route. Those
groups with low diversity in Australia, high diversity in the Old World and specific or generic
identity between the two areas, scem very likely candidates to have utilized this second route.

768 - RICH

Such groups as the Casuariidae, Dromornithidae, Pedionomidae, the Columbidae, the
Psittacidae and many of the highly endemic Australian Passeriformes are good candidates for a
Gondwanan origin - they are highly endemic and their nearest relatives may be in South
America. Other groups such as the Megapodiidae and the Caprimulgiformes are of uncertain
origin and will remain so until a better Southern Hemisphere record is at hand. There are some
worthwhile discussions that favour a northern origin for these groups, but they are not entirely
conclusive because of the possibility of northward dispersal as well as southward dispersal. In
short, one needs a good fossil record to document whether, indeed, parsimony is the best
explanation for what really happened.

SUMMARY AND CONCLUSIONS

The oldest remains of birds on the Australian continent come from Early Cretaceous rift
valley sediments in the southeastern part of the continent. These are only feathers, and it is not
until sometime in the Eocene that a few penguin bones continue this record. Only in the Late
Oligocene or Early Miocene does the record improve on this continent. The mid-Tertiary record
samples mainly fluviatile and lacustrine environments, and as such is highly biased towards
medium-sized to large birds whose living relatives frequent wetland environments.

Only a few groups of birds are useful biostratigraphically in the Australian sequences:
Dromornithidae, Dromaiinae, Pelecanidae and Phoenicopteridae. In the future such groups as
the Anatidae, Burhinidae and Rallidae may also be useful, but further study is needed before this
can be evaluated. The record of the remaining groups, although it may be excellent in the
Quaternary, does not have enough time depth to be of biostratigraphic value.

WINTER SUMMER

PRECIPITATION SEASONAL PRECIPITATION
ARIDITY —>

WINTER

N
&

Yv

|

ARID THRESHOLD
“—~ —~ — Dune building & Salt lakes

Figure 26. Climatic fluctuations characteristic of the last 20 million years on the Australian continent.
(After Bowler 1982).

Climatic change, tied to the northward drift of Australia during the history of birds in the
late Mesozoic and Cainozoic, has had a distinct effect on the composition of the Australian
avifauna. Several groups of birds ( e.g. Phoenicopteridae, Palaelodidae, Dromornithidae) that
prospered in the wetter, more forested conditions of the Miocene and as late as the Pliocene,

MESOZOIC & TERTIARY BIRDS OF AUSTRALIA - 769

became extinct when the fluctuations of the Quaternary (Fig. 26) became the norm rather than
the exception (see Frakes & Vickers-Rich, this volume). Man most likely had an effect on the
final extermination of such groups as the Dromornithidae and the giant Megapodiidae, for there
was certainly time overlap during the Pleistocene. The exact effect is hard to estimate,
however, because of the lack of precise dating of many Quaternary sites and the rarity of sites
where now extinct forms are associated with human activity. Human effects were not only in
direct killing of birds but also in habitat alteration through burning.

_ Although birds are known in Australia in sediments as old as Early Cretaceous, the first
diverse assemblages only occur in the Late Oligocene and Early Miocene, and most of these are
from central or northern Australia, It is, thus, difficult to speculate with much conviction on
the origin of many of Australia's most characteristic birds. Both Gondwanan and
Indomalaysian origins seem viable possibilities, both dispersal and vicariance have played a
role - but until the Palacogene and very latest Mesozoic record of birds on the Australian
continent is better understood, multiple hypotheses should be entertained for many groups,
especially the more endemic members of the avifauna.

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APPENDIX I

Appendix IA. (Page 776). Characteristic bones of avian families present in Australia: tarsometatarsi of A, Dromaius
novaehollandiae (Dromaiinae, emus); B, Podiceps auritus (Podicipedidae, grebes); C, Eudyptes chrysocome
(Spheniscidae, penguins); D, Diomedea sp. (Diomedeidae, albatrosses); E, Puffinus puffinus (Procellariidae, petrels); F,
Pelagodroma marina (Oceanitidae, storm petrels); G, Pelecanoides urinatrix (Pelecanoididae, diving petrels). All scales
except those labelled otherwise are 5 mm in length. Anterior views, with proximal views in D and F. All might
elements except C and G. Drawn by K. Fabb. (After Rich & Thompson 1982).

Appendix IB. (Page 777). Characteristic bones of avian families present in Australia: tarsometatarsi of A, Sula
dactylatra (Sulidae, boobies and gannets); B, Anhinga melanogaster (Anhingidae, darters); C, Phalacrocorax penicillatus
(Phalacrocoracidae, cormorants); D, Nyctanassa violacea (Ardeidae, herons); E, Threskiornis aethiopica (Plataleidae,
ibises); F, Melanitta sp. (Anatidae, ducks); G, Accipiter striatus (Accipitridae, hawks, eagles); H, Falco berigora
(Falconidae, falcons); 1, Lophortyx californicus(Phasianidae, quail); J, Tribonyx ventralis (Rallidae, rails); K, Burhinus
grallarius (Burhinidae, thick-knees). All scales are 5 mm in length. Anterior view in all but D, E, H, J-K, which are
proximal views. All right elements, except K. Drawn by K. Fabb. (after Rich & Thompson 1982).

Appendix IC. (Page 778). Characteristic bones of avian families present in Australia: tarsometatarsi of A, Laridae
(gulls and terns); B-C, Cacatua galerita (Psittaciformes, parrots); D, Ninox novaeseelandiae (Strigidae, hawk owls); E,
Aegotheles cristatus: F, Corcorax malanorhamphos (Corcoracidae, choughs). Humerus of H, Puffinus puffinus
(Procellariidae, petrels), All scales are 5 mm in length. A, B, D-F, anterior views; B, posterior view; H, anconal view.
Drawn by K. Fabb. (From Rich & Thompson 1982).

Appendix ID. (Pages 779-780). Characteristic tarsometatarsi of avian families present in Australia, proximal views:
A, Podiceps auritus (Podicepedidae, grebes); B, Eudyptes chrysocome (Spheniscidae, penguins); C, Puffinus puffinus
(Procellariidae, petrels); D, Pelecanoides urinatrix (Pelecanoididae, diving petrels): E, Sula dactylatra (Sulidae, boobies
and gannets); F, Anhinga melanogaster (Anhingidae, darters); G, Phalacrocorax penicillatus (Phalacrocoracidae,
cormorants); H, Melanitta sp. (Anatidae, ducks), I, Accipiter striatus (Accipitridae, hawks, eagles); J, Lophortyx
californicus (Phasianidae, quail); K, Laridae (gulls end tems); L, Cacatua galerita (Psittaciformes, parrots); M, Aegotheles
cristatus (Aegothelidae, owlet-nightjars); N, Ninox novaeseelandiae (Strigidae, hawk owls); O, Cuculus pyrrhophanus
(Cuculidae, cuckoos); P, Corcorax melanorhyamphos (Corcoracidae, choughs). Scale bar, 10 mm. Drawn by K. Fabb.
(From Rich & Thompson 1982).

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PLATES

Plate 1. A variety of fossil feathers from the Koonwarra Locality, south Gippsland, Victoria. Specimens
are preserved in a lake deposit of Early Cretaceous age. Scale bar, 10 mm.

Plate 2. Dromomithid footprints from the mid-Tertiary deposits of Pioneer, northeastem Tasmania. Scale
bar, approx. 250 mm. (From Rich 1979).

Plate 3. Dromomithid footprint from Pioneer, northeastem Tasmania. Scale showing both inches and
centimetres. (From Rich 1979).

Plate 4. Jaw fragments of Genyornis newtoni from Lake Callabonna, a Pleistocene-aged locale in South
Australia. A, and upper part of D, upper jaws in dorsal and lateral views respectively; B-D (lower), lower jaws
in dorsal and lateral views respectively. Length of B, approx 240 mm. (After Stirling 1913).

Plate 5. Dromomithid cranial remains. A, basicranium showing foramen magnum and occipital condyle of
dromomithid from the Miocene Bullock Creek locality in the Norther Territory,; occipital condyle approx.
15 mm in diameter; B, C, quadrates of dromomithids from same locality in anterior and posterior views
respectively. Scale bar, approx. 10 mm.

Plate 6. Cranial remains of Pleistocene Genyornis newtoni, a dromornithid, from Lake Callabonna, South
Australia. A, skull in side view, approx. 300 mm in total length; B, C, occipital condyle in dorsal and
posterior views; D-G, quadrate in external, external, medial, medial and ventral views respectively. Scale bar,
20 mm, applies to B-G. (From Stirling 1913).

Plate 7. Pelvis of Genyornis newtoni from Lake Callabonna, South Australia (A) and an extant cassowary
(Casuarius). Genyornis pelvis approx. 600 mm in length; cassowary scaled accordingly. (After Stirling
1913).

Plate 8. Dromornis stirtoni, a dromomithid, from the Miocene Alcoota locality, Northem Territory. Atlas
vertebra: A, anterior, B, posterior, and C, lateral views; height of vertebra approx. 65 mm. D, cervico-dorsal
or cervical vertebra, anterior view, approx. 94 mm in height. E, Genyornis newtoni from the Pleistocene
Lake Callabonna locale, left radius, ulna and carpometacarpus, approx. 150 mm in total length. (After Rich
1979).

Plate 9. Dromornis stirtoni, a dromomithid from the Miocene Alcoota locale, Northem Territory. Caudal
vertebra: A, lateral, B, posterior, C, dorsal, D, anterior and E, ventral views. Right carpometacarpus: F,
medial, G, lateral and H, proximal views. A, approx. 35 mm; all others to same scale.

Plate 10. A selection of diagnostic bones of Dromormnithidae from mid- to late Cainozoic deposits of
Australia. Femora of A, Barawertornis tedfordi, Riversleigh, Queensland, Oligo-Miocene (distal width, 87
mm); B-C, Bullockornis planei, Bullock Creek, Northem Territory, Miocene (distal widths 160 mm and >152
mm); D, Dromornis stirtoni, Alcoota, Norther Territory, Miocene (distal width 202 mm); E, //bandornis
woodburnei, Alcoota, Northem Territory, Miocene (distal width 112 mm); F, Dromornis australis, Peak Downs,
Queensland, probably Pliocene (distal width 120 mm). G, Genyornis newtoni, proximal right humerus, Lake
Callabonna, South Australia, Pleistocene (depth from external to internal tuberosity, 25 mm); H-K, //bandornis
sp., characteristic ungual phalanx of pes, Alcoota, Norther Territory, Miocene (total length, 28 mm); L-M,
Dromornis stirtoni, stemum, Alcoota, Northern Territory, Miocene (maximum width across stemocoracoidal
processes approx 225 mm); N, Dromornis stirtoni, scapulocoracoid, Alcoota, Northern Territory, Miocene
(total length >239 mm); O, /lbandornis lawsoni, proximal view of left tibiotarsus, Alcoota, Northem
Territory, Miocene (maximum depth about 88 mm); and P, //bandornis sp., distal end, right tibiotarsus,
Alcoota, Northern Territory, Miocene (distal width, 76 mm). (After Rich 1980).

Plate 11. Femora of Dromornithidae (proximal view) from Australian Cainozoic sediments: A,
Barawertornis tedfordi, Riversleigh Homestead, Queensland, Olio-Miocene; B-C, Biullockornis planei, Bullock
Creek, Northem Territory, Miocene; D, Dromornis australis, Peak Downs, Queensland, Pliocene; E, Dromornis
stirtoni and F, Ilbandornis woodburnei, Alcoota Homestead, Northem Territory, Miocene. See pl. 10 for sizes
of corresponding femora. (After Rich 1979).

782 - RICH

Plate 12. Femora of Dromornithidae (distal end) from Australian Cainozoic sediments: A, Barawertornis
tedforde, B-C, Bullockornis planei,D, Dromornis australis, E, Dromornis stirtoni. See caption for Pl. 11.
(After Rich 1979).

Plate 13. Tarsometatarsi of Dromomithidae (anterior view) from Australian Cainozoic sediments: A,
Barawertornis tedfordi, Riversleigh Homestead, Queensland, Oligo-Miocene; B, Bullockornis planei, Bullock
Creek, Northem Territory, Miocene; C, Dromornis stirtoni, Alcoota Homestead, Northem Territory, Miocene.
Width across distal end of A, 61.4 mm; B, 120.3 mm; C, 150.2mm. (After Rich 1979).

Plate 14. Tarsometatarsi of Dromomithidae (posterior view) from Australian Cainozoic deposits: A,
Barawertornis tedfordi; B, Bullockornis sp.; C, Bullockornis planei, D, Dromornis stirtoni. A, C, D, same
elements as illustrated in Pl. 13. B, Bullock Creek, Northern Territory, proximal width, 75.6 mm. (After Rich
1979).

Plate 15. Tarsometatarsi of Dromornithidae (anterior view) from Australian Cainozoic sediments: A,
Ilbandornis woodburnei, Alcoota Homestead, Northern Territory, Miocene; B-C, //bandornsi lawsoni, Alcoota
Homestead, Northem Territory, Miocene (B, proximal; C, distal ends). A, proximal width, 91.8 mm; _ B, width of
proximal end approx. 67 mm; width of distal end approx. 71 mm.

Plate 16. Tarsometatarsi of Dromomithidae (distal view) from Australian Cainozoic deposits: A, Barawertornis
tedfordi; B, Bullockornis planei; C-D, Dromornis stirtoni; E, Ilbandornis lawsoni, and F, Ilbandornis woodburnei.
Distal end width: A, 61.4 mm; B, 120.3 mm; C, approx. 150 mm; D, 150.2 mm; E, 71.4 mm; and F, same scale
as A in Pl. 15. (After Rich 1979).

Plate 17. Tarsometatarsi of Dromomithidae (proximal view) from Australian Cainozoic sediments: A,
Bullockornis planei and B, Bullockornis sp., Bullock Creek, Northern Territory, Miocene; C, Ilbandornis
woodburnei, Alcoota Homestead, Northern Territory, Miocene. Dromomithidae, phalanx 2 or 3 of digit III or
phalanx 2 of digit IV: D, proximal, E, distal, F, dorsal and G, side views, Leaf Locality, Lake Ngapakaldi, South
Australia, Miocene. Dromornis stirtoni, left plananx 1 of digit II: H, proximal, I, dorsal, J, medial, K ventral and
L, distal views. I/bandornis sp., phalanx 3 of digit III: M, side, N, proximal, O, dorsal, P, ventral and Q, distal
views, Alcoota Homestead, Northem Territory, Miocene. Proximal width A, 116.0 mm; B, 75.6 mm; C, 91.8 mm.
Proximal width: D, approx. 28 mm; D-G, all to same scale. Total length, I-J, 43 mm; H-K, all to same scale.
Width, O-P, approx. 24 mm; M-Q, all to same scale. (From Rich 1979).

Plate 18. Class indeterminate (perhaps dromomithid, but perhaps marsupial), terminal phalanx: A, dorsal; B,
ventral; C, medial or lateral; and D, proximal views; Alcoota Homestead, Northem Territory, Miocene. Length of
A-C, approx. 73 mm; all to same scale. (After Rich 1979).

Plate 19. Egg of a large bird, perhaps dromomithid, found in the coastal dunes between the Scott River and the
Southem Ocean, southwestern Westem Australia, Quatemary. Scale along bottom of photograph is approx. 280
mm (12 inches) in length. (From Rich 1979).

Plate 20. Dromaius gidju from the Miocene deposits of Leaf Locality, Lake Ngapakaldi, South Australia: A-A’,
stereopair of the pes; B-B', stereopair of the proximal view of the tarso metatarsus; right tarsometatarsus in

ventral (C) and dorsal (D) views; distal end of right tibiotarsus in dorsal (E) and ventral (F) views. (After Rich &
Baird 1986),

Plate 21. Casuarius lydekkeri from Quatemary-aged Pureni locality, Papua New Guinea. Pelvis and synsacrum
in lateral (A) and dorsal (B) views; femora, right (CPC26605b) in posterior (C), distal (E) and anterior views; left

(CPC26605c) in posterior (D), distal (F) and anterior (H) views. Scale bar, 10 mm. (After Rich, Plane and
Schroeder 1988).

Plate 22. Casuarius lydekkeri from Quaternary-aged Pureni locality, Papua New Guinea. A, right tibiotarsus,
internal view; B, left tibiotarsus, proximal view; C-D, right tibiotarsi, posterior view; E-F, ibid., external views;
G-H, ibid., anterior views; I, right tarsometatarsus, proximal view; J, right tarsometatarsus, anterior or dorsal
view; K, left tarsometatarsus, anterior or dorsal view; L, right tarsometatarsus, proximal view; M, left
larsometatarsus, anteriror view; N, left tarsometatarsus, distal view. Scale bar. 10 mm. (After Rich, Plane and
Schroeder 1988).

MESOZOIC & TERTIARY BIRDS OF AUSTRALIA - 783

Plate 23. Fossil pelicans of Australia. A, Pelecanus cf. conspicillatus, anterior sternal fragment; B, anterior 3rd
cervical vertebra, C, Archaeocycnus lacustris, 4th cervical vertebra, a pelican, not a swan as originally thought
by de Vis. Cf. P. conspicillatus: D, left quadrate fragment; E, left quadrate; F, distal left humerus; G, right scapula.
H-I, Cf. P. tirarensis, distal right tarsometatarsi. J, Cf. P. conspicillatus, distal left tarsometatarsus. K-L, P. ef.

cadimurka, distal left tarsometatarsus (K); distal right tarsometatarsus (L). | Scapula (G) length, 101.8 mm; all
other specimens to same scale. (From Rich & van Tets 1981).

Plate 24. Fossil pelicans of Australia. A, Pelecanus cf. cadimurka, 4th cervical vertebra; B, proximal nght
humerus thought to be pelican, but Accipitridae. P. cf. conspicillatus: C, left cuneiform; D, right cuneiform; E,
distal left ulna fragmen. F, P. validipes, distal right tarsometatarsus (=P. conspicillatus). F, P. grandiceps, distal
left tarsometatarsus (=P. conspicillatus). H, P. tirarensis, distal right tarsometatarsus. Length of 4th cervical
vertebra (A), 32.4 mm; all other specimens to same scale. (From Rich & van Tets 1981).

Plate 25. Palaclodid material from Neville’s Nirvanah, Lake Palankarinna, South Australia of Oligo-Miocene
age (Al, B1, Cl, D1). Selected views of Australian palaelodids (Al, B1, C1, D1) compared with Palaelodus
crassipes (A2, C2, D2) and P. gracilipes (B2). A, anterior views of tarsometatarsi; B, anterior views of tibiotarsi;
C, proximal views of tarsometatarsi; D, lateral views. Scale bar, 10 mm. (After Rich & Baird 1986).

Plate 26. Stereopairs of palaelodid material from Australia. A-B, distal end of a right tibiotarsus of the
youngest recorded palaelodid (Pleistocene) of Palaelodus n. sp. 2, in anterior (A-A') and distal (B-B') views. C-F,
Palaelodus n. sp. 1, known from Miocene to Pleistocene localities of northem South Australia: C-D, distal end of
left tarsometatarsus in anterior (C-C’) and distal (D-D') views; E-F, proximal end of right tarsometatarsus in
anterior (E-E’) and proximal (F-F’) views. (After Baird & Rich, in press).

Plate 27. Pedionomus torquatus from Fire-hole deposits of late Tertiary or Quaternary age, Morwell, Victoria.
A, partial skeleton in matrix including sternum (st), scapula (sc) and fragments of vertebrae and ribs.
Stereographic pairs: B-B', tibiotarsus (lateral view); C-C', ulna (proximal fragments, palmar view); /d-D', ulna
(distal fragment, anconal view); E-E', femur (anterior view). Proximal width of femur (E), 4.5 mm; all other
illustrations to same scale. (From Rich & McEvey 1980).

Plate 28. Synsacrum and partial skeleton of cf. Pedionomus torquatus from late Tertiary or Quaternary-aged
deposits in a lacustrine fire-hole deposit in coals at Morwell, Victoria. Although the age of this site is not agreed
upon, no differences could be found between this fossil and the living Plains Wanderer. A, in side view; B-B’, in
ventral view. Scale bar, 10mm. (After Rich & McEvey 1980).

Plates 29, 30. Partial skeleton of a primitive aegothelid, Quipollornis koniberi, from the mid-Miocene
lacustrine diatomite near Coonabarabran, eastern New South Wales. Scale bar, 10 mm. (From Rich & McEvey
1977).

784 - RICH PLATE 1

MESOZOIC & TERTIARY BIRDS OF AUSTRALIA - 785

PLATE 2

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786 - RICH PLATE

PLATE 5 MESOZOIC & TERTIARY BIRDS OF AUSTRALIA - 787

PLATE 6

788 - RICH

PLATE 7 MESOZOIC & TERTIARY BIRDS OF AUSTRALIA - 789

PLATE 8

790 - RICH PLATE 9

PLATE 10 MESOZOIC & TERTIARY BIRDS OF AUSTRALIA - 791

792 - RICH PLATE 11

PLATE 12

PLATE 13 MESOZOIC & TERTIARY BIRDS OF AUSTRALIA - 793

PLATE 14

794 - RICH

MESOZOIC & TERTIARY BIRDS OF AUSTRALIA - 795

PLATE 15

796 - RICH PLATE 16

MESOZOIC & TERTIARY BIRDS OF AUSTRALIA - 797

PLATE 17

798 - RICH PLATE 18

PLATE 19

PLATE 20 MESOZOIC & TERTIARY BIRDS OF AUSTRALIA - 799

800 - RICH PLATE 21

PLATE 22 MESOZOIC & TERTIARY BIRDS OF AUSTRALIA - 801

802 - RICH

PLATE

23

PLATE 24 MESOZOIC & TERTIARY BIRDS OF AUSTRALIA - 803

PLATE 25

es
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PLATE 26

804 - RICH

1 cm

MESOZOIC & TERTIARY BIRDS OF AUSTRALIA - 805

27

PLATE

806 - RICH PLATE 28

PLATE 29

PLATE 30 MESOZOIC & TERTIARY BIRDS OF AUSTRALIA - 807

808 - RICH

Anthropornis nordenskjoeldi, a giant, primitive penguin that lived in Eocene times along the
shore of the widening southern rift as Australia split from Antarctica. (From Rich & van Tets
1985, with permission of The Museum of Victoria).

CHAPTER 21

AVIAN FOSSILS FROM THE
QUATERNARY OF AUSTRALIA

Robert F. Baird!

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Australian Avian Palacontologists ............0.0000- 810
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Physical Changes Occurring During
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Prior to the Matuyama/Bruhnes
Magnetic Boundary ................cceeeeees $13
Post-Matuyama/Bruhnes Magnetic
Reversal Boundary .............c0eceeeeeeees 814

Faunal Changes, Biogeography and
Palacoenvironment during the

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Early and Middle Pleistocene ................. 819

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Penultimate Interglacial ................... 819

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Discussion of the Quaternary............. cee 821

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Birds in Quaternary Biostratigraphy ................5. 822
Family Accounts for Australian

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1 Earth Sciences Department, Monash University, Clayton, Victoria 3168 Australia.

810 - BAIRD
INTRODUCTION

The study of Quaternary avian assemblages was once considered only a small part of the
wider field, avian palaeontology. Quaternary birds are now studied by a wide range of people
throughout the world, and because of the depth of know!edge in this subfield, it can now be
regarded as a significant field in its own right. This is especially true for the well populated
continents of North America, Europe and Asia (e.g. China and Japan). It is a relatively new
area of study in Australia, and, therefore, there is a need for an in depth overview of the
information available to this point in time. Towards this end, this chapter gives a brief
overview of the changes in the geography, precipitation, temperature and vegetation within the
Quaternary. It also covers the faunal changes, biogeography and environmental reconstructions
produced through the study of fossil bird assemblages. Further, a discussion of each family is
presented which contains accounts of the fossil record with minimal interpretation.

Only avian fossils from continental Australia and Tasmania are covered in this chapter;
those from islands, both continental and oceanic, are covered in Meredith (Chap. 28, this
volume). The bulk of the data in this chapter are unpublished but are available either from the
author or in Baird (1986c). Similar coverage for the avifauna of the Tertiary of Australasia can
be seen in Rich (Chap. 20, this volume).

Geographic distribution, and specics lists for each of the individual Quaternary localities are
included in Appendix I.

AUSTRALIAN AVIAN PALAEONTOLOGISTS

There have been a sufficiently small number of workers in the field of avian palaeontology
in Australia to give a brief overview of their work.

Baird, Robert F.: Baird arrived in Australia in 1982 to begin a Ph.D. thesis in avian
palaeontology under P.V. Rich at Monash University. The thesis was completed in 1986 and
concentrated on the avian assemblages in cave deposits across southern Australia (Baird 1986c).
Both palaeoenvironmental and biogeographic interpretations were made on these data. Baird has
largely concentrated on avian assemblages in Quaternary cave deposits, including Green
Waterhole (South Australia: Baird 1985), Amphitheatre Cave (Victoria: Baird, submitted b),
and their use in palacoenvironmental analysis (Baird 1989). He has also published on single
species (¢.g. Gallinula mortierii (Tasmanian Native-hen)(Baird 1984, 1986a, submitted)), as
well as coauthored a review of Australian avian palaeontology (Rich & Baird 1986).

de Vis, Charles W.: De Vis arrived in Australia in 1870. He worked mostly on
Quaternary vertebrate material out of the Queensland Museum between the years 1888-1911.
His work with avian fossils was largely restricted to non-passeriformes. It should be noted that
many of deVis’ identifications of avian fossils were incorrect (e.g. van Tets 1974a; Olson
1975, 1977; van Tets & Rich 1980). The reason for the numerous misidentifications, most
frequently put forward, is that the collections of comparative material at that time were poor.
His identifications must all be reviewed and not relied upon.

McEvey, Allen R.: McEvey served as curator of the Department of Ornithology,
Museum of Victoria between 1955 and 1984 where he began work on fossil and subfossil avian
remains with a paper on the Macquarie Island penguin deposits (McEvey & Vestjens 1974).
Since that time, he has been a coauthor on papers describing either single fossil remains (e.g.
Falco berigora (Brown Falcon), Rich et al. 1982) or partial associated skeletons (Pedionomus
(Plains-wanderer), Rich & McEvey 1980).

Meredith, Charles W.: Meredith began his career in avian palaeontology in 1980 when
he commenced his thesis covering the subfossil deposits of Norfolk Island. His works to date
cover ihe fossil faunas of Macquarie and Norfolk islands (Meredith 1985b, Meredith er al. 1985)

QUATERNARY AVIFAUNA OF AUSTRALIA - 811

as well as a summary of fossil avian assemblages from islands (Meredith, Chap. 28, this
volume). Although no longer working full time in this field, Meredith plans on publishing
the information brought forward in his thesis (Meredith 1985b) on the phylogeny of the genus
Pterodroma and the fossil deposits of Norfolk Island.

Miller, Alden: Miller described the material collected by R.A. Stirton, R.H. Tedford and
P. Lawson in the early 1950's from both the Tertiary deposits (in northern South Australia) and
Quaternary deposits (along Cooper Creek). His papers covering Quaternary material include
those on the Dromaiidae (emus: Miller 1962), Pelecanus (pelicans: Miller 1966b), Anhinga
(darters: Miller 1966a) and Phoenicopteridae (flamingoes: Miller 1963). Unfortunately, Miller
suffered an untimely death, leaving further study of this material to others.

Olson, Storrs L.: Olson’s work on Australasian Quaternary material stems from his
visit to Australia for the International Ornithological Congress held in Canberra in 1974. He
has a firm belief that the most productive time searching for new material can be spent in
museum collections. Therefore it is not surprising that his three papers to date are based on
restudies of previously published material, including Gallinula mortierii (Olson 1975), Chionis
(sheathbills: a rebuttal; Olson 1976) and fossil anatids (Olson 1977).

Rich, Patricia V.: Rich first started work on Australian fossil material in 1967 when
she began her Ph.D. at the AMNH on the dromornithid material collected by R. A. Stirton and
collegues. Since that time, she has worked at the Museum of Victoria on a Fulbright
Scholarship and now is Reader in Earth Sciences at Monash University. Rich concentrated her
work on Australian avian fossil non-passerines, with papers covering the dromornithids (Rich
1979), dromaiids (emus: Patterson & Rich 1987), phoenicopterids (flamingoes: Rich et al.
1987), Pelecanus (pelicans: Rich & van Tets 1981), Pedionomus (Plains-wanderers: Rich &
McEvey 1980) and aegothelids (owlet-nightjars: Rich & Scarlett 1977). Rich has also studied
single elements of tytonids (barn-owls: Rich et al. 1978) and falconids (falcons: Rich et al.
1982).

She has also been the senior author on several review papers with van Tets (1982) and Baird
(1986) and the senior editor of two compendiums on Australasian fossil vertebrates (Rich &
Thompson 1982 and this volume).

van Tets, Gerry F.: Van Tets arrived in Australia from Canada in 1963 and has, since
approximately 1970, completed projects on Quaternary cave deposits of Weekes and Victoria
Fossil caves (van Tets 1974b, van Tets & Smith 1974) and taxa occurring into the Quaternary
including Progura (Giant Megapode: van Tets 1974a) and pelicans (Rich & van Tets 1981).
Along with numerous small papers, he has also coauthored chapters on Australian avian
palaeontology (Rich & van Tets 1982) as well as compiling a checklist of extinct Australian
avian taxa (van Tets 1984).

THE QUATERNARY

The Quaternary represents the time during which the reoccurring cycles of glacials and
interglacials occurred at discrete intervals. It is these cycles and the changes in climate
concommittant with them that have influenced the patterns of distribution of Australia's biota
(Fig. 1). Understanding the pattern of these cycles is, therefore, critical to the full
understanding of Australia's flora and fauna. This knowledge may aid in the production of
hypotheses on the current distributions, past ecologies and fossil histories of bird species.

The Quaternary is traditionally broken up into two epochs, the Pleistocene and the
Holocene. For the Pleistocene, convention has placed a lower boundary at 1.85 million years
(Berggren & van Couvering 1974) and the upper boundary at 10,000 years before present (yBP:
Savage & Russell 1983). The upper boundary is defined by the last glacial / interglacial
interface. Consequently the lower boundary of the Holocene is also 10,000 yBP, and it

continues through to the present day.

812 - BAIRD

4

|

W/ZA Irian
Wl Tumbunan
Torresian
[_] Eyrean
_ Bassian

Figure 1: Map of Australian zoogeographic zones.

The localities with associated avian elements dated between 1.85 myBP and 50,000 yBP in
Australia are positioned chronologically through relative dating (i.e. Cooper's Creek, Darling
Downs and Lake Eyre localities) and there are only a handful of these. Much of the Quaternary
avian material is restricted to the last 30,000 yBP with the occasional deposit extending to the
limit of radiocarbon dating at 50,000 yBP Therefore, the record largely concerns the last two
percent of the Quaternary. In most cases, understanding the methods and biases of dating fossil

material can be as important as other sources of information. See Gillespie (1986) for a
background in radiocarbon and Berry (1968) for relative dating.

PHYSICAL CHANGES OCCURRING DURING THE QUATERNARY

The physical changes in the Quaternary are part of a continuity of changes across time, and
only fully understood if placed in this context. The reader is referred to Frakes et al. (1987) for
the sequence of changes in the Australian landmass since the Palaeozoic. In order to understand
the environmental constraints placed upon the avifauna within the past 50,000 years, it is
necessary to have a firm background in the physical changes occurring in that period of time.
The full cycles were approximately 100,000 years in length with cold and dry (glacial) and,

QUATERNARY AVIFAUNA OF AUSTRALIA - 813

warm and wet (interglacial) periods within each cycle. We currently have a problem with
dating many events within the Pleistocene Period (outside of the last 50,000 years) due to the
lack of reliable dating methods for the types of data materials available. I will give a brief
outline of some of these changes concentrating on those changes effecting the southern part of
continental Australia, for it is here where most of the avian assemblages are located.
Information on changes in the physiography and/or vegetation in northern Australia can be
obtained from Kershaw (1986) and Nix (1982). Current physiography and distribution of

vegetation formations can be seen in Jennings & Mabbutt (1977) and Specht (1981)
respectively.

|
\
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\
LAKE BUNGUNNIA

ADELAIDE

Figure 2: The approximate extent of Lake Bungunnia (from Bowler 1979).

Prior to the Matuyama/Bruhnes Magnetic Boundary (1.85 myBP to
700,000 yBP)

The numerous glacial and interglacial cycles typical of this sub-era can be characterized as a
pattern in which: "interglacials were short lived, occupying about 10 percent of each cycle.
The rest of the cycle was taken up by a saw-toothed build-up towards glacial maximum and sea-
level minimum (Broeker and van Donk 1970)" (Williams 1984a).

Williams (1984a) stated that Australia would have been effected by each and every glacial
period recorded in the Northern Hemisphere, which he estimated to be approximately 20.
Unfortunately, due to a number of factors, only the major changes in climatic regimes are

814 - BAIRD

recorded in the Australian terrestrial record. Of these, only the last four are defined with
certainty. Bowler (1979) pointed out that:

"in southeastern Australia, aeolian deposits of the last glacial age correlate well with those
from other parts of the globe. However, the known sequences of episodic deposition and soil
formation are limited to four or five cycles that characterize the loess of Bruhnes age in
Czechoslovakia and China."

Therefore, little is recorded in Australia of the 13 glacial periods known to have occurred in
the Northern Hemisphere prior to the Bruhnes palaeomagnetic period.

Sea-level, Tectonism and Deposition

The major feature influencing avian distribution in southern Australia would have been
Lake Bungunnia. The formation of Lake Bungunnia occurred sometime during the Pliocene by
the uplift of the Mallee Ridge near Overland Corner, South Australia. The presence of this
lake dominated the south eastern landscape during this period (see Fig. 2).

Flora

Less is known of the flora for this time period than that for any other period of time in the
Cainozoic. The only chronologically reliable study to date spanning this period is that of J.
McEwan Mason (Ph.D. thesis, submitted Dec. 1989, Monash University, Clayton). The
vegetation history interpreted from cores in Lake George, New South Walcs suggest an open
forest/woodland with abundant Casuarina, Compositae, Graminaceae and chenopods, with small
amounts of gymnosperms. Just after the Pliocene/Quaternary boundary gymnosperms are no
longer recorded. The remaining elements are continuous to the Matuyama/Bruhnes boundary.

Precipitation and Temperature.

There are no detailed histories of precipitation for the Early Pleistocene of Australia,
although models have been presented to explain the development of environmentally-produced-
landforms currently exposed today. One such model suggested that by the Pliocene/Pleistocene
boundary, sub-tropical high pressure cells would have been pushed north into their present-day
position by the cooling waters between Australia and Antarctica. This would have produced
winter precipitation in southern Australia, and started oscillations in precipitation, which would
increase in amplitude throughout the Quaternary (see Fig. 3).

Post-Matuyama/Bruhnes magnetic reversal boundary (700,000 yBP
to present)

Very little of the physical changes occuring during this period of time are known beyond
the last interglacial/glacial cycle (é.e. 120,000 years). This is the result of several problems
regarding absolute dating, particularly dating past the limit for 14C. Although there are
geomorphic features, fossil animal assemblages and microfloral assemblages which through
their unique compositions are considered to be older than the last interglacial, no absolute date
can be associated with them. Therefore, these proxy data cannot be used as indicators of past
climates in any meaningful sense. Exceptions to the rule are the microfloral assemblages from
Lake George, magnetic reversal dated from 700,000 yBP. The following section will,
therefore, discuss the period between 120,000 yBP and present-day, concentrating on the last
40,000 yBP (regarded as the limit for standard carbon-14 dating).

Sea-level, Tectonism and Deposition.
The loss of Lake Bungunnia is currently considered to have occurred about 500,000 yBP
based upon magnetic reversal ages of less than 700,000 yBP and inferred constant

sedimentaticn rates for the top 4 metres. A date of 400,000 yBP is considered likely for total
desiccation of the lake (Bowler 1979, 1982).

QUATERNARY AVIFAUNA OF AUSTRALIA - 815

WINTER : SUMMER

PRECIPITATION SEASONAL PRECIPITATION

ARIDITY —>|
WINTER

ARID THRESHOLD
“~~ Dune building & Salt lakes

Figure 3: "Diagrammatic summary of changes in humidity experienced by the southern arid and semi-arid
regions of the continent since lower Miocene time. " (From Bowler 1982).

"The disappearance of Lake Bungunnia, presumably by the Murray River cutting an
overflow channel across the Pinaroo Horst, represents the initiation of all subsequent mallee
landforms that formed within the confines of that lake. Thus a wide variety of dune forms,
salinas and fluvial terraces have originated within the past 700,000 years. Indeed, if the
evidence from Lake Tyrrell is representative of the broader region, such events must be placed
within the last 400,000 years" (Bowler 1980).

Since the sea-level high at the time of the last interglacial (approximately 120,000 yBP),
there have been a number of nadirs and peaks (see Fig. 4). Within this broader scale we can fit
the better known sequence of sea-level changes since approximately 40,000 yBP A summary
of these changes is provided in Chappell (1983):

"(i) 40KA: sea level 30 to 40 m below present (Chappell 1974; Bloom et al. 1974).

(ii) 30KA: sea level 40 to 45 m below present (Chappell & Veeh 1978b)

(iii) I8KA: sea level 150 to 160 m below present (north Queensland, Veeh & Veevers
1970; northwest shelf, Jongsma 1970; also undated evidence in northeast Papua New Guinea)

(iv) 9KA to 6.5KA: rising smoothly from about 20m below present, up to present level
(many sites round Australia, Thom & Chappell 1978; northeast Papua new Guinea, Chappell
& Polach 1976)

(v) 6 KA to present: a maximum of around 1 to 2 metres above present reached at coastal
sites about 5.5 to 5.0 KA, falling smoothly to present (Chappell et al. 1982)".

Bowler et al. (1976) regarded the last period of extensive sand dune development to have
occurred during the height of the last glaciation. As a whole, the sand dune development in
Australia is probably as old as 300,000 years (see Fig. 5). This is supported by Williams
(1984a), who mentioned that "most of the depositional features of arid Australia are Tertiary or
younger, and the present duneficlds are probably little older than Late Pleistocene."

Flora
One of the primary assumptions for the biogeography of vegetation in the south-eastern
region is that "...since the Blanchetown Clay underlics such a large area of the mallee, its uppet

816 - BAIRD

age limit provides a reference point for which we may view the evolution of the subsequent
Malle[e] landscapes" (Bowler 1979). This upper age limit is considered to be 400,000 yBP.
Unfortunately no detailed vegetational history is available for this region during that period of
time.

REEF vila Vilb VI Y y eile :
years x 1000, before present
120 80 40 0
w 0 t rn =e 1 n 4 L 1: r 4 4 i i
2 ANS j \ a} om vi
=a / \ fo i \g / } j\e® i
2 -405 / \ / \ / \ A / ‘ 7 My / \ }
a Poss wes \ / Vy T/ \ TV eA
| \

eS i? \TI ; as TY i
D | iT t \

804 ' \ ai
© , ‘ \ Gi
2 \ /
=
& . New Guinea reef crests, dated (this paper): each dot is
Fd 120 from reef heights on traverse of figure 4.

New Guinea reef complexes, low sea level maxima,
undated (Chappell, 1974)

New Guinea reef complexes. dated (Veeh and Chappell, 1970)
Compilation of Steinen, et al (1973) - primarily Barbados data
Sea level minimum not known

rey N 4-4

—— Diagrammatic paleo sea level trace

Sea level using premature emergence correction (see text)

Figure 4: "Late Quaternary palaeo sea levels based on estimates from New Guinea and elsewhere.” (From
Bloom et al. 1974).

If the concept of synchroneity of climatic events can be considered to be valid (Martin
1983), then the last 190,000 years of microfloral changes available for northeastern Australia
may be extrapolated for the rest of Australia (Kershaw 1986). Unfortunately, southern
Australia and northern Australia are influenced by two different climatic patterns (i.e. west-east
anticyclonic fronts and mainly winter precipitation versus west-east cyclonic fronts and mainly
summer precipitation, respectively); this may not be applicable throughout, and, therefore,
only the records provided from southern Australia are considered here. Of the southern records,
only one is considered to extend back to the Matuyama/Bruhnes boundary (Singh & Geissler
1985), but the sequence is fraught with sparse pollen (i.e. in some cases less than 20 grains
were used to determine percentages of all vegetation formations) and barren zones of unknown
duration. Thus, it is here considered largely unreliable for palaeoenvironmental interpretation.

Prior to 40,000 yBP. The vegetation present during the last interglacial, as interpreted from
analyses of pollen assemblages from Lake George, New South Wales and Pulbeena Swamp,
Tasmania, is interpreted to be forest formations. There was an abrupt change just after this
period which is regarded as the change from forest vegetation to herbfield and grasslands, and
subsequently between 65,000 to 40,000 yBP a major interstadial is suggested by the return of
forest vegetation (Kershaw 1981). The latter expansion of forest throughout south-eastern

QUATERNARY AVIFAUNA OF AUSTRALIA - 817

Australia is confirmed by the study of pollen assemblages fi - i
foeka isha 1 p ges from south-east South Australia

@ L. Frome

L forens => Willandra Lake
Seo

_ —> pee
wee wl Geo rge a a

we LL

Kangaroo Is. Se < i
L.Leake = L-Keilambete {(:

Figure 5: "Sand flow lines representing the trends in dunes believed to have been active during the last
glacial maximum" (Bowler 1978).

40,000 - 30,000 yBP. During this period forest vegetation largely gave way to heaths, and
grasslands in the extreme southeast of mainland Australia (Dodson 1974a, 1975; G. Hope
1984). Pollen assemblages to the north, from Lake George, New South Wales, and, to the
south, in Tasmania, suggest that forest formation was present throughout this period.

30,000 - 20,000 yBP. This period is considered a time of large changes in the vegetation,
where forest formation replaced grassland and herbfield vegetation in the Lake George area, and
mallee eucalypt replaced chenopod low scrub formation on the Bunda Plateau (Martin 1973).
Microfloral assemblages from Pulbeena Swamp, Tasmania, and Cave Bay Cave, Hunter Island,
indicate the loss of all taxa representing forests and the dominance of grasses and composites.

20,000 - 10,000 yBP. Little information is currently available for this period of time,
except that from Lake George. The Lake George data suggests that the forest, by this time, had

818 - BAIRD

been completely replaced by grasses and composites (Singh 1983). Those records previously
mentioned, with cores continuous through this period, demonstrate similar flora.

10,000 yBP - Present. Information available from the Snowy Mountains suggest that
treeline had not returned to present day level until 8,600 yBP, prior to which only alpine
herbfield was present from at least 16,500 yBP (Bowler et al. 1976). This period of time is
also characterized by the return of forest to areas where it occurred prior to 40,000 yBP.

Precipitation and Temperature.

Bowler (1982) considered that "...the last time that [climatic] conditions approximated those
of today, that is the last interglacial, occurred about 120,000 years ago (Shackleton & Opdyke
1973)". He then extrapolated this in stating "suffice it to say that the pattern evident in the
past 100,000 years is thought to have been repeated at least four or perhaps five times within
the last 500,000 years". Therefore, the summary presented below may be indicative of a
number of glacial/interglacial cycles.

Prior to 40,000 yBP. Based upon pollen analysis, conditions during the last interglacial
were similar to those at present, after which there was sharp decline in rainfall. Between the
period of 65,000 to 40,000 yBP rainfall increased relative to the period immediately preceding
it, but did not reach present day levels (Kershaw 1981). Geomorphic evidence also suggests
high rainfall, but at the slightly later time of 50,000 and 30,000 yBP. Dating remains difficult
for these events.

40,000 - 30,000 yBP. The coastal and sub-coastal sites of southeastern Australia indicate a
change in climate to temperatures and precipitation that "...were significantly lower than
today", and lower than the period just previous, based upon studies of pollen assemblages
(Kershaw 1981). This is different from the bulk of the geomorphic and palynologic evidence
from inland New South Wales, which suggests that high effective precipitation occurred up
until 25,000 yBP (Bowler et al. 1976, Williams et al. 1986). A possible explanation for the
difference between this interpretation and that presented for more southerly localities is given
by Kershaw (1981). He suggests that "the reason for this difference could lie in the location of
the sites relative to the coast and direction of major rain-bearing winds". He goes on to suggest
that those localities now close to the coast would, at that time of low sea level, have lost the
maritime influences which would have resulted in lower rainfall. The more inland site,
however, would have been little effected by the relative lowering of sea level, and assuming the
temperature was colder than today, the relative evaporation rates would have produced a higher
effective rainfall. In addition, the major pressure belts may have changed in both intensity and
location (Rognon & Williams 1977), which may have also had an effect on this region.

30,000 - 20,000 yBP. Rainfall was similar to the previous period except in New South
Wales where proxy data indicate a decrease in effective precipitation. Bowler (1982) considered
the period between 25,000 and 14,000 yBP to be a time of water deficit, and suggests this is
due to lower precipitation.

20,000 - 10,000 yBP. Effective precipitation reached the lowest levels in the last 100,000
years (see Bowler et al. 1976, Bowler 1982 and Williams e¢ al. 1986) during this period.

10,000 yBP_ - present. By 10,000 yBP precipitation had increased to present day levels in
many areas, and between 7,000 and 5,000 yBP conditions were wetter than today (Dodson
1974b, 1975). A decrease in precipitation occurred after 5,000 yBP (Kershaw 1981), but,
subsequently, precipitation increased to its present day level.

QUATERNARY AVIFAUNA OF AUSTRALIA - 819

In general, the cooling during the last 40,000 yBP peaked around 18,000 yBP, with a
subsequent warming to present day levels by 6,000 yBP (Bowler et al. 1976). Peterson (1971)
indicated that it is unlikely for glaciers to have covered much of the southeastern highlands
during the period of maximum cooling. He favoured, instead, the idea of a small amount of
glaciation surrounding the summit of Mt Kosciusko. The deposits hitherto related to glacial
activity at lower elevations were probably the result of periglacial activity. "Dating of the
maximum of glaciation depends as yet on a single date of 20,200 + 165 yBP (Costin, 1972)"
(Bowler et al. 1976), which has been subsequently recalculated by Galloway & Kemp (1984) to
15,000 yBP. Colhoun & Peterson (1986) regard this recalculated date as "... a more expectable
date for the commencement of deglaciation".

FAUNAL CHANGES, BIOGEOGRAPHY AND PALAEOENVIRONMENT
DURING THE QUATERNARY

Early and Middle Pleistocene

The deposits considered to belong within this period of time have been placed there by
relative dating and, therefore, cannot be associated with any discrete period of time. The
proposed changes in fauna and timing of events should be considered tentative at best.

There are only a few elements considered to be restricted to the Early and Middle
Pleistocene, including the Palaclodidae, Phoenicopteridae (Xenorhynchopsis spp.), the large
form of Cacatua tenuirostris (Long-billed Corella) from Green Waterhole, South Australia
Centropus colossus (Giant Coucal: Pl. 1), Orthonyx hypsilophus (Giant Logrunner) and two
species of unidentified giant eagle. Other elements which occur from this period through to the
Holocene include, Progura (PI. 2) and Gallinula mortierii from Chinchilla Sands, Queensland
(see T.H. Rich et al. 1982).

There is insufficient data for a discussion on the nature of the avifauna of this period. The
emphasis on waterbirds represented in deposits of this age may be attributed to the types of
deposits from which the material comes (i.e. mainly fluvial deposits).

Late Pleistocene

Penultimate Interglacial

The avian assemblages in the late Quaternary can be divided chronologically into those from
the penultimate interglacial, those from the last glacial and those from the last interglacial (or
Holocene). The only deposit which may have an assemblage representing the last interglacial
would be that of Pyramids Cave, Victoria. Because there is no upper date on the assemblage
this assemblage may have been deposited during any of the previous interglacials. The
assemblage consists of a mixture of elements, including extinct species (Pycnoptilus n. sp.
(pilotbird) and Orthonyx sp. (logrunner)), elements which currently occur further north
(Atrichornis rufescens (Rufous Scrub-bird) and species which occur in the area today. The
Pyramids Cave assemblage indicates that Closed forest, particularly Complex Notophy! Vine
Forest was present within 2.5 km of Pyramids Cave. To maintain this type of forest an
increase of approximately 30-50% in summer rainfall and an increase of 1-2°C in mean annual
temperature would have been required sometime predating the oldest part of the Clogg's Cave
assemblage at 22,980 + 2,000 yBP. Therefore this assemblage probably represents an
interglacial phase, based upon these climatic requirements.

820 - BAIRD

Last Glacial

The avian assemblages associated with the height of the last glacial include species that are
now associated with drier habitats in deposits which currently occur in areas characterized today
by woodlands and forests. This phenomenon is recorded in three distinct centres including the
southeast, the Bunda Plateau and the southwest. These species include Pedionomus torquatus
(Plains-wanderer), Melopsittacus undulatus (Budgerigar), Halcyon pyrrhopygia (Red-backed
Kingfisher), Psephotus varius (Mulga Parrot), etc. For example, for the area around Clogg's
Cave, Victoria between 22,980 + 2,000 yBP and 8,720 + 230 yBP a regional habitat of either
very open eucalypt Woodland Formation or Savannah is indicated, suggesting a decrease in
effective precipitation during that period.

The palacoenvironment of the Bunda Plateau region over the past 37,000 years can be
summarized as follows: chenoepod low scrub formation present within 2.5 km of Madura Cave,
Western Australia in all periods of time covered by the deposit, including that of 37,880 +
3,880 yBP, 22,400 + 580 to 15,600 + 250 yBP and 7,470 + 120 yBP, and between 19,300 +
350 and 13,700 + 270 yBP for Koonalda Cave, South Ausiralia. Therefore, no change in
relative precipitation is indicated for this region throughout the last 37,000 years.

The palacoenvironment of the southwestern region of Western Australia for this period can
be summarized as follows: the area around Devil's Lair had climatic conditions similar to those
of today between 37,750 + 2500 yBP and 27,700 + 700 yBP. From sometime between
32,000 and 27,000 years, and 12,000 + 180 yBP the vegetation was a mixture of eucalypt
woodland formation, heathland formation and Acacia tall scrub formation indicating a lower
effective precipitation throughout this period.

Biogeographically there are a number of changes in the avian faunas which may reflect
changes in the environment. With the lowering of the sea level concommitant with the height
of the last glacial, it seems that the arid center expanded and forced the wetter habitats to follow
the sea level outwards.

Extinctions of avian taxa during the height of this last glacial were few. The species to
disappear are of such gigantic proportions, as compared with their modern day counterparts, to
suggest that they were part of the Late Pleistocene megafaunal extinctions and dwarfings
(Marshall & Corruccini 1978). These birds include Progura (giant megapode) and Genyornis
(PI. 3).

Several authors have suggested that immigrants to the continent (e.g. dingoes and
aborigines) were the sole cause of the vast array of continental extinctions in Australia (Jones
1968), as with North America (Martin 1984). Others suggest that these extinctions may not
have been directly caused by aboriginals but through their extensive use of fire. Large areas of
land may have been converted to habitats different from what would occur in the region under
natural fire regimes. Whether this burning was done in sufficient quantity to be the sole cause
of extinction of a number of animals is unknown. This scenario has yet to be sufficiently
corroborated to warrant full support, and a healthy skepticism may be warranted, especially as
additional information has come to the surface demonstrating that the megafauna coexisted with
Aboriginais for several thousand years (Gillespie e¢ al. 1978). It has been suggested that a
change in weather patterns from a stable, regular winter rainfall to an erratic rainfall in tandem
with the introduction of the Dingo (Canis familiaris dingo) may have caused at least the
Tasmanian Native-hen (Gallinula mortierii) to be relict to Tasmania (Baird 1984, submitted a).
It has been suggested that this occurred several times in the Late Pleistocene (Bowler 1982).

Holocene

For the most part assemblages which are believed to be Holocene are composed of
assemblages similar to those of pre-glacial sites but lacking extinct forms. Of the four Holo-
cene assemblages, those from Madura Cave, Devil's Lair and Skull Cave, Western Australia;

QUATERNARY AVIFAUNA OF AUSTRALIA - 821

and Amphitheatre Cave, Victoria are associated with radiocarbon dates. The avian assemblage
from Mabel Cave is considered to be Holocene based on relative dating.

The most interesting of the three radiocarbon-dated assemblages is that from Amphitheatre
Cave, which has one species now restricted to Tasmania (Gallinula mortierii) and two species
whose ranges are currently restricted to eastern Victoria (Ptilonorhynchus violaceus (Satin
Bowerbird) and Dasyornis brachypterus (Eastern Bristlebird: Baird, submitted b). The other
assemblages are comprised of species which occur around the respective sites today, and,
therefore, palaeoenvironmental interpretations of these assemblages lead to a mixture of
vegetation types similar to that which currently occurs in these areas.

@
o

oO e
@ o eo
Ss i) @
c 5)
< @ & e e
Se x S oN iS) oe RJ
wr << 2) Fs >
s So FF KS re pos
@ 4 > ne)
< Ss Oo x
ic) se o 3 OS gr

Wy

Yy
Yy

GY, BD

[Se Rei

Years Before Present (x 1000)

X radiocarbon date A+ relative date. — - conformable sequence uncontormable sec major uncontormity conjectural Coverage

Figure 6: Graphic representation of the proposed vegetations interpreted from the avian assemblages from
the cave deposits included in Baird 1987) (i.e. WI-6le = Devil's Lair, AU-8 = Skull Cave, N-4 = Koonalda
Cave, N-62 = Madura Cave, L-81 = Green Waterhole Cave, G-4 = Curran's Creek Cave, G-4 = Amphitheatre
Cave, EB-1 = Mabel Cave, EB-2 = Clogg's Cave, and M-89 = Pyramids Cave). no palaeoenvironmental
interpretation was available from L-81 due to the non-contemporaneous nature of the deposit. Shading

indicates vegetation formations with black = closed forest formation, grey = forest
formation, cross-hatched = woodland formation, light stipple = Acacia tall scrub
formation and dark stipple = chenopod low shrub formation.

DISCUSSION OF THE QUATERNARY

Climatic conditions across southern Australia interpreted through the study of changes in
the fossil avian assemblages largely parallel results drawn from microfloral assemblages. Both
methods indicate a lowering in effective precipitation around 30,000 years ago, which led to a
drier regime (see Fig. 6). This regime lasted between 30,000 and 10,000 yBP, after which the
effective precipitation increased to present day levels or above. Although minor changes within

822 - BAIRD

these large scale changes may not be discernable in avian assemblages, they do demonstrate that
avian assemblages can mirror gross changes in climate.

The differences of palaeoenvironmental interpretations between regions may result from
variation in the substrates upon which the vegetation grows. For example, no changes are
interpreted for the avian assemblages from Madura and Koonalda caves, even for those times
when other regions had indicated climatic amelioration. A possible reason for this is that the
vegetation of the Bunda Plateau may not reflect the changes in the climate, because there may
be some other limiting factor controlling the vegetation of that region today (i.e. substrate, as
is suggested for the volcanic soils of western Victoria (Dodson 1974a)). Another reason the
vegetation may not change is due to the high evaporation rates of that region, which may keep
the effective precipitation constant, even with a relative increase or decrease of precipitation.

Given that avian assemblages can be used as indicators of climatic change, how does this
method compare with that provided by microfloral analysis? Avian assemblages accumulated
by vertebrate predators have the benefit of being localized by the foraging range of the
accumulators, and, therefore, restricting the area for which the palaeoenvironmental
interpretation is valid. Microfloral analysis, on the other hand, has little control on the
catchment area represented by the assemblage. Unfortunately, avian assemblages are
represented by smaller numbers of individuals and smaller numbers of taxa than that represented
in pollen assemblages. Because of this, only gross changes seem to be reflected in fossil-avian
assemblages, while changes of finer resolution are available from fossil pollen assemblages
(see Baird 1989),

SINCE EUROPEAN SETTLEMENT

From the time of European settlement, range contractions have occurred both through
habitat destruction and introduction of pest species (e.g. Vulpes vulpes (Fox), Orictolagus
cuniculatus (Rabbit), Ovis (Sheep), Bos (Cow), Felis catus (Cat), etc.). Those species which
have been associated with man caused range contractions, include Amytornis textilis (Thick-
billed Grass-wren) on the Bunda Plateau and Calyptorhynchus lathami (Glossy Black-cockatoo)
in Tasmania (Baird 1986b).

Extinctions of avian species associated with the European expansion include Psephotus
pulcherrhimus (Paradise Parrot), Geopsittacus occidentalis (Night Parrot), Dromaius
baudinianus and D, ater.

There are some species which have benefited by the clearing of native habitat. This benefit
is expressed by the range expansions of Ocyphaps lophotes (Crested Pigeon) and Cacatua
roseicapilla (Galah: Forshaw 1969, Parker & Reid 1983, Frith 1982, etc.). These species have
their native habitats in open savannah woodland, Acacia scrub, etc. These habitats are
inherently more open and, therefore, the clearing of land for cultivation and grazing provided
large areas of land which were once unsuitable (e.g. woodland, forest, etc.).

Since European settlement, there have becn a number of introductions to our avian fauna.
These have been well publicized, as it was felt at the time that a little bit of Europe was being
sown every time a new species was successfully introduced. All the introduced species along
with their dates of introduction can be found in Table 1 (adapted from Dickison 1932). Many
of these species have diagnostic postcranial elements aiding faunal dating of the top end of
many deposits.

BIRDS IN QUATERNARY BIOSTRATIGRAPHY

_ To date there has not been sufficient information from avian assemblages to construct any
biostratigraphic scheme in Australia; therefore, biostratigraphy using vertebrate material,

QUATERNARY AVIFAUNA OF AUSTRALIA - 823

especially in Australia, is based largely upon mammals and fish (e.g. Woodburne et al. 1986;
Long, Chap. 12, this volume; Rich et al., Chap. 23, this volume; Turner, Chap. 13, this

volume).

Table 1: Introduced birds of Victoria and the dates of earliest introduction (adapted from Dickison 1932)).

Sturnus vulgaris
Passer domesticus
Padda oryzivora
Acridotheres tristis
Fringilla coelebs
Carduelis chloris
Carduelis carduelis
Emberiza citrinella
Carduelis spinus

Common Starling
House Sparrow

Java Sparrow
Indian Myna
Chaffinches
Greenfinches
European Goldfinch
Yellowhammer
European Sisken

1863 - present
1963 - present
1863 - 1906 (died out
1863 - present
1863 - present
1863 - present
1863 - present
1863 - 1906 (died out)
1864 - 1906 (died out)

Erithacus rubelcula European Robin 1863 - 1906 (died out)
Turdus philomelos Song Thrush 1863 - present
Turudus merula European Blackbird 1864 - present
Emberiza hortulana Hortolan Bunting 1863 - 1907 (died out)
Alauda arvensis Skylark 1863 - present
Tutur communis Turtle Dove 1872 - 1906 (died out)
Streptopelia chinensis Chinese Tutrle dove 1870 - present
Lophortyx californicus California Quail 1863 - 1906 (died out)
Anas platyrhynchus Mallard Duck 1871 - present

8 EEE eee

Local extinctions may be useful for the relative dating of sequences of known taphonomic
history. By knowing the taphonomic history it will be known what could be accumulated, so
that the absence of species from a deposit will mean more, or the presence of certain species in
a deposit will mean more (e.g. Dasyornis brachypteris of south-western Victoria). Relative
numbers of individuals will rarely mean anything about the local population due to biases of
the accumulating agent.

Local events may be useful as indicators of a particuiar time in a closed area. For example,
Hirundapus caudacutus (White-throated Needletail) occurs in the deposits of Mabel Cave. This
species is rarely recorded from land, and when it is, it is largely the result of a special event
such as a bushfire or hailstorm. The Hirundapus material from Mabel Cave is considered to
have occurred as a result of a catastrophic event, and if it occurs in deposits in any of the
surrounding caves in the area, it may be a useful chronologic marker.

A few species are uscd as indicators of the Pleistocene including Gallinula mortierit,
Genyornis, flamingoes and palaclodids. But caution should be exercised in using this method
of dating for given sufficient data then material of several of these species may be found to
extend into the Holocene (e.g. Gallinula mortierii ; Baird, submitted a).

FAMILY ACCOUNTS FOR AUSTRALIAN FOSSIL BIRDS

The familial descriptions include the occurrence of avian species in Australian Quaternary
deposits, and make reference to those deposits of particular intcrest. I will point out changes in
biogeographic ranges, make reference to publications providing characters useful in the
identification of each family, indicate whether the species within the family are useful as
palaecoenvironmental indicators and or of use in biostratigraphy, as well as any other points of
interest [dagger indicates wholly extinct families]. Figs 7 through 9 demonstrate the seven
most common postcranial elements found in deposits. Although the gross shapes for these

824 - BAIRD

elements are similar throughout all birds in detail they can vary extensively (see Baumel et al
1979 and Howard for examples).

I follow the taxonomy of Schodde (1975) and Condon (1975). ‘

Those families not represented in the Quaternary continental record of Australia are not
included here. Summary tables for non-passcrines (Table 2) and passerine (Table 3) families are
included for the major deposits. Numbers within the brackets at the end of each familial
discussion refers to the site numbers where the family is recorded (N.S.W. = New South Wales,
Qld. = Queensland, S.A. = South Australia, Tas. = Tasmania, Vic. = Victoria, and W.A. =
Western Australia).

a ‘ ae (
<= a a i

Figure 7: Postcranial elements from fossil and recent megapodes. A, complete left humerus, Megapodius
reinwardt; B, incomplete left humerus, Progura naracoortensis; C, complete left radius, P. maracoortensis; D,
complete left radius, M. reinwardt; E, distal end left ulna, P. naracoortensis; F, incomplete right ulna, P.
naracoortensis; G, complete right ulna, M. reinwardt; I and I, proximal and distal ends right ulnae, P.
gallinacea (after van Tets 1974a).

CASUARIDAE - EMUS

Emus occur in a wide variety of deposits in Australia, and they are well represented both
geographically and temporally. On continental Australia all of the material studied to date is
referrable to the extant species Dromaius novaehollandiae (Emu: Patterson & Rich 1987). This
includes the fossil species D. patricius, D. 8racilipes and Metapteryx bifrons described by de
Vis (1888, 1891a, 1905). The fossil material ranges widely in size. This variation cannot be
accounted for from samples of modern emus and may be either a result of the small sample
sizes available of the modern specimens, or that the Pleistocene forms may have exhibited a
wider range of sizes than their modern counterparts.

Emu material from the offshore islands, D. ater (King Island Emu) and D. baudinianus
(Kangaroo Island Emu) still retain specific status even after recent accounts of their taxonomy

QUATERNARY AVIFAUNA OF AUSTRALIA - 825

(Parker 1984). [N.S.W. 3, 8, 9, 10, 11, 12, 16; P.N.G, 1: Qld 4,9; S.A. 1, 2, 3, 4, 5, 7, 8,
10, 11, 15, 16, 16a, 16b, 16c, 16d, 16e, 17, 19, 20, 27, 40, 42, 45, 46, 48; Tas. 4, 5, 6, 8, 9,
10; Vic. 2h, 6d, 7; W.A. 1, 3]

+ DROMORNITHIDAE - MIHIRUNGS

There are several species of dromornithids found in Australia, all except Genyornis newtoni
are Tertiary in age. Genyornis occurs into the Quaternary with specimens associated with
radiocarbon dates within the Late Pleistocene. This material ranges from skeletal remains, as
are found at Lancefield, Victoria (Gillespie er al. 1978); eggshells (Williams 1981); footprints
(P.V. Rich, pers. comm.) or gizzard stones (Williams 1981).

Genyornis is one of the avian species which can be considered a member of the megafauna
that became extinct during the Late Pleistocene. [N.S.W. 4, 5; Qld. 2, 10; S.A. 1, 2, 5, 6, 7,
10, 12, 15, 17, 26, 27, 40, 42, 43, 48; Vic. 15; W.A. 7, 9]

PODICIPEDIDAE - GREBES

This material has not been fully studied, although preliminary identification suggest that
there are two different sized species represented (McEvey & Rich, pers. comm.). [N.S.W. 11;
S.A. 5]

SPHENISCIDAE - PENGUINS

Although penguin material is not frequently encountered in deposits on continental
Australia the small amount of material published upon does rate a mention. Material referred
to Eudyptula minor (Little Penguin) has been recorded from Amphitheatre Cave, Victoria,
although it has been suggested that it was not part of the original deposit but instead
incorporated into the collection at some later date (Baird submitted b). As with New Zealand
(Millener, Chap. 27, this volume) and the Australian coastal islands (Meredith, Chap. 28, this
volume), it is expected that this species will occur with some frequency in midden deposits
along the coast. Of greater interest is the material referred to the new genus and species,
Tasidyptes hunteri, from Hunter Island, Tasmania (van Tets & O'Conner 1983). After
reviewing the paper I do not consider either the morphological nor the mensural characters
sufficient to define either a new genus or new species. Instead I would suggest that with a
larger sample size of Eudyptes chrysochome (Rockhopper Penguin) that the fossil material will
be scen to fall within the range of variation for this species (see Meredith 1988; Chapt 28, this
volume). This conclusion is corroborated by Fordyce and Jones where they state "Van Tets &
O'Conner (1983; see Harrison 1984) proposed a new genus and species, Tasidyptes hunteri, for
debatably diagnostic material from a Recent midden at Hunter Island, Tasmania." (Fordyce &
Jones in press). [Tas 3, Vic 6a]

PROCELLARIIDAE - PETRELS AND SHEARWATERS

The procellariids seem to have been an important part of the Aboriginal economy along the
coastlines of Australia (Gaughwin 1978). The species most frequently encountered in these
middens is Puffinus tenuirostris (Short-tailed Shearwater), which is known to breed along the
coast. Puffinus tenuirostris is also abundant as fossils and subfossils on many offshore islands
where it currently breeds (see Meredith, Chap. 28, this volume). Aboriginals may have taken
advantage of this source of food, largely during the breeding season, although the possibility
exists that beach-washed individuals also may have been available for consumption.

826 - BAIRD

Hypotheses concerning material found in inland deposits (e.g. Mabel Cave, Victoria and
Beginners Luck Cave, Tasmania) ranges from stormwreck material opportunistically collected
by vertebrate accumulators (e.g. Sarcophilus (Tasmanian Devil)) to that transported by
Aboriginals to the cave from breeding grounds or shoreline (Baird 1986c). [N.S.W. 2, 6, 7;
S.A. 13f; Tas. 1, 3, 7, 11; Vic. 2c, 6a, 10]

Figure 8: Postcranial elements from fossil and recent megapodes. A, C, complete right carpometacarpus,
Megapodius reinwardt; B, D, incomplete right carpometacarpus, Progura gallinacea; E, incomplete right
coracoid, P. naracoortensis; F, complete right coracoid, M. reinwardt; G, incomplete right coracoid, P.

gallinacea; H, J, humeral end tight scapula, P. gallinacea; I, K, complete right scapula, M. reinwardt (after van
Tets 1974a).

PELECANOIDIDAE - DIVING PETRELS

Other than the numerous records of this species on offshore islands (Meredith, Chap. 28,
this volume), Pelecanoides urinatrix (Common Diving-petrel) is only known from Mabel
Cave, Victoria. As with the procellariids from this deposit, it is not known how the material
was accumulated, discussed above. [Vic. 2c]

QUATERNARY AVIFAUNA OF AUSTRALIA - 827

PELECANIDAE - PELICANS

Representatives of the Pelecanidae from Quaternary deposits are restricted to one
palaeospecies (Pelecanus cadimurka) and one neospecies (P. conspicillatus (Australian
Pelican)). According to Rich & van Tets (1981) only the tarsometatarsus can be used to
separate the two species as all other elements are non-diagnostic. Of the two species, P.
cadimurka is distinctly smaller than P. conspicillatus. [S.A. 5, 6, 7, 19]

PHALACROCORACIDAE - CORMORANTS

Australian fossil material of this family is currently being studied by G. van Tets. To date
there have been no identifications which are confident to species level. De Vis (1905) did
describe a fossil species, Phalacrocorax gregorii, based on a number of different postcranial
elements from northern South Australia. This species is in need of restudy due to the number
of misidentifications made by de Vis. [ S.A. 5, 6, 8, 17, 19, 48; W.A. 9, 10]

ANHINGIDAE - DARTERS

Miller (1966a) reviewed both the material described by de Vis (1888, 1905) and that newly
collected. He concluded that Anhinga parva, a species described as new by de Vis, was a small
cormorant (cf. Phalacrocorax melanoleucus (Little Pied Cormorant)), that A. latipes, another
species described by de Vis, was valid and that the rest of the material be referred to the extant
A. novaehollandiae (Darter). [Qld. 3; S.A. 5, 19, 28]

ARDEIDAE - HERONS

None of the fossil material referred to the Ardeidae has been identified past family level.
[N.S.W. 11; S.A. 5, 16, 28]

CICONIDAE - STORKS

The validity of Xenorhynchus asiaticus (Black-necked Stork) from Darling Downs,
Queensland is not known and should be restudied. [S.A. 19]

THRESKIORNITHIDAE - IBISES AND SPOONBILLS

Theskiornithid material is restricted to specimens from three deposits (i.e. Darling Downs,
Queensland; Weekes Cave, South Australia and Seton Rock Shelter, Kangaroo Island, South
Australia). Of the three elements referred to Platalea subtenuis de Vis from Darling Downs,
two have been identified as the rail, Gallinula mortierii (Olson 1975). The third specimen is
definitely not rallid, according to Olson (1975), but a positive identification as a threskiornithid
has not been made.

Platalea leucorrhoa (Yellow-billed Spoonbill) from Weekes Cave (van Tets 1974 b) consists
of the better part of a whole skeleton which probably entered the cave after a wet period when
water had again become scarce.

Material referred to Threskiornis mollusca (Sacred Ibis) from Seton Rock Shelter (Hope et
al. 1977) is very fragmentary and based upon preliminary re-examination should be referred to
the Anatidae. [S.A. 13f, 39; WA 7i]

828 - BAIRD

Figure 9: Postcranial elements from fossil and recent megapodes. A, incomplete right tarsometatarsus,
Progura naracoortensis; B, complete right tarsometatarsus, Megapodius reinwardt, C, proximal end left
tarsometatarsus; P. gallinacea; D, E, distal end right tarsometatarsi, P. gallinacea; F, proximal end right femur,
P. naracoortensis; G, complete right femur, M. reinwardt; H, complete right tibiotarsus, M. reinwardt; I,
incomplete right tibiotarsus, P. naracoortensis; J, anterior end synsacrum, P. naracoortensis; K, complete
synsacrum, M. reinwardt.

ANATIDAE - DUCKS, GEESE AND SWANS

To date, the fossil anatids have been largely avoided by the few avian palaeontologists
covering Australia. Many of those identified by van Tets from Seton Rock Shelter (Hope et al.
1977) are considered too incomplete for identification beyond familial level. Olson (1977)
attempted to clean up some of the erroneous identifications of anatid material by de Vis (1888,
1889, 1905), which turned out to be Anas superciliosa (Black Duck), Aythya australis
(Hardhead), Anas castanea (Chestnut Teal) and Biziura lobata (Musk Duck). Olson (1977)
indicates that the Pleistocene material of Biziura lobata may be slightly smaller than the living
form.

Of particular interest is new material of a large form of swan from Henschke's Quarry Cave,
S.A. but this material has not been studied. [N.S.W. 10, 11, 15; Qld. 5, 11; S.A. 5, 7, 8,
16f, 19, 33, 45, 48; Vic. 2c, 3, 6a; W.A. 1]

QUATERNARY AVIFAUNA OF AUSTRALIA - 829

ACCIPITRIDAE - HAWKS AND EAGLES

Many of the modern species of accipitrids occurring in southern Australia are represented in
the fossil record (i.e. Hieraaetus morphnoides (Little Eagle), Accipiter fasciatus (Brown
Goshawk), A. cirrhocephalus (Collared Sparrowhawk), Circus sp. (Harrier)). The oldest
modern species represented is Aquila audax (Wedge-tailed Eagle) from Devil's Lair, Western
Australia (> 35,000 yBP). There are two groups of associated elements which are currently
considered to belong to taxa new to science and both represent individuals larger than A. audax.
One group of elements, (i.e. sternum, distal phalanx, proximal end tibiotarsus) comes from
Mair's Cave, South Australia. Another group of associated elements come from Green
Waterhole Cave (L-81) and consists of humeri, ulnae and carpometacarpi. Both of these forms
are not directly comparable for there is no overlap in the elements represented, but both appear
to be significantly larger than Aquila audax.

The Gypacinae material reported in Rich & van Tets (1982) has yet to be substantiated and
should be regarded with scepticism (pers. obs., P. Rich, pers. comm.). {N.S.W. 1; Qld. 3, 5;
oe 5, 9, 14, 16f, 19, 22, 23, 33, 35, 38, 39, 48; Vic. 2a, 2b, 2c, 6a; W.A. Te, 7g, 7k, 71,

FALCONIDAE - FALCONS

Similar to the accipitrids, several falcon species are represented in the fossil record, these
include Falco berigora (Brown Falcon), F. cenchroides (Australian Kestrel) and F’. peregrinus
(Peregrine Falcon: Baird 1985, 1986c; Rich et al. 1982). All three of these species are known
to use caves for both roosting and nesting, and, therefore, their remains would be expected to be
incorporated into cave deposits. They could be accumulators of vertebrate material as well,
although no deposits have been attributed to these species (see Baird, Chap. 10, this volume).
[N.S.W. 10; S.A. 5, 14, 23, 33, 37, 39; Tas. 4; Vic. 2a, 2c, 6d]

MEGAPODIIDAE - MOUND-BUILDERS

The endemic group of Australian fossil megapodes were first recognised by van Tets
(1974a), who described two species from the limited fossil material then available, Progura
gallinacea and P. naracoortensis (see Pl. 2), Olson (1980, 1985) has made statements on the
possibility of two separate genera involved in this complex but failed to state specifically
which characters led him to come to these conclusions. Since that time, additional material has
suggested that the two species may in fact be one which exhibits sexual size dimorphism (van
Tets 1984). This is based both on the lack of characters which consistently separate the two
species (other than size) and the fact that both species occur in every deposit from which the
genus is known. The genus extends into the Tertiary where it is recorded from the deposits of
Chinchilla (van Tets 1974a).

The family is also represented by Leipoa (Malleefowl), which occurs in one Late
Pleistocene deposit, Victoria Fossil Cave, South Australia. [N.S.W. 10, 13, 16; Qld. 3, 7;
S.A. 44, 45, 46, 47]

PHASIANIDAE - PHEASANTS, QUAIL, ETC.

Coturnix material is commonly found in Quaternary fossil deposits and can make up a large
proportion of the total avian assemblage (e.g. Clogg's Cave, Victoria). Their susceptability to
capture probably relates to their size and habits, which include a terrestrial lifestyle and periodic

830 - BAIRD

population booms. Therefore, they are more likely to be collected by Tyto alba and be
represented in natural traps than other species (see Baird, Chap. 10, this volume).

There are three species of Coturnix which occur in southern Australia, C. pectoralis
(Stubble Quail), C. australis (Brown Quail) and C. chinensis (King Quail). Currently, only
Coturnix chinensis can be identified with certainty, for the other two species are similar both in
morphological and mensural characters. C. chinensis, on the other hand, is much smaller than
the other two species and can be distinguished on most of its postcranial elements.

All of the Quaternary material studied to date has demonstrated no statistically significant
deviation in size over modern samples. [N.S.W. 1; Qld. 1, 8, 10; S.A. 14, 16f, 33, 46; Vic.
2a, 2c, 2g, 6a, 6d; W.A. 1, 5, 7e, 10]

TURNICIDAE - BUTTON QUAIL

In southern Australia turnicid fossils are frequent in cave deposits and can make up a large
proportion of the avian assemblage. The reasons for this are the same as those of the Coturnix
spp. in this region, and relates to their natural histories (see Baird, Chap. 10, this volume).

Of the three species of Turnix which occur in southern Australia (e.g. Turnix varia (Painted
Button-quail), T. pyrrhothorax (Red-chested Button-quail) and T. velox (Little Button-quail)),
only 7. varia can be identified with certainty. Both 7. pyrrhothorax and T. velox overlap
extensively in size, and no morphological characters have been found to distinguish the two.
These species are sometimes referred to as probable identifications largely based on
biogeographic probability. 7. varia on the other hand, is much larger than either T.
pyrrhothorax or T. velox. [N.S.W. 1, 16; Qld. 6, 8, 10; S.A. 14, 16f, 27, 29, 30, 32, 33, 34,
39, 45, 46; Vic. 1, 2a, 2c, 2g, 4, 5, 6a, 6d; W.A. 7b, 7c, 7d, 7e, 7f, 71, 10]

PEDIONOMIDAE - PLAINS-WANDERERS

This monotypic family could be used quite effectively as an palaeoenvironmental indicator,
because it is currently restricted to areas of "savannah" grassland where there is a wide diversity
of grass species (Harrington et al. 1988). Its very diagnostic elements (see Olson & Steadman
1981 and Rich & McEvey 1980) have shown up in a number of deposits from Victoria
including Clogg's Cave (Victoria, Baird 1986c), Harman's Cave (Victoria: Baird unpubl. data)
and Morwell (Rich & McEvey 1980).

The material from Victoria Fossil Cave, South Australia described by van Tets & Smith
(1974) has been reviewed in Rich & McEvey (1980) and is considered to be of some other
charadriiform, not Pedionomus. [Vic. 2a, 4, 8]

RALLIDAE - RAILS

Members of this family are prevalent throughout all types of deposits in Australia. This is
related to their life histories which can include being both hydrophilic and terrestrial, and
crepuscular to nocturnal in their habits, therefore making them vulnerable to capture by
nocturnal predators (see Baird, Chap. 10, this volume). This makes rails susceptable to
becoming interred in fluvial deposits, trapped in pit falls and captured by nocturnal cave
dwelling predators, thereby covering most of the possible ways of being incorporated into the
fossil record.

Almost all of the rallid species, whose distributions encompass areas with fossil avian
assemblages, have been identified as part of these assemblages. The list of species includes
Rallus philippensis (Buff-banded Rail), Rallus pectoralis (Lewin's Rail), Porzana fluminea

QUATERNARY AVIFAUNA OF AUSTRALIA - 831

(Australian Crake), P. tabuensis (Spotless Crake), Gallinula mortierii (Tasmanian Native-hen)
and G. tenebricosa (Dusky Moorhen).

The species that has received most attention is Gallinula mortierii (now endemic to
Tasmania). Olson (1975) showed that a number of specimens from the Plio-Pleistocene
Chinchilla deposits (de Vis 1888a, 1892) were all referrable to the extant species Gallinula
mortierit, but due to the specimens’ small demeanour he suggested a separate subspecies for
them, G. m. reperta. Baird (1984), using a larger sample provided by Late Pleistocene fossils
from localities in the southeast of Australia, demonstrated the large overlap in the
measurements of G. m. reperta and G. m. mortierii and, therefore, referred the de Vis Chinchilla
material to the nominate form.

Additional material from the Darling Downs area of southeastern Queensland, dated at
between 30,000 yBP and 24,000 yBP (Gill 1978) demonstrated that the species once occurred
in the well-watered Murray-Darling catchment (Baird 1986a). The species is also known from
Wyandotte Creek, northeastern Queensland (McNamara & Baird, in press). It has been
suggested that Gallinula mortierii, like the flamingoes, was excluded from the Australian
mainland through the change in environmental regimes during the last glacial. But material
dated recently from Amphitheatre Cave, Victoria yielded a mid-Holocene date, requiring a
review of this hypothesis. The review (Baird submitted a) concluded that both the environment
and the dingo may have had a role to play in the extermination of Gallinula mortierii from the
mainland. The species is also known from cave deposits in Tasmania (van Tets 1978).

Material from many of the other extant rail species of Australian has been found in the
caves of the southeast (Baird, unpubl. data). Many of the inland lake deposits contain rail
species, but as of yet, no identifications have been given this material.

The large rail identified in Fox Cave, South Australia (U-22) by G. F. van Tets and
mentioned in Rich & van Tets (1982) was misidentified to order; see Alcedinidae. [N.S.W. 10;
Vic. 2a, 2c, 3, 4, 6a, 6b, 6c, 6d, 7, 11; Tas. 4, 9; Qld. 5, 11; S.A. 9, 10, 13, 14, 16f, 19, 23,
27, 44, 46]

OTIDIDAE - BUSTARDS

The specimen referred to this family is comparable with the modem species, Ardeotis
australis (Australian Bustard). [S.A. 6]

CHIONIDAE - SHEATHBILLS

The only fossil record of sheathbills in the world came from Victoria Fossil Cave (South
Australia: van Tets & Smith 1974). Unfortunately, the record is incorrect and "appears to have
decided similarities to the Passeriformes" (Olson 1976).

BURHINIDAE - THICKNEES

Fragmentary material referred to the extant Burhinus magnirostris (Bush Thick-knee) has
come from Seton Rock Shelter (Kangaroo Island, South Australia: Hope ef al. 1977) as well as
some undescribed material from Green Waterhole (South Australia: Baird, unpubl. data). [SA
13f, 20]
LARIDAE - GULLS AND TERNS

There are a few continental records of this family from coastal or estuarine deposits.
Amphitheatre Cave (Victoria) and Seton Rock Shelter, for example, both contain Larus

832 - BAIRD

novaehollandiae (Silver Gull). Although this species is not restricted to the coast, it is most
abundant there. I would expect that more material of this family would be discovered in the
deposits of Cooper Creek and some of the inland lake systems. [S.A. 13f; Vic 6a]

Sterna sp. cf. S. nereis (Fairy Tern) is also found in Seton Rock Shelter, Kangaroo Island,
South Australia. [S.A. 13f]

CHARADRIIDAE - PLOVERS

Although not particularly abundant, species in this family do occur with some regularity in
cave deposits right across Australia (i.e. Erythrogonys cinctus (Red-kneed Dotterel: Cloggs
Cave, eastern Victoria), Peltohyas australis (Inland Dotterel: Victoria Fossil Cave, southeastern
South Australia: van Tets & Smith 1974) and Calidris sp. (Madura Cave, southeastern Western
Australia)). [SA 27; Vic 2a; WA 7f]

SCOLOPACIDAE - SANDPIPERS

A number of scolopacids occur in Quaternary deposits including Gallinago hardwickii
(Latham's Snipe) from the deposits at both Seton Rock Shelter, Kangaroo Island, South
Australia (Hope et al. 1977) and Victoria Fossil Cave, South Australia (van Tets & Smith
1974). Tringa glareola (Wood Sandpiper) and Calidris ruficollis (Red-necked Stint) also have
been reported from Victoria Fossil Cave. [S.A. 13f, 27]

PHOENICOPTERIDAE - FLAMINGOES

Flamingoes had a long history on the Australian continent dating back to the Miocene
(Rich & Baird 1986, Rich, Chap. 20, this volume). Their remains are present in many of the
Pleistocene lake deposits of South Australia and western New South Wales. There are
currently recognized four (there are more in Tertiary deposits) species in Australia (i.e.
Xenorhynchus tibialis, X. minor, Phoenicopterus ruber and Ocyplanus proeses, Rich et al.
1987). Only the two Xenorhynchopsis species are found in localities of Quaternary age,
although both Phoenicopterus and Ocyplanus are kncwn from several localities of unknown
age.

If, as Rich et al. (1987) suggest, the unknown localities prove to be Quaternary, then it was
probable that at some point in time there were up to four species of flamingoes living
sympatrically.

The hypothesis for their disappearance from the Australian continent as caused by
"...increased aridity accompanied by the disappearance of reasonably permanent shallow lakes,
and of feeding and breeding grounds, resulted during the Quaternary in major extinctions" (Rich
et al. 1987) may be plausible in light of other palacoenvironmental information. [N.S.W. 10;
S.A. 5, 18, 19, 25]

seins Distribution of families of nonpasserine birds across the major Quatemary-aged localities in
ustralia.

QUATERNARY AVIFAUNA OF AUSTRALIA - 833

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834 - BAIRD
+ PALAELODIDAE

The only Quaternary material referrable to the Paleolodidae in the world comes from
Kudnampirra (Locality 4) Cooper Creek Lower, South Australia (Baird & Rich, in press). The
single specimen is referred to a new species of Palaelodus, which is restricted to the Australian
continent, and is the youngest record for the family in the world. [SA 5]

TYTONIDAE - MASKED OWLS

This is one of the only groups of predatory birds to use caves with any frequency in
Australia. Because of this fact and that owls of the genus Tyo have been demonstrated to have
low pH stomach acid and, therefore, produce bones in their ejectamenta of remarkable
completeness, they are probably the most important accumulators of small mammals and birds
in Australia (see Baird, Chap. 10, this volume). This dubious distinction has done little to
enliven interest in the natural histories of the four species in the genus nor has it produced any
work on the differences in the taphonomy between the two most important species, Tyto alba
(Barn Owl) and T. novaehollandiae (Masked Owl). The only references to these animals are
restricted to anecdotal accounts of the probable size of prey species for each. Because of this
most of the deposits are attributed to T. novaehollandiae with little or no corroborating
evidence (see Baird, Chap. 10, this volume, for more information).

The only two species recorded in the fossil record are T. alba and T. novaehollandiae. They
occur throughout the deposits of southern Australia. Most importantly, both species have been
recorded from the cave deposits of the Nullarbor Plain, where in historical times only T.
novaehollandiae had been recorded with any confidence (Parker 1977). [S.A. 5, 9, 14, 33, 37,
39, 46; Vic. 2a, 2g; W.A. 4, 7e, 7i, 71]

STRIGIDAE - HAWK-OWLS

Members of this family are rarely recorded as fossils. Only a single deposit, McEachern's
roan Victoria has yielded specimens referrable to Ninox connivens (Barking Owl: McNamara
1981). [Vic. 6]

PODARGIDAE - FROGMOUTHS

The Podargidae are only known from a few fossil assemblages (Scotts River, Western
Australia; Clogg's Cave, Victoria; and Aiyenu Cave, Western Australia). All specimens have
been referred to Podargus strigoides (Tawny Frogmouth) but are largely represented by
fragmentary material, except in McEacherns Cave, where a number of associated elements are
found. [Vic. 6; W.A. 4, 8]

AEGOTHELIDAE - OWLET-NIGHTJARS

Aegothelids occur regularly in cave deposits and are considered autochthonous speleophyles
(regular cave dwellers); therefore, the inclusion of their elements in any cave assemblage should
be considered to be more a result of periodic deaths of aged individuals rather than the result of
any other accumulating agent (unless evidence can prove otherwise).

All of the specimens recorded to date are referrable to the extant species Aegotheles cristatus
(Australian Owlet-nightjar). The Australian specimens have been compared with modern
species and show none of the gigantism seen in the New Zealand genus, Megaegotheles (Rich

QUATERNARY AVIFAUNA OF AUSTRALIA - 835

& Scarlett 1977), now placed in Aegotheles by Olson et al. (1987 Id. 10; S.A. 9; Vic. 2
2c, 2g, 6b; WA 1, 7f, 7i, 10] Eg ert

APODIDAE - SWIFTS

Swifts have been recorded from one Quaternary deposit in Australia, Mabel Cave, Victoria.
A number of specimens of Hirundapus caudacutus (White-throated Needletail) were found in the
cave. This was considered unusual because these Swifts rarely alight on land; therefore, the
cause of the concentration was hypothesised to have been either a bushfire or thunderstorm
forcing them to earth where the accumulator of the deposit could collect them.

It is expected that Collocalia spodiopygia (White-rumped Swiftlet) will occur in the cave
deposits in northern Australia as it is currently resident there and is recorded from caves in the
eee = (New Ireland (Baird, unpubl. data) and New Caledonia (Balouet & Olson 1989)).

ic.

COLUMBIDAE - PIGEONS

There have been a wide variety of pigeons recorded as fossils in Australia including
Leucosarcia melanoleuca (Wonga Pigeon), Phaps chalcoptera (Brush Bronzewing) and P.
elegans (Common Bronzewing). Pigeons are recorded from both fluviatile (e.g. Darling
Downs, Queensland) and cave deposits (e.g. Mabel Cave, Victoria).

The smaller species (e.g. Ptilinopus spp.) are expected from Tyto-accumulated cave
assemblages in northern Australia as they are excavated because species of this size have been
recorded from islands in the Pacific (e.g. New Ireland, Baird, unpubl. data).

After reviewing the types of Nyroca effodiata de Vis 1905, Leucosarcia proevisa de Vis
1905 and Lithophaps ulnaris de Vis 1891, van Tets & Rich (1980) concluded that all three were
referrable to the extant genus Phaps. The large size of the fossil material suggested an
association with either P. chalcoptera or P. histrionica (Flock Bronzewing), but it was too
large to be P. elegans. [N.S.W. 11; Qld. 3, 10; S.A. 5, 12f, 19; Tas. 2; Vic. 2a, 2c, 6a, 6b;
WA 1, 7d, 7e, 7f, 10]

CACATUIDAE - COCKATOOS

Until the material from Green Waterhole was reported by Baird (1985), there had been no
cockatoos recorded as fossils in Australia. Since then, there have been reports of fossil
cockatoo material from several deposits, but none are as rich in number of species and
individuals as Green Waterhole (see Baird, in press b).

The extant species recorded in the fossil record include Calyptorhynchus lathami (Glossy
Black-cockatoo), C. funereus (Yellow-tailed Black-cockatoo: Mabel Cave, Victoria), C.
magnificus (Red-tailed Black-cockatoo), Callocephalon fimbriatum (Gang Gang), Cacatua
tenuirostris (Long-billed Corella), Cacatua roseicapilla (Galah)(Baird 1985, 1986b). Of these
C. lathami from Green Waterhole represents a range extention (see section on biogeography,
this chapter) and the Cacatua tenuirostris from Green Waterhole represents an example of
Pleistocene gigantism where the fossil material is on average 4% larger than the extant C.
tenuirostris. [Vic. 2c, 6a; S.A. 19]

LORIIDAE - LORIKEETS

Lorikeets can be the most abundant and conspicuous birds when eucalypt species are
flowering abundantly. Their relative abundance can also be quite high in some avian

836 - BAIRD

assemblages. For example, in Devil's Lair Glossopsitta porphyrocephala remains make up
42% of the total avian assemblage (based on minimum numbers of individuals: Rich & Baird
1986). All of the southern species are represented in avian assemblages, including Glossopsitta
concinna (Musk Lorikeet), G. porphyrocephala (Purple-crowned Lorikeet), G. pusilla (Little
Lorikeet) and Trichoglossus haematonotus (Rainbow Lorikeet).

The three species of Glossopsitta can be separated by size, with G. concinna the largest, G.
porphyrocephala of middle size and G. pusilla the smallest. Trichoglossus is larger still than
G. concinna. [Vic. 2a, 2c, 2g, 6b; W.A. 1, 10]

PLATYCERCIDAE - BROAD-TAILED PARROTS

Species of the broad-tailed parrots are some of the best indicators of changes in the
environment of Australia in the past 30,000 years. For example, the range extensions of
Psephotus varius (Mulga Parrot: Devil's Lair, Western Australia) and Melopsittacus undulatus
(Budgerigar: Clogg's Cave, Victoria) along with other data are indicative of drier climates
during the height of the last glacial period. Species in this family may also be good indicators
of disturbance of habitats since European settlement, as in the case of Geopsittacus occidentalis
(Night Parrot: Madura Cave, W.A.) on the Roe Plain and Pezoporus wallicus (Ground Parrot:
Seton Rock Shelter, Kangaroo Island; Victoria Fossil Cave, South Australia and Clogg's Cave,
Victoria) in southeastern Australia (Baird in press b).

There have been a wide range of species in this family recorded from late Quaternary
assemblages in Australia. These include Alisterus scapularis (King Parrot), Polytelis
anthopeplus (Regent Parrot), Pezoporus wallicus, Geopsittacus occidentalis, Melopsittacus
undulatus, Lathamus discolor (Swift Parrot), Purpureicephalus spurius (Red-capped Parrot),
Platycercus elegans (Crimson Rosella), P. icterotus (Western Rosella), P. eximius (Eastern
Rosella), Barnardius zonarius (Port Lincoln Parrot), Psephotus varius, Northiella haematogaster
(Bluebonnet), Neophema chrysostoma (Blue-winged Parrot), N. elegans (Elegant Parrot) and N.
slendida (Scarlet-chested Parrot)

Characters for the identification of these genera and species are largely unpublished, but
many can be accessed in Baird (1986c). [NSW 16; S.A. 12f, 19, 24; Vic. 2a, 2c, 2e, 2g, 4, 6a,
6b; W.A. 1, 7e, 7f, 10]

CUCULIDAE - CUCKOOS

There are both extinct and extant species of cuckoos recorded in Australia. The extinct
species was described as Centropus colossus (Baird 1985) and provides both a range extension
and the largest example for the genus. The extant species include most of those species within
the prey size range for Tyto alba (i.e. Cuculus pallidus (Pallid Cuckoo), C. pyrrhophanus (Fan-
tailed Cuckoo), C. variolosus (Brush Cuckoo), Chrysococcyx lucidus (Shining Bronze-cuckoo),
etc.) All of these species can be identified by size of the postcranial elements except material
falling in the area of mensural overlap between Chrysococcyx basalis (Horsfield's Bronze-
cuckoo) and C. lucidus. [S.A. 19; Vic. 2a, 2c; WA 7d, 7e, 71, 10]

ALCEDINIDAE - KINGFISHERS

_ All of the southern species of alcedinids have been found in the fossil record. The list
includes: _Ceyx azureus (Azure Kingfisher), Halcyon sancta (Sacred Kingfisher), H.
pyrrhopygia (Red-backed Kingfisher) and Dacelo novaeguineae (Laughing Kookaburra). These
species are easily identified by their size (Baird 1985, 1986c).

QUATERNARY AVIFAUNA QF AUSTRALIA - 837

Halcyon Pyrrhopygia is recorded from Clogg's Cave, Victoria at the height of the last
sees and is considered part of the suite of species indicative that the region was drier than

y.

The record of Dacelo novaeguineae from Fox Cave (SAM P.19041) was originally

incorrectly reported as a large rail (Rich & van Tets 1982). [N.S.W. 10: Qld. 10: S.A. 19:
Vic. 2a, 2c, 6b, 6d; W.A. 8] ). [N ; Qld. 10; S.A. 19;

a
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PASSERINES = 3 Sas lsdie/S8/815 o/ s/o] 3/2] 2/8} e}mrfoelse!s 5
= 2 =3 S;EleElS Sislslal] sis) s/s] Fis/slsl/szl ez] sisz
es SB (BELO ISELS/SlS FLEE} LIS! slo; S| 2/SlFB/2/ 28] s18 se
=o = of aa gsicelas o/s S|/o}/e2} 2/0} 2}]a]/o}/E/€E 3 |e] s (8s
o> = aa} ®/ES| 2 3 = £/2 2 Qo rs ale a = Q P= oS b= 2 2/02
ae | £ OL XP ISZ,SE/ ES EIS} se} a] uj} ele} o;/ao;/2/8 l/c] & lf] olae
<6 = aos}/m)jz22, a /rj/Szlojn} Si Slalasyol ni slasyo}la«el}|o}alo fas
= 1 Le || { Sj" jrjpsihi’ H{ojfN {=f |
Mabel Cave, Vict <10 MF |MC| © | 15 144 | ria} a)2 114 | 1|2 a} 4a]4
—— a ion 5 A ~ _ +
Skull Cave, WA 2.92.08| 3, chlo} +) 5) 4] 4 nok Ta
a Nt = | es ee } ae 1 SS Se eel
Madura Cave, WA (Upper level) 5+ 12 yey 8 . 2 1 1 |
a —— =— + os t—j—} + 4} ff ff
Amphitheatre Cave.Vict 46 |yey -B/ P| 1 6 | 1 2 rfa fa
Seton Rock Shel SA ose rely tle bd a Tal | V4 il | | i} | f 2 | 1
ton ock elter e, © |
Keck lag ab j16.12.1 |0F Md) at EY tt | | Ls me +] 4
we | | | ‘ |
Clogg's Cave, Vict Eaters weg chl | + | 16 r/1}/2]a]2}e2]4 2/4 1 1 1 |
—. - —_—. ~~ + + + i oD + _+—_} _}_}_ ff __4 __}_
P 15.62.25) 7 | ] T T i
Madura Cave, WA, (Middle level) a2atsaleee 2 | O | * | 4 1 1 1 1
esd eet |} +} a I j 4 os Cs es = -
.
Koonalda Cave. SA Be lete ch} oO | 5 | 10] ol aol (Ae 2 1 | 1 [
A =o Trerz2s|@ h/iow) eT) ac a | =F 7 ro)
Victoria Fossil Gave, S.A 19:24:26) 35 ChiO?! 2 1 1
Mictoria Fossil Cave > 25 9°" 8B MC2 @ | | | iz eI h ye
McEachern’s Cave, Vict. | 2.08 1815. -chipiel "| 4 TT | 1 ] 1 2
ce ave. Vic |
— —_|zereea5/9°F | et ets pooh Stee | ||
. |
Devil's Lair, WA aoe | peg CHIME! | 7\4 i 1 | 1|4 1 1
- eae Freee TETE i A a = 4 + + + +
a5.q224) 7 _. 5 | 1
Henschke's Bone Dig, SA 1.8lge,-Chl P 1 | |
= intial oe ldo _| >ss'"|wie @{}_j | 1 | | | a Abe bere Ded
Madura Cave, W.A. (Lower level) [37 9:35\y%y 8 OO} * | 1 | 1 I
Cites ss kame ope ient ARE nhs 4 AE ee Se eb T { oj} + + +
. rT] ]
Green Waterhole, SA > 40 AF b* | 5 1 1 1 } 1 1
alee 2 | 4 a Poe OS Set a el + +} + —
Pyramids Cave, Vict(Lower level)| > 40 AF O}e i t]4 3 | | 1 1 I 1 |
= 6 | ak Es Bees {oo jt _ =e

Table 3: Distribution of families of passerine birds across the major Quaternary-aged localities in Australia.

MENURIDAE - LYREBIRDS

It is curious that Menura is absent from the fossil record, especially as several other
terrestrial birds have been recorded from cave deposits. The factors considered most important
for the apparent lack of this taxon from cave deposits is that there are few pitfall traps within
the range of either species (i.e. Menura novaehollandiae (Superb Lyrebird) and M. alberti
(Albert's Lyrebird)), and that Menura is capable of flying vertically. The latter would enable
individuals to free themselves from pitfall caves where many other terrestrial vertebrates would

be trapped.

ATRICHORNITHIDAE - SCRUB-BIRDS

Both species of Atrichornis, A. clamosus (Noisy Scrub-bird) and A. rufescens (Rufous
Scrub-bird), have been recorded in the fossil record. Atrichornis clamosus material is restricted
to two caves in the southwest of Western Australia. Although the caves are not within the
species’ current distribution, the historical distribution of the species probably included the
whole southwestern corner up as far north as Perth. The A. rufescens fossil material from
Pyramids Cave, Victoria is smaller than the northern representatives of the species, although it
is uncertain whether a species cline or palaeospecies occurred. The latter is believed most

838 - BAIRD

likely as the species demonstrates a north-south cline today (Ferrier 1984, Baird, in prep. b).
[Vic. 22; W.A. 1, 10]

HIRUNDINIDAE - SWALLOWS

Baird (1985) considered all the taxa within the family Hirundinidae to be inseparable due to
the large amount of intraspecific variation in their postcranial osteological characters. The
family can be easily identified, but determination, at the moment, should be left at gen. et sp.
indet. (see Baird, submitted b).

The family has several species which frequently nest in caves (e.g. Hirundo neoxena
(Welcome Swallow), H. rustica (Barn Swallow), Cecropis ariel (Fairy Martin)), and, therefore,
considered to be speleophiles. Because of this their material should be removed from the
discussion of the taphonomy of otherwise allochthonous deposits (see Baird, Chap. 10, this
volume). [S.A. 12f, 19; Vic. 2a, 2c, 2f, 6a, 6b; W.A. 1, 7e, 10]

ORTHONYCHIDAE - LOGRUNNERS, QUAIL-THRUSHES, WHIPBIRDS,
ETC.

Osteologically the members of this family are very diverse. A far more satisfactory
taxonomic arrangement is that proposed by Sibley & Ahlquist (1985) where all members,
except Orthonyx are removed from the family and distributed elsewhere. For the time being I
will follow tradition but cover the component parts individually.

Orthonyx - Logrunners

This interesting group of birds has not been let down by its fossil record. There are several
occurrences of the genus or similar genera in southeastern Australia. The first to be described
was Orthonyx hypsilophus (Baird 1985) from an incomplete pelvis in Green Waterhole,
southeastern South Australia. The individual described was larger than the largest extant
species O. spauldingi (Chowchilla) and believed to form a grade in the developement of using
the hindlimb for foraging. Additional material has come to light which has altered this view,
and the new species is currently believed to belong to a separate genus. A second new species
is recorded from Pyramids Cave, eastern Victoria (Baird 1986c) and is believed io be similar to,
but smaller than, O. temminckii (Logrunner). As there is no obvious cline in the extant
species (as is found with A¢trichornis rufescens), the material will be described as a new species.
[S.A. 19, Vic. 22]

Cinclosoma - Quail Thrushes

Several species of quail-thrush have been found in cave deposits including Cinclosoma
punctatum (Spotted Quail-thrush), C. alisteri (Nullarbor Quail-thrush) and C. cinnamomeum
(Cinnamon Quail-thrush). Although the amount of comparative material is not extensive, the
basic species groups can be delineated via mensural characters.

C. alisteri can be useful in determing palacoenvironment, as the species is currently
restricted to the treeless plain of the the Bunda Plateau,

It is interesting to note that the osteological features which I have used to define

Cinclosoma also occur in Ptillorhoa (Baird, pers. obs.). [S.A. 9; Vic 2a, 2c, 2 '
7b, 74, 7e, 70) E rel ; Vic 2a, 2c, 2g, 6a, 6b; W.A.

QUATERNARY AVIFAUNA OF AUSTRALIA - 839
Psophodes/Sphenostoma - Whipbird/Wedgebill

Psophodes olivaceus (Eastern Whipbird) occurs in several deposits in eastern Victoria.
Although cave deposits occur in the appropriate areas, P. nigrogularis (Western Whipbird) has
not yet been recorded. [Vic 2a, 2g]

Sphenostoma has only been found from cave deposits on the Bunda Plateau. They have
been referred to Sphenostoma sp. and are most likely Sphenostoma occidentalis (Chiming
Wedgebill) based upon biogeographic probability. [S.A. 7e]

SYLVIIDAE - OLD WORLD WARBLERS

Cincloramphus is readily identified on two of its postcranial elements, the humerus and
tarsometatarsus. Both species have been recorded in the fossil record and are separable on
mensural characters, with C. cruralis (Brown Songlark) larger than C. mathewsi (Rufous
Songlark). Also both species exhibit sexual size dimorphism, with the males significantly
larger than the females. There is an area of mensural overlap where female C. cruralis cannot
be separated from male C. mathewsi, and, therefore, only the male C. cruralis and female C.
mathewsi can be identified with certainty.

Cinclorhamphus material in both Weekes Cave (van Tets 1974b) and Seton Rock Shelter
(Hope ef al. 1977) has been restudied and referred to cf. Lichenostomus virescens (Singing
Honeyeater) and indeterminate Passeriformes, respectively.

Megalurus timoriensis has been recorded from Mabel Cave, eastern Victoria, which is an
extension of its range to the south. This deposit is considered to be Holocene, and unless it
represents the early Holocene then there is no clear correlation between a time of climatic
amelioration and the presence of this species. [SA 12f; Vic 2a, 2c, 4; WA Te]

MALURIDAE - FAIRY WRENS, GRASS WRENS

Both the humerus and femur are easily identifiable to this family and form the basis for
most of the records discussed below.

Malurus - Fairy-wrens

Material referred to this genus has been found in a number of avian assemblages from caves.
None of the fossil material has been identified to species level due to the lack of morphological
characters and the extensive overlap in mensural characters among the extant species. [Vic. 2a,
2c; W.A. 1, 10]

Amytornis - Grass-wrens

Amytornis material has been found on the Bunda Plateau (Baird 1986c), Flinders Ranges
(e.g. Mair's Cave, South Australia) and Kangaroo Island, South Australia (Baird, unpubl. data).

That from the Bunda Plateau is identified as A. textilis (Thick-billed Grass-wren) based
upon both morphological and mensural characters and believed to have formed the bridge
between eastern and western populations of that species (Baird 1986c). The other material has
not been fully studied.

Amytornis species, because of their restricted range of habitats, are considered useful
indicators of palaeoenvironments. [S.A. 9, 13f; W.A. 7b, 7d, 7e, 7f]

840 - BAIRD

Stipiturus - Emu-wrens

The smallest of the malurids, the genus Stipiturus and its species are easily identified by
size. Only S. malachurus (Southern Emu-wren) has been recorded, but these specimens come
from a wide range of localities, including Kangaroo Island (i.e. Seton Rock Shelter, Baird,
unpubl. data) and Clogg's Cave, Victoria (Baird 1986c). [S.A. 13; Vic 2a]

ACANTHIZIDAE - BRISTLE BIRDS, AUSTRALIAN WARBLERS, TITS

Acanthiza - Thornbills

Acanthiza has been recorded as fossil in cave deposits (¢.g. A. chrysorrhoa (Yellow-rumped
Thornbill)). Only one specimen has been identified to species level, because the bulk of the
species overlap so extensively in mensural characters and identification to species level is not
considered possible. [Vic 2c; WA 7f]

Pycnoptilus - Pilotbirds

Pycnoptilus has always been considered monotypic based on the extant representative.
Interestingly, there has now been found material referrable to this genus which is significantly
smaller than the modern species P. floccosus (Pilotbird) and occurs contemporaneously in an
avian assemblage with the modern species (i.e. Pyramids Cave, eastern Victoria). [Vic 2a, 2c,
2g]

Dasyornis - Bristlebirds

All three modern species of bristlebirds have been recorded in the fossil record and one is
represented by a number of postcranial elements. The species are distinct from one another in
size with D, broadbenti (Rufous Bristlebird) the largest and D. longirostris (Western Bristlebird)
the smallest (see Baird 1985, 1986c). D. longirostris, currently Testricted to the extreme
southwest of Western Australia, is only found in the cave deposits of that area (e.g. Devil's
Lair and Skull Cave). In the southeast of Australia there are currently two parapatric species
(D. brachypterus (Eastern Bristlebird) and D. broadbenti ), with D. broadbenti restricted to
western Victoria and D. brachypterus to the east coast from eastern Victoria to southern
Queensland. Parapatry in these two species seems to be a relatively recent phenomenon, for D.
brachypterus is found in cave deposits in western Victoria (Amphitheatre Cave and Curran's
Creek Cave) and D. broadbenti is found in a cave in eastern Victoria (Clogg's Cave) (Baird,
submitted b; Baird 1986c). [Vic. 2a, 2c, 6, 7,9; S.A. 22, 23; W.A. 1, 10]

NEOSITTIDAE - SITTELAS

Neositta chrysoptera (Varied Sittella) is identifiable by its humerus and recorded from Mabel
Cave. [Vic 2c]

CLIMACTERIDAE - TREE-CREEPERS

The Climacteridae is one of the more unusual and, therefore, easily identified families in
Australia. The species in this family share the unique character of the tarsus of having two
very deeply excavated sulci on the trochleae for digits III and IV (Baird, in press c). Along with
numerous other characters, the family can be identified based on at least three of its seven long

QUATERNARY AVIFAUNA OF AUSTRALIA - 841

bones. Unfortunately, species level determination is not considered possible at this time due to
extensive size overlap of their postcranial elements. [Vic. 2a, 2c, 2g; WA 10]

MELIPHAGIDAE - HONEYEATERS

The family is known in almost every Quaternary cave deposit, but due to the morphological
uniformity within the group, generic separation is not currently considered possible except
where unique characters provide additional information as with the mandible of Melithreptus
(Morioka & Bock 1971). [S.A. 12f, 19; Vic 2c; WA 4, 7e, 7f, 71]

PARDALOTIDAE - PARDALOTES

Material referred to the Pardalotidae is only identified to genus and is restricted to Cloggs
Cave [Vic. 2a]

PLOCEIDAE - FINCHES

Postcranial elements of finches are found in a number of deposits but for the most part
cannot be separated to generic or specific level. The cranial material from Weekes Cave, South
pale has been referred to Poephila guttata (Zebra Finch: van Tets 1974b). [Vic 2a, 6b;

A 71, 10]

PTILONORHYNCHIDAE - BOWERBIRDS

The bowerbirds are currently restricted to eastern Australia but occur across a variety of
habitats. The only species reported in fossil assemblages is Ptilonorhynchus violaceus (Satin
Bowerbird). Historically, the species has only been recorded as far to the west as the Otway
Ranges, Victoria. Baird (submitted b) has noted the species as fossil as far as the Glenelg
River, on the South Australian/Victorian border. The cave deposit is mid-Holocene, but it has
been suggested that the species has since left the region because the climate has since changed
making the habitat unsuitable to the species which prefers a range of habitats between wet
sclerophyll and rainforest. [Vic 2c, 2g, 6a]

ARTAMIDAE - WOOD SWALLOWS

There are three species of wood-swallows which are restricted to the drier parts of inland
Australia (Artamus cinereus (Black-faced Woodswallow), A. leucorhynchus (White-breasted
Woodswallow) and A. superciliosus (White-browed Woodswallow)). They can be used in
faunal analysis as indicators of this type of environmental regime. Artamus personatus
(Masked Woodswallow) occurs both in the wetter sections of the continent and the drier
sections. A. minor (Littke Woodswallow) is restricted to northern Australia, and A.
cyanopterus (Dusky Woodswallow) occurs in only the wetter portions of the continent.

The family can be identified by both cranial and postcranial characters. Interspecific
identification relies on mensural characters, but only given a combination of elements where
the variation in proportions of postcranial elements is sufficient to offset the overlap of ranges
of individual elements (Baird 1986c).

The species thus far identified from fossil avian assemblages include A. cyanopterus in both
southwestern Western Australia (e.g. Skull Cave) and eastern Victoria (e.g. Clogg's Cave and
Mabel Cave), and A. cinereus on the Bunda Plateau (e.g. Koonalda Cave).

842 - BAIRD

The three skulls identified as A. leucorhynchus from Weekes Cave (van Tets 1974b) have
been restudied and appear to belong to both Cinclosoma alisteri (two skulls) and Artamus sp.
(one skull). [Vic. 2a, 2c; W.A. 1, 7d, 7e, 7£, 71)

GRALLINIDAE - MUDLARKS

Although there are two published occurrences of Grallina cyanoleuca (Australian Magpie-
lark) as fossil (i.e. Seton Rock Shelter (Hope et al. 1977) and Victoria Fossil Cave (van Tets
& Smith 1974)), one is considered a misidentification (Baird, unpubl. data), and the other lacks
diagnostic characters for identification and needs to be restudied. [SA 12f, 26]

CRACTICIDAE - BUTCHERBIRDS, MAGPIES, CURRAWONGS

The cracticids are a group peculiar to Australasia but have some links with the corvids
(Sibley & Ahlquist 1985). Today the family has representatives in every terrestrial habitat,
mainly serving as scavengers or omnivores, although Sibley & Ahlquist (1985) have included
the aerial insectivorous genera Artamus and Peltops in their tribe Cracticini. The family can be
distinguished from the Australian corvids by many characters, some of which have been pointed
out in Baird (submitted b).

There are several fossil deposits with members of this family, including Seton Rock Shelter
(Hope et al. 1977), Devils Lair (Baird 1986). Larger cracticid material, which includes Strepera
versicolor (Grey Currawong), S. graculina (Pied Currawong) and Gymnorhina tibicen
(Australian Magpie), overlap in size where versicolor and graculina covers G. tibicen and
therefore identification is only confident outside of the area of overlap.

Identification to species level is based upon measurements of total lengths of elements.
Distal and proximal ends have been found to overlap too extensively in their size ranges
between species to be reliable. Because of this, the material from Seton Rock Shelter has been
relegated back to the Cracticidae due to its fragmental nature, until some other species specific
characters can be demonstrated. [S.A. 12f, 26; Vic. 4, 6a, 6d; W.A. 2, 10]

CORVIDAE - CROWS

This family ranges across Australia, with the crows occurring in the north (Corvus orru
(Torresian Crow), C. bennetii (Little Crow)) and the ravens (C. tasmanicus (Forest Raven), C.
coronoides (Australian Raven), C. mellori (Little Raven)) in the south. To date, there have
been no fossil species of crow identified with certainty. The problem with species
identification is that pre-1970 the exact number and identification of the Australian species had
not been studied in detail. With Rowley's paper on the genus (Rowley 1970), the classification
has settled, but because of the unsurity of identification of material collected pre-1970 these
specimens can only be used as guides. Secure determinations will require new series of each
species to be collected.

The genus has been recorded in a number of fossil localities, mainly in the southeast, but
this is probably biased by the number of deposits studied there. This group would benefit from
a concerted osteological analysis.

Specimens from Victoria Cave (van Tets & Smith, 1974) of Corvus tasmanicus have been
reassigned to Corvus sp. until sufficient amounts of comparative material of all the species
occurring in the Naracoorte area are available. [S.A. 12f, 19, 26; Vic 2a, 6a]

QUATERNARY AVIFAUNA OF AUSTRALIA - 843
ACKNOWLEDGEMENTS

Thanks are extended to the following people and their respective institutions who kindly
loaned much of the fossil material included in this chapter (in alphabetical order): Dr Ken
MacNamara, curator of Palaeontology at the Western Australian Museum; Mr Neville Pledge,
curator of Palaeontology at the South Australian Museum; Dr Tom Rich, curator of Fossil
Vertebrates at the Museum of Victoria.

I would also like to express my appreciation for the numerous loans of modern comparative
material made possible through the help of the following people and their respective
institutions Gin alphabetical order): Mr W. Boles, assistant curator of Omithology at the
Australian Museum; Ms Belinda Gillies, assistant curator of Ornithology at the Museum of
Victoria; Dr G. Ingram, curator of Ornithology, and Mr W. Longmore, assistant curator of
Ornithology at the Queensland Museum; Mr S. Parker, curator of Ornithology at the South
Australian Museum; Dr G. Storr, curator of Ornithology, and Mr R. Johnstone, assistant
curator of Ornithology at the Western Australian Museum; Dr G. van Tets, senior research
scientist at the C.S.I.R.O.; Dr F. Vuillemiur, head of Ornithology at the American Museum of
Natural History.

Special thanks are extended to Dr Charles Meredith for reading various drafts of this
manuscript.

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Thompson, eds., Monash University Offset Print. Unit, Clayton: 526-572.

ROGNON, P. & WILLIAMS, M.A.J., 1977. Late Quaternary climatic changes in Australia and North Africa: a
preliminary interpretation. Palaeogeog. Palaeoclim. Palaeoecol. 21: 285-327.

ROWLEY, I., 1970. The genus Corvus (Aves: Corvidae) in Australia. CS/RO Wildl. Res. 15: 27-71.

SAVAGE, D.E. & RUSSELL, D.E., 1983. Mammalian Paleofaunas of the World. Addison-Wesley Publishing
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SCHODDE, R., 1975. Interim List of Australian Songbirds. Passerines. Aust. R. Om. Union, Melbourne.

SCOTT, H.H., 1932. The extinct Tasmanian Emu. Paps. R. Soc. Tas. 1931: 108-110.

848 - BAIRD

SHACKLETON, N.J. & OPDYKE, N.D., 1973. Oxygen isotope and palaeomagnetic stratigraphy of equatorial
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SIBLEY, C.G. & AHLQUiST, J.E., 1985. The phylogeny and classification of the Australo-Papuan
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SINGH, G., 1983. Late Quaternary vegetation and lake level record from Lake George, New South Wales: 18
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SINGH, G. & GEISSLER, E.A.,1985. Late Cainozoic History of vegetation, fire, lake levels and climate at
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SPECHT, R.L., 1981. Major vegetation formations in Australia. In Ecological Biogeography of Australia,
A. Keast, ed., Dr W. Junk. The Hague, 165-297.

STIRLING, E.C. & ZIETZ, A.H.C., 1896. Preliminary notes on Genyornis newtoni, a new genus and species
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QUATERNARY AVIFAUNA OF AUSTRALIA - 849

APPENDIX I

Radiocarbon dates for selected Australian Pleistocene Sites bearing fossil birds.

Locality Dates Material Codes
(Upper & Lower)
(x 1000 yBP)

Queensland
King Creek 23.6+0.6//41.5+6.1
South Australia

{13f] Seton Rock Shelter 10.9+0.16//16.1+0.1
[27] Henschke's Bone Dig 33.8+2.4/-1.8//>35

Victoria Fossil Cave 16.74.25//>25
[7e, WA] Koonalda Cave 13.740.27//23.740.85
Baldina Creek 34.141.44 Molluscs SUA1065
Hookina Creek 33.27477//41.342?
Tasmania
[3] Cave Bay Cave 7.18+0.9//18.55+0.6 Shell ANU1552
Stockyard Site 760+0.07 Shell
JF-78 12.6+0.2 R5001/4
MU-206 10.14+0.2
Victoria
[2a] Clogg's Cave 8.742.3//22.27422
[6d] McEachems Cave 2.8+0.08//28.6+0.85
[7] Lancefield 26.6+0.65
{11] Spring Creek 19.8+0.39
Bushfield Bone Site 6.605+0.19

Western Australia

Devil's Lair 0.32+0.08//35.2+1.8
Skull Cave 2.9+0.08//7.875+0.1
Madura Cave 7.5+0.12//37.94+3.5AU-28

AU-24 430+0.16

850 - BAIRD ET AL.

APPENDIX II

Localities Yielding Avian Assemblages of Quaternary Age in
Australia

RE. Baird!, P.V. Rich!’ & GF. van Tets>

INTRODUCTION

The following appendix is designed to provide information on the geographic location,
species recorded and site information for Quaternary aged localities in Australia (i.e. radiocarbon
dates, etc.).

Identifications are derived from the literature, personal communications or personal
observations of the senior author. Alterations from the listings in Rich & van Tets (1982) are
real and determined primarily through the research of RFB.

The organization of each section revolves around current political boundaries. The states
are in alphabetical order, beginning with New South Wales and ending with Western Australia.
There are no known fossil localities yielding avian material of Quaternary age in the Australian
Capital Territory or the Northern Territory.

The figures noting the geographical distributions of the deposits also make the distinction
between cave deposits (open stars) and other deposits (closed stars: see Baird 1988a: this
volume). These deposits were placed in alphabetical order and subsequently numbered. Cave
names and numbers follow Matthews (1968).

The species lists are exactly that, and stratigraphic or chronologic divisions of the material,
if available, can be retrieved from the selected references. Organization and nomenclature in
each species list follows the taxonomy presented in Condon (1975) and Schodde (1975).
Common names for the various taxa can be found in the abovementioned references.

1 Department of Earth Sciences, Monash University, Clayton, Victoria 3168, Australia.

2 Departments of Botany and Zoology, Monash University, Clayton, Victoria 3168, Australia.

3 Division of Wildlife and Rangelands Management, CSIRO, Lyneham, Australian Capital Territory 2606,
Australia,

| Broken Hill
e

NEW SOUTH WALES

(see

Fig. I-1, below)

© Bourke <3

Armidale @

® Cobar

*10

Pani 4

44
#15
#13
289 SYDNEY

4
yy 16 2

x 6
x CANBERRA e I oe
ar nha
* cave deposits aT mace Da § } 7
* all other deposits ) / .
Q__100km * No
— — State borders ta 4 e

[1] Ashford Caves Area (UCMP Locs.

V-67237 & V-5545).

bone breccia (Rich 1975, Rich & van Tets

1982).

cf. Haliaeetus leucogaster (Accipitridae),
cf. Coturnix, (Phasianidae),

Tumicidae,

Psittaciformes,

Passeriformes.

[2] Bass Point * (Bowdler 1970, Gaughwin

[3]

[4]

[5]

1978).
Puffinus tenuirostris (Procellariidae),

Aves undetermined.

Bingara (Marcus 1975).
Dromaius sp. cf. D. novaehollandiae

(@romaiidae).

Canadian Gold Lead (Gulgong/Mudgee)
(Etheridge1889, Owen 1879b (unnamed
sediments), Rich 1975).

Dromomithidae (not Genyornis).

Cuddie Springs (Anderson & Fletcher
1934).
Genyornis sp. cf. G. newtoni

QUATERNARY AVIAN LOCALITIES - 851

(Dromomithidae).

[6] Currarong (Lampert 1971, Gaughwin 1978).
Puffinus tenuirostris (Procellariidae),

Aves undetermined.

[7] Durras North (Lampert 1966, Gaughwin
1978).
Puffinus tenuirostris (Procellariidae).

[8] Fisherman's Cliff (Marshall 1973).
Moorna Fm. (Fisherman's Cliff Local
Fauna).

Dromaiidae Undet.

[9] Frenchman's Creek (Marshall 1973).
Rufus Fm. (Frenchman's Creek Local Fauna).

Dromaiidae Undet.

[10] Lake Menindee (Tedford 1967, Rich 1975,
Rich et al. 1982 Patterson & Rich 1987, Rich &
van Tets 1982).

Site 1 (UCMP V-67233).
Dromaius novaehollandiae
(Dromaiidae),

Anatidae,

cf. Megapodiidae.

Site 2 (UCMP V-5371, =V-67185).

Dromaius novaehollandiae
(Dromaiidae),

Falco berigora (Falconidae).

Gallinula sp. (Rallidae)

Halcyon sp. (Alcedinidae)

Site 3 (UCMP V-67186).

Dromaius sp. cf. D. novaehollandiae
(Dromaiidae),

cf. Cygnus atratus (Anatidae),

Rallidae,

Alcedinidae.

Site 4 (UCMP V-67187).
Anatidae,

Rallidae,

Burhinidae,

852 - BAIRD ET AL.

Phoenicopteridae,

Passeriformes.

[11] Lake Tandou (Allen 1972, Rich 1975).
Dromaius sp. (Dromaiidae),
Podiceps sp. (Podicipedidae),
cf. Ardeiformes,
cf. Cygnus atratus (Anatidae),
cf. Columbidae.

[12] Lake Victoria (Marshall 1973).
Lake Victoria Sands

cf. Dromaius (Dromaiidae).

[13] Walli (Canowindra/Mandurama) (van Tets
1974a).
unknown cave
Progura gallinacea (Megapodiidae).

[14] Wellington (Mitchell 1839, Miller 1962,
Rich 1975, van Tets 1974a).
Bone Cave
Mitchell's Cave
Phosphate Mine

15] Willandra (Garpung A.).
Anatidae (large sp.).

[16] Wombeyan (Goulbum).
Broom Breccia (UCMP V5537) (Hope
1982).

Aves undet.

Guineacor Cave (W-121).

Progura naracoortensis
(Megapodiidae),

Turnix sp. (Turnicidae),

Platycercus sp. cf. P. elegans
(Platycercidae),

Passeriformes.

Wombeyan Quarry Cave (no longer
exists) (Hope 1982).

Dromaius sp. (Dromaiidae),

Progura gallinacea (Megapodiidae).

[17] Yessabah (near Kempsey).

unknown cave

Passeriformes.

PAPUA NEW GUINEA
[1] Pureni.
Casuarius sp. (Casuariidae).

QUEENSLAND
(see Fig. I-2, below)

Ww cave deposits
Cairns * all other deposits
Ai State bo
<u! a '@ borders
*Burketown

6 1%

to} 160km
_=—a

@ Winton
8x
@ Rockhampton
Dundo a4
e BRISBANE
| 3 *5
-_ =) keer). ae eA ns a
7» 10 rr

[1] Chillagoe.
Royal Arch Cave (CH-9) (Rich & van
Tets 1982).
Coturnix sp. (Phasianidae).

[2] Diamantina (general locality - could be only
of several places along the Diamantina River)
(Rich 1979).

Genyornis sp. (Dromomithidae).

[3] Darling Downs, Eastern (general locality)
(de Vis 1888b, 1891b, 1911; van Tets
1974a, van Tets & Rich 1980, Rich &

QUATERNARY AVIAN LOCALITIES - 853

van Tets 1982).

Columbidae,
Anhingidae, Psittaciformes,
Gypaetinae (= Taphaetus branchialis de Vis, cf. Aegothelidae,
Taphaetus lacertosus de Vis), Alcedinidae,
Progura gallinacea (Megapodiidae), Passeriformes,
(x) Palaeolestes gorei (Accipitriformes ?), Aves, undet.
Phaps indeterminate, either chalcoptera or
histronica (= Lithophaps ulnaris, de Vis), [11] Wyandotte Fm.
(Columbidae). Wyandotte Local Fauna * (McNamara
& Baird, pers. comm.)

[4] Darling Downs, near Dalby (Condamine Anhinga novaehollandiae (Anhingidae),
River) (Rich & van Tets 1982). Anseranas semipalmata (Anatidae),
Dromaius novaehollandiae (Dromaiidae). Anas sp. cf. A. superciliosa,

of. Anas.

[5] Darling Downs, King Creek (near Gallinula sp. cf. G. mortierii (Rallidae)
Ellinthorp) * (de Vis 1890, 1891a, 1905; van
Tets 1974c, Gill 1978, Baird 1986c). SOUTH AUSTRALIA
Buteonidae indet. (= Uroaetus brachialis, (see Fig. I-3, below)

de Vis, Taphaetus brachialis de Vis),
Gallinula mortierti (Rallidae), 7 7 Jk. Je
G. tenebricosa, 2 :
cf. Cygnus atratus (Anatidae). see tie & > a a
! eg hs

[6] Floraville Crossing (Floraville Fauna). wali aA
cf. Anseranas (Anatidae), a Maree ar
Tumicidae. * OoWsea pe AS

\ | €? |
if. os, AS g ¢/ “h17 |

[7] Gore (Longman 1945, van Tets 1974a, RR ohly “a
Bartholomai 1977). iA tee QR Sina ;
unnamed cave * allother deposits e ah 5
Progura naracoortensis (Megapodiidae). ee eee Aid 3 { ; f

panes, ME af oot

[8] Rockhampton District. ; ae BR rit I
Olsen's Tourist Cave (O-1). f NT og , oe 25)
Coturnix sp. (Phasianidae), , “aii syy 29.
Turnix sp. (Tumicidae). Tas“ Vie aN,

[9] Thorlindah (Paroo River) (Patterson & Rich [1] Baldina Creek (near Burra) (Patterson &
1987, Rich 1979). Rich 1987, Rich 1979).

Dromaius novaehollandiae (Dromaiidae), Dromaius novaehollandiae (Dromaiidae),
Genyornis sp. cf. G. newtoni (Dromomrnithidae). cf. Genyornis newton (Dromomithidae).

[10] Victoria River. [2] Billeroo Creek (Curnamona) (Williams
Russenden Cave (VR-2) * (upper bone-rich 1980).
pocket wall) (Rich in Archer 1978). Eurinilla Fm.

Coturnix sp. (Phasianidae), Dromaius sp. (Dromaiidae),

Turnix sp. (Tumicidae), Genyornis newtoni (Dromomnithidae).

854 -

[3]

[4]

[5]

BAIRD ET AL.

Burra (= Bute) (Patterson & Rich 1987).

Dromaius novaehollandiae (Dromaiidae).

Coffin Bay.

Brother's Island (Patterson & Rich 1987,
Johns 1966).

beach sand aeolianite.

Dromaius novaehollandiae (Dromaiidae),

Aves, undet.

COOPER CREEK (includes several localities
collected originally by J.W. Gregory and later
by R.A. Stirton, R.H. Tedford & colleagues,
see Rich 1975).

UNIVERSITY OF CALIFORNIA LOCALITIES

(Miller 1963 - 1966a-b, Rich 1975, Rich &
van Tets 1982, Rich et al. 1987).

Katipiri Sands (Malkuni Fauna).

Site 1 (UCMP V-5377).
Rallidae.

Site 2 (UCMP V-5378 = Unkimilka Waterhole).
Dromaius sp. (Dromaiidae),
Dromomnithidae,

Phalacrocorax sp. (Phalacrocoracidae).

Site 3 (UCMP V-5379) (between White

Crossing and Site 2).

Dromaius novaehollandiae (Dromaiidae),

Phalacrocorax sp. (Phalacrocoracidae;
middle-sized sp. and large sp.).

Site 4 (UCMP V-5380).

cf. Dromornithidae,

Anhinga novaehollandiae (Anhingidae),

Phalacrocorax sp. (Phalacrocoracidae;
middle-sized sp. and large sp.),

Anatidae,

Xenorhynchopsis minor
(Phoenicopteridae).

Site 5 (UCMP V-5381 = Pirranna Soakage).

Dromomithidae,

Phalacrocorax sp. (Phalacrocoracidae;
middle-sized sp.),

Anatidae.

Site 6 (UCMP V-5382 = Marconi locality =
Malkuni Waterhole) (Rich et al. 1987,
Rich & van Tets 1981).
Podiceps sp. (Podicipedidae),
Pelecanus conspicillatus (Pelecanidae),
Phalacrocorax sp. (Phalacrocoracidae),
Dromomithidae,
Anatidae,
cf. Aquila (Accipitridae),
Tyto sp. cf. T. novaehollandiae

(Tytonidae).
Site 7 (UCMP V-5859) (Rich & van Tets 1981).
Phalacrocorax sp. (Phalacrocoracidae;

middle-sized sp.),

Pelecanus conspicillatus (Pelecanidae),
Ardeidae,
Anatidae,
Accipitridae,

Phoenicopteridae.

Site 8 (UCMP V-5860) (Patterson & Rich 1987,

Rich & van Tets 1981).

Dromaius sp. (Dromaiidae),

Dromomithidae,

Pelecanus conspicillatus (Pelecanidae),

P. cadimurka,

Phalacrocorax sp. (Phalacrocoracidae;
large sp.),

Ardeidae,

Anatidae,

Xenorhynchopsis tibialis
(Phoenicopteridae).

Site 9 (UCMP V-5861 = Katapiri or Kuttipurra
Waterhole) (Rich & van Tets 1981).
Dromaius sp. (Dromaiidae),
Dromomithidae,
Pelecanus cadimurka (Pelecanidae),
Phalacrocorax sp. (Phalacrocoracidae;
middle-sized sp.),
Anatidae.

Site 10 (UCMP V-5869).
Anhinga novaehollandiae (Anhingidae),
Phalacrocorax sp. (Phalacrocoracidae),

Ciconiidae.

Site 14 (UCMP V-5866).
Podicipedidae,

Anhinga novaehollandiae (Anhingidae),
Pelecanus conspicillatus (Pelecanidae),
Phalacrocorax sp. (Phalacrocoracidae),
Ardeidae,

Anatidae,

Xenorhynchopsis minor

(Phoenicopteridae).

Site 16 (UCMP V-5868) (Rich & van Tets
1981).

Pelecanus conspicillatus (Pelecanidae),
Phalacrocorax sp. (Phalacrocoracidae),

cf. Cygnus atratus (Anatidae).

Site 18 (UCMP V-6147) (Rich & van Tets
1981).

Dromaius sp. (Dromaiidae),

Pelecanus conspicillatus (Pelecanidae),
Phalacrocorax sp. (Phalacrocoracidae),
Plataleidae,

cf. Cygnus atratus (Anatidae),

Columbidae.

Locality unknown (Miller 1966a-b, Rich &
van Tets 1981).
Anhinga laticeps (Anhingidae),
Pelecanus conspicillatus (= P. grandiceps,
de Vis; Pelecanidae).

H.Y.L. Brown Locality (Cutipirra) (Rich
& van Tets 1982).

Phalacrocoracidae.

J.W. GREGORY LOCALITIES (de Vis,
1888-1905, Gregory 1906, many are synonyms
of UCMP localities).

East of Pirani (Olson 1977).
Biziura lobata (= "Biziura exhumata",;

Anatidae).

Emu Camp (= Malkuni Waterhole = UCMP
V-5382 = Markoni). (Olson 1977, Rich

QUATERNARY AVIAN LOCALITIES - 855

1979, Rich & van Tets 1981).

cf. Genyornis newtoni (Dromomnithidae),

Pelecanus conspicillatus (Pelecanidae),

(x) Phalacrocorax gregorti
(Phalacrocoracidae),

(x) P. vetustus,

Anas castanea (= A. gracilipes, DeVis;
Anatidae),

(x) Archeocygnus lacustris,

(x) Chenopsis nanus.

Kalamurina (Rich & van Tets 1982).
cf. Gypaetinae (questionable identification,

P. V. Rich, pers. comm. )

COOPER CREEK, LOWER (Patterson &
Rich 1987, Rich et al. 1982, Rich & van Tets
1982, Rich et al. 1987).

unspecified locality.

Dromaius novaehollandiae (Dromaiidae),

Anhinga novaehollandiae (Anhingidae),

Pelecanus conspicillatus (= P. grandiceps
de Vis and = P. proavis, de Vis;
Pelecanidae),

(x) Phalacrocorax gregorit
(Phalacrocoracidae),

(x) P. vetustus,

(x) Archeocygnus lacustris (Anatidae),

(x) Chenopsis nanus,

Anas castanea (= Nettapus eyrensis, de Vis
and A. gracilipes, de Vis; Anatidae),

Xenorhynchopsis tibialis
(Phoenicoptertidae).

Locality 2 (Mudlamarukupa) (Patterson & Rich
1987). Dromaius sp. (Dromaiidae),
Dromomithidae.

Locality 3 (Tantatuluru) (Patterson & Rich
1987).
Dromaius sp. (Dromaiidae).

Locality 4 (Kudnampirra) (Patterson & Rich
1987, Baird & Rich, in press).
Dromaius novaehollandiae (= D. patricius,

de Vis; Dromaiidae),

856 - BAIRD ET AL.

Palaelodus sp. (Palaelodidae).

Locality 5 (Eli Hartigs Soak) (Rich et al.

1982).

Falco berigora (= Asturaetus "Plioaetus"
furcillatus, de Vis; Falconidae).

Locality 6 (Patara Mordu, Pataramordu,
Pataruwordu).
(x) Aviceda gracilis (Leptodontidae).

COOPER CREEK, PIRANNA SOAKAGE.
(Piaranni, Piranni, Pijari; see UCMP

V-5381) (Rich & van Tets 1982,

Rich et al. 1987).

Unduwumpa (= Undusoumpa).

(x) Archeocygnus lacustris (Anatidae),

(x) Chenopsis nanus,

Xenorhynchus minor
(Phoenicopteridae).

Wurdulmandkula (= Wurdulmankula)
(Patierson & Rich 1987, van Tets & Rich
1980).
Dromaius novaehollandiae (= D. patricius,
de Vis; Dromaiidae),

(x) Phalacrocorax gregorii (Phalacrocoracidae),
(x) P. vetustus,
(x) Archeocygnus lacustris (in part = Anatidae

and Phoenicopteridae),

(x) Chenopsis nanus (in part Phoenicopteridae),
Phoenicopterus ruber (Phoenicopteridae),
Ocyplanus proesus,

Phaps sp. (either P. histronica or P.
chalcoptera = Leucosarcia proevisa
and Nyroca effodiata, DeVis;
Columbidae).

[6] CURRAMULKA (York Peninsula) (Rich & van

Tets 1982).

Cora Lynn Cave (Y-1).
Tytonidae.

Town Cave (Y-2).
of. Ardeotis australis (Otididae).

[7]

[8]

Dempsey's Lake (Williams 1980).
Dromaiidae,

Dromomithidae,

Anatidae.

Devon Downs (near Mannum) (Williams

1980).

Dromaius novaehollandiae (Dromaiidae),

Anas sp. (Anatidae),
Cygnus atratus,

Biziura lobata,

Aquila audax (Accipitridae).

Diamantina River (see Queensland).

[9] FLINDERS RANGES REGION.

Mair's Cave (F-3) (Baird 1984).
Accipitridae,

Falco cenchroides (Falconidae),
Coturnix sp. (Phasianidae),
Turnix sp. (Tumicidae),

Gallinula mortierii (Rallidae),
Columbiformes,

Psittaciformes,

Tyto alba (Tytonidae).

Aegotheles cristatus (Aegothelidae),
Cinclosoma sp. (Orthonychidae),
Amytornis sp. (Maluridae),
Passeriformes, unident.

[10] Gawler (Rich & van Tets 1982).

[11] Hookina Creek (Parachilna) (Williams

Rallidae.

1980).
Pooraka Fm.
Dromaius sp. (Dromaiidae),
Genyornis sp. (Dromomithidae),

Aves undet.

[13] KANGAROO ISLAND REGION.

unknown locality.

Dromatius baudinianus (Dromaiidae).

[13a] Cape du Couedic (Parker 1984).
Dromaius baudinianus (Dromaiidae).

[13b] Eleanor River (Parker 1984).
Dromaius baudinianus (Dromaiidae).

[13c] Emu Four Hole Cave (K-20) (Williams
1980, Parker 1984).
Dromaius baudinianus (Dromaiidae).

[13d] Kelly Hill Cave (K-1) (Williams 1980,
Parker 1984).

Dromaius baudinianus (Dromaiidae).

[13e] Rocky River (Williams 1980).
swamp deposit.

Dromaius sp. (Dromaiidae).

[13f] Seton Rock Shelter (K-30) * (Hope et
al. 1977).
Pachyptila sp. cf. P. salvini
(Procellariidae),
Puffinus sp.,
Threskiornis sp. cf. T. molucca
(Threskiomithidae),
cf. Anseranas semipalmata (Anatidae),
Tadorna sp. cf. T. tadornoides,
Anas sp. cf. A. superciliosa,
A. sp. cf. A. castanea,
Malacorhynchus membranaceus,
Hieraaetus morphoides (Accipitridae),
Falco berigora (Falconidae),
Coturnix sp. cf. C. pectoralis
(Phasianidae),
Turnix varia (Tumicidae),
T. velox,
Rallus philippensis (Rallidae),
R. pectoralis,
Porzana sp. cf. P. fluminea,
Gallinula mortierit,
G. ventralis,
Burhinus magnirostris (Burhinidae),
Gallinago hardwickii (Scolopacidae),
Larus novaehollandiae (Laridae),
Sterna sp. cf. S. nereis,
Ocyphaps lophotes (Columbidae),
Pezoporus wallicus (Platycercidae),

QUATERNARY AVIAN LOCALITIES

Lathamus discolor,

Hirundinidae undet.,

Meliphagidae,

Cincloramphus cruralis (Sylviidae),
Grallina cyanoleuca (Grallinidae),
Gymnorhina tibicen (Cracticidae),
Strepera sp.,

Corvus sp. (Corvidae),

Passeriformes, unident.

[14] Lake Callabonna.
(Holocene).
Phalacrocorax sp. (Phalacrocoracidae).

(Diprotodon level) (Patterson & Rich 1987,
Stirling & Zietz 1896-1913, Rich 1979).
Dromaius novaehollandiae (Dromaiidae),

Genyornis newtoni (Dromomnithidae).

[15] Lake Eyre area (exact locality unknown)
(Rich & van Tets 1982).
Phoenicopteridae (="Ocyplanus proeses"
de Vis).

[16] Lake Kanunka (Patterson & Rich 1987,
Rich & van Tets 1982).
Dromaius novaehollandiae (Dromaiidae),
Phalacrocoracidae,
Anatidae,
Accipitridae,
Charadriiformes,

Phoenicopteridae.

Site 1 (UCMP V-5772) (Miller 1963, Patterson
& Rich 1987, Rich & van Tets 1982, Rich et
al. 1987).

Dromaius novaehollandiae (Dromaiidae),
Anhinga novaehollandiae (Anhingidae),
Ciconiidae,

Anatidae,

Grus sp. (Gruidae),

Charadriiformes,

Ocyplanus proesus. (Phoenicopteridae),

Phoenicopterus ruber,

Xenorhynchopsis minor,

- 857

858 - BAIRD ET AL.

Site 2 (Rich & van Tets 1981, Rich & van Tets
1982).
Pelecanus cadimurka (Pelecanidae),
P. conspicillatus,
Accipitridae,

Passeriformes.

South (Rich & van Tets 1982).
Ardeidae,
Anatidae,
Accipitridae,
Rallidae,
Otididae,
Phoenicopteridae.

[17] Lake Kittakittaooloo (Rich & van Tets
1982).

Dromaius novaehollandiae (Dromaiidae).

[18] Lake Millyera area (Callen 1984).
Eurinilla Fm.
cf. Dromaius (eggshell: Dromaiidae).

[19] Lake Palankarinna (Williams 1980).
Katipiri Sands (Channel Sand Locality,
UCMP V-5854).

Aves undet.

[20] LOWER SOUTH EAST REGION
unknown sinkhole (Mt. Gambier Range),
(Lydekker 1891, Owen 1879a).
cf. Genyornis newtoni (Dromomithidae).

Goulden's Hole (L-8).
Aquila audax (Accipitridae),
Phalacrocorax melanoleucus
(Phalacrocoracidae),
Passeriformes, unident.

"Green Waterhole"(L-81) (Williams 1980
Baird 1984, Baird 1985).

Accipitridae,

Falco sp. cf. F. berigora (Falconidae),
Gallinula mortierii (Rallidae),

Phaps chalcoptera (Columbidae),

Phaps sp.,

Calyptorhynchus magnificus (Cacatuidae),

C. lathami,

Callocephalon fimbriatum,

Cacatua tenuirostris,

Platycercus sp. (Platycercidae),
Centropus colossus (Centropodinae),

Dacelo novaeguineae (Alcedinidae),

Orthonyx hypsilophus (Orthonychidae),

Hirundinidae,

Dasyornis broadbenti (Acanthizidae),

Manorina melanocephala (Meliphagidae),

Corvus sp. (Corvidae).

The Pines (L-61) (Baird 1985).
Cacatua tenuirostris (Cacatuidae).

Tantanoola Cave (L-12?) (Rich & van Tets
1982).

Aves, undet.

[21] Manku (Kalakoopah Creek) (Rich et al. 1987).
Phoenicoptendae, indet..

[22] Mt. Gambier.
cf. Genyornis newtoni (Dromomithidae).

Mair's Cave (see Buckalowie).

[23] Normanville (= Salt Creek) (Patterson &
Rich 1987).
Dromaius novaehollandiae (Dromaiidae),
Genyornis newtoni (Dromornithidae),
Tumicidae,
Rallidae,
Passeriformes,
Aves, undet.

[24] Penola (Woods 1866, Rich 1979)
swamp deposit.
Dromomithidae.

[25] Pondalowie Bay (Williams 1980).

Dromaiidae.
Port Lincoln (see Brother's Island)

[26] Salt Creek (Barker) (Williams 1980).
swamp deposit.

Dromaius sp. (Dromaiidae),
Genyornis newtoni (Dromomithidae),

Aves undet.

Tantanoola Cave (see Lower South East Region).

[27] UPPER SOUTH EAST REGION.
Blanche Cave (U-4) (Rich 1979).

cf. Genyornis newtoni (Dromomithidae).

Dogs Prohibited Cave (U-?) (Williams
1980).

Aves undet.

Fox Cave (U-22).

Progura naracoortensis (Megapodiidae),

Dacelo sp. cf. D. novaeguineae
(Alcedinidae).

Henschke's Bone Dig (U-91) * (Patterson
& Rich 1987, van Tets 1974a).
Dromaius novaehollandiae (Dromaiidae),
Anatidae,

Progura gallinacea (Megapodiidae),
P.. naracoortensis,

Coturnix sp. (Phasianidae),

Turnix varia (Tumicidae),

Rallidae undet.

Gallinula mortierii

Psittaciformes undet.
Passeriformes undet.

Corvus sp. (Corvidae).

Victoria Fossil Cave (U-1) * (van Tets &
Smith 1974, Olson 1976, Patterson & Rich
1987).

Dromaius novaehollandiae (Dromaiidae),
Progura naracoortensis (Megapodiidae),
Leipoa ocellata,

Coturnix australis (Phasianidae),
Coturnix sp.,

Turnix varia (Tumicidae),

cf. Pedionomus torquatus (Pedionomidae),
Rallus philippensis (Rallidae),
Peltohyas australis (Charadriidae),
Tringa glareola (Scolopacidae),
Gallinago hardwicki,

QUATERNARY AVIAN LOCALITIES - 859

[28]

Calidris ruficollis,

Pezoporus wallicus (Platycercidae),
Tyto novaehollandiae (Tytonidae),
Gymnorhina tibicen (Cracticidae),

Grallina cyanoleuca (Grallinidae).

Wombat Cave (U-58).
Progura naracoortensis (Megapodiidae).

WARBURTON RIVER LOCALITIES
(Rich & van Tets 1981, Patterson & Rich 1987).
Cassidy Locality (UCMP V-5539).

cf. Genyornis sp. (Dromomithidae).

Green Bluff Locality (UCMP V-5775)

(Patterson & Rich 1987, Rich 1975, Rich &
van Tets 1982).

Dromaius novaehollandiae (Dromaiidae),

Dromomithidae,

Phalacrocorax sp. (Phalacrocoracidae),

Anhinga sp. (Anhingidae),

Anatidae,

Accipitridae,

Passeriformes.

Kalamurina (Patterson & Rich 1987, Rich

1979, Rich & van Tets 1981, Rich & van Tets

1982).

Dromaius novaehollandiae (Dromaiidae),

Dromomithidae,

Pelecanus conspicillatus (= P. validipes de Vis:

Pelecanidae),

(x) Phalacrocorax gregorii (Phalacrocoracidae),

(x) P. vetustus, Anas castanea (= "Anas
gracilipes" de Vis: Anatidae),

?Gypaetinae (="Taphaetus lacertosus" de Vis:
Accipitridae).

Lookout Locality (UCMP V-5776) (Rich
1975).
Phalacrocorax sp. (middle-sized sp., large sp.)

(Phalacrocoracidae).

Marcus Locality (Rich 1975, Rich & van Tets
1981, Rich & van Tets 1982).

Pelecanus conspicillatus (Pelecanidae),
Phalacrocorax sp. (middle-sized sp., large sp.)

860 - BAIRD ET AL.

(Phalacrocoracidae)
Ardeidae.

nr. Lake Miamiana (UCMP V-5539)
(Williams 1980).

Dromomnithidae.

Purdrakadrinna Soakage (UCMP V-5777)

Rich 1975).

Phalacrocorax sp. (Phalacrocoracidae).

TASMANIA
(see Fig. I-4, below)

YON

“ \

¢ )

Te,

4° ~8ee

nC

10aYOe—_, —,
nye ot gros ee

28 Launceston

\ \7

\ Y es

\ &
\ Jo

cave deposits \ steatngordon HOBART nods
% N
« aotherdeposts (OR

Y

° 35 70k \ ci > p) UKM
mM A

—— esa ?

Lae a ; ey

[1] Cape Grim (Gaughwin 1978).

Puffinus tenuirostris (Procellariidae).
[2] Flowery Gully (Gill 1968).

Quarry (FG-202).

Phaps sp. (Columbidae).

[3] HUNTER ISLAND.

Cave Bay Cave * (Bowdler 1975, Gaughwin

1978).
Puffinus tenuirostris (Procellariidae),
undetermined Aves.

Muttonbird Midden * (Bowdler 1979,
Gaughwin 1978).

Puffinus tenuirostris (Procellariidae).

Stockyard site * (O'Conner 1980, Geering
1981, van Tets & O'Conner 1983,
Gaughwin 1978).

Tasidyptes hunteri (Spheniscidae),

Puffinus tenuirostris (Procellariidae),

undetermined Aves.

[4] JUNEE-FLORENTINE VALLEY (Goede &

[5]

[6]

Murray 1979, van Tets 1978).
Beginners' Luck Cave (JF-78) *.
Puffinus griseus (Procellariidae),
Gallinula mortierii (Rallidae).

Emu Cave (JF-154).
Dromaius sp. (Dromaiidae).

Titans Shelter (JF-97) *.
Falco sp. (Falconidae).

Irishtown (Scott 1932).

Dromaius diemenensis (Dromaiidae).

King Island.

Dromaius minor (Dromaiidae).

[7] LOUISA BAY.

[8]

[9]

Louisa River 1 (Vanderwal & Horton 1984,
Gaughwin 1978).
Puffinus tenuirostris (Procellariidae),

undetermined Aves.

Louisa River Cave 2 (Vanderwal & Horton
1984, Gaughwin 1978).

Puffinus tenuirostris (Procellariidae).

Mole Creek (Scott 1932).
Unknown Cave.

Dromaius "diemenensis" (Dromaiidae).

Montague * (Murray & Goede 1978).
Pleisto Scene Cave (MU-206) * (van Tets
1978).

Dromaius sp. (Dromaiidae),

Gallinula mortierii (Rallidae),

Aves, undet.

[10] SMITHTON.

[10a] Mowbray Swamp(Scott 1932, Murray &
Goede 1977).
Dromaius "diemenensis" (Dromaiidae),
Aves undet.

[10b] Pulbeen Swamp * (Murray & Goede 1977,
Colhoun et al. 1977).
Dromaius sp. (Dromaiidae),
Aves undet.

[10c] Scotchtown Cave (mostly quarried away)
(Gill & Banks 1956).
Aves, undet.

[11] West Point (Gaughwin 1978).
Puffinus tenuirostris (Procellariidae).

(see Fig. I-5, below)
7 4 NT
; [WA 5
» 1
*Ouyen te
=
Horsham
7. \ x
5%, .
i) MELBOURNE Ge= b ?
sat & ey oH Me, A Baiinsdalesy -— ——~ mers
Sd ree — 3a Warrnambool PAL 8° vA eed aig
SRS on ~ a * all other deposits
~v, \ =
Q___100km LF ~ state borders

[3 Limestone

[1] Bat's Ridges.
Velvet Bush Cave (BR-65).
Tumicidae,

Passeriformes.

[2] BUCHAN DISTRICT.

[2a] Clogg's Cave (EB-2) * (Baird 1986c).
Accipiter sp. cf. A. fasciatus (Accipitridae),
Falco berigora (Falconidae),

Coturnix sp. (Phasianidae),
C. chinensis,
Turnix varia (Tumicidae),

T. sm. sp.,

QUATERNARY AVIAN LOCALITIES

Pedionomus torquatus (Pedionomidae),
Rallus philippensis (Rallidae),
R. pectoralis,
Porzana fluminea,
P. tabuensis,
Erythrogonys cinctus (Charadriidae),
Phaps chalcoptera (Columbidae),
P. elegans,
Trichoglossus haematodus (Loriidae),
Glossopsitta sp. cf. concinna,
Glossopsitta sp. cf. pusilla,
Pezoporus wallicus (Platycercidae),
Melopsittacus undulatus,
Platycercus elegans,
P. sp. cf. P. eximius,
Neophema chrysostoma,
Cuculus pallidus (Cuculidae),
C. variolosus,
C. sp.,
Chrysococcyx lucidus,
C. sp,
Tyto alba (Tytonidae),
Podargus strigoides

(Podargidae),
Aegotheles cristatus (Aegothelidae),
Halcyon sancta (Alcedinidae),
H. pyrrhopygia,
Hirundinidae,
Anthus novaeseelandiae (Motacillidae),
Petroica sp. (Muscicapidae),
Pachycephala sp.,
Psophodes olivaceus (Orthonychidae),
Cinclosoma punctatum,
Cincloramphus mathewsi (Sylviidae),
C. sp.,
C. cruralis,
Malurus sp. (Maluridae),
Stipiturus malachurus,
Dasyornis brachypteris (Acanthizidae),
D. broadbenti,
Pycnoptilus floccosus,
Climacteris sp. (Climacteridae),
Pardalotus sp. (Pardalotidae),
Zosterops sp. (Zosteropidae),
Ploceidae indet.,
cf. Emblema temporalis,

Artamus sp. (Artamidae),

- 861

862 - BAIRD ET AL.

A. cyanopterus,
A. sp. cf. A. cyanopterus,
Corvus sp. (Corvidae).

[2b] Kingsley Cave).
Aquila audax (Accipitridae).

[2c] Mabel Cave (EB-1) * (Baird 1986c).
Puffinus sp. (Procellariidae),
Pelecanoides sp.,

Phalacrocorax sp. cf. P. melanoleucus,
(Phalacrocoracidae)

Anatidae,

Circus sp. (Accipitridae),

Falco cf. peregrinus (Falconidae),

Coturnix chinensis (Phasianidae),

Coturnix sp.,

Turnix varia (Tumicidae),

Rallus philippensis (Rallidae),

Vanellus miles (Charadriidae),

Leucosarcia melanoleuca (Columbidae),

Phaps chalcoptera,

P. elegans,

Calyptorhynchus lathami (Cacatuidae),

C. funereus,

Callocephalon fimbriatum,

Cacatua roseicapilla,

Trichoglossus haematodus (Loriidae),

Glossopsitta concinna,

G. cf. G. pusilla,

Alisterus scapularis (Polytelidae),

Platycercus elegans (Platycercidae),

P. cf. P. eximius,

Neophema chrysostoma,

Chrysococcyx sp. (Cuculidae)

Aegotheles cristatus (Aegothelidae),

Hirundapus caudacutus (Apodidae),

Halcyon sancta (Alcedinidae),

Hirundinidae,

Anthus novaeseelandiae (Motacillidae),

Cinclorhamphus cruralis (Sylviidae),

Megalurus timoriensis

Cinclosoma punctatum (Orthonychidae),

Malurus sp. (Maluridae),
Dasyornis brachypteris (Acanthizidae),
Pycnoptilus floccosus,

Acanthiza sp.

Daphoenositta chrysoptera,

Climacteris sp. (Climacteridae),

cf. Anthochaera carunculata (Meliphagidae),

cf. A. chrysoptera,

Zosterops sp.,

Ptilonorhynchus violaceus
(Ptilonorhynchidae),

Artamus cyanopterus (Artamidae),

Cracticidae.

[2d] The Maze (M-30?).
Aves undet.

[2e] M-28.
Platycercus cf. eximius
(Platycercidae),

Passeriformes.

[2f] NG-2.
Hirundinidae,

Aves, undet.

[2g] Pyramids Cave (M-89) (Baird 1986c).
Coturnix sp. (Phasianidae),
Turnix varia (Tumicidae),
Glossopsitta sp. (Loriidae),
Alisterus scapularis (Polytelidae),
Platycercus elegans (Platycercidae),
P. cf. P. elegans,
P. sp.,
Chrysococcyx sp.,
Tyto novaehollandiae (Tytonidae),
Aegotheles cristatus (Aegothelidae),
Caprimulgus mysticalis (Caprimulgidae),
Atrichornis rufescens (Atrichomithidae),
Orthonyx sp. (Orthonychidae),
Psophodes olivaceus,
Cinclosoma punctatum,
Pycnoptilus floccosus (Acanthizidae),
P.n. sp.,
Climacteris sp. (Climacteridae),
Ptilonorhynchus violaceus

(Ptilonorhynchidae).

[2h] Trogdip Cave (EB-10).
Dromaius novaehollandiae (Dromaiidae).

[3] Bushfield Bone Site (near Warrnambool,
Merri River; Baird 1984).
Anatidae,
Gallinula mortierii (Rallidae).

[4] BYADUK.
Church Cave (H-15) (Baird, in press d).
Coturnix sp. (Phasianidae),
Turnix varia (Tumicidae),
Psittaciformes,
Aegotheles sp. cf. A. cristatus
(Aegothelidae),
Passeriformes.

Flower Pot Cave (H-19) (Baid, in press d).

Tumicidae,
Barnardius barnardi (Plaqtycercidae).

Harman's Cave (H-12) (Baird, in press d).
Coturnix sp. (Phasianidae),

Turnix varia (Tumicidae),

Rallidae,

Pedionomus torquatus (Pedionomidae),
Platycercus eximius (Platycercidae),
Psittaciformes,

Cincloramphus mathewsi (Sylviidae),
Gymnorhina tibicen (Cracticidae),

Passeriformes.

[5] Gisborne.

Gisborne Cave (H-27) (Baird, in press d).

Tumicidae,

Passeriformes.

[6] GLENELG RIVER DISTRICT.

[6a] Amphitheatre Cave (G-2) (Baird 1984,
1986c, in press a).
Eudyptula minor (Spheniscidae),
Pachyptila sp. (Procellariidae),
Phalacrocorax sp. (Phalacrocoracidae),
Anatidae sm. sp.,
Coturnix sp. (Phasianidae),
Turnix varia (Turnicidae),
Rallus philippensis (Rallidae),
Gallinula mortierii,
Larus novaehollandiae (Laridae),
Phaps chalcoptera (Columbidae),

QUATERNARY AVIAN LOCALITIES - 863

P. elegans,

Cacatua tenuirostris (Cacatuidae),

C. roseicapilla,

Platycercus elegans (Platycercidae),

Hirundinidae,

Cinclosoma punctatum (Orthonychidae),

Dasyornis broadbenti (Acanthizidae),

D. brachypteris,

Ptilonorhynchus violaceus
(Ptilonorhynchidae),

Cracticidae,

Corvus sp. (Corvidae).

[6b] Curran's Creek Cave (G-4) (Baird 1984,

1986c).

Coturnix sp. (Phasianidae),

Gallinula mortierii (Rallidae),

cf. Porzana,

Phaps elegans (Columbidae),

Glossopsitta pusilla (Loriidae),

Pezoporus wallicus (Platycercidae),

Platycercus sp. cf. P. eximius,

Neophema sp. cf. N. chrysostoma,

Ceyx azurea (Alcedinidae),

Halcyon sp.,

Aegotheles sp. cf. A. cristatus
(Aegothelidae),

Hirundinidae,

Cinclosoma punctatum (Orthonychidae),

Malurus sp. (Maluridae),

Dasyornis broadbenti (Acanthizidae),

Ploceidae indet.

[6c] Fern Cave (KB-1).

Gallinula mortierii (Rallidae).

[6d] McEachern's Cave (G-5) (McNamara 1981,

Baird 1984).

Dromaius novaehollandiae (Dromaiidae),

Falco sp. cf. F. berigora (Falconidae),

Turnix sp. cf. T. varia (Tumicidae),

Coturnix sp. (Phasianidae),

Ninox sp. (Strigidae),

Dacelo sp. cf. D. novaeguineae
(Alcedinidae),

Podargus sp. (Podargidae),

Dasyornis sp. (Acanthizidae),

864 - BAIRD ET AL.

Gymnorhina sp. cf. G. tibicen (Cracticidae),
Strepera sp. cf. S. versicolor

Passeriformes, undet.

[7] Lancefield * (Gillespie et al. 1978, Baird
1984).
Dromaius sp. (Dromaiidae),
cf. Genyornis (Dromomithidae),
Gallinula mortierii (Rallidae).

[8] Morwell (Rich & McEvey 1980).
Pedionomus torquatus (Pedionomidae).

[9] Mount Eccles.
Natural Bridge (H-10).
Aves undet.

[10] PHILLIP ISLAND.
Forrest Caves (Gaughwin 1978).

Puffinus tenuirostris (Procellariidae).

Cape Woolamai (Gaughwin 1978).

Puffinus tenuirostris (Procellariidae).

Point Grant (Gaughwin 1978).

Puffinus tenuirostris (Procellariidae).

Cat Bay (Gaughwin 1978).

Puffinus tenuirostris (Procellariidae).

[11] Spring Creek * (Baird 1984, Flannery &
Gott 1984).
Gallinula mortierii (Rallidae).

[12] Warrnambool District.
Thunder Point Cave (W-8) (Rich & Gill
1976).
cf. Dromomithidae.

WESTERN AUSTRALIA

(see Fig. I-6, below)

[1] AUGUSTA.

Bone Cave (AU-2) (Patterson & Rich 1987).
Dromaius novaehollandiae (Dromaiidae).

Deepdene Cave (AU-1) (Rich & van Tets

1982).
Biziura lobata (Anatidae),

Passeriformes.

Labryinth Cave (AU-16).

Aves, undet.

rm
a | NT :
a au
Ss
Der
Tas. =) Vic i |
|
y \
Ie |
rut
Fe |
|
WA ISA 0 roo] |
——=
km |
| |
| | |
n* : o i
|

[2]

~~ 6 cave deposits
Esperance
5 * all other deposits
° 300km - state borders
—

Skull Cave (AU-8) * (Porter 1979, Baird
1986c).

Coturnix sp. (Phasianidae),

Phaps elegans (Columbidae),
Glossopsitta porphyrocephala (Loriidae),
Platycercus icterotus (Platycercidae),
Aegotheles cristatus (Aegothelidae),
Atrichornis clamosus (Atrichomithidae),
Hirundinidae indet.,

Malurus sp. (Maluridae),

Dasyornis longirostris (Acanthizidae),

Artamus cyanopterus (Artamidae).

AU-24 * (Archer & Baynes 1972).

cf. Passeriformes.

Coolup Bore 46-48 feet.

Cracticidae.

[3] East Moore (Patterson 1984).
possibilities

Caladenia Cave(EM-17)WAM dig, early 1900's

Dromaius novaehollandiae (Dromaiidae).

[4] ENEABBA.
Aiyenu Cave (E-9).
Podargus strigoides (Podargidae).

Facts of Life Cave (E-12).
Meliphagidae.

Weelawadji Cave (E-24).
Tyto sp.

{5] Hunter River (Butler & Merrilees 1971).
sand hills 2.5 miles north east of
Bremer Bay.

Coturnix sp. (Phasianidae),
Psittaciformes,

Passeriformes.

[6] Jurien Bay.
Kjeldahi Cave (J-5).
Passeriformes.

[7] NULLARBOR PLAINS REGION.
[7a] Abrakurrie Cave (N-3).
Psittaciformes.

[7b] Bildoolja Cave (N-31) (Baird, on press b).

Turnix sp. cf. T. velox (Tumicidae),

Hirundinidae indet.,

Cinclosoma sp. cf. C. alisteri
(Orthonychidae),

Amytornis sp. (Maluridae).

Horseshoe Cave (N-59).

Turnix sp. (Tumicidae),

[7c]
Passeriformes.

[7d] Koomooloobuka Cave (N-6) (Baird, in
press b).
Turnix sp. cf. T. velox (Tumicidae),
Geopelia sp. cf. G. striata (Columbidae),

Psittaciformes,

QUATERNARY AVIAN LOCALITIES - 865

Chrysococcyx sp. cf. C. basalis (Cuculidae),

Oreoica gutteralis (Muscicapidae),

Cinclosoma sp. cf. C. alisteri
(Orthonychidae),

Amytorns sp. (Maluridae),

Artamus sp. (Artamidae).

[7e] Koonalda Cave (N-4) * (Baird 1986c).

[7f]

Anatidae sm. sp.,

Accipiter sp. cf. A. fasciatus (Accipitridae),
A. cirrhocephalus,

Circus sp.,

Falco berigora (Falconidae),

F. cenchroides,

Coturnix sp. (Phasianidae),

Turnix sp. cf. T. velox (Tumicidae),
Phaps elegans (Columbidae),
Ocyphaps lophotes,

Polytelis anthopeplus (Polytelidae),
Melopsittacus undulatus (Platycercidae)
Psephotus haematogaster,

Neophema sp. cf. N. splendida,
Chrysococcyx sp. (Cuculidae),

Tyto alba (Tytonidae),

T. novaehollandiae,

Caprimulgus guttatus (Caprimulgidae),
Hirundinidae,

Anthus novaeseelandiae (Motacillidae),
cf. Oreoica gutteralis (Muscicapidae),
Sphenostoma sp. (Orthonychidae),
Cinclosoma alisteri,

Cincloramphus cruralis (Sylviidae),

cf. Cincloramphus,

Amytornis textilis (Maluridae),

cf. Lichenostomus virescens (Meliphagidae),
Meliphagidae, indet.,

Artamus cinereus (Artamidae).

Madura Cave (N-62) * (Baird 1986c, in
prep.).

Turnix varia (Turnicidae),

T. sp. of. T. velox,

Calidris sp. (Charadriidae),

Charadrii indet.,

Ocyphaps lophotes (Columbidae),
Geopsittacus occidentalis (Platycercidae),
Melopsittacus undulatus,

866 - BAIRD ET AL.

Aegotheles cristatus (Aegothelidae) [8] Scott River (Butler 1969, Rich 1975).
Colluricincla sp. cf. C. harmonica coastal dunes.

(Muscicapidae), ?7Dromomithidae or ?Aepyornithidae
Cinclosoma alisteri (Orthonychidae), cf. Dacelo novaeguineae (Alcedinidae),
Pomatostomus superciliosus (Timaliidae), cf. Podargus strigoides (Podargidae).
Amytornis textilis (Maluridae),

Acanthiza chrysorrhoa (Acanthizidae), [9] West Bullsbrook (Rich & van Tets 1982).
Acanthizidae undet., peat swamp.
cf. Lichenostomus virescens (Meliphagidae), ?Phalacrocoracidae.

Artamus sp. (Artamidae).
[10] Witchcliffe.

[7g] New Cave (N-11). Bride Cave (WI-24).
Accipiter sp. cf. A. cirrhocephalus Phalacrocorax sp. (Phalacrocoracidae),
(Accipitridae), Passeriformes

Passeriformes, undet.
Devil's Lair (WI-61E ajoins Nannup Cave

[7h] Old Homestead Cave (N-83). (WI-60)) * (Cook 1960, Balme et al.
Passeriformes. 1978, Baird 1986c).
Aquila audax (Accipitridae),
[7i] Thylacine Hole (N-63) (all specimens Accipiter cirrhocephalus,
mummified). Coturnix sp. (Phasianidae),
Falco cenchroides (Falconidae), Turnix varia (Tumicidae),
F. berigora, Rallus philippensis (Railidae),
Tyto alba (Tytonidae), Porzana sp. cf. P. fluminea,
T. novaehollandiae, Phaps elegans (Columbidae),
Aegotheles cristatus (Aegothelidae) Glossopsitta porphyrocephala (Loriidae),
Melopsittacus undulatus (Platycercidae),
[7j}] Warbla Cave (N-1). Purpureicephalus spurius,
Falconidae. Platycercus icterotus,

Barnardius zonarius,
[7k] Weebubie Cave (N-2). Psephotus varius,
Neophema sp. cf. N. elegans,
Tyto novaehollandiae (Tytonidae),
Cuculus pyrrhophanus (Cuculidae),

Accipitridae,

Passeriformes(songbirds).

Aegotheles cristatus (Aegothelidae),
[71] Weekes Cave (N-15) (van Tets 1974b,

Baird 1986c).
Platalea flavipes (Threskiomithidae), Anthus novaeseelandiae (Motacillidae),

Accipiter fasciatus (Accipitridae), Malurus sp. (Maluridae)
Falco cenchroides (Falconidae),

Atrichornis clamosus (Atrichomithidae),

Hirundinidae, indet.,

Dasyornis longirostris (Acanthizidae),

Turnix sp. cf. T. velox (Turnicidae), Climacteris sp. (Climacteridae),

Chrysococcyx sp. cf. C. basalis (Cuculidae), Ploceidae, indet.

Tyto sp. cf. T. novaehollandiae (Tytonidae), Strepera versicolor (Cracticidae).

Meliphagidae, Aves, undet.
Poephila guttata (Ploceidae),
Artamus sp. (Artamidae), Mammoth Cave (WI-38) * (Lundelius 1960,

Passeriformes, undet. Merrilees 1968, Archer et al. 1980).

QUATERNARY AVIAN LOCALITIES - 867

Merrilees 1968, Archer ef al. 1980).
cf. Dromaius (Dromaiidae),
Dromomnithidae,

Aquila audax (Accipitridae).

small cavern north of Nannup Cave
(WI-60).

Accipiter cirrhocephalus (Accipitridae),
Turnix sp. (Tumicidae),

Psittaciformes.

[11] Yanchep.
Orchestra Shell Cave (YN-130) * (Archer
1974).
Hirundinidae,

Aves undet.

PLATES

Plate 1. Restoration of the Giant Coucal (Centropus colossus) from the Pleistocene of Australia (Rich &
van Tets 1985).

Plate 2. Restoration of the giant megapode (Progura gallinacea) from the Pleistocene of Australia (Rich &
van Tets 1985).

Plate 3: Restoration of a mihirung (Genyornis newtoni) from the Pleistocene of Australia (Rich & van
Tets 1985).

868 - BAIRD ET AL. PLATE 1.

PLATE 2 QUATERNARY AVIAN LOCALITIES - 869

870 - BAIRD ET AL. PLATE 3

CHAPTER 22

FOSSIL EGGS FROM THE
TERTIARY AND QUATERNARY
OF AUSTRALIA

Dominic L. G. Williams! and Patricia Vickers-Rich2

IMILOMUGHON 7.42. FEM a. oe aes tasy ies <0 theee 872
Egg Shell Structure and
PBeriMinOlO PYyng arematg thet Pest ogee a8 872
Methods= sctade res oes. heat ie hia een shies 874
Curvature AnalySIS.........cccceeeeee neers 874
Shell Thickness and Egg
DimMeNSIONS.6...4.3 caps eee ee eee os 876
TEhiteeS CCtrOnIN Pee siege se 8 bet feet szes 877
Shell Porosity and Gas Conductance
Othe ag se. busst st Tete vee East 877
Fossil Eggshell from the Quaternary of
AUISCEAIA’. 205. dis cp Se Beeler tose ees 878
TMICKNESS:, chtatvran tes oh dooseeenan pee yte'st 878
EEG SIZE even sshcec oe dae ginentedenie seg dateen es 878
Shell:Structure:) Emuscs.cg.0. seoes sete 878
Shell Structure: Mihirung
(GENYOFNIS) 2.2... 0eceeeeeneeceeeteeenees 879

a

Taphonomy of Fossil Eggshell......... 880
Radiocarbon Dating of
Fossil Eggshell ..............:.seeeeee 881
Other Egg Types from the Quaternary
of Astraliav.sv:..2.3 2 fscecee tense. 08 881
Fossil Eggshell from the Tertiary of
AUIS traits. .3) Bo coves de natiie e's ide ooeasiauss 882
Method of AnalySis .............eceeeee eee 882
Eggo (SIZC heaves scestetsetteieeders sampiaionsee 882
Shell Structurc.........c.ceee cee eeece neers 884
Shell Porosity ............ceseeeee eee ee eee eee 885
SUMMALY......2..0.sscecoeecsceneesseeeeeeeeee 886
GONCIUSIONS:.. oo. nscaie spec cee gees es enesaes cates 886
Acknowledgements ............0eeeeeee sence eens 886
References........cccesersesccsecernccsreseeeetenenee 886
BlateSs © tesa deve eepthlabwieneden seta tseetetle weber 888

1 Deceased; formerly of Department of Biogeography and Geomorphology, Research School of Pacific Studies,
Australian National University, Canberra, Australian Capital Territory 2601, Australia.

2. Earth Sciences and Botany/Zoology Departments, Monash University, Clayton, Victoria 3168, Australia.

872 - WILLIAMS & RICH

INTRODUCTION

The study of fossil eggs provides insights into the taxonomy and nesting biology of species
that might otherwise be known only from skeletal remains, or not at all. Fossil eggs are rarely
preserved intact, but shell fragments are relatively common and may be locally abundant.
Eggshell belonging to chelonians, crocodilians, lizards and birds has been identified in the
fossil record, in sediments as old as Mesozoic (Hirsch 1983, Hirsch & Packard 1987).
Dinosaur eggs have aroused special interest, because well-preserved shell, and even whole
clutches of eggs, have been discovered in various parts of the world (Andrews 1926, Dughi &
Sirugue 1966, Kerourio 1981, Horner 1982). Much effort has gone toward identifying egg
types, discovering how dinosaurs nested, and in trying to solve the puzzle of their disappearance
(Erben ef al. 1979, Seymour 1979, Williams et al. 1984). In some cases it has been possible
to analyse organic materials still present in the fossil shell (Voss-Foucart 1968, Kolesnikov &
Sochava 1972).

Cainozoic deposits yield mostly avian eggshell, and most studies have involved the
description and identification of large egg remains, particularly those of ratites (Dughi &
Sirugue 1962, Sauer & Sauer 1978, Dughi & Sirugue 1978). The term "ratite" is used here
only to group the large, flightless birds (most with palaeognathous palates), and no
phylogenetic implications are intended. The following discussion covers some principles of
eggshell identification and analysis, as applied to the Australian arid-zone deposits of Cainozoic
age.

Identification and analysis of fossil eggshell relies on studies of macro- and microstructure
and an understanding of egg physiology. Shell fragments are usually the only material
available to the palacontologist, but they can yield a surprising amount of information. Shell
thickness and curvature measurements allow estimation of egg size and shape, even when the
relationship of fragments to the original egg (or eggs) is lost. Shell microstructure is very
distinctive for the higher vertebrate taxa (Sadov 1970, Hirsch 1983, Hirsch & Packard 1987),
and it is also useful at lower taxonomic levels in birds (Board e¢ al. 1977). This is particularly
so for ratites, including extinct species known from Quaternary deposits (Tyler & Simkiss
1959).

The shell of an egg does more than provide physical protection for the embryo. It also acts
as a barrier to pathogenic organisms, and at the same time it must allow exchange of oxygen
and carbon dioxide between the embryo and the nest environment. Much of the resistance to
gascous diffusion is provided by the eggshell (Wangensteen et al. 1970/71), so the total area of
the pores that pass through the shell must be sufficient for the needs of the hatchling at the end
of incubation. Shell porosity cannot be too great, however, for in most species evaporative
water loss must be limited to about 15% of egg mass (Rahn & Ar 1974). Shell porosity is,
therefore, a compromise between allowing free exchange of respiratory gases and regulating
water loss. In the case of eggs incubated underground, or in nesting mounds of decaying
vegetation, water loss is not a problem, because of high nest humidity, but respiratory gas
exchange is hampered by low oxygen and carbon dioxide levels within the nest. Consequently,
these types of eggs tend to have thinner, more porous shells than eggs which are incubated
above ground (Seymour & Ackerman 1980). The relationships between egg size, shell
thickness and shell porosity are well known for living birds (Ar et al. 1974, Rahn & Ar 1974),
and this serves as a basis for interpreting the physiology of fossil eggs.

EGG SHELL STRUCTURE AND TERMINOLOGY

It is convenicnt to use geographical terms for egg morphology, even though few eggs
approach a spherical shape. Thus, an egg has an equator, two poles, an axis and regions of

AUSTRALIAN FOSSIL EGGS - 873

high and low latitude, Most eggs are ellipsoidal, and the two hemispheres are not symmetrical.
Therefore, additional terms such as egg elongation (length/width ratio) and asymmetry index
have been defined (Hoyt 1976). A considerable amount of work has been done to develop
empirical formulae to estimate egg volume, surface area and weight from egg dimensions
cece 1965, Besch et al. 1968, Preston 1968, 1969, Shott & Preston 1975, Tatum

Basic shell structure and terminology are shown in Fig. 1, together with the characteristics
of some vertebrate shell types. With the exception of chelonian eggshell, which consists of
aragonite (Hirsch 1983), vertebrate eggshell is composed of calcite with organic material
permeating the shell. The calcite in eggshell is organized as columnar crystals with their long
axes perpendicular to the shell surface (Fig. 1). These crystals form initially as small spherical
nuclei (mammillary cores) of radially oriented crystals, attached to the outer membrane
surrounding the egg. This membrane becomes detached from the shell in the later stages of
incubation, when calcium absorption by the embryo erodes the mammillary cores. As shell
growth proceeds, the outward-facing crystals grow longer and wider, eventually coming into
lateral contact to form the rigid sheil structure. The inner, or mammillary, surface of most
eggshell consists of a uniform array of mammillary knobs, which represent the original
nucleation sites for shell growth.

CHELONIAN AVIAN CROCODILIAN
PORE APERTURE
Z

N
i

PORE CANAL ——

"SPONGY LAYER"
/MAMMILLARY LAYER
/

|
me
|
|
\

|) CRYSTAL COLUMN |

MAMMILLARY SURFACE /

MAMMILLARY ~
CORE

Figure 1. Three types of vertebrate eggshell structure, external surface at top. (After Hirsch 1983).

Different shell types vary in the structure of their shell units, and in the degree of lateral
fusion of these units. Chelonian shell units, with simple abutting contact, are composed of
radiating spicular aragonite crystallites, while crocodilian shell units are composed of wedge-
shaped crystals and form a complex, interdigitating contact. Soft-shelled reptilian eggs are
insufficiently calcified for contact to occur between the crystal units, and the "shell" remains
flexible. Avian eggshell has relatively long, parallel-sided crystal units, whose margins appear
to be fused. Their boundarics are visible under a polarising microscope.

874 - WILLIAMS & RICH

Pore canals form between the crystal units as the shell grows. Depending on the species,
their shape varies from simple, straight tubes to complex branching networks. Many kinds of
ratite eggshell have branching pores with multiple apertures (Tyler & Simkiss 1959). Large
eggs, with a shell thickness of 1 mm or more, have typical pore diameters of up to 0.25 mm
(Board & Tullett 1975). In some shells the pores may be filled with organic material, or the
apertures may be capped or partly plugged by wax. It is not clear how such deposits modify
gaseous diffusion through the shell (Board ef al. 1977), but experiments show good agreement
between measured gas conductance and that predicted by pore geometry alone (Rahn & Ar
1974).

METHODS

Some of the diagnostic characters of large egg fragments can be seen with a hand lends, or
with the naked eye. Gross shell structure, colour, surface morphology and taphonomic details
are often fairly obvious. Other observations are best done in the laboratory, such as light and
electron-microscope studies, as well as elemental and amino acid analyses. Shell properties
such as thickness, pore density and curvature vary considerably over the shell surface.
Consequently, it is desirable to examine enough shell fragments to ensure that the sample is
representative. This is particularly so for curvature analysis, which relies for its success on
sampling fragments from all parts of the egg.

CURVATURE ANALYSIS

Egg size can be estimated from the curvature of shell fragments (Williams 1981). Viewed
from either end, eggs have a circular cross-section. In lateral view, egg shapes range from
almost circular to elliptical, depending on the species. Any single shell fragment, therefore,
has two components of curvature: one corresponding to the circular cross-section at that point
(Cw), and another related to its longitudinal curvature (CL) (Fig. 2A). Longitudinal curvature
is perpendicular to, and less than, lateral curvature. Thus, by measuring the two curvature
components, a fragment can be oriented with respect to the original egg, and the egg diameter
at that point can be calculated. At the poles of an egg the two curvatures are equal (but neither
corresponds to a circular curve), Providing that fragments from the widest part of the egg are
sampled, egg width can be estimated.

Shell curvature is measured with a modified Geneva Lens Measure (Sauer 1968) shown in
Fig. 2B. This operates with two fixed probes, spaced to fit the width of most shell fragments
in the sample, and a central moving probe connected to a mechanical dial gauge. A flat surface
in contact with all three probes gives a dial reading of zero, but a circular surface displaces the
measuring probe by an amount related to the diameter of the curve. In practice, the gauge is
calibrated against known circular curves, and a regression is calculated of the form:

Diameter = k (Curvature reading) “1
(where k is a constant)

Practical measurement of egg fragments requires that the shell has a smooth surface. The
inner surface is usually smoother than the external. To take a pair of readings, the shell
fragment is gently pressed against the probes and rotated to find the maximum and minimum
values, A degree of judgement is required due to the effect of surface irregularities. Distorted
fragments must be excluded from measurement.

AUSTRALIAN FOSSIL EGGS - 875

It is convenient to convert paired curvature values to diameter, although it must be noted
that longitudinal "diameter" has no physical reality. The pairs of diameter values (D., Dw) are
plotted (Fig. 3) to form a scatter of points. Polar regions plot close to the line DL, = Dw, and
equatorial regions plot furthest from the origin. Estimation of egg width involves selecting the
largest diameters, allowing for random scatter of points. Using about 95% of the largest value
is suggested.

CL

Egg width Dial gauge

Measuring
probe

Eggshell Gakkai

| |

8mm

Figure 2. A. Egg geometry, showing curvature components C;, Cw and egg width, Dw. B. Construction
of shell curvature gauge.

Egg length and degree of symmetry are inferred indirectly from the plotted points, but at
present there is no quantitative solution. Likely limits for egg elongation are imposed by the
observed range from 1.1 to 1.7 in living birds (Hoyt 1976). For a nearly circular egg, the
bivariate curvature plot would reveal a tight cluster of points (Fig. 3A), whereas an elongated
shape would produce an elongated distribution, modified by the degree of asymmetry between

the egg hemispheres (Figs 3B-C).

876 - WILLIAMS & RICH
SHELL THICKNESS AND EGG DIMENSIONS

Shell thickness measurements should include a representative sample of unabraded
fragments, free of encrusting sediment.

There is a relationship between shell thickness (L in cm) and egg weight (W in g) for living
species of birds, due to the compromise between physical protection for the embryo and its
respiratory requirements. The relationship is:

L = 5.126 x 10-3 (w0-456) (Ar et al. 1974)

Egg weight estimated from this relationship can be checked against egg size estimated from
shell curvature, since:

W = 0.51 x egg density x elongation x (egg width)?
(Hoyt 1979)

The density term is approximately 1.08 g/ cm-> for eggs of about 1 kg. There is only a
slight dependence of density on egg size (Paganelli et al. 1974). Egg elongation is likely to lie
in the range of 1.1 to 1.7 (Hoyt 1976), and a value inferred from the curvature arising from
faulty data or from unusual egg characteristics can be revealed by this check. For example,
mound-nesting birds, such as the Australian megapodes, produce very thin-shelled eggs, which
weigh about 200 g (Seymour & Ackerman 1980). However, the shell thickness (0.3 mm)
would indicate a weight of only 50 g. If fossil fragments were the only available material, a
comparison between weight estimates derived from curvature analysis and shell thickness could
reveal the unusual nature of these eggs.

>
ee)
‘?)

LONGITUDINAL CURVATURE

EGG WIDTH (Dy) EGG WIDTH (Dy) EGG WIDTH (Dw)

Figure 3. Hypothetical curvature distributions for: A, a nearly spherical egg, diameter approximately 100
mm; B, a symmetrical ellipsoidal egg, width approximately 100 mm; C, an asymmetrical, ellipsoidal egg,
width approximately 100 mm.

AUSTRALIAN FOSSIL EGGS - 877

THIN SECTIONING

Thin sections of eggshell are extremely useful in displaying crystal structure, pore

morphology, organic inclusions and in estimating shell porosity. Sections are made both
parallel to the shell surface (tangential sections) and perpendicular to it (radial sections). In
preparing a tangential section, it is possible to select the level within the shell which will be
displayed.
_ Thin sections are made using normal petrological techniques. Shell fragments are
impregnated with plastic (e.g. low viscosity epoxy resin; see Tompa 1980), perhaps using
vacuum impregnation, and a flat surface is ground on the embedded specimen. The specimen is
then mounted on a glass slide, and ground to a thickness of about 50 microns. It is usually not
necessary to grind extremely thin shell sections to gain useful information, but the section
should appear translucent. Grinding can be done by using several grades of carborundum paper
(ranging between 100 and 1200 grit) on a thick sheet of backing glass. Each tangential section
requires one shell fragment, but multiple radial sections can be made by sawing thin slivers
from a number of fragments and mounting them as a stack on a single slide.

SHELL POROSITY AND GAS CONDUCTANCE OF THE EGG

The functional measure of shell porosity is the capacity of the whole egg to exchange gases
by diffusion through the shell pores. For convenience this is expressed as water vapour
conductance for whole eggs. Conductance to other gases, such as carbon dioxide and oxygen,
can be estimated by substituting appropriate values for the diffusion coefficients, which depend
on molecular weight (Rahn & Ar 1974).

Assuming a constant external environment, gas conductance is a function of pore diameter,
pore density, effective pore length and total shell area of the egg. The first three of these are
estimated, in the case of fossil eggshell, by measuring the porosity of tangential thin sections.
Total shell area is related to egg volume by:

Ash = 4.928 x (egg volume)9-668 (Hoyt 1976)

Total pore area, therefore, is:

Ap = mean porearea x poredensity x Agh

Effective pore length is measured from radial thin sections, counting only the portion of
restricted pore diameter, and excluding the funnel-shaped apertures where pores meet the shell

surface.
Water vapour conductance (G20) expressed in mg of water lost per egg per day per torr of

partial pressure difference is given by:
GH20 = 23.4 x Ap x 1! (Aretal. 1974)

The value obtained can be checked against the value expected of a normal bird egg of similar
size:

GH20 = 0.432 x W278 (Aretal. 1974)

878 - WILLIAMS & RICH

If estimated G is much greater than predicted G, for example by a factor of 2 or more, then
underground incubation of the eggs is a possibility (Williams et al. 1984).

FOSSIL EGGSHELL FROM THE QUATERNARY OF
AUSTRALIA

Pleistocene sand dunes in arid Australia are a source of abundant fossil eggshell (PI. 1).
Significant finds have also been made in fluviatile and lacustrine sediments, but such
discoveries are less common.

Experience from the arid regions of central Australia indicates that the most common types
of eggshell to be found in old dune deposits belong to the emu (Dromaius novaehollandiae) and
the extinct mihirung (Genyornis newtoni) (Williams 1981). Identification of these two shell
types is simple, providing that a few well-preserved fragments are available. Other kinds of
shell may also occur, such as waterbird shell in dunes close to former sites of permanent water,
but such fragments obviously come from smaller eggs. In some areas, such as the Lake Eyre
Basin, fluviatile deposits contain a variety of small egg remains, probably belonging to
waterbirds.

Large shell fragments are easily seen as they weather from sand on eroding dune surfaces
(Fig. 4). Many fragments are as large as 20-30 mm across, but despite their size, they are
easily transported by the wind and may form a trail downwind of the source. Fossil shell is
usually buff in colour, although newly broken surfaces are white. Fresh emu eggshell is dark
green, bleaching in the sun to greenish-gray or white. Modern shell is also distinguished from
fossil shell by the dull sound as pieces are moved against each other. Fossil shell usually
tinkles like pieces of fine china.

THICKNESS

Emu and mihirung shell both have similar mean thicknesses of 1.14 0.1 mm (Table 1).
This immediately distinguishes them from other shell types in Australia. In some areas there
is the possibility of finding remains of ostrich eggs, as there are feral populations left from a
period of ostrich farming earlier this century. Ostrich shell is easily identified by its thickness
(about 2 mm) and by the distinctive depressions on the shell surface. Otherwise, the only other
large eggs in the modern Australian environment are those of the cassowary (Casuarius
casuarius), which are similar to emu eggs, but would not be expected to occur in arid
environments.

EGG SIZE

The mean size of emu eggs is about 90 mm x 135 mm. They weigh on average about
0.65 kg. Curvature estimates on mihirung shell indicate an egg size of about 125 x 155 mm
(for Genyornis newtoni). Mihirung eggs (Pl. 2) were probably less elongated than emu eggs,
and weighed approximately 1.0 to 1.3 kg (Williams 1981).

SHELL STRUCTURE: EMU

Emu eggshell has one of the most distinctive structures of any bird (Pl. 3B). The normal
columnar crystalline layer is overlain by two additional layers that together make up about one

AUSTRALIAN FOSSIL EGGS - 879

fourth of the total shell thickness. The middle layer consists of highly porous calcite that
forms a labyrinth of tubules into which the underlying shell pore canals connect. This layer is
about 10% of shell thickness. The remaining 15% of the shell is a superficial layer of
discontinuous calcite nodules. These nodules are dark green in a newly laid egg, and are
responsible for the “orange peel" texture of emu eggs. Although the nodules are very dense and
non-porous, the discontinuous nature of this layer means that the underlying layer is exposed,
providing a path for gaseous diffusion for the embryo.

The three-layered structure of emu shell is clearly visible even to the naked eye. However,
the middle layer is a plane of weakness, so that sand-blasting in a dune environment can strip
the two outer layers, leaving only the columnar layer of the shell. This process is rarely
complete, and remnants of the outer layers usually remain. Confusion with Genyornis shell
is, therefore, unlikely, particularly as eroded emu shell is only about 0.8 mm thick.

The adaptive significance of emu shell structure is not clear. Total shell thickness is
consistent with egg weight using the regression of Ar et al. (1974), so that the outer two layers
cannot be considered to contribute unnecessary material. Perhaps it is one solution to the
problem of isolating the embryo from the nest environment. It is unlikely to be an adaptation
to an arid environment, as the eggs of the cassowary also have the three-layered structure,
although the outermost layer is more sparsely distributed.

ST ___..._ eee

Table 1. Comparison of fossil and modern egg dimensions of Genyornis and Dromaius.
*Estimate from direct measurement; **values derived from estimates. (From Williams 1981).

Shell Egg Egg Elongation Egg
thickness width length weight
Genyornis _1.15+0.12 125mm* 155mm** 1.24** 1.33**
(fossil) mm (N=278) N=219 kg
Dromaius 1.09 + 0.12 90-95mm - - -
(fossil) mm (N=72) N=20

Dromaius 1.09 + 0.09 mm 90.5 + 3.9 mm 133.8 + 6.3 1.48 + 0.04 0.61 + 0.08 kg
(fossil) (N=348 eggs) (N=32 eggs) (N=32eggs) (N=26 eggs)

oo —————————————OOSS— —ON0MNMBMBB

SHELL STRUCTURE: MIHIRUNG (GENYORNIS)

The shell of Genyornis newtoni consists of a simple columnar crystal layer (Pl. 3A).
Shell pores open at the surface in short grooves, which tend to be paralled, and are, in fact,
aligned with the egg axis. Each groove is associated with a single pore, but there may be
several apertures per groove, due to branching of the pores.

There is no evidence that mihirung shell had additional calcite layers. Field identification,
therefore, relies upon measurement of shell thickness and the absence of additional shell layers.
Confusion with emu eggshell is most unlikely, and there appear to have been no other large
dromornithids that survived into the Late Pleistocene (Rich 1979). The youngest Genyornis

880 - WILLIAMS & RICH

is approximately 26,000 yBP (Williams 1981), so the presence of its eggshell in a deposit may
be an indicator of pre-Holocene age.

TAPHONOMY OF FOSSIL EGGSHELL

Shell fragments from dunes tend to have rounded edges due to sand-blasting by the wind. If
wind-abraded fragments are encrusted by pedogenic carbonate, then the fragments were broken
and exposed to weathering prior to burial. Otherwise, sand-blasting could have occurred during
a recent episode of dune deflation. Fragments with sharp edges indicate that other fragments
may be close by on the surface, or that more material lies just below the surface. Partially
complete eggs preserved in sand dunes usually consist of many small fragments still in their
relative positions, and unless considerable care is taken during collection, these fragments
become separated. We caution those who might attempt actual reconstruction of an egg. Even
when it appears that most of the shell is present, vital fragments will be missing, and it often
turns out that less than half of the shell is collectable. Egg reassembly is unlikely to be a
rewarding experience!

The inside surface of eggshcll may reveal the stage of incubation reached by the egg. Ina
newly-laid egg, the mammillary cores are attached to the shell membrane, so that if the egg is
buried and the organic constituents decay, the mammillary surface consists of fairly uniform,
rounded knobs. However, an egg which has been incubated to the stage of skeletal calcification
is likely to display cavities at the apices of the mammillary knobs, the sites of calcium
absorption by the developing embryo. Ina well preserved shell fragment, the two conditions
are usually discernable using a hand lens.

Egg predation by mammals and large reptiles seems to have been a constant drain on the
reproductive output of emus and mihirungs. Partially complete eggs are commonly found in
which sections of shell are folded back on each other, or even nested together. This is likely to
happen when a predator breaks an egg, and sections of the shell remain together, held in place
by the shell membranes. Predator activity is confirmed by the occasional discovery of shell
fragments bearing scratches, gouges and small puncture marks left by the teeth of the predator.
Puncture marks are characterized by the removal of a conical plug of shell from the inner
surface of the egg, forming a hole with a bevelled edge. In some cases, fragments are found
with a pair of puncture marks corresponding to the canine teeth of the predator. In the
Pleistocene of Australia, the most likely egg predators are the thylacine (Thylacinus
cynocephalus) and the devil (Sarcophilus harrisi), and possibly small dasyurids such as
Dasyurus spp. (Williams 1981). Large varanid lizards may also have taken the eggs of emus
and mihirungs. Both Thylacinus and Sarcophilus have sufficient strength, and jaw gapes
large enough, to carry eggs for some distance. Even the large potoroid Propleopus might have
engaged in egg eating. Egg stealing from these large birds must have been a dangerous activity
at some stages of the nesting season, and perhaps predators carricd their booty to sand dunes
away from distant nesting sites.

So far, only the remains of isolated eggs are known from dune deposits. These could have
been transported by predators, and thus it is difficult to assert that mihirungs nested on dunes.
However, it is clear that mihirungs nested in the vicinity of dunes, and, judging by the high
proportion of mihirung shell to emu shell at some sites, emus may even have been the less
common species, or the two birds may have preferred different nesting sites.

RADIOCARBON DATING OF FOSSIL EGGSHELL

Datable material is scarce in Quaternary deposits of arid Australia. In the deserts, vegetation
tends to be insubstantial and is poorly preserved in sedimentary deposits. Where datable plant

AUSTRALIAN FOSSIL EGGS - 881

remains are found, it is not always certain whether they are contemporaneous with the
sediments being dated, or whether they are later intrusions. Mollusc shells are useful for dating
some deposits, usually dunes associated with nearby salt lakes. However, even when mollusc
shells are available in sufficient quantities for dating, the result may be affected by encrusting
pedogenic carbonate, which is difficult to remove.

Fossil eggshell is a useful adjunct to the range of datable materials. Where it occurs in
dunes at all, it is often possible to collect a usable sample (50 g), and the shell of emus and
mihirungs is robust enough to allow mechanical removal of pedogenic carbonate. It is also
easy to select only mihirung shell for dating, thereby eliminating the possibility of
contamination by bleached fragments of modern emu shell. Eggshell also has the advantage
that its stratigraphic origin is usually clear, and it is unlikely to be intrusive.

The suitability of eggshell for radiocarbon dating is becoming better accepted. The
carbonate which forms eggshell is derived from metabolic carbon dioxide in the parent female
(Sturkie & Mueller 1976), and is thus in equilibrium with environmental carbon. Carbonate of
inorganic origin, such as that ingested by birds as gizzard stones, is unlikely to affect the
radiocarbon age of eggshell. Feeding trials with domestic fowl (Long et al. 1983) demonstrated
that little or no correction to shell carbonate dates is necessary, provided that the shell has not
been altered during burial. The degree of alteration may be tested by examination of thin
sections in polarized light. If the regular radial crystal structure is present, then there is no
reason to suspect contamination by extraneous carbonate. If, however, thin sections reveal
non-radial crystals, lacking the columnar structure of normal eggshell, or if there are opaque
patches, discoloured haloes around pore canals or signs of corrosion, then contamination should
be assumed. Eggshell from sand dunes that we have examined tends to be very well preserved,
although there is often adhering pedogenic carbonate. When preparing a shell sample for
dating, each fragment should be examined closely, and superficial carbonate should be removed
mechanically. Acid pretreatment is less satisfactory, because it penetrates the shell and
dissolves part of the sample.

In some cases, the likelihood of contamination can be checked if the sediment contains a
few aquatic mollusc shells. These mollusc shells form as aragonite, and this changes to calcite
during recrystallisation. X-ray diffraction measurement will determine the ratio of aragonite to
calcite, and thereby indicate the probability of recrystallisation in the eggshell.

OTHER EGG TYPES FROM THE QUATERNARY OF
AUSTRALIA

Eggs of other species of ratites are known from Quaternary and late Tertiary (see below)
sites in Australia. Perhaps the most tantalizing specimen of these is the Scott River egg from
southeastern Western Australia (Rich 1979), an almost complete egg of very large size (276
mm long). The surface of this egg is rough and nodular, like that of the emu, but much of this
texture could be due to post-depositional alteration and sand-blasting. Speculation that this is
the egg of an Elephant Bird (Aepyornis sp.), Somehow transported intact from Madagascar is
difficult to accept because Aepyornis shell is smooth (Tyler & Simkiss 1959). The
possibility should be considered that the Scott River egg belongs to an extinct species of
dromaiid, so far unknown from skeletal remains. However, until this specimen is further
studied in detail, no decision can be made concerning its affinities.

Dunes in western New South Wales provide specimens which suggest the former existence
of another species of dromaiid. A few weathered fragments of emu-like shell have been

882 - WILLIAMS & RICH

collected from dunes at Lake Menindee and Lake Mungo. This shell differs from that of the
emu in that the two outer layers are thinner than those of emu shell, and the outermost layer is
more uniform and finely textured than the nodular layer in Dromaius novaehollandiae (Pl. 4).
More specimens are needed to determine whether these differences are significant. However,
even in early historical times the emus were a more diverse group than the single surviving
species. Islands in Bass Strait and Kangaroo Island supported morphs which were smaller than
the mainland species (Rich & van Tets 1982, Parker 1984, Patterson & Rich 1987). It would
not be surprising to find evidence of other Pleistocene species of Dromaius.

FOSSIL EGGSHELL FROM THE TERTIARY OF AUSTRALIA

Perhaps the most interesting eggshell fragments from the Tertiary of Australia, and the only
ones really studied in any detail, come from Snake Dam, a site on the Clayton River to the
southeast of Lake Eyre (Williams & Rich in press, Rich 1979). One fragment (Pl. 2) of the
pair has been analysed to determine the size of the original egg, and to identify its taxonomic
origin. The shell fragment is extremely thick, comparable to that of the extinct elephant bird
(Aepyornis) of Madagascar. Elephant bird eggs were as large as 35 cm in length (Berger et al.
1975). The Snake Dam egg was probably even larger. The shell structure of the Snake Dam
egg is unique in our experience, matched in its unusual structure only by the triple-layered shell
of the emu.

METHOD OF ANALYSIS

Analysis on the Snake Dam fragment was carried out as described above and by Williams
et al. (1984). Thickness was measured with vernier calipers, taking the mean of 13 readings.
Curvature measurements were made at several points on the shell fragment, on the inner
surface, which was smooth. The curvature gauge was as described above and in Williams
(1981).

A single tangential thin section from about the mid-level of the shell was scored for
porosity (Williams et al. 1984). Shell area was estimated by optical enlargement of the thin
section, using a planimeter. Pore transverse sections were measured on 38 pores, using a
calibrated eyepiece fitted to a petrological microscope. Water vapour conductance of the egg
was estimated using data from Ar et al. (1974). Chemical analysis and X-ray diffraction
analysis was performed on a 96 mg sample of the shell.

EGG SIZE

Curvature readings gave values of egg width as large as 118 mm, including shell thickness.
Longitudinal curvature was too small to be measured with the curvature gauge, but the contrast
between the two curvature components shows that the fragment does not originate close to the
poles of the egg. In the light of other data, the fragment is not from the equatorial region, so
the value of egg width determinated from this approach is a minimum.

A mean value of 4.07 + 0.05 mm was obtained from 13 readings. Using the regression of
Ar et al. (1974) based on shell thickness (W = 1.05 x 10® shell thickness 2-19 ), the Snake
Dam egg would have weighed 14.7 kg when laid. This would be one of the largest bird eggs
known. Ar et al. (1974) used total shell thickness for their regression, which in the case of
most shell types is equivalent to functional pore length (Pl. 4). However, the Snake Dam

AUSTRALIAN FOSSIL EGGS - 883

shell has a rugose surface, and the pores are effectively only 94% of measured shell thickness.
If shell thickness is adjusted to 94% of 4.07 mm, a value of 3.8 mm is obtained, yielding an
estimate of 12.6 kg for egg weight (Tables 2,3).

} Fa
TATA

EAC Tc}

Figure 4. Drawings of radial sections of (top) possible new type of dromaiid eggshell from Lake Menindee
in westem New South Wales, compared with shell of the living Dromaius novaehollandiae (bottom). See Fig.

1 for abbreviations.

Using the estimate 12.6-14.7 kg for egg weight, and assuming a possible range of 1.1 - 1.7
for egg elongation (Hoyt 1976), then the dimensions of the Snake Dam egg can be calculated:
a) egg elongation = 1.1, then egg weight of 12.6 kg indicates an egg with dimensions of 27.3
cm x 30.1 cm; egg weight of 14.7 kg indicates an egg 28.8 cm x 31.7 cm; b) egg elongation
= 1.7, then egg weight of 12.6 kg indicates an egg 23.6 cm x 40.2 cm; egg weight of 14.7 kg
indicates an egg 24.9 cm x 42.3 cm. This gives a possible size range of 24-29 cm x 42-32 cm

for the Snake Dam egg.

884 - WILLIAMS & RICH
SHELL STRUCTURE

The outer surface of the Snake Dam fragment is very unusual for bird eggshell in that it
consists of fine ridgelets lying close together, separated by deep fissures. Most bird eggshell
has a relatively smooth surface, marked only by tiny pits where the pores communicate to the
surface. Curvature analysis shows that the ridgelets on the Snake Dam shell are aligned with
the egg axis, which is typical of many egg types bearing elongate surface features. Thin
sections show that the shell pores connect with the fissures near the outer shell surface (PI. 4).
Fissuring extends about one-third of the way through the shell, possibly affecting the strength
of the egg and the ease with which the chick could have escaped upon hatching.

Tangential sections through the shell reveal that the crystal columns, the basic units that
make up the shell, are in close contact with interdigitating boundaries (Fig. 4). Both radial and
tangential sections, viewed with crossed polars, show that the crystalline structure of the shell
is intact.

Table 2. Characteristics of the Snake Dam fossil shell fragment, cf. Dromornis (UCR 17877).

Shell area examined, mm? 59.6
Pore density, per mm? 1.7
Mean pore area, mm? x 102 3.2 +2.7/-1.9
Mean pore size, major axis 79.6 + 30.2
Mean pore size, minor axis 46.4 + 14.5
Major axis/minor axis 1.7
% pore area 0.54

Radial sections reveal prominent "growth lines" parallel to the shell surface. These lines
are convex outward near the inner shell surface, where they conform to the concentric structure
of the mammillary cores. Inclusions, such as these "growth lines", are possibly the remains of
organic material incorporated during shell growth. The mammillary cores are eroded in
appearance, suggesting that the Snake Dam egg had been incubated at least to the point of
skeletal calcification of the chick (Erben et al. (1979). Pores are rarely solitary, tending to
branch into pairs or triplets, and branching occurs parallel to the egg axis.

In some cases blocked pores have been observed (Fig. 4). Secondary calcite and fine detrital
sediment were not seen, and there is no sign of alteration due to fossilization. Chemical
analysis of a small sample of shell revealed that it consists of low-magnesium calcite with a
trace of silica. The composition is (Cap.99Mgo,.11) CO3, equivalent to 39.02% calcium by
weight and 0.27% magnesium. It is noted that the host sediment, thought to be Etadunna
Formation (Miocene or Late Oligocene), is likely to be high-Mg dolomitic claystone,
suggesting that little chemical exchange has occurred between shell and sediment.

_ The structure of the Snake Dam shell, although unusual for bird eggshell, is comparable in
its uniqueness with the multi-layered shell of the emu. The form of the crystal columns, and

AUSTRALIAN FOSSIL EGGS - 885

their intimate contact with each other, combined with the highly-organized branching pore
structure are entirely consistent with bird eggshell, but not typical of reptile shell.

SHELL POROSITY

Porosity was measured on one 60 mm2 tangential section from about the mid-level of the
shell. This was a compromise, because the number and size of pores varies at different levels
within the shell. The data obtained are, therefore, an approximation of the true value for the
egg.

Water vapour conductance is a measure of the potential gas exchange of the whole egg, and
relates to the requirements for respiratory gas exchange between embryo and the nest
environment, as discussed above. Results obtained from a single shell fragment are only an
approximation to the actual value, but the result obtained from porosity data may be checked
against regressions based on egg weight, measured for living bird species (Ar ef al. 1974).

Using estimated egg weight of 14.7 kg and egg density of 1.10 g/ cc (Paganelli e¢ al.
1974), egg volume can be calculated. From this, egg surface area can be estimated (Table 3).
Applying the relationship GyH20 = 23.42 x Ap X L-! (Ar eral. 1974), water vapour conductance
would be 917 mg/torr/day. Using only the estimated egg weight in the relationship, GH20 =
0.432 W°-78, a value of 769 mg/torr/day is obtained, which is in good agreement with the first
estimate. If egg weight was closer to the lower estimate (12.6 kg), then the corresponding
values would be 826 and 683 mg/torr/day, respectively. Thus, the estimated range of egg size,
12.6-14.7 kg, is considered to be reasonably close to the actual fresh weight of the egg.

owe

Table 3. Characteristics of the Snake Dam eggshell fragment (UCR 17877) derived from shell
properties. *Estimate based on shell porosity, pore length; **prediction based on egg weight,
using fresh bird eggs.

Egg width, cm 24 - 29
Egg length, cm 42 - 32
Egg volume, cm? 11470 - 13400
Egg weight, g 2600 - 14700
Shell thickness, cm 0.407 + 0.005
Functional pore length (L), in cm 0.038
Egg surface area, cm? 25002780
Yopore area 0.535
Total pore area (A,), cm? 13.4 - 14.9
*Estimated Gyy29, mg/torr/day 826 - 917
**Predicted avian Gyo, mg/torr/day 683-769

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886 - WILLIAMS & RICH
SUMMARY

The Snake Dam shell fragment is the remains of an extremely large egg, with an estimated
weight of 12-15 kg. Its shell structure, while unusual for bird eggshell, is clearly bird-like.
Physiologically, the egg was probably not unusual, its porosity being consistent with above-
ground incubation. The shell structure is unlike that of any other bird, and there are no
obvious similarities with eggshell of any existing large ground birds in Australia, or for that
matter, anywhere else. It, likewise, does not resemble the eggshell of the only other large
ground bird known from eggshell in the Australian, Genyornis newtoni. Consequently no
phylogenetic conclusion can be reached, but the attribution to Dromornis sp., the largest
known fossil bird of the late Cainozoic of Australia (Rich 1979) is likely correct.

CONCLUSIONS

Fossil eggshells are known from numerous localities in the Quaternary and Tertiary of
Australia. Thus far, the only ones that have received detailed attention have been those of the
large ground birds, the mihirungs and emus, entirely through the work of D. L. G. Williams.
Emu eggshells are unique in possessing three distinct layers, and although they are of much the
same thickness, can easily be distinguished in the field from those of the mihirung Genyornis
whose shell possesses a single layer only. The eggshell fragments of cf. Dromornis from the
Tertiary of northern South Australia possesses a unique structure for birds and shows no clear
alliance with any group of birds. There is much scope for further work on fossil eggshell, not
only for identification of the taxa involved but also for providing insights into nesting
behaviour and physiological adaptations of birds in the Australian Cainozoic.

ACKNOWLEDGEMENTS

The majority of the work in this paper was carried out by D. L. G. Williams, as a
continuation of the work began as his Ph.D. which he received from Flinders University under
the guidence of Dr R. T. Wells. Dom died far too early in his career, and his loss, especially in
the areas of Pleistocene palaeomammalogy and avian oology, is great indeed. William's work
and Rich's has been supported generously by the Australian Research Grants Scheme (now the
Australian Research Council), and by funds from Flinders University, the Australian National
University, and Monash University. Draga Gelt provided much of the drafting expertise, and
Steve Morton and Ian Stewart provided photographic and electronmicrographic assistance. Dr
M. O. Woodburne and Dr. T. H. Rich kindly allowed us to study the only two specimens
known of cf. Dromornis ; Neville Pledge (South Australian Museum) and Ron Scarlett
(Canterbury Museum) provided us with comparative material. Thanks to G. Miller (University
of Colorado) for his review of this article, and especial thanks to Karl Hirsch for his useful
comments and guidance on several drafts of this manuscript and throughout the study.

REFERENCES

ANDREWS, R.C., 1926. On the Trail of Ancient Man. Putnam, New York and London.

AR, A., PAGANELLI, C.V., REEVES, R.B., GREENE, D.G. & RAHN, H., 1974. The avian egg: water vapor
conductance, shell thickness, and functional pore area. Condor 76: 153-158.

BERGER, R., DUCOTE, K., ROBINSON, K. & WALTER, H., 1975. Radiocarbon date for the largest extinct
bird. Nature 258: 709.

BESCH, E.L., SLUKA, S.J. & SMITH, A.H., 1968. Determination of surface area using profile recordings.
Poult. Sci. 47: 82-85.

AUSTRALIAN FOSSIL EGGS - 887

BOARD, R.G. & TULLETT, S.G., 1975. The pore arrangement in the emu (Dromaius novaehollandiae)
eggshell as shown by plastic models. J. Microscopy 103: 281-284.

BOARD, R.G., TULLETT, S.G. & PERROTT, H.R., 1977. An arbitrary classification of the pore systems in
avian eggshells. J. Zool, 182: 251-265.

DUGHI, R. & SIRUGUE, F., 1962. Distribution verticale de oeufs d'oiseaux fossiles de I'Eocene de Basse-
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cab & SIRUGUE, F., 1966. Sur la fossilisation de oeufs de dinosaures. C. R. Acad. Sci. 262: 2330-

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ERBEN, H.K., HOEFFS, J. & WEDEPOHL, K.H., 1979. Paleobiological and isotopic studies of eggshells
from declining dinosaur species. Paleobiol. 5: 380-414.

HIRSCH, K.F., 1983. Contemporary and fossil chelonian eggshells. Copeia 1983: 382-397.

HIRSCH, K.F. & PACKARD, M.J., 1987. Review of fossil eggs and their structure. Scanning Microscopy
1(1): 383-400.

HORNER, ILR., 1982. Evidence of colonial nesting and "site fidelity” among omithischian dinosaurs. Nature
297: 675-676.

meet hh 1976. The effect of shape on the surface-volume relationships of birds' eggs. Condor 78: 343-

HOYT, D.F., 1979. Practical methods of estimating volume and fresh weight of birds eggs. Auk 96: 73-77.

KEROURIO, P., 1981. Nouvelles observations sur le mode de nidification et de ponte chez les dinosauriens
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LONG, A., HENDERSHOTT, R.B. & MARTIN, P.S., 1983. Radiocarbon dating of fossil eggshell.
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PAGANELLI, C.V., OLSZOWKA, A. & AR, A., 1974. The avian egg: surface area, volume, and density.
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WANGENSTEEN, O.D., WILSON, D. & RAHN, H., 1970/71. Diffusion of gases across the shell of the hen's
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WILLIAMS, D.L.G. & RICH, P.V. (in press). Giant fossil egg fragment from the Tertiary of Australia.
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PLATES

Plate 1. Fossil Genyornis egg weathering from a dune near Port Augusta, South Australia. Although this
egg appeared remarkably complete, attempts to reassemble it were unsuccessful.

Plate 2. Fragment of a very large egg ( UCR [Univ. Calif. Riverside] 17877, loc. RV-7237) of cf.
Dromornis , from Snake Dam, Clayton River, South Australia; Miocene in age. A, A', stereo pair of

external aspect of shell; B, iintemal aspect of shell; least width measurement (left margin in illustrations A,
A’), 24.9 mm.

Plate 3. Broken edges of eggshell, A, Genyornis newtoni; B, Dromaius novaehollandiae illustrating the
internal structure of the shell. CL, columnar/continuous layers; ML, mammillary layer; P, pore; PL, porous
(middle) layer; and SL, superficial layer.

Plate 4. Thin sections of the Snake Dam eggshell fragment (UCR 17877): A, radial section, perpendicular
to egg axis. Pores visible as deep fissures. Crystal columns appear narrow in this plane, but are elongated
when viewed parallel to egg axis; B, tangential section about one third way from inner shell surface; pores
occur in linear arrays approximately parallel to the egg axis; C, enlarged view of pores with calcite crystals
partially infilling some. Shell thickness, 4 mm; mean pore size, 80 microns x 46 microns.

AUSTRALIAN FOSSIL EGGS - 889

PLATE 1

PLATE 2

A A\

890 - WILLIAMS & RICH PLATE 3

AUSTRALIAN FOSSIL EGGS - 891

PLATE 4

892 - WILLIAMS & RICH

ect Satan. gt

A baby Diprotodon optatum, bogged in a swamp at Lancefield, Victoria 26,000 years ago, is
watched helplessly by its mother. During episodes of drought conditions thousands of Grey
Kangaroos also died and were buried in this same swamp when it was one of the few remaining
sources of water in the district. Diprotodon spp. were the last members of a diverse group of
four-footed herbivorous marsupials, the diprotodontids, that were common in Australia during the

Late Cainozoic. They had evolved by the Late Oligocene and persisted until at least 19,000
years ago.

CHAPTER 23

MONOTREMES,
PLACENTALS, AND

MARSUPIALS:

THEIR

RECORD IN AUSTRALIA
AND ITS BIASES

Thomas H. Rich!

INMOMUCHOM) 05.8 cadets Wide chi hs va0e eye one’ 894
What are Mammals, When and Where

did they Develop?............ccceeeeeeeeeee 895
Dental Nomenclature and Dental Formulae

of Therian Mammals..................0... 898
A Tale of Marsupials and Placentals ....... 902

Chronological Framework of Australian
Terrestrial Fossil Mammal Sites....... 914

The Record of Mammals in Australia... 922
Order: Monotremata..............cccccee 922
Supercohort: Marsupialia............00.. 924
Order: Dasyuromorphia.................. 925
Order: Peramelemorpha.................. 929
Order: Notoryctemorphia................ 931
Order: Diprotodontia................066005 932

Suborder: Vombatiformes............ 932
Infraorder: Phascolarctomorphia.. 932
Infraorder: Vombatomorphia ...... 935
Infraorder: Phalangerida............. 944

Superfamily: Phalangeroidea.... 944
Superfamily: Macropodoidea.... 946
Superfamily: Burramyoidea...... 953

Superfamily: Phalangerida

INCETLAE SCCIS....ccceeseececeeeeeess 957
Order: Yalkaparidontia.................... 957
Supercohort: Placentalia................... 958
Order: Chiroptera ...............ccceceeeees 958
Order: Rodentia...........cccccceeeecceeeeees 959
Order: Primates............ccecccesseceeeeeee 961
Order: Carnivora ...........ccceceeceeece ees 963
Family: Canidae.................. 963

History of the Terrestrial Mammals of
AAISHA A oo DOU ces cated tevessg hts 963
Introduced Mammnal..................c002ceee eee 963
Acknowledgemenlts..........cccccceseeeeeeeeeees 963
IRCICTENCES-. stub Sos roland cates hiwehsive bee dveeses 972

Appendix I: Australian Mesozoic and
Tertiary Terrestrial Mammal
LG CANES) ces est cage gsabiiegs'ghanteniiee 1005
Appendix II: Literature References to the
Fossil Terrestrial Mammals of
PAUSU AIA os a nseaid an ote Seka’ tena dat 1057

1 Museum of Victotria, Melbourne, Victoria 3000, Australia.

894 - RICH

INTRODUCTION

During the Cainozoic, at least, Australia's terrestrial mammalian fauna evolved in virtually
complete isolation from its nearest relatives on other continents. This is quite in contrast to
the contemporaneous events in North America, Asia and Europe, where interchange between
the continents was almost a continuous process. Africa was somewhat isolated during the
Cainozoic, but faunal interchange was frequent enough to be a major factor in the evolution of
the terrestrial mammals there. Only South America approaches Australia in the degree of
isolation of its mammalian fauna from that of the rest of the world during the Cainozoic, and
even there, it was not as pronounced. Terrestrial placental mammals were always an important
part of the South American fauna during the Cainozoic, and in the middle of that era, two
additional placental orders entered the continent and radiated widely. In the late Cainozoic, there
was further widespread interchange between South and North America with the establishment of
the Panamanian landbridge.

In Australia, there was little or no interchange after the marsupials evolved in the
Cretaceous. With the exception of bats, the Australian terrestrial mammalian fauna was solely
made up of monotremes and marsupials until the appearance of rodents in the Pliocene. It has
been asserted that the entire Australian marsupial fauna could have descended from a single
species that entered the continent in the late Mesozoic or early Cainozoic. More than a single
species might have reached Australia, presumably across Antarctica from South America at that
time, but if more than one did so, certainly there is no evidence that the number was sig-
nificantly greater than that. The monotremes appear to be the sole survivors of a radiation of
mammals that began no later than the Late Jurassic. As the oldest undoubted marsupials
anywhere are Late Cretaceous and an Early Cretaceous monotreme is known from Lightning
Ridge, it is quite likely that monotremes were in Australia long before marsupials.

This history implies that in a sense, Australia's mammalian fauna represents the product of
an almost independent, simultancous experiment in the evolution of that class. Revelation of
this history could provide insights into mammalian evolution that are otherwise unobtainable.
Just as astronomers might learn much about the formation and history of planetary systems if
it were possible to make detailed observations on a second one, the opportunity to observe an
independent history of mammals provides a possible pathway to isolate factors related to
accidents of history from those inherent in the nature of this group that determined how its
evolution proceeded. Unfortunately, with the exception of two specimens from the Early
Cretaceous and a small collection from near the Palaeocene/Eocene boundary yet to be
described, the record of terrestrial mammals from Australia does not begin until the latest
Oligocene, by which time 88 percent of mammalian history was over.

What is known of the history of Australia’s mammalian fauna has been discussed in the
literature on a number of previous occasions. Among these reviews are Anderson (1933),
Abbie (1941), Ride (1964), Keast (1968, 1972), Archer & Bartholomai (1978), and Archer
(198 1a, 1984b). Although his hypotheses concerning the evolution of Australian mammals
were based almost exclusively on recent material, the classic work of Bensley (1903) cannot be
ignored by any serious student of the topic. To summarize the previous work on Australian
fossil mammals, a list of references pertaining to each terrestrial mammalian genus with a
fossil record in either Australia or New Guinea is given in the second appendix to this chapter.

Aplin & Archer (1987) list and evaluate eight principal classifications of marsupials that
have been proposed beginning with Simpson (1945) and then give a ninth of their own. A
further classification has recently been gencrated (Marshall, Case & Woodburne 1989). On first
inspection, these classifications appear to differ greatly from one another. Many of the
differences, however, are not owing to fundamental disagreements about the relationships of

AUSTRALIA'S MAMMALIAN RECORD - 895

Figure 1. Comparison of the medial sides of the jaws of A, a lizard Varanus; B, a mammal-like reptile
Cynognathus; and C, a marsupial, Thylacinus, x 3/5. Note the reduction of the number of bones in the jaw
to a single element in the typical mammalian condition of Thylacinus as compared with those of the two
reptiles. (A, after Romer 1956; B, after Broili and Schréder 1934; C, after Flower 1885).

major groups but to a ripple effect caused by adherence to the logic of cladistic classifications
that is inevitable when workers differ in the recognition of even a single dichotomy.

Many new higher categories of marsupials from subfamilies to orders, have been proposed
during the past decade. Frequently these of necessity have been based solely on dental remains.
One is struck by how often these new taxa can be characterised as intermediate between two
higher groups in fundamental characters; e.g. the bulungamayines are potoroids on the basis of
jaw and lower incisor morphology with typically macropodid molars (Flannery, Archer &
Plane 1983, 1984, but see Woodburne 1984). Thus, the inclusion of the new taxa necessarily
into only one of the higher groups is quite likely to be subject to revision as additional
specimens are found. For this reason, as well as the different viewpoints of the various
workers involved, marsupial classifications are likely to remain in a state of flux for the
foreseeable future, particularly owing to the ripple effect.

For the purposes of this chapter, the classification of Aplin & Archer (1987) will be
followed. Only with a perspective from a few decades hence will this choice be seen as
perspicacious or not. However, just as the shape of the bottle has little to do with the taste of
the wine, the choice of a particular current classification utilised is not vital to understanding
the fundamental aspects of the groups involved nor their gencral relationships to one another.

WHAT ARE MAMMALS, WHEN AND WHERE DID THEY
DEVELOP?

Mammals derive their name from the mammae (Latin, breasts, teats) with which they
suckle their young. They are also characterised by the presence of hair, endothermy or warm-
bloodedness, and the nature of the articulation between the jaw and skull. It is only this latter
feature which is capable of preservation in the fossil record.

In the reptilian ancestors of mammals, the articular bone of the jaw contacted the quadrate of
the skull (see Figs 1, 2A). By the Late Triassic, 200 myBP, the earliest mammals had
appeared. In these animals, the reptilian pattern of jaw-skull contact was maintained, but, in
addition, the dentary of the jaw articulated with the squamosal of the skull. In more advanced
mammals, the bones which had formed the reptilian jaw-skull articulation became restricted to

896 - RICH

\ incus
| (=quadrate)
articular layingane
quadrate (=angular)
¢ stapes
|
malleus
retroarticular (= articular)
gon tympanum = Process of
feuedied articular i
ih ie manubriu
weenie’ [| (= retroarticular
process)

Figure 2. Comparison of the position of the articular (= malleus), quadrate (= incus) and angular (=
tympanic) in A, the mammal-like reptile Thrinaxodon; and B, the marsupial Didelphis. (After Crompton &
Jenkins 1979).

functioning as two of the three ossicles that linked the ear drum to the inner ear; the articular
became the malleus; the quadrate became the incus (see Fig. 2). Although it played no direct
part in the jaw-skull articulation of reptiles, the angular is another element of the reptilian jaw
that became incorporated into the structure of the mammalian ear. There it became the
tympanic ring, which supports the ear drum. With the quadrate and articular removed from the
jaw-skull articulation, the dentary-squamosal articulation provided all contact between the jaw
and skull. Understanding of this transition between reptiles and mammals in the function of
these two bones did not have to await the discovery of the Late Triassic and Jurassic specimens
which display the double articulation in the adults. Study of the embryological development of
mammals demonstrated more than a century ago that Meckels cartilage which develops along
the inside of the jaw of the embryo in the same position as the articular of an adult reptile,
became the malleus of an adult mammal.

In addition to the three major mammalian groups extant at present, the monotremes,
marsupials, and placentals, there were three other groups that appeared in the Mesozoic and
were all extinct by the end of the Oligocene.

The first of these were the triconodontans. As suggested by their name, their tecth have
three major cusps, which are arranged in an anteroposteriorly directed line (see Fig. 3), It is
generally held that the other mammalian groups evolved from ancestors with a dental pattern
similar to that of primitive triconodontans. A long-lived group, they persisted from the Late

AUSTRALIA'S MAMMALIAN RECORD - 897

Figure 3. Intemal view: A, of the jaw; and B, an isolated molar of Priacodon, a Late Jurassic triconodont.
(After Simpson 1929).

Triassic to the Late Cretaceous in both the Southern and Northern hemispheres but are as yet
unknown in Australia.

Figure 4. Occlusal view: A, of an isolated right upper molar, x 13; and B, of an isolated right lower
molar, x 16, of Docodon, a Late Jurassic docodont (After Simpson 1929).

The second group were the Docodonta, which are characterised by a unique pattern of
quadrate molars adapted for puncturing and slicing (see Fig. 4). Although only 4 genera have
been recognised, the Docodonta are rather widely represented in Middle and Late Jurassic
deposits of the Northern Hemisphere.

By contrast, there were at least forty-eight genera of the third group, the Multituberculata.
They appear in the Late Triassic, if the enigmatic haramyids are included, or the Late Jurassic
and persisted until the Early Oligocene. With enlarged lower incisors and molars suitable for
grinding, these animals were presumably rodent-like in their habits (see Fig. 5). This
similarity and the time of their demise coming soon after the appearance of the rodents in the
Late Palaeocene, has led to the obvious suggestion that the multituberculates were displaced by
rodents. Although they lasted for a long time and were numerous and widespread in the
Northern Hemisphere, there is only a single, tentative record in the Southern Hemisphere,
Ferugliotherium windhauseni from the Late Cretaceous of Argentina (Bonaparte 1987).

Rowe (1988) provides a fascinating discussion of what a mammal is, "...the most recent
common ancestor of extant Monotremata and Theria...," and analyses 158 osteological and
dental characters which diagnose the Class. By his analysis, the earliest true mammal is
Phascolotherium bucklandi, a triconodontan from the Middle Jurassic of England.

898 - RICH

¥

\\li\

Figure 5. A, largely hypothetical restoration of a ptilodontid multituberculate skull; B, right upper
dentition of the Late Cretaceous multituberculate Meniscoessus robustus, x 4.75. (A, after Clemens & Kielan-
Jaworowska 1979; B, after Clemens 1963).

DENTAL NOMENCLATURE AND DENTAL FORMULAE OF
THERIAN MAMMALS

The Theria are characterised by the shape of their molars. In the primitive Theria including
the monotremes, the molars have a triangular shape in occlusal view so that those in the upper
jaw interlock with those in the lower (see Fig. 6). This is quite distinct from the pattern seen
in triconodonts and although docodonts mimic this pattern, theirs developed from a marshalling
of independently evolved cusps (Butler 1986). Although more advanced therians have dental
patterns quite unlike reversed interlocking triangles, all evolved from ancestors that did.

Figure 6. Diagram showing the occlusal relationships between upper and lower molars of a primitive
member of the Theria, Kuhneotherium praecursoris. Areas on upper and lower molars with the same number
occlude against one another. (After Bown & Kraus, 1979).

Cheek teeth of therian mammals are complicated structures, and a detailed system of naming
the cusps and crests evolved. The effort has been motivated by the fact that because teeth are
both complicated and durable structures, they are often the only part of the skeleton of a
mammal available for analysis. A major step in the evolution of this nomenclatural system
was provided by Osborn (1907). A guide into the more recent literature can be found in Bown
& Kraus (1979). Figs 7-8 give terminology applied to molars of two major marsupial groups.

AUSTRALIA'S MAMMALIAN RECORD - 899

StE

mec.
Ant —>
me. Ling

P.prcer. - |

pr.

B
mecd
*
end \ med

Figure 7. A, upper cheektooth of Sminthopsis (Dasyuridae): a.c., anterior cingulum (or precingulum);
a.prcr., preprotocrista; me., metacone; mec., metacrista; pa., paracone; pac., paracrista; p.prcr., post-

protocrista; pr., protocone; st A, position of stylar cusp A; st B-E, stylar cusps B, C, D, and E. B, lower
cheektooth of Sminthopsis (Dasyuridae): a.c., anterior cingulum (or precingulid); c.o., cristid oblique; end,

entoconid; hycd, hypocristid; hyd, hypoconid; hyld, hypoconulid; med, metaconid; mecd, metacristid,; pacd,
paracristid; pad, paraconid; pastd, parastylid; p.c., posterior cingulum (or postcingulid); prd, protoconid.
(After Archer 1981b).

900 - RICH

me.bucc.basin pa.bucc.basin
posparacr.

A premetacr. pas.
postmetacr. Bec
me. t p.-l.butt pa.
_-L.butt.me. pel.
f mel. ant.cing.
post.cing: \y pr.
posthypocr. iy preprotocr.
Prey pert: postprotocr.
preentocd postmetacd
med
B end parametcd

postentocd
posthypocd

pad
preprotocd
prd

prd
protostylid

Figure 8. Schematic presentation of cusp and crest nomenclature of generalised koala (Phascolarctidae)
molars. A, right M4: B, right Mg; C, right M2. Abbreviations: ant. cing., anterior cingulum; c.o.,
cristid obliqua; end., entoconid; hyp., hypocone; hyd., hypoconid; me., metacone; me. bucc. basin, metacone
buccal basin; med., metaconid; mesd., metastylid; mc/., metaconule; pa., paracone; pa. bucc. basin, paracone
buccal basin; pad., paraconid; parametacd, para-metacristid; pas., parastylare comer of tooth; pcl.,
protoconule; p.-l. butt. me., postero-lingual buttress of metacone; p.-l. butt. pa., postero-lingual buttress of
paracone, post. cing., posterior cingulum; postentocd., postentocristid; posthypocd., posthypocristid;
posthypocr., posthypocrista; postmetacd., postmetacristid; postmetacr.; postmetacrista; postparacr.,
postparacrista; postprotocd., postprotocristid; postprotocr., postprotocrista; pr., protocone; prd., protoconid;
prehypocr., prehypocrista; premetacr., premetacrista; preentocd., preentocristid; preparacr., preparacrista;
preprotocd., preprotocristid; preprotocr., preprotocrista; protostylid, protostylid. (After Archer 1978b).
Tedford & Woodburne (1987) conclude, based on the analysis of upper molars of Ilaria, Madakoala, and
Miralina, that the metaconule as used above has no counterpart in the Placentalia and designate a new name,
neometaconule, for it. Likewise, they regard the hypocone as used above as homologous with the metaconule
of the Placentalia and therefore the proper name for that cusp.

AUSTRALIA'S MAMMALIAN RECORD - 901

19/4 C!/; P3/3 M4/q is the basic adult dental formula of the marsupials and 13/3 C!/, P4/4
M3/3 is that of placental mammals. Most late Mesozoic and Cainozoic members of the two
groups have the respective formula or have reduced the number of teeth from them. Where
increases have occurred, the postcanine teeth are all alike in most instances; i.e. homodont as
in porpoises, armadillos and honey possums. Where there are more teeth than the usual
formula and the difference in tooth pattern between premolars and molars is maintained, then
only one or two cheekteeth (= molars + premolars) are added beyond the usual seven.

Individual teeth are identified by a numerical system, where the numbering begins with 1 at
the most anterior of a given tooth type and proceeds posterior: i.e. I! designates the most
anterior upper incisor; I), the most anterior lower incisor. With one important exception, there
is general agreement about the system for designating the individual teeth of Australasian
mammals. That exception is the cheektecth of marsupials. Most workers have adopted the
following system, Pl P2 P3 DP3 M1 M2 M3 M4, where DP3 is the deciduous premolar
which is displaced by the M1, the permanent first molar. An alternative followed by many
workers is to regard these same teeth as P1 P2 P4 dP4 M1 M2 M3 M4, where the dP4 is the
deciduous premolar which is displaced by the M1. The rationale of substituting P4 for P3 and
dP4 for DP3 is the assumption of the hypothesis that marsupials have lost the P3 of placental
mammals.

Archer (1978a) puts forward an alternative scheme for enumerating the cheek teeth of
marsupials that is used here. Ontogenetic evidence in a few marsupials that have been studied
thus far indicates that there are two families of cheekteeth, P1 P2 P3 and M1 M2 M3 M4 M5.
The tooth here designated M1 is the dP4 or DP3 of the other two systems. In appearance, it is
like the molars behind it rather than the premolars. Thus, by adopting the system of Archer
(1978a), all the cheekteeth of each family are united by a common morphology and quite
separate from the other. P3 does not bud off the dental lamina beneath M1 as would be
expected if P3 was the permanent replacement of M1. Rather, P3 buds off the dental lamina
between P2 and M1.

Discussions of tooth enumeration systems in therians and particularly marsupials can be
found in Mahoney & Ride (1975), Bown & Kraus (1979) and Clemens (1979a).

Figure 9. Left lower molar of the symmetrodont Tinodon bellus, A, intemal view; and B, occlusal view.
(After Cassiliano & Clemens 1979).

The most primitive therians have essentially no more to their molar structure than the
reversed interlocking triangles. One such group of therians are the symmetrodonts (see Fig. 9);
the monotremes are another. Near the end of the Jurassic, an elaboration of the molars took
place which was to have a profound effect on the course of mammalian evolution. This was
the development of a basin (or talonid) behind the principal triangular pillar (or trigonid) on the
lower molar and a new cusp on the upper molar, the protocone, which occluded in the talonid
basin in much the way a pestle acts in a mortar to grind objects to powder. This innovation,
which has been termed the tribosphenic molar (see Fig. 10), apparently laid part of the essential
groundwork for the rise of the marsupial and placental mammals. Except for the three genera

902 - RICH

Figure 10. Diagram showing the occlusal relationships between upper and lower molars of a tribosphenic
therian, Didelphodus sp. Contrast with Fig. 8. Note particularly the presence of a basined talonid on the
lower molar and the protocone on the upper in contrast to the condition in Kuehneotherium praecursoris.
(After Bown & Kraus 1979).

of monotremes, all living mammals fall into one of these two groups. The two groups differ-
entiated from one another about 100 million years ago near the boundary between the Early and
Late Cretaceous.

A TALE OF MARSUPIALS AND PLACENTALS

It is the structure and mode of function of the female reproductive system that forms the
most obvious difference between the marsupials and placental mammals. Unfortunately, there
is little evidence from the fossil record about the evolution of this system. The question
remains moot as to whether the marsupial condition represents a stage through which the
placental mammals passed or if the common ancestor of the two groups had a reproductive
system in the females closer to that of the egg-laying reptiles from which they ultimately
arose.

If one accepts the premise that the marsupials represent a structural intermediate between the
reptiles and placentals, there are two problems that must be explained. First, in marsupials
there is a pseudovagina present in the females which has no counterpart in placental mammals.
Second, the ureters pass lateral to the vagina in placental mammals and medial to the twinned
vaginac in marsupials (see Fig. 11).

On first sight, it would appear that the pseudovagina of marsupials is homologous to the
true vagina of placental mammals. Both occupy the same position relative to the ureters and
both are medial structures. However, examination of embryos shows that the true vaginae are
formed by the posterior growth of the Miillerian ducts from the ovaries. In placental
mammals, the two Miillerian ducts fuse to form the medial vagina while in marsupials, they
form the lateral vaginae, fusing only for a short distance (see Fig. 12). In marsupials, from the
point of fusion of the Miillerian ducts, an outpocketing forms that grows posteriorly to form
the pseudovagina. The structure is not linked directly to the urogenital sinus until the time of
the first birth. It is most likely that the pseudovagina is an innovation that appeared
subsequent to the separation of the marsupials and placental mammals. Although most young

AUSTRALIA'S MAMMALIAN RECORD - 903

— Uterus

Z -Uterus — Lateral vagina
vin Vestigial w
olffian ;
duct ~ Birth canal

(Median Vagina)

Vestigial Wolffian

Urogenital duct

Vagina Sinus

Figure 11. Urogenital tract of a female eutherian (left) and marsupial (right). (After Tyndale-Biscoe 1973).

\ Metanephric Degenerated mesonephric
.., kidney duct

wt
it
im

Mullerian duct

/ Lateral vaginal fp viinar
Ureteric bud Metanephric canal ~ —— é Sal
(grows dorso-mediad) j mass | Fg Incipient urethra
: Ureter Urogenital si
> Median vagina nll 9 ae
| ~ Genital tubercle
Allantois | Sexually indifferent Stage Female Specialization

\ ¢ —-Cloaca

Cloacal membrane
wor
© hey

Degenerated mesonephric duct

\
1
1
f

Uterus
Incipient urethra

Ureteric bud
(grows dorso-laterad)

Median ‘ :
Ly vagina Urogenital sinus
Eutherian
Figure 12. "Diagrammatic illustrations comparing development of differences in spatial relationships
observed in female urogenital ducts between marsupial and placental mammals (ventral view)." (After

Lillegraven 1969).

marsupials pass through the pseudovagina during the birth process, not all do so. Some utilise
the lateral vaginae.

In the development of the urogenital system in female mammals, the ureters grow forward
from the base of the bladder towards the kidneys, while the Miillerian ducts, which will become
the uterus and vagina of the adults, grow posteriorly from the ovaries. Topologically, whether
one or the other is the more lateral is simply a matter of how these two growing tubes meet

and pass one another. Therefore, it does not seem a major transition in developmental pattern

904 - RICH

to imagine an ancestral placental mammal switching from one arrangement to the other if it
was advantageous to do so.

Exactly what the respective advantages are of the two reproductive methods is unclear
(Kirsch 1977a, Parker 1977). The marsupial young are born at a very immature stage and must
crawl from the opening of the urogenital sinus to a teat where it firmly attaches itself. This all
happens within the normal duration of a single estrous cycle. By contrast, in the placental
mammal, through secretion of progesterone, the estrous cycle is halted at a stage where the
uterus is able to accept the implantation of the embryo and remains in that condition until the
time of birth long after the normal duration of the estrous cycle.

Each reproductive method may have its advantages without one necessarily being
consistently superior to the other. However, it is of interest to note that although the
developmental differences between the two structural patterns do not seem to be great, the
dichotomy is rather clearcut. One does not find within a family or order some species that have
the marsupial pattern and others that have the placental one. It seems that once a particular
reproductive strategy was chosen, the commitment to it is such that the cost of switching is
too great to enable this to occur in response even to major differences in habitat preference and
mode of life between members of the same family or order.

A hres

een
Saas “An Coin! gE
Ay, Oe.

yh Ps
C ype Ve

AT ae
unio (oN
on aft

Figure 13. Alphadon lulli, didelphid from the Late Cretaceous of North America. A, left M23, occlusal
view, x12; B, occlusal view of right mandible, x 10.8; C, labial view of right mandible, x 10.8. (After
Clemens 1966).

AUSTRALIA'S MAMMALIAN RECORD - 905

It may be that the critical difference lies in the amount of parental effort invested in the
growth of each offspring. Lillegraven et al. (1987) have argued that by maintaining the young
in utero until they are more advanced, eutherians generally have a correspondingly shorter
lactation period prior to the time of weaning than marsupials, lactation being more energy
inefficient in terms of physical development.

Although Holoclemensia from the late Early Cretaceous was initially regarded as a
marsupial, doubt has been recently placed on this allocation. The first undoubted marsupials
are from the Late Cretaceous of North America. The central group then was the Didelphidae, a
morphologically conservative family that persists to the present day and includes 70 of the 78
living marsupial species in the Americas (see Fig. 13).

.)

nage
ays

Figure 14. Pediomys cooki, pediomyid from the late Cretaceous of North America. A, right maxilla with
p3_m4, x 12; B. occlusal view of right mandible, x 12; C, labial view of right mandible, x 12. (After
Clemens 1966).

Out of the Didelphidae arose two additional families in the Late Cretaceous. One was the
Pediomyidae, which was characterised by a reduction in the stylar cusps along the outside or
buccal surface of the upper molars (see Fig. 14). This pattern was repeated later, independently

906 - RICH

Aut ul

fp sf

SPL. S44,

\y mM

/ ¥
wah, ie wt Mi od

Figure 15. Didelphodon vorax, stagodontid from the Late Cretaceous of North America. A, left M?,
occlusal view, x 7.2; B, occlusal view of left mandible, x 2.4; C, labial view of left mandible, x 2.4; D,
lingual view of left mandible, x 2.4. (After Clemens 1966, 1968).

AUSTRALIA'S MAMMALIAN RECORD - 907

by the Australian thylacines and possibly independently by the South American borhyaenids as
well. Possession of this feature by these three groups appears to be an example of parallel
evolution in which from the same stock, the Didelphidae, similar structures appeared more than
once 1n response to similar selection pressures. The protocone and talonid basin are reduced in
the thylacines and borhyaenids and enlarged in the pediomyids from the condition in didelphids,
suggesting that the similarities of the buccal regions of the molars were acquired independently,

The third North American Late Cretaceous marsupial family was the Stagodontidae (see
Fig. I 5). Known from only a few specimens, these animals had evolved to become specialised
carnivores. They were larger than other contemporary marsupials, reaching the size of a sea
otter or large native cat. Their jaws were short, deep and heavy, and the dentition was well
adapted for crushing prey such as shell-fish.

As the Cretaceous drew to a close in North America, the marsupials declined in number of
species until only a few didelphids were left. At the same time, placental mammals were
increasing in diversity there, perhaps as a result of immigration from eastern Asia where they
may have originated somewhat earlicr. Whether the rise of one group brought about the decline
of the other is an open question. Alternatively, it has been suggested that the marsupials were
for the most part members of what has been termed "the Triceratops community" which
included dinosaurs (Van Valen & Sloan 1977). This was displaced by another Community in
which placental mammals were numerous and marsupials, although present, were rare, “the
Protungulatum community." The Triceratops community was associated with an open
country flora, while the Protungulatum community was associated with forests. With
climatic cooling at the end of the Cretaceous, the forests spread towards the equator. Thus,
under this model the change in abundances of placentals and marsupials would not be related to
direct interaction, but rather was owing to their linkage with different communities.

Marsupials persisted in North Amercia as a minor element of the fauna until they became
extinct there in the Middle Miocene. Never again were they to achieve on that continent the
diversity they had in the Late Cretaceous. At the end of the Pliocene, with the establishment
of the Panamanian isthmus, North America was re-invaded by marsupials entering from South
America. Five species in four genera are found today at least as far north as southern Mexico
but only one ranges into the United States and Canada: Didelphis virginiana.

Late Cretaceous mammals are known in Europe from only a single published specimen,
However, there are three European Palacocene faunas with large samples. In these there are no
marsupials, so it seems likely that when they do appear in the record there in the Early Eocene,
this was close to the time of their actual entry into that continent. This is quite likely on
another ground. The Early Eocene was a time of extensive interchange of terrestrial mammals
between North America and Europe. Fifty-seven percent of the Early Eocene mammals in the
Paris Basin are also known from North America while the Late Palacocene and Middle Rocene
faunas share few genera in common (McKenna 1975, Savage & Russell 1943), The most
probable route of entry was via Greenland before sea floor spreading had completed the
separation between North America and Europe (McKenna, 1983). Only the durable,
conservative Didelphidae reached Europe. The evolutionary patiern there was much as it was in
North America during the Tertiary. Only three genera have been recognised in the European
Tertiary, and the differences between these generalised omnivores is not marked (Crochet 1940).
They finally became extinct in the Middle Miocene.

Until 1984, marsupials were unknown in Africa. Almost simultaneously, didelphids were
recognised in the Early Eocene of Algeria (Crochet 1984) and the Early Oligocene of Egypt
(Simons & Bown 1984). In that same year, a didelphid was also reported from the Early
Oligocene of Kazachstan (Gabunia et al. 1984). Although evidently present on both
continents, marsupials were never an important clement of the mammalian fauna of either,

Kielan-Jaworowska (1982) recognised the order Deltatheroida as the sister-group of the
Marsupialia within the metatheria. Deltatheroidans are known from Asia and North America

908 - RICH

Figure 16. Mandibles of caenolestoids. A, Caenolestes sp. (Caenolestidae), Recent; B, Eudolops carolo-

ameghinoi (Polydolopidae), Early Eocene; C, Abderites meridionalis (Caenolestidae), Early Miocene, D,
Epidolops ameghinoi (Polydolopidae), Late Palaeocene. Length of scale 10 mm. (B after Paula Couto 1952; A,
C, D after Marshall 1980).

AUSTRALIA'S MAMMALIAN RECORD - 909

30mm

Figure 17. Lycopsis longirostrus (Borhyaenidae). A, lateral view of right upper dentition; B, occlusal
view of right upper dentition; C, lateral view of right mandible; D, occlusal view of right mandible. (After

Marshall 1977a).

and have a dental formula of three premolars and four molars and an alisphenoid component to
the auditory bulla as in marsupials but differ from them in having higher trigonids on the lower

910 - RICH

molars and wider stylar shelves and smaller protocones on the upper molars (Kielan-
Jaworowska & Nessov 1990).

In South America, the history of marsupials was much different from that on all the
continents previously mentioned (Patterson & Pascual 1972, Simpson 1980, Marshall 1982,
Marshall & Muizon 1988). Prior to their arrival in South America along with the placentals at
the end of the Cretaceous or the beginning of the Tertiary, the mammalian fauna there was
dominated by an assemblage of non-tribosphenic forms (triconodonts, eupantotheres, paratheres
[= edentates sensu lato] and possibly multituberculates), which apparently evolved in isolation
on the Gondwana continents since the Middle to Late Jurassic. This assemblage is documented
by the fauna from Los Alamitos Formation from Patagonia (Bonaparte 1987, 1990).

The Tiupampa local fauna from Bolivia and the Laguna Umayo local fauna from Peru are
dated as Maestrichtian (latest Cretaceous) (Marshall & Muizon 1988) or early Palaeocene (Van
Valen 1988a-b). Somewhat younger than the assemblage from the Los Alamitos Formation,
these sites differ strikingly from it in faunal composition. These younger sites indicate that at
about the Cretaceous-Tertiary boundary, there was a major turnover in the South American
mammalian fauna with the disappearance of all the previous elements except the paratheres and
the arrival of the placentals and marsupials from Laurasia where they are well known during the
latter half of the Cretaceous. The marsupials represented are peradectids, microbiotheriids,
didelphids, carloameghiniids and borhyaenids. As in North America, the pediomyids became
extinct near the end of the Cretaceous in South America. The others continued into the
Cainozoic there, with the microbiotheriids and didelphids persisting to the present day.

During the Cainozoic, a minimum of seven marsupial families arose in South America
from the didelphids or didelphid-like forms. The only one of these to survive into modern
times are the didelphids themselves and the Caenolestidae, a group of generalised omnivores
characterised by forward projecting lower incisors and didactylous rather than syndactylous hind
feet, unlike the Australian diprotodontians, which they otherwise resemble (see discussion
below). This family appeared in the Early Eocene. Some of the early members were among
the most specialised in the group, having enlarged the M2s into shearing blades (Fig. 16B).
However, it was the dentally unspecialised which continue to survive (Fig. 16D).

Somewhat earlier, Late Palaeocene, are the first known records of the Polydolopidae, a
family closely related to the caenolestids and even more dentally specialised. These forms
developed elaborate molars and enlarged the P3s into serrated blades (Fig. 16A, C). This
specialised group became extinct after the Early Oligocene, even before the relatively
specialised caenolestids disappeared.

The principal South American terrestrial carnivores for much of the Tertiary were the
Borhyaenidae. Patene is the most primitive member of the family, being somewhat didelphid-
like in its features. This genus is known from the oldest deposits (Late Palaeocene) containing
borhyaenids. Along with it are more advanced forms indicating that the group had evolved
from the didelphids somewhat earlier. Borhyaenids are distinguished from didelphids by the
reduction in the outer margin or stylar shelf of the upper molars, the tendency of the paracone
and metacone to be closer to one another, and the increase in size of the molar series posteriorly
so that the M4 and M9 are the largest in the row, the M> being reduced (Fig. 17). Borhyaenids
persisted until the Early Pliocene. It has been suggested that the decline of the larger
borhyaenids in the late Tertiary was brought about by competition from large carnivorous
ground birds, the phororhacoids (Patterson & Pascual 1968, 1972, Marshall 1978).

During the Middle to Late Pliocene, there existed a highly specialised derivative of the
borhyaenids, the Thylacosmilidae. This family of large carnivores is most noteworthy for the
development of enlarged canines (Marshall 1976). These bear an uncanny resemblance to those
of the placental Machairodontinae, the so-called "saber-tooth cats" (Fig. 18). These quite
distinct groups independently evolved striking saber-teeth, an excellent example of convergent
evolution.

AUSTRALIA'S MAMMALIAN RECORD - 911

_ The Argyrolagidae are a family of marsupials that converge in skeletal structure with
bipedal, Jumping and leaping rodents elsewhere in the world such as the kangaroo rats of North
America (Heteromyidae), jerboas of Africa and Asia (Dipodidae) and hopping mice (Notomys)
of Australia. As in all of these arid-adapted forms, the hind-limbs are elongated and the skull
has inflated chambers surrounding the middle ear (Fig. 19). One feature is shared with a group

Figure 18. Skulls of (A) the placental machairodontine felid Eusmilus sicarius and (B) the marsupial
thylacosmilid Thylacosmilus atrox, x 0.25. (After Riggs 1934).

of marsupials that otherwise are quite unlike them, the wombats: they have ever-growing,
columnar, hypsodont molars. Argyrolagids appear in the Early Oligocene and became extinct
in the Early Pleistocene. Their relationships with other South American marsupials are not
well understood (Simpson 1970a, b, Wolff 1984).

The last three South American marsupial families are extremely enigmatic, all being known
from very little material. The Early Miocene Necrolestidae bear some resemblance to placental
moles (Fig. 20). Patterson (1958) tentatively allied the family with the Borhyaenoidea. Archer
(1984a), however, because they have none of the unique features of the marsupials, removes
them from the group. The Grocberidae are known from only a single mandible fragment and an
unassociated skull fragment, which are superficially rodent-like (Fig. 21). Their record is
restricted to the Early Oligocene, approximately contemporaneous with the appearance of the
rodents in South America (Simpson 1970c). Perhaps the Groeberidae were displaced by the
invading rodents (Simpson 1980). The Late Oligocene Patagonia peregrina is the sole
representative of another rodent-like marsupial family, the Patagoniidae (Fig. 22). Known only
from a few mandible fragments and isolated teeth, it is quite similar to the Groeberidae except
for the presence of a lower canine.

The Antarctic record of marsupials, and all terrestrial mammals for that matter, is restricted
to a single site on Seymour Island, which is south of the southern tip of South America. Of
Late Eocene age, all the specimens have been referred to Antarctodolops and Eurydolops, both
in the Polydolopidae (Case, Woodburne & Chaney 1988, Woodburne & Zinsmeister 1984),

It is singularly unfortunate that Australia, with the most diverse assemblage of marsupials,
has the poorest record of Mesozoic and Tertiary terrestrial vertebrates of all continents except
Antarctica (Fig. 23). Although mammals appeared elsewhere in the world by the Late Triassic,
their earliest Australian record is based solely on two specimens from the Early Cretaceous
(Archer et al. 1985, Rich, Flannery & Archer 1989) when the history of mammals was half

912 - RICH

Figure 19. Argyrolagus scaglii. A, latcral view of skull; B, ventral view of skull; C, lateral view of
jaw, x 2.3. (After Simpson 1970a).

over. Another major break follows for the next oldest, securely-dated occurrence of terrestrial
mammals on the continent and the earliest record of marsupials is Late Oligocene (Tedford et
al. 1975) by which time 88 percent of the history of the class had already passed. Recently, a
few dozen isolated marsupial teeth have been found at the Tingamarra site in southeastern
Queensland, which may well prove to be of an age in the Tertiary prior to Late Oligocene (see
Appendix I). What it has so far been possible to document in the history of this group in
Australia is primarily evolution within modern families and the barest traces of a number of
extinct families, many of which are closely allied to others still extant. Except possibly for
the macropodoids and phalangerids, the time of emergence of the various families was mostly
long over when the earliest glimpses of this fauna are afforded to us. To discover early Tertiary

AUSTRALIA'S MAMMALIAN RECORD - 913

ors 20. Necrolestes patagonensis, right lateral view of snout region of skull, x 2.3. (After Patterson

Figure 21. Groeberia minoprioi mandible. A, occlusal view; B, ventral view; C, lateral view; x 5.75.
(After Simpson ef al. 1962).

and additional Mesozoic sites is obviously a first priority research objective for understanding
how the pattern of descent of the marsupial groups occurred on this continent.

However, by working with what clues are preserved in the modern animals that suggest the
condition of their common ancestors as well as studying fossils of forms that are structurally
intermediate between modern groups, it is possible to begin postulating about how the
evolution of the Australian mammals took place. The structurally intermediate forms that have

914 - RICH

been found as fossils cannot be regarded as actual ancestors. They occur too late in time for
that, because their presumed descendants were living alongside them.

Not only is the pre-Miocene record of Australian mammals virtually non-existant, the
number of Neogene sites is low relative to other continents. Less than one hundred and fifty
Late Oligocene through Pliocene sites are known in Australia, while in North America there
are more than two thousand for the same time interval (Savage & Russell 1983). The quality
of preservation at many of these sites is not the best. However, it has been possible to use
them to put together a general outline of the Neogene and Quaternary evolution of Australian
mammals. One of the first needs to be met to accomplish this has been the establishment of a
sequence of the sites in both relative terms and, where possible, in physical time units.

Cz alveolus

Ht tesen y+

apy N /

Figure 22. Patagonia peregrina mandible. A, lateral view; B, occlusal view. Length of scale 2 mm.
(After Pascual & Carlini 1987).

CHRONOLOGICAL FRAMEWORK OF AUSTRALIAN
TERRESTRIAL FOSSIL MAMMAL SITES

The chronological framework in which the various sites where fossils of terrestrial
mammals occur is based on a combination of contributions from several disciplines.

Physical stratigraphy is used in the few instances where it can be shown that one
assemblage of fossils is from a rock body that overlies, and thus succeeds, another in time,
which has also yielded a suite of fossils. This is the most direct approach for demonstrating

AUSTRALIA'S MAMMALIAN RECORD - 915

NORTH SOUTH
A
MY AMERICA AMERICA EUROPE AFRIC ASIA ANTARCTICA | AUSTRALIA
ie}

10 oi
20
30
40 a =
50 :
ee
60

70

CENOZOIC

80

90

100

110

CRETACEOUS

120

130

140

150

oO
no 4
of, 8 =? =
{a}
<5
1704 &
= +— aa eee
ake J
180
190
LATE
TRIASSIC an?

Figure 23. Occurrences of mammals through geologic time. (Modified from Clemens, Lillegraven ef al.

1979).

916 - RICH

that one type of fossil is younger than another. However, it does not provide any measure of
how great the time difference is, only a chronological ordering of biological events.

Biostratigraphy, the relative ordering of rock units in time, is done in two quite different
ways at sites where fossils of terrestrial mammals are found in Australia. In the first approach,
the relative ranges of various fossil taxa found at the sites in question are determined by
reference to their recorded vertical ranges in more-or-less continuous sections of rock elsewhere.
As more such sections are examined, confidence in the relative ranges of the taxa increase. In
practice, this means that the taxa useful for this type of biostratigraphy in Australia are not the
fossil mammals themselves, because they do not occur through any significant vertical
thickness of rock. Rather, they tend to be restricted to single sites of no practical thickness.
So, it is the organisms associated with the fossil mammals which are also much more
widespread in rock units that can be utilised for biostratigraphic purposes in this manner. Such
forms include marine mollusca, foraminifera, spores, and pollen, All of these occur frequently
enough in continuous sedimentary sequences such as most commonly occur in marine deposits
that the relative scale of their respective vertical ranges in rock can be accurately assessed.

The second method of determining the relative ages of deposits with fossils is utilised only
when there is no other available criteria. This approach is to analyse the degree of evolutionary
advancement of one or more taxa at a given site relative to closely related taxa at another. It
has been termed the stage-of-evolution approach. In order to carry out such an analysis, one
must already have an idea as to the direction of the trends of structural change among the
organisms analysed. By having a few instances where two or more suites of fossils can be
dated relative to one another or by comparing fossil taxa with modern members of the same
groups, the general direction of morphological change can be inferred. By then assuming that
change occurred in only one direction, it is possible to estimate how relatively advanced two
species are with respect to one another and hence, the relative ages of the rocks from which
they came. The more taxa for which this can be done, the more confidence that can be placed
in the resulting analysis. It is only in this way that to some extent one of the greatest
weaknesses of this method can be overcome, the assumption that the changes occur in only one
direction. In detail, this assumption is certainly false but with broader sampling of taxa, the
likelihood increases that a fauna with more advanced members will be younger. This approach
is discussed in further detail and defended by Savage (1977). A critique of the approach is given
in the same volume by Eldredge & Gould (1977).

Stirton et al. (1968) and Stirton, Woodburne & Plane (1967) relied heavily upon this
method to order the Australian Tertiary mammal sites. In many instances, there was no other
basis available to estimate the ages of the fossil sites. They utilised diprotodontoids
exclusively, but more recently other groups have been analysed for the same purpose; é.g.
pseudocheirids (Woodburne, Tedford & Archer 1987),

Where possible, an estimate of the age for a fossil site in terms of physical time units is
made. In practice this means utilising one of a number of different dating procedures that rely
on the constant rate of decay of radioactive isotopes no matter to what physical conditions they
have been subjected .

The best known of these radiometric dating schemes is based on the decay of carbon
fourteen, 14C. With a half-life of 5,730 years, this technique is only adequate for samples less
than about 35,000 years old. New techniques based on detecting the !4C remaining in the
sample rather than noting the decay of that isotope might eventually permit age determinations
two to three times the present limit.

Any carbon that is incorporated into living tissue can, in principle, be utilised to date when
an organism lived. !4C is formed primarily in the atmosphere owing to the conversion of
nitrogen fourteen, !4N, to that isotope owing to cosmic ray bombardment. Approximately 70
tonnes of nitrogen is annually converted to !4C in the earth's atmosphere in this manner. The

AUSTRALIA'S MAMMALIAN RECORD - 917

carbon subsequently combines with oxygen to form carbon dioxide, some of which is then
incorporated into the structure of living plants by the process of photosynthesis. Once the
plant dies, the intake of '*CO, into its structure ceases and the amount of 14¢C begins to
decrease as that element decays back to !4N at a constant rate. In the same fashion, when an
animal dies, the intake of !4C into its body structure ceases because ultimately the source of
that isotope is the plants on which it is directly or indirectly dependent.

Provided there is no addition or deletion of carbon to the remains of an organism subsequent
to its death, the amount of !4C remaining gives a reliable measure of the time of cessation of
metabolism or death. Experience has shown that some types of fossils give more reliable dates
than others. Charcoal, for example, can be treated chemically so that virtually all the !4C that
may have entered a sample from the atmosphere subsequent to its being burnt can be removed,
making possible a reliable date. On the other hand, samples based on bone collagen are always
regarded as minimum dates. This is because !4C from the atmosphere can be incorporated into
the sample long after the animal's death in such a manner that it is impossible to separate it in
any way from the !4C present at the time of death.

Potassium forty, 4°K, decays with a half-life of about 1,250 million years. In principle, as
a tool for determining physical dates, this decay scheme is no different than !4C to 14N. The
longer half-life means that rocks as old as the solar system can be dated. In fact, one of the two
real problems that prevents using 4°K decay more frequently is that unless the sample in
question is at least 500,000 years old, not enough 49Ar accumulates to make it possible to
detect. The other major problem with applying this dating scheme to the problems of the
chronology of Australian Cainozoic mammals is that ideally what is dated is the last time the
rock was heated to a high enough temperature that any 4°Ar previously incorporated in the
crystal lattice of the mineral grains examined was able to escape. Once the mineral grains
cooled below a critical temperature, any 49Ar subsequently produced by the decay of 4°K
remained trapped in the crystal unless subsequent weathering or other chemical processes breaks
down the lattice. It is for these reasons that what is needed to carry out a meaningful 4°K-49Ar
dating is a fresh sample of an igneous rock that was formed penecontemporancously with the
fossils to be dated. Thus far in Australia, such dating has been done only on basalt flows that
cover or bracket fossil sites. On other continents, however, volcanic ash deposits or tuffs have
also proven to be similarly useful. This is particularly true where, as often happens, the
fossils occur within the dated tuff.

Because the earth's magnetic field reverses polarity from time-to-time and the orientation of
the field at the time of compaction of sediments can be determined, palacomagnetic data has
been widely used in recent years for assessing geologic age (e.g. Lindsay et al. 1980).

A major stumbling block to the application of this technique in Australia is that only a few
terrestrial mammal fossil sites have been found to be in a geologic context where the method
could be utilised; e.g. Bone Gulch and the Fisherman's Cliff local faunas (Bowler 1980,
Woodburne ef al. 1985) and Portland Local Fauna (MacFadden et al, 1987). Ideally, what is
wanted is a series of sites distributed over a considerable vertical thickness of rock. In Australia

Figure 24. Correlation of the Australian and New Guinean Tertiary terrestrial mammal faunas and local
faunas. Where no other method is indicated by a symbol, stage-of-evolution of the entire mammalian
assemblage but particularly the diprotodontoids, was the basis for the correlation, The age assignments for
the faunas and local faunas generally follow Woodbume et al. (1985). Archer et al. (1989) regarded the
Northern and Central Australia sites listed below Alcoota on the righthand panel to be significantly older.
Riversleigh upper System C is regarded by them as latest Middle Miocene (about 10 myBP) and the base of
Riversleigh System, latest Oligocene (about 25 myBP). See Appendix I for discussion of individual

mammalian faunas and local faunas.

918 - RICH

CORRELATION OF NEW GUINEA & AUSTRALIAN
New Guinea & Eastern & South-eastern Coastal Australia

ee Sranen
Eeeie. Nae Curramulka, Krui
River
Lake Tyers,

Sunlands
F orsyth’s (@)
Bank

Canadian
Lead

Geilston Bay

Tingamarra

Basis for Correlation

@ Marine Invertabrates oo Radiometric ‘@) Magnetic
Chiroptera Fy Pollen

AUSTRALIA'S MAMMALIAN RECORD - 919

TERTIARY MAMMAL-BEARING FAUNAS, LOCAL FAUNAS AND SITES
Northern & Central Australia

Bone Gulch, Fisherman's Cliff, Floraville,
Palankarinna, Rackham's Roost, Quanbun

Riversleigh upper :
System C Bullock SS
Creek
= Riversleigh lower se <n : ;
: System C cs eeameoneas
<-| — Kangaroo + @ Kutjamarpu
Riversieigh System B Well ~ “|
— ete aia Etadunna
Riversieign D-site SSS, : : Faunal Zone E Ericmas
Equivalents cs . Seeses
.. & Etadunna
Riversleigh ee “*S 4 Faunal Zone D
Caves : z Sie
et Etadunna
BARS eet = ~ Faunal Zone C

lan's Prospect

Wadikali

. we
: gee Etadunna a.
Riversleigh : : Spee Faunal eee a
System A Ss
Etadunna
Faunal Zone A

Tarkarooloo, Yanda,

oy
@

Correlations between units at Riversleigh and South Australia suggested by Archer, Hand, Godthelp &
Megirian (1989).

920 - RICH

f Bullock Creek (IM)

A Quanbun
(?mP-Pleist.)

|
| Rackham's Roost (P)
|

1

Alcoota (IM) 4

a

Mogorafugwa (P)

Floraville (7IP) &
Riversleigh, Nooraleeba (mM) A

Tara Creek (P?)

A
Bluff Downs (eP) &

Tropic of Capricorn 7
Kangaroo Well (?mP) &

— eee

YQ |

Etadunna Faunal Zones C-E, Kutjamarpu, (lIO-mM)

t

Tingamarra (eT?) &
Chinchilla (?mP)4&
Kanunka (?IP)

Palankarinna, (?mP), Ngapakaldi Fauna, Etadunna Faunal Seats Los we .
Zones A,B,D, & E (IO-mM) lan's Prospect Lightni 5 Sten arn cae le oe ae
| ((0-mM? > @ ! ightning Ridge (eK) &
ad ik i Talyawalka
lands (eP) | ? ;
Sunlands (@P) | AVP eist) A Krui River,
i i& Bone Gulch, Bow (?mP)
Finpa, vanea, K\ Fisherman's Clif & cared oh tes
Hementon t (7IP ore Pleist) 9 Big Sink (e-mP)
ricmas p ;
Wadikali (O-mmy) os | sa a
Curramulka | id. g&pimadai (?)/ Great Buninyong
+$ Estate Mine (P),

Town Well (M?-P?)

Forsyth's Bank, ——

Hamilton (eP)

Batesford Quarry (M),

Dog Rocks (P)

a Lake Tyers (P)
Bunga Creek (P)
iN Morwell (P?)

Beaumaris (IM-eP)

Wynyard (eM)
Geilston Bay (IO)

Figure 25. Localities of Mesozoic and Tertiary terrestrial mammal faunas and local faunas in Australia and

New Guinea.

Smeaton (7/P or e Pleist)

AUSTRALIA'S MAMMALIAN RECORD - 921

144
1as¥ 1% w36

¥v Open sites $8 oo
@ Cave
# Unknown

Figure 26. Quaternary terrestrial mammal localities in Australia (After Horton 1984).

where there is more than one fossil site in an area, all seem to be clustered at nearly the same
stratigraphic level. There is no instance where sites are scattered through hundreds of metres of
vertical thickness of rock in which deposition was continuous or nearly so over several million
years as in the studies of Lindsay ef al. (1980) in the late Cainozoic Siwalik deposits of the
southern foothills of the Himalaya Mountains. Such a geological setting is prerequisite for
such investigations in order that a number of magnetic reversals can be recorded.

Stirton et al. (1968) provide an excellent summary of the Tertiary terrestrial mammal sites
of Australia and New Guinea. They list the fauna from each locality and discuss the geology

922 - RICH

and basis for correlation. In addition, they provide a correlation chart showing the chronologic
relationships between localities and a map of where they are located. Fig. 24 is a revised
correlation chart and Fig. 25 a revised map of those sites with those added that have been found
subsequently. The information in Stirton et al. (1968) is updated in Woodburne er al. (1985)
and below in Appendix I.

Williams (1980) provides a map of the Quaternary mammal localities of South Australia
and the fauna associated with each. Merrilees (1968, 1979a) presents much the same kind of
information for Western Australia. Horton (1984) has a map of Australian Pleistocene
mammalian fossil sites (Fig. 26). Both he and Murray (1984b and this volume) present maps
of Australia with the distribution of most of the larger Pleistocene marsupials.

As a generality, with unfortunately all too few exceptions, it may be said that Australian
Quaternary terrestrial mammal sites fall into one of two age groups. Either they are younger
than 35,000 years before present and capable of being dated by the 14C-technique or they are
regarded as "Quaternary, beyond the range of 140"; i.e, somewhere in the first 98 percent of
Quaternary time. It is quite possible that most of these latter localities are no older than a few
hundred thousand years. As yet there is no firm basis for distinguishing an Early from a Late
Pleistocene site on faunal grounds. Not enough is yet known about the chronology cf
mammalian faunal events within the Australian Quaternary to make possible a synthesis of it
as has been done for the corresponding period of time in Europe (Kurtén 1968) or North
America (Kurtén & Anderson 1980). However, given the much greater number of Quaternary
sites as compared with Tertiary ones (compare the number of Pleistocene sites in Australia
alone (Fig. 26) with the Mesozoic and Tertiary ones for Australia plus New Guinea (Fig. 25),
it seems that a detailed Quaternary chronology of mammalian faunal events is a more feasible
goal than a Tertiary one in Australia and New Guinea.

THE RECORD OF MAMMALS IN AUSTRALIA

Order: Monotremata

Living monotremes are divided into two quite different families, the platypus
(Ornithorhynchidae) and the echidnas (Tachyglossidae). Because the morphological and adaptive
differences between these two families of primitive, egg-laying mammals are so great,
Darlington (1957) proposed that they represent two widely separated branches of a large,
otherwise unknown, radiation of monotremes.

Until recently, the accepted view was that monotremes were the last remnant of a major
radiation of mammals including the triconodontans, docodonts and multituberculates, which
were grouped together as the Prototheria (Clemens 1979b). Within the Prototheria, on the
basis of similarities in the structure of the braincase, some workers contended that the
multituberculates were the group most closely related to monotremes (Kielan-Jaworowska 1971
1974, Kermack & Kielan-Jaworowska 1971),

Kemp (1983) has argued that the similarities seen between monotremes and possibly
multituberculates as well with the other prototherians are either non-existent or primitive
(plesiomorphic) character states and hence of little use in demonstration phylogenetic
relationships. Rather, he advocated allying the monotremes and, tentatively, the
multituberculates, with the therians.

Corroborative evidence for this viewpoint has come from assessment of the dental pattern of
the Early Cretaceous Steropodon galmani. Unlike the living platypus, Ornithorhynchus
anatinus, S. galmani has fully enamelled cheek teeth (Archer et al. 1985). These teeth have the
reversed triangle pattern that is the hallmark of the molars of therians. Archer et al. (1985)

AUSTRALIA'S MAMMALIAN RECORD - 923

gh >
ANN
AY
il
aay" py
“ie
Uf

‘i
| |
( ull i |
\ SS "
i iN Li) 4)
)

\ \ | 4
p\ | Me he
y\ MH
if? J

Figure 27. Obdurodon insignis. Isolated left lower molar: A, occlusal view; B, posterior view; C,
anterior view; D, buccal view; E, lingual view. (After Clemens 1979b). Isolated left upper molar: F,
occlusal view.

924 - RICH

went further than this and regarded the dentition of Steropodon galmani as having a
tribosphenic pattern. However, Kielan-Jaworowska et al. (1987) on the basis of wear facet
analysis have argued that the pattern is not tribosphenic and predicted that the unknown upper
molars lack a protocone.

There are only two Tertiary monotremes known, the Early to Middle Miocene Zaglossus
robusta and the Late Oligocene-Middle Miocene Obdurodon insignis. O. insignis is virtually
identical to the living platypus, Ornithorhynchus insignis, as far as known postcranially
(Archer et al. 1978). However, unlike the modern platypus, it had fully enamelled cheek teeth
which closely resemble those of Steropodon galmani (Woodburne & Tedford 1975). The upper
molar of O. insignis lacks a protocone and thus corroborates the viewpoint of Kielan-
Jaworowska et al. (1987) that monotremes do not have a tribosphenic dentition (Fig. 27). This
low monotreme diversity at a time when the mammalian record is reasonably good in Australia
coupled with the close resemblance between the cheekteeth of S. galmani and the 85 million
year younger O. insignis, suggests that perhaps Darlington's idea of an extensive monotreme
radiation in the late Mesozoic and early Cainozoic may be erroneous.

Bonaparte (1990) has recognized a resemblance between the dentition of Obdurodon and
Steropodon with the South American Cretaceous “eupantotheres” and the North American
dryolestoids. On this basis, he tentatively proposes a relationship between these groups of
non-tribosphenic therians.

Both echidnas and platypus are known from Quaternary deposits, but their remains are not
abundant. These and the one Tertiary echidna are not markedly different from their living
counterparts, except that some specimens of Zaglossus were much larger than the living
Zaglossus bruijni and occurred in southern Australia, while the modern occurrence of the genus
is exclusively in New Guinea (Murray 1978b).

SUPERCOHORT: MARSUPIALIA

With a few possible exceptions, the Australian marsupials appear to be a united group. By
this is meant, it is likely that all of them could be descended from a single species that reached
the continent from Antarctica.

Based on the degree of similarity in the structure of proteins in the various groups, it
appears that the Australian marsupials are more closely related to one another than the
didelphids and caenolestids of South America are to one another (Kirsch 1977b, Lowenstein et
al. 1981, Sarich pers. comm. to Archer 1986 in Aplin & Archer 1987, Sarich pers. comm. in
Marshall et al. 1989). Assuming that the average rate of change in protein structure is constant
and varies stochastically, this implies that the differentiation of marsupials began later in
Australia than in South America. It supports the hypothesis that a marsupial immigrated into
Australia from South America after the caenolestids and didelphids differentiated from one
another, an event that probably occurred in the Late Cretaceous.

Further supporting a natural taxonomic division between the marsupials of Australia and
those elsewhere is the work of Biggers & De Lamater (1965). They demonstrated that
spermatozoa frequently formed pairs once outside the testes in all American marsupials
examined, while the phenomenon was not observed among Australian marsupials. In addition,
the work of Szalay (1982) suggests there is almost a perfect division in foot structure between
a primitive condition of tarsal structure found in all but one of the American marsupials and an
advanced one in all Australian marsupials (Fig. 28). The single exception is the South
American Dromiciops, the one extant genus of the Microbiotheria. Consequently, Australian
marsupials plus microbiotheres are grouped together as the Australodelphia, while all other
marsupials are placed in the Ameridelphia.

AUSTRALIA'S MAMMALIAN RECORD - 925

Case (1989) further supports this hypothesis with an explanation that the diversification of
the Australian marsupials did not take place until the early Cainozoic, because only then did
habitats become more diverse. In the Late Cretaceous-early Cainozoic, rainforests dominated
the continent. Only as Australia drifted into lower latitudes as the Cainozoic progressed did a
more varied flora become established, opening the way for the radiation of the marsupials there.

Tibia & Fibula
A facets Ss) eae e
Sustentacular &
Tibia &
Sustentacular 7 eC) Fibula facets
facet

ay

Figure 28. Contrast between the condition of the calcaneum articulations in the Ameridelphia (A) and the
Australidelphia (B). Left calcaneum in dorsal view. Note that the sustentacular facet is separate from the facets for
the astragalus and calcaneum in the ameridelphian (Glironia), while the three facets are contiguous in the
australidelphian (Thylacinus). (After Szalay 1982).

In this context, it is tantalising to note that the preliminary identifications of the possibly early
Tertiary Tingamarra Local Fauna suggest that it is dominated by a variety of "polyprotodonts"
strongly reminiscent of American didelphids (see Appendix I).

For many years, the great similarity in structure of the skull of the Australian thylacines
and some of the South American borhyaenids supported an hypothesis of a special relationship
between those two groups. At present, however, it seems the similarities are owing to a
remarkable case of convergence (see further discussion under Family: Thylacinidae).

The caenolestids of South America have in the past been allied to the Australian
diprotodontians by some workers. The similarity is largely based on the development of an
enlarged, most anterior lower incisor, the diprotodont condition. With this as the primary
feature linking the two groups, it is readily explained as a condition independently evolved in
the two groups. Furthermore, Ride (1962b) has shown it is likely that the lower incisor,
which is enlarged in the two groups, is not the homologous tooth. Unlike all diprotodontians,
caenolestids lack the syndactylous condition on their hind feet, where digits 2 and 3 are
enveloped in a single sheath of skin.

Marshall (1980) has reviewed the evidence for the origin of the Australian marsupial fauna,
and its relationship to the South American marsupials.

Order: Dasyuromorphia

Family: Dasyuridae
The dasyurids, which include the native cats and native mice, are the Australian marsupials
closest morphologically to the didelphids of the Americas (Fig. 29), The only consistent
dental difference between the two families is that the Didelphidae have five upper incisors and
four lower incisors, while the Dasyuridae have four uppers and three lowers. Presumably the
dasyurids arose from the didelphids after an episode of faunal interchange between the Americas

926 - RICH

Figure 29. Dasyurus maculatus. A, lateral view of skull and right mandible; B, dorsal view of skull; C,
ventral view of skull; x1. (After Green 1983)

AUSTRALIA'S MAMMALIAN RECORD - 927

and Australia across Antarctica. Among the oldest dasyurids known from the Late Oligocene-
Middle Miocene of South Australia is Ankotarinja, which might be more properly placed in the
didelphids (Archer 1976c). At present, this genus is too poorly known to be sure, but such a
relationship would be consistent with the idea that the didelphids were here in Australia and that
the dasyurids evolved from them on this continent. Ankotarinja as a didelphid would also be
consistent with the idea that the bandicoots or peramelids evolved directly from the didelphids
rather than through the dasyurids (see below). Certainly, if Ankotarinja had been found in
South America instead of South Australia, undoubtedly it would have been placed in the
didelphids on the basis of the known structure of the molars, premolars, mandible and maxilla.
When the dasyurids are first recorded in the fossil record, presumably long after the time of their
origin, except for this somewhat enigmatic possible didelphid, they fit easily in the Dasyuridae.
There is little fossil evidence of how they arose, for even the possible didelphid from the Late
Oligocene-Middle Miocene would be the descendant of the actual didelphid ancestor of the
dasyurids; an actual ancestor cannot live alongside its descendant, and contemporaneous with
Ankotarinja was Keeura, an undoubted dasyurid.

Isolated teeth informally identified at present as “polyprotodont" from the Tingaburra Local
Fauna of southeast Queensland are strikingly didelphid-like in their appearance (Godthelp, pers.
comm. 1989), This site may be significantly older than the South Australian sites that have
produced Ankotarinja.

Modern members of the Dasyuridac can be divided into eight or ten groups, but the
likelihood appears to be that some of the genera with numerous species such as Antechinus
will in future be subdivided, requiring a further assessment of the basic divisions of the family.
The extinct genera Ankotarinja, Keeuna and Wakamatha do not fit neatly into any present
division of modern dasyurids.

Dasyurids, judged by the sizes of the placental Carnivora, were small to medium-sized
carnivores. Even regarding the Tasmanian Wolf or Thylacine as part of this group (although it
is placed in its own family, the Thylacinidae), there are no known lion-sized carnivores among
the Australian mammalian fauna. It has been suggested that xiphodont crocodiles and the giant
varanid Megalania prisca filled this role. Certainly, there were large marsupial prey species
that could have been successfully attacked only by large carnivores.

Archer (1982a) has reviewed the fossil history of the family in detail.

Family: Myrmecobiidae

The numbats (Fig. 30D) are represented by only a single species, Myrmecobius fasciatus,
an anteater with a reduced dentition somewhat reminiscent of the Mesozoic triconodontans.
Their fossil record is brief, being confined to Holocene deposits on the Nullarbor Plains
(Lundelius & Turnbull 1978, 1989). However, on the basis of serological evidence, they
appear to have been a lineage distinct from the dasyurids since the time of the initial radiation
of that group (Kirsch 1977b). On the other hand, because of the structure of the basicranium,
it is clear that the sister-group of the myrmecobiids are the dasyurids (Archer & Kirsch 1977),

Family: Thylacinidae

When they first appear in the record in the Late Oligocene-Middle Miocene deposits at
Riversleigh in northwestern Queensland, the thylacinids are a rather diverse group including
Thylacinus sp., Nimbacinus dicksoni and three additional, as yet unnamed taxa. This
diversity at Riverslcigh is particularly remarkable because the group is all but unknown from
the contemporaneous fossil assemblages in South Australia.

Subsequently, the Thylacinidae are represented by only two species, the Late Miocene
Thylacinus potens, and the Quaternary Thylacinus cynocephalus. This Late Cainozoic decline
of the thylacinids occurred when the larger dasyurids such as Dasyurus and Sarcophilus

appeared,

928 - RICH

Figure 30. Thylacinus cynocephalus. A, lateral view of skull and right mandible; B, dorsal view of skull;
C, ventral view of skull; x0.5. (After Green 1983). Myrmecobtus fasciatus, D, ventral view of the skull, x

Because of their remarkable structural similarity to some of the South American
Borhyaenidae (compare Figs 17, 30A-C), they provide one of the most fascinating
biogeographic problems to be found among the Australian mammalian fauna. Archer (1982b)
provides an excellent review of this controversy.

The thylacinids are characterised by a tribosphenic dentition in which the stylar cusps have
been reduced from the conditions found in most didelphids and dasyurids (Fig. 30A-C).

AUSTRALIA'S MAMMALIAN RECORD - 929

The remarkable similarity between Thylacinus cynocephalus and some of the extinct
Borhyaenidae has been explained by basically two different hypotheses. In one, the similarity
has been interpreted as Owing to a close phylogenetic relationship between the two groups
implying interchange between South America and Australia of forms that had reached a
ee level of organization (Bensley 1903, Sinclair 1906, Wood 1924, Archer

»C).

Under the alternative hypothesis, the similarities were explained as owing to a remarkable
example of convergent or parallel evolution from a common ancestor that was a didelphid.
Didelphids are thought of as giving rise directly to the Borhyaenidae and the Dasyuridae. In
a: Thylacinidae arose from a dasyurid stock (Simpson 1941, 1948, Tate 1947, Marshall

These previously cited analyses were based on interpretation of dental and cranial characters.
However, recently elements of the tarsal skeleton have been analysed (Szalay 1982) and in
addition, albumin from Thylacinus has been compared to that from a number of different
marsupials (Sarich et al. 1982).

Szalay (1982) has found that the tarsals of Thylacinus have the typical pattern of the
australidelphians. This supports the view that Thylacinus was derived from dasyurids in
Australia independent of borhyaenids, which were derived from didelphids in South America.
However, the anomaly of the tarsal structure of the South American Dromiciops having the
derived Australian tarsal pattern provides a warning that this seemingly straight-forward
interpretation of thylacine-borhyaenid relationships may be modified by future work.

The albumin data indicate that Thylacinus is closely allied with dasyurids and progressively
more remote from myrmecobiids, peramelids, diprotodontians, and finally didelphids. Unless
one entertains the idea that borhyaenids were derived from dasyurids, a hypothesis thus far not
susceptible to testing by albumin or collagen structural studies, these results appear to again
indicate that Thylacinus is not the sister-group of the Borhyaenidae. Ironically, as pointed out
by Archer (1982b), the albumin results are somewhat too strong. Assuming a uniform
stochastic divergence in albumin structure, Thylacinus diverged from Dasyurus and
Dasyuroides 7 million years ago (Sarich et al. 1982). However, a variety of undoubted
thylacines have been found in the Late Oligocene-Middle Miocene Upper Site Local Fauna of
the Riversleigh District, at the very minimum 12 million years old and the better known
Thylacinus potens from the Late Miocene Alcoota Fauna is very similar to modern Thylacinus
cynocephalus, supporting the hypothesis that the separation between thylacines and dasyurids
occurred much earlier than indicated by the evidence of albumin structure. The slight
differences between the oldest and youngest described species of Thylacinus are also concordant
with the view that during the late Cainozoic, the genus evolved very slowly.

The last known living individual of Thylacinus cynocephalus died in the Hobart Zoo in
1933. Their numbers in Tasmania declined drastically with the advent of European settlement.
Dogs were then introduced to the island, and the Tasmanian government offered a bounty for
the destruction of the thylacines from 1888 to 1909 (Ride 1970). On mainland Australia, the
youngest remains of the species that have been firmly dated radiometrically are 3090 + 90 yBP
(Archer 1974). However, thylacine remains in the Kimberley district of Western Australia have
been less securely radiometrically dated at 0 + 80 yBP (Ibid.).

Dawson (1982b) provides a review of the systematics of Thylacinus in the Late

Quatemary.

Order: Peramelemorpha

Bandicoots are divided into two separate families. Most are accommodated in the
Peramelidae, but the extant rabbit-eared bandicoot, Macrotis, is placed in the family

930 - RICH

Figure 31. Comparison between didelphid, dasyuird, peramelid, and thylacomyid upper molars indicating
how structural evolution from a tribosphenic form (didelphid and dasyurid) to the quadrituberculate forms seen
in peramelids may have occurred. A, Didelphis virginiana (Didelphidae); B, Dasyurus viverrinus
(Dasyuridae); C, Jsoodon obesulus (Peramelidae); D, Macrotis leucura (Thylacomyidae); x 5S.
Abbreviations: me, metacone; mel, metaconule (= neometaconule of Tedford & Woodbume, 1987, see
caption of Fig. 7 for explanation); pa, paracone; pr, protocone; st B, stylar cusp B; st D, stylar cusp D.

Thylacomyidae along with the extinct Pliocene form Jschnodon. Several independent lines of
evidence support this separation. The suggestion of the division was first made by Bensley
(1903) and strongly advocated by Archer & Kirsch (1977). Bensley first noted that in evolving
from an ancestor with a tribosphenic dentition, the upper molars of bandicoots had changed
from a triangular to a quadrate outline in two different ways (see Fig. 31). In the peramelids
this was accomplished by the appearance of a prominent hypocone behind the protocone.
These two cusps together with the paracone and metacone on the buccal side of the tooth are
the four principal cusps on the peramelid upper molar. In the thylacomyids, there are four
principal cusps as well, but what has happened is that the metacone has shifted lingually to
form the principal posterointernal, rather than posteroexternal, cusp as in peramelids. The two
principal external cusps in thylacomyids are formed by enlarged stylar cusps, which in
peramelids are relatively as small as in dasyurids and didelphids. The paracone is present but is
not a large cusp in thylacomyids, and the hypocone only occurs on an occasional specimen,
then only as a dimunitive cusp.

The division between the two families has been further supported by comparative serology
(Kirsch 1977b) and comparisons of chromosome structure (Martin & Hayman 1967, Hayman
& Martin 1974),

AUSTRALIA'S MAMMALIAN RECORD - 931

Archer, on the basis of dental (1976b) and basicranial (1976e) features, has hypothesised
that peramelids arose directly from didelphids, rather than through a dasyurid intermediate.
However, many of the features shared by didelphids and peramelids to the exclusion of
didelphids are primitive; e.g. a higher number of incisors. Thus, if peramelids do share a later
common ancestor with dasyurids than didelphids as indicated by serological studies (Kirsch
1977b), it would suggest that this common ancestor retained many primitive traits found today
only in didelphids and peramelids.

It has been proposed that the peramelid dentition provides a structural intermediate between
the tribosphenic didelphid and dasyurid dentitions on the one hand and the bunodont,
selenodont, and lophodont patterns in the diprotodontians on the other. Bensley (1903)
advocated that his hypothetical “Properamelidae" gave rise separately to bunodont and
selenodont diprotodontians. In turn, the bunodont forms were ancestral to the lophodont
diprotodontians in Bensley's view. Archer (1976b), too, regarded the peramelids as central to
the radiation of the diprotodontians but places emphasis on the ease with which it is possible
to derive a selenodont molar from that of a peramelid to advocate the view that peramelids gave
ris€ to selenodont diprotodontians which in turn gave rise to all other diprotodontians by fusion
of adjoining cusps where necessary to form ridges and crests.

Another viewpoint is that bandicoots represent a radiation from dasyurids or didelphids that
had nothing to do with the diprotodontians and which arose independently from the same
tribosphenic stock, perhaps more than once (e.g. Ride 1971, Simpson 1945). On the basis of
comparative serology, brain structure, and possession of a superficial thymus gland,
i> ins are no closer to perameloids than to dasyurids (Kirsch 1977b, Baverstock et al.
1987).

A variety of small peramelids is represented in the Late Oligocene-Middle Miocene deposits
at Riversleigh. In the similar-aged deposits of the Etadunna and Namba formations in South
Australia only an unnamed genus also present at Riversleigh and Perameles are represented. In
the slightly younger Kutjamarpu Local Fauna, C. Campbell has recognised Peroryctes,
Echymipera, Isoodon, and Perameles (pers. comm. to Archer, cited in Archer 1981a). All
four are extant genera, and this is also the case for all peramelid records in the younger Tertiary
and Pleistocene faunas. If these Miocene records are correct, the differentiation of the
peramelids was completed to the generic level by the end of that epoch. However,
reconsideration of this material suggests that it may be premature to assign it to known
peramelid genera.

In contrast, the sole Tertiary record of the Thylacomyidae consists of a single lower jaw
bearing the name /schnodon australis, an extinct genus and species. /. australis is from the late
Cainozoic Palankarinna Local Fauna and in the structure of the molars is intermediate between
peramelids and Macrotis, the only extant genus of the family. This supports the suggestion
that thylacomyids evolved from peramelids during the late Tertiary.

Order: Notoryctemorphia

Family: Notoryctidae

The only fossil record of the marsupial moles is from Late Oligocene-Middle Miocene
deposits at Riversleigh, northwestern Queensland (Aplin & Archer 1987). Because the living
animals are restricted to deserts, the discovery of these animals at Riversleigh was something of
a surprise, as during the mid-Tertiary, it was an area of lush rainforests.

Serologically, notoryctids clearly belong to the Australian radiation of marsupials but
otherwise stand apart from them (Kirsch 1977b). Their different morphological features
suggest conflicting relationships for these highly modified fossorial animals. On dental
evidence, they appear to be allied with dasyurids and peramelids (Fig. 32). Notoryctids have

932 - RICH

been frequently described as syndactylous like peramelids and diprotodontians, but not all who
have investigated these enigmatic animals agree that this is the case. The auditory region
shows some similarities to the diprotodontians.

Figure 32. Notoryctes typhlops. A, occlusal view of upper molar; B, occlusal view of lower molar; C,
lingual view of lower molar. (After Bensley 1903).

Order: Diprotodontia

The numerous families of the Order Diprotodontia are united by the twin features of a
syndactylus foot and the diprotodont structure of the lower incisors, which gives the order its
name. The roof of the tympanic region of the skull is formed at least in part by the
epitympanic wing of the squamosal (Aplin 1987). Furthermore, the brain of all
diprotodontians has a unique connection between the right and left cerebral hemispheres, the
fasciculus aberrans (Abbie 1937, Smith 1902). Serological work by Kirsch (1977b) supports
the view that diprotodontians are a close knit group, well separated from all other marsupials,
and further provides evidence for groupings of many families within the order. Also,
diprotodontians possess a superficial thymus which is not found in other marsupials
(Symington 1898, Yadav 1973, Kirsch 1977b).

By the Late Oligocene-Middle Miocene when the fossil record of terrestrial mammals in
Australia is first well documented, the diprotodontians were a highly diversified order with all
but two of the eighteen families represented. Of these, only the macropods underwent a major
adaptive radiation in the late Cainozoic, the others being approximately at least as diverse in
the Late Oligocene-Middle Miocene as subsequently. These two observations suggest that the
order had a significant early Tertiary history.

Suborder: Vombatiformes

This suborder is characterised by large, relatively specialised forms in contrast to the
typically smaller, more generalised phalangerimorphians. This is a group which appears to
have flourished in the Palaeogene and to be on the decline when first encountered in the fossil
record in the Late Oligocene-Middle Miocene. Seven families are formally recognized in the
classification of Aplin & Archer (1987), and they note that there are three more from the Late
Oligocene-Middle Miocene yet to be named. Only two of the ten families are extant. This
contrasts markedly with the other diprotodontian suborder, the phalangeridians which including
one family yet to be named from the Late Oligocene-Middle Miocene has four extinct and eight
extant families. This at least equally ancient group does not appear to have declined during the
Neogene but rather radiated widely.

Infraorder: Phascolarctomorphia

Family: Phascolarctidae
Phascolarctids or koalas are characterised dentally by the presence of selenodont molars as
are ilariids and pseudocherinies. In the case of pseudocheirines, the angles formed by the outer
crests are obtuse rather than acute as in koalas (see Fig. 33), and there are secondary cusps

AUSTRALIA'S MAMMALIAN RECORD - 933

Figure 33. Upper (A) and lower (B) molars of a modem koala, Phascolarctos cinerus, x 4.

between the four principle ones; e.g. a protoconule between the protocone and paracone. The
distinction of this feature with ilariids is less marked, and there are forms such as Koobor
which could be in either group (see below).

Although there is only a single extant species of koala, Phascolarctos cinerus, the presence of
three genera with two species each (Litokoala, Madakoala, Perikoala) (Figs 34, 35) in Late
Oligocene-Middle Miocene deposits of South Australia (Woodburne, Tedford, Archer & Pledge
1987, Springer 1987) suggest a moderate diversity for the phascolarctids at that time. Later
Tertiary sites contain evidence of at most three more species based on four specimens. These
are Koobor notabilis from the Chinchilla Fauna, Koobor jimbarratti from the Bluff Downs

Figure 34. Right maxillary fragment of Perikoala palankarinnica from the Ditjimanka Local Fauna, South
Australia.

934 - RICH

Hillis

Lyn

\

Figure 35. Isolated right upper molar in (A) labial and (B) occlusal view of Litokoala kutjamarpensis
from the Kutjamarpu Local Fauna. (After Stirton, Tedford & Woodbume 1967).

Local Fauna, and Phascolarctos maris from the Sunlands Local Fauna. [Koobor may be an
ilariid (Pledge 1987c; Tedford & Woodburne 1987).] Given the paucity of material, it seems
likely that the full diversity of phascolarctids has not yet been adequately gauged. However, the
slim amount of fossil evidence is consistent with the idea based on serology and other
neontological evidence that phascolarctids are a group which originated and radiated well prior
to Late Oligocene-Middle Miocene and have subsequently declined in diversity. The only major
change now apparent within the family since the Late Oligocene-Middle Miocene has been the
reduction of the stylar cusps on the upper molars.

Figure 36. Check teeth of representatives of the Zygomaturinae. Zygomaturus trilobus, A, left P3_M2; B,

rght P3-M2. Kolopsis torus; C, left p3-M2: D, right P3-M2; x 1.16. (After Stirton, Woodbume & Plane
1967).

AUSTRALIA'S MAMMALIAN RECORD - 935

Infraorder: Vombatomorphia

Families: Diprotodontidae and Palorchestidae——"diprotodontoids"

Diprotodontids include the largest marsupial that ever lived, Diprotodon optatum, an animal
the size of a living rhinoceros. Like macropodids (kangaroos) these herbivorous forms are
characterised by the presence of two transverse lophs on their molars (the bilophodont
condition) (see Figs 36-39). Unlike them, however, diprotodontids were quadrapedal and lacked
the masseteric foramen passing from one side of the mandible to the other below the posterior
end of the tooth row (see Fig. 40). In general form their dentitions suggest similarities to the
browsing, rather than grazing, kangaroos.

.

Figure 37. Cheek teeth of representatives of the Diprotodontinae. Diprotodon optatum: A, left p3_M2;

B, right P3-M2; Pyramios alcootense: C, left P3-M?; D, right P3-M2. x 1.16 (After Stirton, Woodbume &
Plane 1967).

On the basis of the structure of the most posterior upper premolar (P3), the Diprotodontidae
may be divided into two subfamilies. The Zygomaturinac have a complex P? with four or five
cusps present. The Diprotodontinae typically have a single cusp on the P3, although
Diprotodon has a somewhat more complex condition with a horseshoe-shaped loph developed.

Formerly, two other subfamilies were placed in the Diprotodontidae, the Nototherinae and
Palorchestinae; ¢.g. Stirton, Woodburne, & Plane (1967). Archer (1977a) has shown that
members of the Nototheriinae are closely related to Diprotodon, and, therefore, that subfamily
was united with the Diprotodontinae. Archer & Bartholomai (1978) considered the
Palorchestinae to be so distinct from the other members of the family that they placed it in a
family of its own, the Palorchestidae. As noted by Stirton, Woodburne, & Plane (1967),
palorchestids are distinguished from the other diprotodontoids by the presence of a large
epitympanic fenestra immediately anterior to the ear region on the base of the skull (see Fig.
41). In this feature they bear a striking resemblance to wombats, which is unlike the condition
found in any diprotodontid. For reasons such as this, Aplin & Archer (1987) consider it likely
the two families are not particularly closely related and thus reject diprotodontoids as a formal
taxonomic catagory.

936 - RICH

\ ga S : Z , ~
Pers. BO 5 990 eo a
\ 5) B PA A \uli(fauiens . ™ Z i :

OS ESS OF Qe #2

Figure 38. Skull of the palorchestid Ngapakaldia tedfordi. A, lateral view; B, occlusal view; x 0.6.
(After Stirton 1967).

Diprotodontoids are known from the Late Oligocene Geilston Bay fauna from near Hobart,
Tasmania (Tedford et al. 1975).

In both families and both subfamilies of the diprotodontids, there is a marked increase in
body size during the Neogene, greater than that experienced by any placental group of similar
habits (Stirton, Woodburne, & Plane 1967). A similar trend occurred among mac-opodids
which at any one time were significantly smaller. Perhaps the selection pressure for this
monotonic size increase among diprotodontoids was in part, at least, owing to competition
from the ever larger macropodids.

Diprotodontoids finally became extinct in the Late Pleistocene, the cause of which is still
unresolved. Most theories for the extinction of the superfamily centre on either the marked
climatic changes at the end of the Pleistocene or the effect of the appearance of humans at least
34,000 years before their extinction (Wright 1986), or a combination of both factors.

AUSTRALIA'S MAMMALIAN RECORD - 937

Figure 39. Jaw of the palorchestid Ngapakaldia tedfordi. A, occlusal view; B, lateral view.

Family: Wynyardiidae

The type specimen of Wynyardia bassiana was collected from the Early Miocene Fossil
Bluff Sandstone near Wynyard, Tasmania. For a long time, it was the only pre-Pliocene
marsupial known from Australia. Much of the skeleton is preserved, but unfortunately the
teeth were destroyed. Although first regarded as a kangaroo (e.g. Johnstone 1888), when
Spencer (1901) named Wynyardia bassiana and published the first detailed description and
analysis, he regarded it as intermediate between other marsupials and diprotodontians. Wood
Jones (1931) restudied the specimen and allied it close to the Phalangeridae. Ride (1964)
examined the auditory region for the first time and concluded that while undoubtedly a
diprotodontian, there were many features reminiscent of non-diprotodontians and favoured
Osgood's (1921) placement of the species in its own family. Haight & Murray (1981) studied
the endocast of the brain and concluded that the closest affinities were with phalangeroids,
particularly phalangerids. Aplin (1987) re-examined the specimen yet once again and like Ride,
emphasized the auditory region in his analysis. While Ride regarded W. bassiana as a primitive
diprotodontian, he was impressed by its didelphoid features, whereas Aplin considered it clearly
diprotodontian and the didelphoid features as primitive and, therefore, of no phylogenetic
significance. Among diprotodontians, Aplin tentatively considered W. bassiana to show its
greatest affinities with vombatimorphs. His uncertainty was owing primarily to the formation
of the anterior wall of the tympanic cavity by the tympanic process of the alisphenoid rather
than the tympanic process of the squamosal, a condition more like in the phalangeridians.

After a hiatus of nearly a century, additional material tentatively referrable to the
Wynyardiidae was recovered from three different sites in Late Oligocene-Middle Miocene
deposits of South Australia. The most complete of these was a pair of articulated skeletons
found curled together at Lake Palankarinna in the Ditjimanka Local Fauna. These have been
named Muramura williamsi and given a preliminary description by Pledge (1987b).

Pledge refers the specimens Tedford et al. (1977) identified as wynyardiid in the Pinpa fauna
to Muramura. This latter material was assigned to the Wynyardiidae primarily on the similar

938 - RICH

structure of the hind limbs to the holotype of Wynyardia bassiana, there being little else
comparable between the Tasmanian and South Australian material. The dentition of the Pinpa
specimen appears to represent a stage intermediate between the selenodont condition, interpreted
as the most primitive among diprotodontians by Archer (1976d), and the more advanced
bilophodont condition.

A massateric foramen

anterior
_ mental foramen

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; mandibular
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posterior masseteric

mental foramen crest

a

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coronoid process + mandibular notch

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7

. angular
ascending ramus
9 © process
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x Ke) eminence
Mi Mp _M3 &
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— digastric process

Figure 40. Extemal views of a macropod mandible (A), Sthenurus (Simosthenurus) occidentalis), x 0.75,
illustrating the location of the masseteric foramen, a structure not present on the mandible of a
diprotodontoid (B). (A, after Tedford 1966a; B, after Stirton 1967).

On the basis of the similarity with the molar structure of the Pinpa specimen, Rich &
Archer (1979) tentatively assigned a partial skull they named Namilamadeta snideri from the

AUSTRALIA'S MAMMALIAN RECORD - 939

Tarkarooloo Local Fauna to the Wynyardiidae (Fig. 42). It appears to be most closely related
to vombatids but also shares many derived features with diprotodontoids and macropodids.

The wynyardiids as recognized here may prove to be an unnatural group when better known.
But the forms placed in the family here do all seem to be Structurally primitive diprotodontians
that may have been closely allied with the ancestors of one or more of the better known
families of the order, particularly the vombatids, that persisted into the late Cainozoic.

posterior lacerate for
fenestra rotunda

fenestra ee) '
ete 7 }bs anterior entocarotid for.
a ‘ i

posterior entocarotid for.
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promontorium
Squamosal lamina

——» ANTERIOR

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enestra incisura tympanica groove

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posterior

epitympanic

fossa

postglenoid process

Figure 41. Ventral view of the left half of the basicranium of the palorchestid Ngapakaldia tedfordi. Note
large epitympanic fenestra. (After Stirton 1967).

Family: Mariidae

The cheek teeth of this family were low crowned, had closed roots, and a crown pattern
similar to that of unworn teeth of living wombats. In concordance with the ideas of Winge
(1941), Ride (1971) and Archer (1976d) the crown pattern of these teeth supports the
hypothesis that it was derived from a selenodont condition (Tedford & Woodburne 1987).
Precisely what relation these vombatomorphians may have had with the Vombatidae is unclear
at the present time, but the sparse record does hint that there may have been a radiation of these
forms in the early to mid-Tertiary with perhaps the vombatids descending from some part of it.

The ilariids are superficially somewhat koala-like in the structure of their upper molars (see
Fig. 43A). Unlike koalas, however, the angle of the crests linking the paracone and metacone
to stylar cusps are higher and the angles formed between them more acute. Likewise, the lower

940 - RICH

a5 Pe
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Figure 42. Skull fragment of Namilamadeta snideri, a possible wynyardiid. A, lateral view; B, occlusal
view; x 1.5. (After Rich & Archer 1979).

molars differ from those of koalas and all other marsupials for that matter in having two small
circular basins formed lingual to the protoconid and paraconid (see Fig. 43B and Plate 1A).

Despite this superficial resemblance of their teeth to koalas, there is compelling evidence in
the auditory region that ilariids are vombatimorphians rather than phascolarctomorphians.
Most vombatimorphians including Ilaria illumidens have the anterior part of the tympanic
cavity formed by the tympanic process of the squamosal rather than the tympanic process of the
alisphenoid (Tedford & Woodbume 1987).

Illaria from the Late Oligocene-Middle Miocene of South Australia is the one unquestioned
member of the family. Two other genera which may belong here are Kuterintja from the
somewhat younger South Australian Late Oligocene-Middle Miocene Ngama Local Fauna and
Koobor from the Pliocene of Queensland, which was originally described as a koala and may
yet be found to belong there.

Family: Vombatidae
Two genera of modern wombats are recognised: Vombatus, the common wombat, and
Lasiorhinus, the hairy-nosed wombat. Among living marsupials, they are the only ones with

AUSTRALIA'S MAMMALIAN RECORD - 941

Figure 43. /laria illumidens. A, occlusal view of left p3-MD; B, occlusal view of right Ij-Ms5; x 0.9.

hypsodont or high-crowned cheek teeth (Fig. 44). Unworn cheek tecth show a faint selenodont
pattern, indicative of derivation from a selenodont ancestor. The teeth lack roots and continue
to grow throughout the life of the individual. In this manner, despite an appreciable amount of
abrasive grit in their diet, wombat's teeth never wear out. The one other marsupial group to
have hypsodont cheek teeth are the extinct Argyrolagidae of South America.

Although the skeletons of living wombats are rather sturdy, where known in Miocene sites,
vombatids are represented solely by rare, isolated teeth. Only one of these has yet been
formally named, Rhizophascolonus crowcrofti. Based on an isolated P? with the primitive
condition of closed roots, this species is from the Late Miocene Kutjamarpu Local Fauna
(Stirton, Tedford, & Woodburne 1967). A Late Oligocene-Middle Miocene isolated molar from
the Tarkarooloo Local Fauna was illustrated but not described or named by Rich & Archer

(1979).

942 - RICH

A B

TUE D4

Wt

TT

ff

qed

fa

Figure 44.

Isolated left upper molar of Vombatus ursinus. A, lateral view; B, occlusal view; C, anterior
view; x 15.

Pliocene records of wombats are equally sparse although the quality of the material is better

with an entire mandible of Ramsayia lemleyi known from the Bluff Downs Local Fauna
(Archer & Wade 1976).

Phascolonus gigas from the Pleistocene was the true giant among wombats, being twice
the height and length of any living wombat (Fig. 45). Stirling (1913) was of the opinion that,

Figure 45. Skull of Phascolonus gigas (above) compared to skull of Vombatus ursinus (below) drawn to
same scale.; x 0.3.

AUSTRALIA'S MAMMALIAN RECORD - 943

if anything, the humerus, radius, and ulna of P. gigas suggested that it was even a better
adapted fossorial animal than living wombats. If SO, it was certainly one of the largest
fossorial animals that ever lived. The other larger Pleistocene wombats are all included in
Ramsayia by Dawson (1981). A single molar fragment from the Early Pliocene Hamilton
Fauna suggests the presence of Lasiorhinus or Vombatus. Otherwise, these two smaller
genera of wombats are restricted to the Late Pliocene to Recent.

Warendja wakefieldi is a late Quaternary gracile wombat. All other known jaws of
wombats are massive and Support cheek teeth that are wide relative to their length. This
animal, by contrast had ever-growing hypsodont cheek teeth in a mandible no bulkier than that
of a red kangaroo (Hope & Wilkinson 1984), Unfortunately, although part of the skull and
upper dentition of this tantalising animal is now known (Flannery & Pledge 1987) none of its
postcranial skeleton has been recognised. It is likely that it was more similar in appearance to
what Miocene wombats were like than are the more familiar living ones.

Family: Thylacoleonidae

Among mammals generally, evolutionary change of dietary preference has been from
carnivorous to omnivorous to herbivorous. One of the few exceptions to this generalisation
are the thylacoleonids. Presumably, their ancestors had a dentition similar to the omnivorous
modern phalangerids. By enlarging the P3's and reducing the molars, the dentition became
more suitable for a strictly carnivorous mode of life, so much so that Van Valen (1969)
regards Thylacoleo carnifex by one measure, the ratio of the length of the carnassial shearing
surface to the total cheektooth length, as the most extreme carnivore that ever lived among
mammals (Fig. 46). In addition to the enlarged P3's, the upper molars of thylacoleonids have
only the three principle cusps, protocone, paracone, and metacone, lacking the metaconule and
any stylar cusps, in contrast to all other marsupials.

Figure 46. Thylacoleo carnifex, lateral view of skull; x 0.3. (After Lydekker 1887).

The way in which the carnivorous specialisations came about illustrates the principle that
evolution can only act on what is available for natural selection to modify and therefore does
not necessarily produce an optimum result. As the canine was reduced in the ancestral stock,
the enlarged incisors appear to have become adapted to the functions normally assumed by the
canines among most mammalian carnivores. The development of the principle shearing of the
dentition between the P3 and P3 + M2 was simply an enlargement of a system already extant
in phalangerids. This is in contrast to the principle shearing being between the last upper

944 - RICH

premolar and the first lower molar (and sometimes the ith upper molar and the ith + 1 lower
molar(s) as well) as occurs elsewhere in carnivorous mammals.

The foot structure and limb proportions provide strong support to the notion that
thylacoleonids evolved from a phalangerid condition into one suitable for grasping prey as well
as capable of climbing trees (Finch 1971, Wells & Nichol 1977).

Thylacoleonids have long been regarded as derived from phalangeroids; e.g. Owen (1840).
However, on the basis of cranial evidence which to date has only been partially presented,
Aplin & Archer (1987) have related them to vombatimorphians. Amongst the most prominent
features suggesting this allocation is the presence of a squamosal epitympanic process rather
than an alisphenoid epitympanic process forming the anterior wall of the tympanic cavity in
Thylacoleo. Murray, Wells & Plane (1987), however, noted that there is a double layer of
bone in this area in Wakaleo vanderleuri, one component formed of squamosal and the other of
alisphenoid. They tentatively advocated a derivation from a phalangeroid, most likely a
primitive burramyid on the basis particularly of dentition and skull structure.

Three different genera of Thylacoleonidae are known. Priscileo from the Late Oligocene-
Middle Miocene of South Australia is both the oldest and with the M> present, the most
primitive member of the family known (Rauscher 1987). Wakaleo is an exclusively Late
Miocene form that in reduction of the premolars is advanced over Thylacoleo, a larger Pliocene
and Pleistocene genus (Murray, Wells & Plane 1987). This suggests a second lineage leading
to Thylacoleo must have existed by the Late Miocene (Archer & Rich 1982).

—
= f=/4
SE

Figure 47. Trichosurus vulpecula mandible: A, lateral view; B, occlusal view; length of scale 10 mm.
(After Marshall 1980).

Infraorder: Phalangerida
Superfamily: Phalangeroidea

Family: Phalangeridae

Today this family is split into two subfamilies, the monospecific Ailuropinae with
Ailurops ursinus from Sulawesi, and the Phalangerinae, which is furth:r divided into two
tribes. Of these, the Trichosurini to which Trichosurus and Strigocuscus belong, are the
most widespread, being found in virtually all the habitats present today in Australia and New
Guinea as well as islands between and to the west. The Phalangerini, which includes Phalanger
and Spilocuscus, is restricted to forests and thick scrub of the Cape York Peninsula of
Australia, the Celebes on the west through New Guinea to the Solomon Islands on the east.

AUSTRALIA'S MAMMALIAN RECORD - 945

Molars of phalangerids are bunodont and among the Australian fauna can be easily confused
with some potoroids. The premolars are readily separated from those of all potoroids except
Propleopus by the greater mediolateral width of the base of the tooth relative to its length and
on the P3, there is not a horizontal blade developed, but rather the crest of the tooth descends
posteriorly at a noticeable angle (see Fig. 47). Finally, phalangerids lack the masscteric
foramen present on the mandible of potoroids below the posterior end of the tooth row.

The oldest record of the phalangerids may be from the Late Oligocene Geilston Bay Local
Fauna from near Hobart, Tasmania (Tedford et al. 1975). The earliest undoubted phalangerids
are Strigocuscus reidi and Trichosurus dicksoni from the Late Oligocene-Middle Miocene
Riversleigh locality (Flannery & Archer 1987a). These are both members of the Trichosurini,
the most derived clade of the phalangerids. The existence of this most derived clade at that time
implies that the other clades of the family must have existed as separate entities by the Late
Oligocene-Middle Miocene as well. Neither of these Riversleigh forms are strikingly different
from modern species of their respective genera, suggesting that no major changes have taken
place since the Late Oligocene-Middle Miocene. Evidently, the major phyletic events which
produced the Phalangeridae were completed by the Late Oligocene-Middle Miocene with only
modest changes occurring subsequently. Trichosurus, in particular, seems to be a generalist in
its dietary preferences and capable of living successfully in a wide variety of conditions. Thus,
the pattern of conservative morphological evolution in Australian phalangerids since the
Miocene appears quite explicable in terms of behavioural flexibility.

Family: Miralinidae

Thus far, Miralina, with at least two and perhaps as many as four species, is the only genus
known of this family. The genus occurs at a number of Late Oligocene-Middle Miocene sites
in the Etadunna and Namba Formations of South Australia.

All that is known of the species of Miralina are dental remains. These are remarkably
reminiscent of phalangerids and differ from them only in matters of detail. For example, the
P3 is mediolaterally compressed and turned inward rather than bulbous and tumed outward as in
phalangerids. Likewise, the M2 trigonid is compressed and blade-like rather than nearly as wide
as the talonid as in phalangerids (Woodburne, Pledge & Archer 1987).

The lower molars are intermediate in complexity between phalangerids and ektopodontids
with their pattern of transverse lophs broken up into a series of anteroposteriorly directed
cristids. If the P3's of Miralina were bulbous, rather than blade-like, it would be tempting to
interpret them as ektopodontids somewhat more primitive than Chunia.

Family: Ektopodontidae

Isolated molars were the first evidence of the former existence of this intriguing family (see
Fig. 48). Their appearance was so peculiar that initially they were thought to be
multituberculates. Later, it was realised that the appression facettes formed by the movement
of adjacent teeth in a row required that the ridges of cusps were oriented in the living animal at
90 degrees to the anteroposterior oricntation found in multituberculates. Ektopodon serratus
was the first species to be described and it was then tentatively regarded as a monotreme
(Stirton, Tedford, & Woodburne 1967). This initial discovery was made in the Late Miocene
Kutjamarpu Local Fauna of South Australia.

Subsequent discovery of the more primitive ektopodontid genus Chunia_ in Late Oligocene-
Middle Miocene deposits Ied to the interpretation that the family was a phalangeroid marsupial
(Woodburne & Clemens 1986c, Woodburne 1987a). Archer (1976d) pointed out how the
ektopodontid molar pattern could be regarded as structurally intermediate between what he
regards as the primitive condition in diprotodontians, selenodonty as occurs in pseudocheirines
and particularly phascolarctids, and the more advanced conditions of _bunodonty as in
phalangerids and potoroids and lophodonty as in diprotodontids and macropodids,

946 - RICH

Two species are now recognised in each of the Miocene genera Chunia and Ektopodon. In
the Early Pliocene, there is a single species, Darcius duggani, and a single unnamed tooth from
the Portland Local Fauna carries the range of the group into the Early Pleistocene. Thus, the
group appears to have declined through the late Cainozoic, perhaps its demise being brought
about by the arrival of rodents in Australia during the Pliocene.

Figure 48. Isolated left upper molar in: A, occlusal; B, ventral; C, posterior, and D, ventral views of
Ektopodon serratus from the Kutjamarpu Local Fauna, South Australia, x 4.5. Arrow indicates anterior tip of
tooth.

Superfamily: Macropodoidea

Macropodoids or kangaroos in the broadest sense are divided into two families, the
Macropodidae (kangaroos in the narrow sense, wallabies, euros, and wallaroos) and the
Potoroidae (rat-kangaroos). So long as only extant species are considered, it is quite easy to
separate the two familics morphologically. Potoroids have low crowned, bunodont molars, a
large masseteric foramen passing from one side of the dentary to the other below the posterior
end of the tooth row, the frontal bone in contact with the squamosal on the side of the
braincase (Fig. 49), and the ventral surface of the mandible is noticeably deeper below the
posterior molars than at either end (Fig. 50). Macropodids have higher, bilophodont molars, a
relatively smaller masseteric foramen, the alisphenoid bone in contact with the parietal on the
side of the braincase, and the ventral border of the mandible is not markedly convex downward.

Figure 49. Relationships of the bones on the side of the braincase in macropodoids. Frontal and
squamosal are stippled; note that these two bones are separated by the parietal and alisphenoid in the
Macropodidae, A, and contact one another in the Potoroidae, B. (After Flannery, Archer & Plane 1984).

AUSTRALIA'S MAMMALIAN RECORD - 947

Figure 50. Differences in mandibular profile and distribution of enamel on Ij in macropodids and
potoroids. Note that ventral surface of the macropod mandible, A, is equally deep along the entire length of
the row of cheek teeth in contrast to that of the potoroid, B, which is markedly deeper below the M3_4 than
anteriorly or posteriorly. In addition, the entire buccal surface of the Ij of the macropod is covered with
enamel (stippled) whereas the potoroid has a dorsal band of dentine exposed (white area above the stipple).
(After Flannery, Archer & Plane 1984).

Macropodoids are, without doubt, members of the phalangeriformes. The only major
question about their relationships is whether they are closely related to phalangerids as the
structure of their dentition suggests or are the most primitive phalangeriformes with no close
relationship to any particular other member of that group as their auditory region implies
(Flannery 1987). The evidence which suggests macropodoids are closely related to phalangerids
within the phalangeriformes are phalangerid-like features that occur at least in primitive
macropodoids, if not all of them. These include a blade-like or plagiaulacoid P3 with at least
some fine grooving; the cristid obliqua on Mo terminating anteriorly on the protostylid rather
than the protoconid; the cristid obliqua on M3_5 with a distinct, bucally convex kink; and all
molars advanced in being bunodont rather than sclenodont, the ancestral condition of
phalangeriformes as exemplified in pscudocheirids (Archer 1978a), The feature of the auditory
region that militates against this view is that the most primitive macropodoids have an
ectotympanic that is only loosely connected to the squamosal, while all other known
phalangeriformes have a more advanced condition where the two bones are fused together and in
addition, the basicranium is pneumatized (has a complex of chambers). Because auditory
characters are so complex, Flannery (1987) argued that they are less likely to have occurred
twice in the same manner than the dental features, and hence he favours the hypothesis that the
macropodoids are the most primitive phalangeriformes.

Utilising serological data for the same problem, Kirsch (1977b) concluded that
macropodoids were extremely close to phalangerids.

Family: Potoroidae
Potoroids have been regarded as representing an intermediate grade or even an actual stage in
the evolution from phalangerids to macropodids (e.g. Bensley 1903). Although bipedal, the

948 - RICH

syndactylous digits two and three are not as reduced relative to digit four, and thus the hind foot
is not as specialised as in macropodids. One potoroid, Hypsiprymnodon, even retains digit
one. Potoroid molars are so similar to phalangerids that it is often difficult to decide to which
of the two families isolated specimens should be assigned. Most potoroids possess
plagaulacoid or blade-like P3's with finer, more evenly spaced grooves present on the side than
any found in macropodids. The size of the P3's varies greatly between genera with the Late
Oligocene-Middle Miocene Wakiewakie having the longest plagaulacoid tooth of any mammal
relative to its size (Fig. 51). The structure of the brain of potoroids is more phalangerid-like
than macropodid-like. Serologically, potoroids are much more closely linked to macropodids
than to phalangerids (Kirsch 1977b).

Figure 51. Wakiewakie lawsoni mandible fragment in lateral view; x 2.5. (After Woodbume 1984).

If potoroids had masseteric foramina no larger than those of macropodids, there would be no
major stumbling blocks to the hypothesis that phalangerids gave rise to potoroids which in
turn gave rise to macropodids, However, potoroids do have much larger masseteric foramina
than macropodids, and phalangerids lack them entirely. So, in order to accept the hypothesis, it
is necessary to assume that the masseteric foramina have been secondarily reduced in macropods
after having appeared and become enlarged in the evolution from phalangerids to potoroids.

At least four, and possibly five, subfamilies of potoroids are known. Restricted to the Late
Oligocene-Middle Miocene, the Bulungamayinae have macropodid-like lophodont, rather than
bunodont, molars in a potoroid-like dentary with a large opening into the dental canal.
Although too young to be an actual intermediate between the two families, this subfamily does
demonstrate that such structually intermediate forms did exist.

The remaining potoroid subfamilies all have bunodont molars. Only one genus is placed in
the Hypsiprymnodontinae, //ypsiprymnodon, which has a single modern species and one from
the Late Oligocene-Middle Miocene, the two differing only slightly. All the remaining living
rat kangaroos are members of the Potoroinae. Generally the hypsiprymnodontines are more
primitive than the potoroines. The most obvious difference between them is in the hind feet,
where the hypsiprymnodontines possess all five digits while the potoroines have lost digit I.

The feet of the extinct Propleopinae are unknown but they can be seen to differ quite
markedly from the remaining potoroines in the structure of their molars and premolars. The
molars have a prominent cingulum around them not seen in any other macropodoid, and the
P3's are tall, prominent teeth with a broad, mediolatcrally expandea base. While other
macropodoids are known tc occasionally attack and take animal prey, propleopines appear to
have been highly specialised for a carnivorous niche, the cingula on the molars serving to
protect against damage to the gums by bone splinters and the P3's hypertrophied for a grasping
function (Archer & Flannery 1985 but see Sanson, this volume). The extinct propleopine

AUSTRALIA'S MAMMALIAN RECORD - 949

Propleopus, is the only member of the family to have reached a size as great as the larger
macropodids (Fig. 52).

The extinction of Propleopus may have had a more profound change on the terrestrial
mammalian fauna than the disappearance of a single rare kangaroo genus might at first sight
suggest. Other than Propleopus, the late Cainozoic terrestrial mammalian fauna of Australia
lacks an obvious large, cursorial predator.

Figure 52. Propleopus oscillans. Maxilla fragment: A, lateral view; B, occlusal view; x 1. Mandible
fragment, C, occlusal view; D, lateral view; x 1. (After Woods 1960).

950 - RICH

Studies of the anatomy of Thylacoleo carnifex by Finch (1982) and Thylacinus
cynocephalus by Smith (1982), demonstrate that the length of the femur relative to the tbia is
too great for a cursorial animal. Marshall (1982) makes the same comment about all the
several members of the South American Borhyaenidae for which the proportions of the hind
limb are known. A number of these extinct marsupial carnivores were in the size range of
coyotes and wolves. In South America, Marshall (1978, 1982) postulates, the adaptive zone of
the large terrestrial cursorial predator was filled not by the borhyaenids but by the flightless
phororachoid birds until the appearance of the larger fissiped carnivores on that continent in the
Early Pleistocene.

Despite the wide variety of marsupials known from Australia and South America, none of
any kind, carnivore, omnivore, or herbivore, was or is a large cursorial quadraped. In contrast,
cursorial quadrapedal placental herbivores and carnivores evolved repeatedly during the
Cainozoic. Given their mode of reproduction in which the near helpless neonate must be
continuously attached to an external nipple of the mother, the only possible evolutionary
pathway to rapid locomotion for a marsupial may have been bipedal hopping. If so, the "killer
kangaroo" Propleopus may have taken virtually the only evolutionary route to this adaptive
zone available to a marsupial carnivore. Likewise, the inability of the quadrapedal
diprotodontoids to evolve cursorial species may in part underlie their apparent disadvantage in
competition with the swift macropodoids as the grassland and open spaces began to dominate
the continent in the late Cainozoic.

Figure 53. Lateral view of the left mandible of the Late Oligocene-Middle Miocene propleopine Ekaltadeta
ima. (After Archer & Flannery 1985). Natural size.

The earliest of these carnivorous kangaroos is Ekaltadeta from the Late Oligocene-Middle
Miocene (Fig. 53). Propleopus itself appears in the fossil record at the beginning of the
Pliocene along with the earliest grazers. Like them, the species of Propleopus became larger
through the Pliocene and Pleistocene. The disappearance of Propleopus at the beginning of
the Pleistocene would have left the large cursorial predator adaptive zone open until the time of
appearance of the dingo by 3,450 years ago. As several of the more common elements of the
extinct megafauna (Diprotodon, Protemnodon, Procoptodon, and Sthenurus) have been reported
recently at 6,000 years ago by Gorecki et al. (1984) and Wright (1986), i’ is not inconceivable
that the much rarer Propleopus survived to that time as well. If so, it is tempting to consider
that Propleopus overlapped chronologically with the dingo, so that the large cursorial predator
adaptive zone was never empty during the late Cainozoic of Australia. [While there are literally
tens of thousands of individually identifiable specimens, particularly isolated teeth, of

AUSTRALIA'S MAMMALIAN RECORD - 951

Diprotodon, Protemnodon and Sthenurus in Australian museum collections, the total for
Propleopus is less than 100.]
_ The entire record of the fifth and problematical subfamily, the Paleopotoroinae, is fourteen
isolated teeth of one species, Paleopotorous priscus Flannery & Rich (1986), from a single
locality of Late Oligocene-Middle Miocene age. It is more primitive than any other potoroid
and can only be allocated to this family rather than the phalangerids, which it otherwise closely
resembles because among other things, the trigonid is not noticeably taller than the talonid.
_Potoroids presumably radiated earlier in the Tertiary than the macropodids. Among Late
Oligocene-Middle Miocene macropodoids, nine different potoroid genera have been named while
only two macropodids have been. More indicative than their numbers, however, these mid-
Tertiary potoroids were not markedly different than their Quaternary or living descendants.
Among the macropods, on the otherhand, there were no larger forms in the mid-Tertiary such as
were to form the most prominent element of the family at the end of the Cainozoic, In
addition, most living potoroids are found in moist forests, a habitat which was more widespread
in Australia during the early and mid-Tertiary than today. Thus, the meagre fossil evidence and
the apparently greater opportunities for potoroids in the past are consistent with an early or
mid-Tertiary radiation for the family, significantly earlier than the late Cainozoic radiation of
the macropodids.

Family: Macropodidae

When the macropodids first appear in the record in the Late Oligocene-Middle Miocene
about 15-25 million years ago, they are represented primarily by isolated teeth (Flannery &
Rich 1986). They are small forms similar in size and molar morphology to the modern
Setonix. It is not until the Late Miocene that larger species appear (Woodburne 1967b,
Flannery, Archer & Plane 1983). Beginning at that time, the macropodids diversified with a
bewildering rapidity into a variety of genera and species (Bartholomai 1972a). The most
extreme example of this diversification is to be found in Macropus.

Fourteen valid extant species were recognised in Macropus by Kirsch & Calaby (1977).
Unfortunately, no single worker has reviewed the fossil species of Macropus recently. But by
amalgamating the taxonomic conclusions of a number of workers in the last two decades
(Bartholomai 1975, 1978a, Flannery 1980, Tedford 1966b) if the same limits to the genus are
applied to fossils as to the extant forms, there appears to be about fourteen valid, extinct
species recognized, No Miocene specimens of Macropus are known, and it appears that they
arose and differentiated in a matter of a mere five million years into at least twenty-eight
species. This adaptive radiation of grazing forms has been attributed to the increase in the
extent of grasslands in the late Tertiary and modifications to the structure of the dentition
(Sanson 1978, 1980). The dental changes include an increase in the development of lophs or
links between the principle lophs on the molars, reduction or loss of the most posterior
premolar, and curvature of the plane of the occlusal surface on the lower teeth resulting in the
simultaneous occlusion of only one or two pairs of upper and lower molars (Fig. 54).

Although, in part, the appearance of large macropodids in the Pliocene may have been a
response to the spread of grasslands, this cannot be the entire answer for several of the large
forms that appeared at this time were browsers. Many of the genera and species that evolved as
part of this Plio-Pleistocene radiation are now extinct. The recency of this Plio-Pleistocene
radiation is reflected by the fact that the boundaries between many of the living species and
even genera are often difficult to recognize. Interbreeding is possible in several cases between
members of nominally different species and even genera (Van Gelder 1977), and serologically,
the macropodids are more homogeous than other marsupials (Kirsch 1977b).

Macropodids may be divided into three subfamilies. The Balbarinae, restricted to the Late
Oligocene-Middle Miocene, are among the most primitive macropodoids, only the potoroid
Hypsiprymnodon is comparable in this respect. Known only from dental remains, their low,

952 - RICH

compressed Mb) trigonid, straight molar tooth row and a twisted dentary are all primitive
macropodoid features. With a small, but definite, masseteric canal in the mandible, they are
clearly macropodids and not potoroids.

Figure 54. Mandibles of a browsing macropod (A, Protemnodon) and grazing macropod (B, Macropus) in
lateral view. Note the much more prominent P3 in Protemnodon and the straight rather than curved profile to a
line drawn through the tips of the molar cusps.

AUSTRALIA'S MAMMALIAN RECORD - 953

With the exception of Lagostrophus, sthenurines, the second subfamily, have reduced all the
digits of the hind foot except for the fourth. In some species of Sthenurus the reduction of the
side toes is greater than that in the modern horse. Macropodines, the third and sole surviving
subfamily, have a less reduced hind foot with the fifth digit functionally important and
parame! large. Also unlike sthenurines, the enamel on their molars is smooth rather than
crenu :

Figure 55. A, occlusal view of maxilla fragment with P3-M2 of the sthenurine Procoptodon rapha; B,

occlusal view of maxilla fragment with M24 of Procoptodon goliah; C, occlusal view of mandible fragment
with P3, M3-5 of Procoptodon goliah; x 1. (After Stirton & Marcus 1966).

The largest kangaroo that ever lived was a sthenurine, Procoptodon. This animal stood
approximately 3 m high and was, thus, well adapted to browsing in trees. The molars had an
extremely complicated pattern of lophs and crests, which apparently served to increase the
mastication efficiency of the tooth per unit area (Fig. 55). Such an increase would have been
important for a large animal to overcome the problems imposed by the fact that with linear
enlargements in size, surfaces increase only as a function of the square of the linear dimensions
whereas volume and hence mass increase as the cube. Sanson (1978) regards the dental pattern
of Procoptodon as indicative of a grazing rather than browsing animal.

Superfamily: Burramyoidea
Family: Burramyidae

Burramys parvus has the rare distinction of having been first recognised as a fossil (Broom
1896b) and only much later, found to be still alive (Anon. 1966b).

954 - RICH

Classically, burramyids have been regarded as the most primitive members of the
Diprotodontia (Bensley 1903, Kirsch 1977b). Archer (1976d) however, argues that their molar
pattern may have been derived by reduction of the crests from that of a Diprotodontian with a
selenodont condition as now occurs in pseudocheirines (Fig. 56).

Figure 56. Burramys parvus. A, palatal view of skull; B, occlusal view of mandibles; C, lateral view of
mandible; approximately x 2.7. (After Ride 1956).

Two additional species of Burramys have been described, one from the Pliocene Hamilton
Local Fauna (Turnbull, Rich & Lundelius 1987c) and the other from the I ate Oligocene-Middle
Miocene Ngama Local Fauna (Pledge 1987e). Except for slight differences in size and the
number of roots on the MS, the three species are much alike. Burramys has also been found at
Riversleigh. It, thus, appears to be not only one of the longest lived mammalian genera along
with the phalangerids Trichosurus and Strigocuscus and the macropodoids Hypsiprymnodon

AUSTRALIA'S MAMMALIAN RECORD - 955

and Bettongia, but also quite widespread during the Tertiary. Tedford et al. (1975) tentatively
allocated a lower incisor from the Late Oligocene Geilston Bay fauna to the Burramyidae.

Burramys has been frequently reported from Quaternary cave deposits across southern
Australia, Many of these sites are in places such as the Nullarbor which is quite unlike its
present restricted habitat in the Southern Alps.

Except for their smaller size, there is no consistent morphological difference for separating
all the modern burramyids or Pygmy possums from the Petaurinae and Dactylopsilinae. All
have bunodont molars. Many burramyids, but not all, have only three molars, rather than four,
whereas the other two groups always have four. In most, but not all, burramyids, the P3 is as
high as the M9; in the other two groups, the P3 is markedly lower.

Superfamily: Petauroidea

Family: Pseudocheiridae
Living petauroids can be clearly divided into two groups on the basis of their molar
structure. Dentally the more primitive, the Pseudocheiridae have selenodont molars (Fig. 57)
the pattern Archer (1976d) hypothesized as ancestral to the other conditions found in
diprotodontians. Under this hypothesis, the W-shaped ectoloph on the upper molars of
pseudocheirids was in turn derived from an ancestor with a peramelid-like dentition.

Figure 57, Pseudocheirus peregrinus. A, occlusal view of upper molar, B, occlusal view of lower molar;
xT.

Because phascolarctids, too, have selenodont molars, they were considered as closely allied
to pseudocheirids by many workers. More recent work, however, has shown on the basis of
chromosome structure and serology, that phascolarctids are more closely related to vombatids
and presumably the condition of selenodonty with phascolarctids is the retention of a primitive
character state within the diprotodontians,

Pseudocheirids are relatively common in the Tertiary as well as the Quaternary fossil record.
This abundance promises to make them useful tools for biostratigraphy as diprotodontoids have
proven in the past (Woodbume, Tedford & Archer 1987).

To date, three genera and seven species have been described in the Late Oligocene-Middle
Miocene of South Australia (Pledge 1987d, Woodbume, Tedford & Archer 1987), and two
genera and three species occur in the Early Pliocene Hamilton Local Fauna (Turnbull, Rich &
Lundelius 1987b). There are many more undescribed Tertiary pseudocheirids from the
Riversleigh district awaiting description and analysis (Woodburne, Tedford & Archer 1987).

The earliest well known pseudocheirids, the Late Oligocene-Middle Miocene genera, are all
rather distinct from one another. This is concordant with a long prior history for the family.
Between these earliest pseudocheirids, which are not directly related to the modern ones, and the
Early Pliocene forms, which are, is a significant gap in the evolutionary history of the family.
Pseudocheirids apparently are a family like macropodids that underwent a major radiation in the
late Cainozoic in response to the rapidly changing Australian environment as the continent
moved northward and became drier.

956 - RICH

Family: Petauridae
Petaurine petaurids include only two living genera, Petaurus and Gymnobelideus.
Dactylopsila, the common striped possum, have been placed in a subfamily of its own,
Dactylopsilinae, by Kirsch (1977b). Dentally, the two subfamilies are quite similar, both
having relatively simple bunodont molars (Fig. 58), which were probably derived from
ancestors with a selenodont dentition similar to pseudocheirids. The only Tertiary record of

Figure 58. Petaurus breviceps. A, occlusal view of upper molar, B, occlusal view of lower molar; x 8.5.

petaurids are rare specimens from the Late Oligocene-Middle Miocene Etadunna Faunal Zone C
of South Australia, some material from Riversleigh "...marginally more plesiomorphic
[primitive] than the modern genus [Petaurus]" (Aplin & Archer 1987) and nine isolated molars
and molar fragments from the Early Pliocene Hamilton Local Fauna of Victoria. The
Hamilton specimens appear to belong to two or three different, perhaps extant, species of
Petaurus (Turnbull, Rich & Lundelius 1987a).

Figure 59. Tarsipes spencerae skull in: A, left lateral; and, B, palatal view; x 4.25. (After Parker 1890).

AUSTRALIA'S MAMMALIAN RECORD - 957

Superfamily: Tarsipedoidea
Family: Tarsipedidae
The cheek teeth of honey possums are modified into a series of tiny pegs (Fig. 59). On the
other hand, they are far more numerous than the cheek teeth of other marsupials.
Serologically, tarsipedids stand apart from all other diprotodontians, but morphologically
their affinities clearly lie with that group (Kirsch 1977b). Although apparently separated from
other diprotodontians since the time of origin of the order, by the Palacogene at the latest, there

is only a single fossil specimen recorded of this group, a pelvis from a Quaternary cave deposit
in Western Australia (Archer 1972).

Family: Acrobatidae
Acrobates is known from a number of Quaternary sites but is unrecorded from earlier
deposits.
Both the molecular evidence and a wide variety of morphological evidence supports the idea
of a close relationship with the Tarsipedidae (Aplin & Archer 1987).

Superfamily: Phalangerida incertae sedis
Family: Pilkipildridae

Were it not for the enlarged P3's, this family would probably be placed with some difficulty
in the petaurids (Plate 1B). The other features besides the P3's which would make such an
allocation questionable are the phalangerid- and miralinid-like upper molars. On the other hand,
the low, basined lower molars are petaurid-like in their morphology. Although there is little
doubt that this family belongs in the Diprotodontia, with such a mixture of features, it cannot
be confidently allied with any particular other group in the order.

The family is known from the Late Oligocene-Middle Miocene of South Australia as well
as Riversleigh. Its common occurrence in both areas provides a basis for correlating between
them. To date, two genera with two species each have been named. At no site is the family
common, and in total, only half a dozen specimens, all dental remains, have been described
(Archer, Tedford & Rich 1987).

Order: Yalkaparidontia

Family: Yalkaparidontidae

When first discovered, the zalambdodont teeth of Yalkaparidon were so peculiar that the
animal was thought to be a placental rather than a marsupial. Subsequently a mandible with
the inflected angle preserved was found, establishing that this animal was, in fact, a marsupial.
Examination of the ultrastructure of the enamel of I1 and a premolar of Yalkaparidon cohent
corroborated the marsupial affinities of this species (Lester ef al. 1988).

Even now, its affinities within the Marsupialia are uncertain. The highly reduced molars
are reminiscent of those of Notoryctes typhlops, the marsupial mole. The base of the skull has
a combination of characters which although peramelemorph or bandicoot-like in general
aspects, display significant differences (Archer, Hand & Godthelp 1988). The incisors are most
un-bandicoot-like, being more reminiscent of diprotodontians. The lower incisor is compressed
from side to side, enlarged, procumbant, and has an enamel band confined to the
anteroventrolateral surface as is common in many diprotodontians. It is just possible that the
rostrum and dental portion of the type specimen of Yalkapariodon cohent is from a
diprotodontian and the back of the skull is from a bandicoot. However, this is quite unlikely,
and even if it eventually proves to be the case, the two taxa represented are each quite peculiar
in their own right.

958 - RICH

To date, only one genus of this order is recognized, Yalkaparidon, and the total amount of
material listed by Archer, Hand & Godthelp (1988) consists of five specimens found at four
different sites in the Late Oligocene-Middle Miocene deposits at Riversleigh, northwestern
Queensland. The present state of knowledge of this group is similar to that of the
Ektopodontidae when first described. Only the discovery of further specimens is likely to
firmly establish the affinities of this enigmatic group and affirm or deny its status as a separate
order of marsupials.

SUPERCOHORT: PLACENTALIA

Order: Chiroptera

The earliest known bat is Jcaronycteris from the Middle Eocene Green River Formation of
Wyoming, U.S.A. Except for a lesser degree of fusion of the carpal elements (Jepsen 1966), it
is not strikingly different from a living microchiropteran. Apparently bats evolved from
insectivores in the Palacocene or Late Cretaceous. The conventional view is that the
Chiroptera are a monophyletic group, but it has been proposed that mega- and
microchiropterans evolved independently from different insectivore stocks (Jones & Genoways
1970) or the megachiropterans from primates and the microchiropterans from insectivores
(Pettigrew 1986, Pettigrew & Jamieson 1987). Pettigrew & Jamieson base their hypothesis
on the sharing by primates and megachiropterans of a pattern of nerve connection between the
retina and the brain that is not found in other vertebrates. Wible & Novacek (1988) on the
basis of cranial and postcranial osteological characters plus others of the fetal membrances,
reject the primate affinities of the megachiropterans and conclude that they belong in the same
order Chiroptera with the microchiropterans. Implicitly, they also reject the idea that the two
groups independently evolved from different insectivoran stocks.

Six families of bats occur today in Australia: Pteropodidae, Emballonuridae,
Megadermatidae, Rhinolophidae, Vespertilionidae, and Milossidae (Hall 1981). All have
Palacogene fossil records on other continents, but none do in Australia (Koopman & Jones
1970).

Pteropodidae (megachiropterans or fruit bats) are characterised by simple, basin-shaped
molars. The other five familics are in the microchiroptera and have a basically tribosphenic
molar pattern with a pronounced W-shaped ecctoloph and only three molars maximum, one less
than most marsupials and the usual case in placentals (Plate 2).

Bats were the first terrestrial placental mammals to reach Australia. The oldest record of the
Chiroptera on this continent is a profusion of specimens and taxa (about twenty-five new
species with records already published for the Megadermatidae [Hand 1985], Molossidae [Hand
1990] and Hipposideridae [Sigé, Hand & Archer 1982]) from the the Late Oligocene-Middle
Miocene Riversleigh Fauna of northwest Queensland plus an isolated rhinolophid molar from
the Ngapakaldi Fauna of South Australia (Archer 1978c).

The work of Hand and her colleagues on the Riversleigh material has found the greatest
similarity with the European mid-Tertiary chiropteran faunas. However, this is probably in
part an artefact owing to the tremendous amount of work that has been put into the study of the
fossil record of the group there. By contrast, the Asian (Legendre et al. 1988) and South
American (McKenna 1980) fossil records of chiropterans are all but unkown, and the African
is only moderately better (Butler 1984). When these latter are better known, presumably it will
be the Asian that will be found to show the closest affinities with Australia, for that is the case
with the modern fauna. It is generally hypothesised that bats entered Australia from the north

AUSTRALIA'S MAMMALIAN RECORD - 959

during the Cainozoic; e.g. Simpson (1961), Hand (1985). However, Hershkovitz (1972) argued
for an origin on one of the Gondwana continents with subsequent dispersal northward.

As there is no published record of non-marine mammals in Australia in the Late Cretaccous
and Cainozoic prior to the Late Oligocene, it is uncertain how much earlier bats may have
entered the continent. Various groups of Australian bats show marked differences in the
amount of endemicity, suggesting that not all of them entered the continent at once from but
rather arrived at different times during the Cainozoic (Simpson 1961).

Unlike other mammalian orders, bats reached an adaptive plateau early in their history and
have evolved only slowly since. Once in Australia, they continued to follow this evolutionary
pattern: although a number of endemic species and a few endemic genera evolved, there were
no major structural innovations in the late Cainozoic of Australia.

Bats are present in virtually every Australian Quaternary cave site.

Hall & Richards (1979) provide a useful key to the identification of modern Australian bats.

Order: Rodentia

Rodents were among the last major placental orders to appear. Their earliest record is from
Late Palaeocene deposits of North America. By the Oligocene, they had reached all continents
except Australia and Antarctica. One of the most diverse mammalian orders, Wood (1966)
recognized a total of forty-three families.

Rodents are characterised by the presence of a single pair of evergrowing incisors in the
skull and lower jaws, which have a band of enamel on the anterior side only. Behind the
incisors is a diastema, and unlike any marsupials, there are no canines or premolars developed
between the incisors and molars in any murid rodents (Fig. 60).

Unlike all the other continents where the order is known, rodents in Australia are a
relatively undiverse group. This may be owing to their relatively late entry into Australia
(about Early Pliocene, see below) or that marsupials or birds, particularly the psittaciformes
(parrots in the broadest sense), may have already occupied many of the kinds of niches rodents
entered successfully on other continents.

All Australian rodents belong to one family, the Muridae, which originated in Southeast
Asia and spread out from there. Australian murids are divided into two subfamilies, the
Hydromyinae, restricted to New Guinea and Australia, and the nearly ubiquitous Murinae
represented by the single genus Rattus. Because of their restriction to the New Guinea-
Australia area and their earlier appearance in the fossil record there than the murines, the
Hydromyinae are frequently referred to as "The Old Endemics". The Hydromyinae, in tum, are
split into three tribes, the basically New Guinea groups Hydromyini and Uromyini and the
Australian Conilurini. Although all three tribes are represented on both land masses, the
contrast in the relative numbers of species of the various groups is quite marked. The New
Guinea Hydromyini are almost exclusively rainforest dwellers, while the Australian ones are
almost exclusively non-rainforest dwellers. This suggests that the Australian Hydromyini did
not enter Australia from New Guinea but may have bypassed it, occupying drier habitats as are
to be found on Timor in moving from southeast Asia (Flannery 1988).

The oldest records of rodents in Australia are the Early to Middle Pliocene Chinchilla Local
Fauna (Pseudomys vandycki) of southeastern Queensland (Godthelp 1990), the Early Pliocene
Bluff Downs Local Fauna of northeastern Queensland (Archer & Wade 1976), the Pliocene
Rackham's Roost Local Fauna of northwestern Queensland (Godthelp 1987, Archer, Godthelp,
Hand & Megirian 1989), and the Neogene Dog Rocks Local Fauna of southern Victoria
(Whitelaw 1989). Because the approximately contemporaneous Hamilton Local Fauna of
western Victoria totally lacks rodent remains although small mammals are abundant there, it
appears that the Early Pliocene was probably about the time the rodents reached this

960 - RICH

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AUSTRALIA'S MAMMALIAN RECORD - 961

continent, having entered from southeastern Asia via New Guinea. An earlier entry may have
been impossible Owing to the fact that the Australia-New Guinea land mass was further south
and thus further from a potential source of rodents.

The oldest detailed record of rodents available in Australia is the Pliocene Rackham's Roost
Local Fauna of northwestern Queensland. In it are exclusively Conilurini with a mixture of
living genera including Pseudomys and Zyzomys and other, extinct genera that have yet to be
named (Godthelp 1988). It appears that the Conilurini differentiated rapidly soon after entering
Australia and have evolved little since then.

In contrast, the oldest Australian and New Guinea records of the Murinae and the
Hydromyini and Uromyini are no more than 40,000 years. The difficulties in separating
modern species of Rattus from one another combined with establishing a workable diagnosis
for the genus is concordant with the view that speciation is currently underway rather than a
long completed process in this genus (Baverstock et al. 1977, Musser 1981).

Evidently, although interchange was possible between Australia and New Guinea during the
Pleistocene with lowering of sea level at various times, the Hydromyinae did not utilise the
route frequently. By contrast, species pairs of Rattus on the two land masses indicate at least
five episodes of interchange between Australia and New Guinea (Lee et al. 1981).

Watts & Aslin (1981) provide a listing of skull and dental features useful in distinguishing
the modern species of Australian rodents from one another as well as a gencral introduction to
the group including a summary of its history.

Order: Primates

The earliest known primate is Purgatorius ceratopsis from the latest Cretaceous of North
America. During the Cainozoic, the order diversified on every continent except Australia and
Antarctica (Szalay & Delson 1979). Only a single species of this order reached Australia,
Homo sapiens.

The earliest firm record of Jomo sapiens is equal to or greater than 40,000 years ago at
Upper Swan, Western Australia (Pearce & Barbetti 1981).

The occurrence at Kow Swamp, Victoria, of undoubted Homo sapiens, which none-the-less
bear a striking resemblance to Middle Pleistocene Homo erectus, raises the issue of whether
there were multiple entries into Australia of H. sapiens or a single group that diversified
morphologically once the continent was reached. The Kow Swamp specimens have been dated
at between 9,500 and 13,000 yBP (Thorne & Macumber 1972). Much older specimens from
Lake Mungo, New South Wales radiometrically dated from 24,500 to possibly as old as 30,000
yBP (Bowler et al. 1972, Bowler & Thorne 1976) are much closer to modern H. sapiens in the
structure of the skull. Several explanations have been offered to explain this (Kirk & Thorne
1976). One is that there were at least two separate migrations of //. sapiens into Australia, an
advanced group which had arrived by 30,000 yBP and an archaic group that arrived possibly as
late as 10,000 years before present. A second explanation is that there was only a single
migration of this species and that the archaic population evolved from a more advanced, gracile
stock.

The literature concerning Homo sapiens in Australia is extensive. A summary article
relating to this topic appeared in Thorne (1981).

Figure 60. Rodent skulls and jaws. A, Rattus lutreolus (Murinae); B, Hydromys chrysogaster
(Hydromyinae, Hydromyini); C, Conilurus albipes (Hydromyinae, Conilurini); D, Melomys cervinipes
(Hydromyinae, Uromyini). (After Watts & Aslin 1981).

962 - RICH

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Phylogeny of Australian terrestrial mammals primarily after Aplin & Archer (1987) and

Novacek (1986).

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(9) Microchiroptera

61.

Figure

AUSTRALIA'S MAMMALIAN RECORD - 963

Order: Carnivora

Family: Canidae
The dingo or native dog is regarded as a subspecies of the domesticated dog, Canis familiaris
dingoensis. Although a subspecies of the domesticated dog, it is unclear whether dingos were
introduced to Australia by humans or entered the continent independently. The oldest firm date
for the dingo in Australia is 3,450 + 95 yBP (Milham & Thompson 1976). However, Smyth
(1869) reported dingo remains associated with the volcanic ash deposits at Tower Hill which
have been dated at 7,300 yBP (Gill, 1973c).

HISTORY OF THE TERRESTRIAL MAMMALS OF AUSTRALIA

Fig. 61 is an attempt to illustrate the history of the various groups of Australian terrestrial
mammals. The solid vertical bars give the known range of cach group, the heavy dashed lines,
the phylogenetic relationships between them.

INTRODUCED MAMMALS

Although Europeans began introducing mammals to Australia only two centuries ago, it is
frequently the case that their remains are encountered in a context where they might be
mistaken for Pleistocene fossils. It, therefore, behooves the mammalian palacontologist
working in Australia to be familiar with the more commonly preserved elements of these
animals (Figs 62-70).

ACKNOWLEDGEMENTS

Timothy F. Flannery of the Australian Museum, Sydney, read a preliminary draft of this
chapter and provided many useful comments. Robert Hodge of Fredericksburg, Virginia,
United States of America brought the record of dingo at Tower Hill in Smyth (1869) to my
attention. Jorge V. Crisci of the Museo de La Plata, Argentina, introduced me to the NTSYS-
pc computer programme.

For the use of figures, I wish to thank the following individuals and organisations:
American Elsevier, New York, Fig. 11; American Museum of Natural History, New York,
Figs 7, 16b; American Philosophical Society, Philadelphia, Fig. 18; University of Arizona
Press, Fig. 26; Australian Museum, Sydney, Fig. 55; British Museum (Natural History),
South Kensington, Fig. 46; Bureau of Mineral Resources, Geology, and Geophysics, Canberra,
Figs 36-39, 40B, 41; Field Museum of Natural History, Chicago, Fig. 22; Harvard University
Press, Cambridge, Figs 19-21; University of Kansas Press, Lawrence, Fig. 12; Linnean
Society of New South Wales, Mossman, Fig. 8, 28; Macmillan, London, Fig. 1C; Museum
of Northern Arizona Press, Figs. 16A, C, D, 47; Queen Victoria Museum and Art Gallery,
Launceston, Figs. 29, 30A-C, 64-48, 70; Queensland Museum, Fortitude Valley, Fig. 53;
Society of Economic Paleontologists and Mineralogists, Figs. 49-52; South Australian
Museum, Adelaide, Figs 34, 35, 48; University of California Press, Berkeley, Figs 2, 5, 6, 9-
10, 13-15, 17, 23, 27A-E, 40A, 54A; University of Chicago Press, Fig. 1A; C.H.S. Watts
and H.J. Aslin, Fig. 60; Yale University Press, New Haven, Figs 3, 4; Zoological Society of
London, Fig. 56. D. Gelt prepared most of the original illustrations in this article, N. Day
many of the line drawings, and A. Bennett prepared Pl. 1. S. Morton provided all of the
photographic reproduction. The article was written on a Macintosh SE personal computer

964 - RICH

provided by Computer Knowledge (Melbourne) through the good graces of M. Smart and the
expert technical assistance of B. Hogan. The work was carried out at Monash University,
where printing was accomplished on the facilities in the Earth Sciences Department.

Figure 62. Metatarsals of a cow, A, sheep, B, horse, C, compared to a kangaroo, D, Macropus giganteus;
x 0.44.

Figure 63. Ferret dentition. Left upper tooth row, A, occlusal view; B, lateral view; right lower jaw, C,
occlusal view; left lower jaw, D, lateral view.

AUSTRALIA'S MAMMALIAN RECORD - 965

Figure 64. Canis, dog, A, lateral view of skull and right mandible, B, dorsal view of skull, and, C, palatal

view of skull; x 0.5. (After Green 1983).

966 - RICH

Figure 65. Cat, Felis, A, lateral view of skull and right mandible; B, dorsal view of skull; and, C, palatal
view of skull; x1. (After Green 1983).

AUSTRALIA'S MAMMALIAN RECORD - 967

Figure 66. Horse, Equus, A, lateral view skull and right mandible; B, dorsal view of skull; and, C,
palatal view of skull; x 0.2. (After Green 1983).

968 - RICH

Figure 67. Sheep, Ovis, A, lateral view of skull and right mandible; B, dorsal view of skull; and, C,
palatal view of skull; x 0.4. (After Green 1983).

AUSTRALIA'S MAMMALIAN RECORD - 969

Figure 68. Cow, Bos A, lateral view of skull and right mandible; B, dorsal view of skull; and C, palatal
view of skull; x 0.2. (After Green 1983).

970 - RICH

Figure 69. A, Sus, domestic pig, upper (max.) and lower (mand.) teeth; B, Homo sapiens, right lower
(top) and upper (bottom) human dentition. (A, from Schmid 1972; B, from Gregory 1920).

AUSTRALIA'S MAMMALIAN RECORD - 971

Figure 70. Hare, Lepus, A, lateral view of skull and right mandible; B, dorsal view of skull; and C,

palatal view of skull; x 0.94. (After Green 1983])

972 - RICH

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1002 - RICH

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AUSTRALIA'S MAMMALIAN RECORD - 1003

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1004 - RICH

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interpretation of late Cainozoic history. J. geol. Soc. Aust. 17: 39-51.

WYRWOLL, K.H. & DORTCH, C.E., 1978. Stone artifacts and an associated diprotodontid mandible; from
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YADAV, M., 1973. The presence of the cervical and thoracic thymus lobes in marsupials. Aust. J. Zool.
21: 285-301.

ADDENDUM

The following are valuable recent summaries of the respective areas indicated:

Baverstock, P.R., Flannery, T., Aplin, K., Birrell, J. & Kries, M., 1990. Albumin immonological
relationships of the bandicoots [Perameloidea: Marsupialai} --- a preliminary report. In
Bandicoots and Bilbies, J. H. Seebeck, P. R. Brown, R. L. Wallis & C. M. Kemper, eds. Surrey
Beatty & Sons Pty. Ltd., Sydney: 13-18. Relationships of the Perameloidea.

Flannery, T. F., 1989. Phylogeny of the Macropodoidea; a study in convergence. In Kangaroos,
Wallabies and Rat-kangaroos, G. J. Grigg, P. Jarman & I. Hume, eds., Surrey Beatty & Sons
Pty. Ltd., Sydney: 1-46. Relationships of the Macropodoidea.

Flannery, T. F., 1990. Mammals of New Guinea. Robert Brown & Associates Pty. Ltd., Carina, Qld.
Summary of the geological history and mammalian fossil record of New Guinea.

AUSTRALIA'S MAMMALIAN RECORD - 1005

APPENDIX I

AUSTRALIAN MESOZOIC AND TERTIARY TERRESTRIAL
MAMMAL LOCALITIES

Thomas H. Rich

Museum of Victoria Melbourne, Victoria 3000

Australia

Michael Archer, Suzanne J. Hand, Henk Godthelp & Jeanette Muirhead

School of Biological Sciences
University of N.S.W.
Kensington, N.S.W. 2033
Australia

Neville S. Pledge

South Australian Museum
Adelaide, South Australia 5000
Australia

Timothy F. Flannery
Australian Museum

Sydney South, N.S.W. 2000
Australia

Michael O. Woodburne & Judd A. Case
Department of Earth Sciences
University of California

Riverside, California 92521

U.S.A.

Richard H. Tedford

Department of Vertebrate Paleontology
American Museum of Natural History
New York, New York 10024

U.S.A.

William D. Turnbull
Department of Geology
Field Museum

Chicago, Illinois 60605
U.S.A.

Ernest L. Lundelius, Jr.
Department of Geological Sciences
University of Texas

Austin, Texas 78713

U.S.A.

Leaellyn S. V. Rich

Wesley College

Glen Waverley, Victoria 3150
Australia

Michael J. Whitelaw

Florida Museum of Natural History
University of Florida

Gainesville, Florida 32611

U.S.A.

Anne Kemp

Queensland Museum

P. O. Box 300

South Brisbane, Queensland 4101
Australia.

Patricia V. Rich

Earth Sciences Department
Monash University
Clayton, Victoria 3168
Australia

1006 - RICH, ET AL.

INTRODUCTION

"Much has been learned in recent years about Paleogene faunas on all the continents (save Australia)---even

Antarctica has yielded fossil mammals," (Gingerich 1988).

Gingerich'’s comment accurately reflected the relative state of knowledge about the history of the
terrestrial mammals on the Australian continent in 1988. At that time, although there was then available a
reasonably adequate overview of their Neogene Australian history, the Palaeogene was only known at the
very end, and the entire Mesozoic record consisted of two specimens. However, the recent discovery of a site
near the Palaeocene/Eocene boundary in southeastern Queensland promises to shed light on this earlier
history of Australian mammalian faunas that until now has been so poorly known.

Although a few specimens of terrestrial mammals of Tertiary age had been found in Australia prior to
1953, no concerted earlier effort had been made explicitly to discover such fossils. In that year, R.A. Stirton
and his colleagues began to systematically explore Tertiary deposits in Australia for remains of terrestrial
mammals (see chapter by R.H. Tedford, this volume). During the next thirteen years, as a result of their
discoveries, a broad outline of the evolutionary history of terrestrial mammals during the Neogene and latest
Palacogene emerged. An appropriate closing to an era was the publication two years after Stirton's death of a
summary of the state of knowledge of the Tertiary terrestrial mammals (Stirton, Tedford & Woodbume 1968).

Additional Tertiary mammal sites have been found since 1968, and knowledge of the faunas at those
previously known increased. Most of this effort has taken place in northeastern South Australia and
northwestern Queensland with lesser amounts being carried out in New Guinea, the Northern Territory, and
southem Victoria.

In 1978, Archer and Bartholomai reviewed Australian Tertiary mammals (Archer & Bartholomai 1978).
However, because their main intention was to summarize the evolution of mammals during the Tertiary as
documented by the fossil evidence, many details conceming the faunal composition and geological setting of
individual sites in the style of Stirton, Tedford & Woodbume (1968) were not repeated there. Without going
into the details presented in Stirton, Tedford & Woodburne (1968), Woodburne et a/. (1985) updated and
summarised much of the same information presented in 1968 and included a discussion of selected Pleistocene
sites.

The general conclusions drawn by Stirton, Tedford & Woodbume (1968) have withstood subsequent
testing rather well.

Elsewhere in the world, mammals appeared by the Late Triassic in North America, western Europe,
southern Africa and eastern Asia (Lillegraven ef al. 1979, Clemens 1986). Thus far, two Cretaceous
specimens (Archer et al. 1985, Rich, Flannery & Archer 1989) are the only evidence of what was probably
the first three-fourths of the time mammals were present in Australia, Evidently during this "dark age" of
Australian mammalian history, the major Australian marsupial and monotreme family-level radiations
occurred.

Use of the terms fauna and local fauna follows recommendations set forth in Tedford (1970). A local
fauna is regarded as a geologically contemporaneous group of fossils from a single site or series of sites
having limited geographic and stratigraphic distribution. The contemporaneity is established by the
taxonomic similarity of the various assemblages of fossils assigned to the local fauna and by relevant
geological data. A fauna "... represents the maximum geographic and temporal limits of a group of
organisms sharing a suite of common species ..." (Tedford 1970, p. 684). A fauna may be composed of one
or more local faunas. Where single specimens are recorded, these are designated by the site names without
the appellation fauna or local fauna, following the recommendation of Hibbard (1958).

Owing to the often fragmentary nature of most of the fossils, species-level identifications of the
elements in the various fossil assemblages often cannot be accurately made. To convey an accurate
impression of the precision of taxonomic identification warranted, the following conventions have been
employed. Usage of the abbreviation "¢f." follows the standard English translation of the Latin “conferre,"
to compare. When placed before the name of a taxon, "cf." is intended to denote that one or more specimens
exist which appear to be closely ailicd to the species of that taxon but as yct, no diagnostic characters have
been recognised that either enable specific separation or synonomy with the species of that taxon, e.g. cf.
Beltongia. A generic name followed by "sp." implies that there are specimens which can be confidently
identified to generic level but the species assignment is uncertain; e.g. Prolemnodon sp. "New genus" or "n.
sp." implies that a new laxon has been recognised but to date is unpublished.

Several new genera of mammals which have not yet been formally established, are known from the
Australian Tertiary. Where these are known to occur in more than one fauna or local fauna, they are
designated with a capital letter in two or more lists to indicate where a common taxon is recognized, e.g.
"Genus B".

AUSTRALIA'S MAMMALIAN RECORD - 1007

The discussion of individual faunas, local faunas, and sites can best be broken down into three groups:
Cretaceous and Tertiary faunas of Papua New Guinea plus easter and southeastem coastal Australia, central

and northem Australian Oligocene? through to Middle Pliocene faunas; and ?Late Pliocene - ?Early
Pleistocene faunas (Figs 24-25).

Cretaceous and Tertiary Faunas of Papua New Guinea plus eastern and southeastern
coastal Australia

Lightning Ridge Local Fauna, p. 1014
Tingamurra Local Fauna, p. 1014
Geilston Bay Local Fauna, p. 1015
Wynyard, p. 1036

Batesford Quarry, p. 1036

Canadian Lead, p. 1036

Beaumaris Local Fauna, p. 1037
Sunlands Local Fauna, p. 1038
Forsyth's Bank, p. 1039

Hamilton Local Fauna, p. 1039

Big Sink Local Fauna, p. 1040

Bluff Downs Local Fauna, p. 1041
Tara Creek, p. 1043

Lake Tyers, p. 1043

Bunga Creek, p. 1044

Great Buninyong Estate Mine, p. 1044
Awe Fauna, p. 1044

Mogorafugwa, p. 1045

Ian's Prospect, p. 1045

In this category all faunas can be dated to some degree by either radiometric and (or) marine macro- or
microfossils. These faunas are, therefore, of primary importance in serving as chronological benchmarks
against which to calibrate the sites in the other two categories. Unfortunately, only three of the faunas are
extensive, and all of these are Pliocene: Hamilton, Bluff Downs, and Awe.

Central and Northern Australian Oligocene?-Early Pliocene sites:

Ngapakaldi Fauna, p. 1016
Etadunna Formation Faunal Zones A-E

A, p. 1017

B, p. 1018

C, p. 1018

D, p. 1019

E, p. 1020
Pinpa Fauna, p. 1021
Yanda Local Fauna, p. 1022
Tarkarooloo Local Fauna, p. 1023
Wadikali Local Fauna, p. 1024
Ericmas Fauna, p. 1025
Kutjamarpu Local Fauna, p. 1025
Kangaroo Well Local Fauna, p. 1027
Riversleigh District, p. 1027
Riversleigh District System A local faunas, p. 1028
Riversleigh District D-Site Equivalent local faunas, p. 1029
Riversleigh District Cave Assemblages local faunas, p. 1029
Riversleigh District System B local faunas, p. 1030
Riversleigh District lower System C local faunas, p. 1032
Riversleigh District upper System C local faunas, p. 1033
Bullock Creek Fauna, p. 1035
Alcoota Local Fauna, p. 1037
Rackham's Roost Local Fauna, p. 1042

1008 - RICH, ET AL.

[FevavileLocal Fauna |

Floraville Local Fauna

Riversleigh System A

Etadunna Faunal Zone B

LATE OLIGOCENE -
MIDDLE MIOCENE

Cone Sees 6m ee

Cree eee Ee

Cc eee
fEuryzygoma TL T TUT TUT TTT TTT TTT TT TTT

jAcrobates TT PT TT TTT TT TT ee

Ankotarinja

[Buiungamaya | TM | | dT TT TE

Cnn BREE eee

Brachipposideros

Antechinus
Betiongia

Table I-1. Australian Tertiary terrestrial mammalian genera recorded at more than a single
locality at sites where three or more such genera are known.

AUSTRALIA'S MAMMALIAN RECORD - 1009

Gumardee
Hypsiprymnodon
agorchestes
Lasiorhinus
Nambaroo
Namilamadeta
Ngapakaldia
Obdurodon
Osphranter
Palaeopotorous
Palorchestes
Petaurus
Petramops

1010 - RICH, ET AL.

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AUSTRALIA'S MAMMALIAN RECORD - 1011

Vombatus
[Wabularoo
Wakaleo

1012 - RICH, ET AL.

0.00 0.20 0.40 0.60 0.80 1.00

Etadunna Faunal Zone B
Pinpa Fauna

Ericmas Fauna

Wadikali Local Fauna
Tarkarooloo Local Fauna
Etadunna Faunal Zone C
Etadunna Faunal Zone D
Etadunna Faunal Zone E
Kutjamarpu Local Fauna
YandaLocal Fauna
Riversleigh System A
Riversleigh System B
Riversleigh upper System C
Riversleigh lower System C
Kangaroo Well Local Fauna
Bullock Creek Fauna ——s

East of Lakes Eyre and Frome,
South Australia
ate Oligocene - Middle Miocene

Riversleigh
N.W. Qid.

Alcoota Fauna Late Miocene
Beaumaris Local Fauna ig ‘
Awe Fauna New Guinea

Sunlands Local Fauna nn Pia
Quaternary & Recent

Curramulka Local Fauna

Dog Rocks Local Fauna
Floraville Local Fauna

Bluff Downs Local Fauna
Quanbun Local Fauna
Kanunka Local Fauna
Chinchilla Local Fauna
Comadai Local Fauna
Hamilton Local Fauna

Bow Local Fauna

Rackham's Roost Local Fauna
Big Sink Local Fauna

Bone Gulch Local Fauna
Fisherman's Cliff Local Fauna

Pliocene & Quaternary

b+ +++

0.00 0.20 0.40 0.60 0.80 1.00

Simpson's Coefficient of Faunal Similarity (C/N1) where C is the number
of taxa in common between two areas being compared and N1 is the
number of taxa in the smaller assemblage.

Figure I-1. Dendrogramme showing similarities between Australian Tertiary terrestrial mammalian sites based
on the number of genera shared in common. See text for explanation.

AUSTRALIA'S MAMMALIAN RECORD - 1013

The basis for dating the faunas and local faunas here regarded as Late Oligocene-?Early Pliocene in
central and northem Australia are tenuous. In most instances, the physical relationships between sites are
uncertain.

Only in two instances are sequences of faunal assemblages established by the time-honoured principal of
superposition. The first of these is the Etadunna Formation Faunal Zones A-E plus the overlying Kutjamarpu
Local Fauna. The second is the Ericmas Fauna above the Pinpa Fauna. To date, all attempts to correlate
other assemblages within this interval have relied on assessment of the mammals alone.

In an effort to gain further insights into the temporal relationships between sites, the occurrences of all
Australian Tertiary terrestrial mammalian genera recorded at more than a single locality at sites where three or
more such genera are known were tabulated (Table I-1). Using this data as a starting point, the coefficient of
faunal resemblance suggested by Simpson (1947) was calculated for every pair of sites. The coefficient is
given by the formula C/N, where C is the number of genera in common between two sites and Nj] is the
number of genera recorded at the site with the fewest number. This matrix of similarities was then converted
into a dendrogramme (Fig. I-1) utilising the SAHN (Sequential, Agglomerative, Heirarchical, and Nested)
clustering routine of the NTSYS-pc Numerical Taxonomy and Multivariate Analysis System of computer
programmes (Rohlf 1989).

The particular algorithm employed, UPGMA for Unweighted Pair-Group Method, Arithmatic Average,
simply finds the first pair of sites which share the highest Simpson coefficient. Then the two sites are
clustered and treated as one by calculating the arithmetic average of the Simpson coefficient for the two sites
with respect to all remaining sites. Then the process is repeated until all sites are so clustered. This routine
is not designed to arrange sites in order by geological age, rather it groups them solely by the degree of
similarity as measured by the Simpson coefficient.

Because the data are quite incomplete and the studies of many faunas and local faunas are still in a
preliminary stage, the application of this algorithm provides only a general overview, not a detailed picture,
of the degree of faunal similarity between them. For example, at Riversleigh, the dendrogramme in Figure I-
1, generated solely on the basis of shared mammalian genera, suggests that System B, lower System C, and
upper System C are equally close to one another. However, the presence of wynyardiids and ilariids in
System B and their absence from both the lower and upper local faunas of System C altematively suggests
that the latter are closer faunally to one another than either is to System B.

Inspection of Fig I-1 suggests that the Late Oligocene-Middle Miocene fauna has a unity sharply
distinguishing it from from the younger faunas. Although this overall pattern is made highly evident by this
approach, attempting to group sites solely on the basis of shared mammalian genera appears to demonstrate
primarily that Late Oligocene-Middle Miocene sites within a given region such as Riversleigh are generally
more similar to one another than they are to particular sites in more distant regions. This approach provides
only a modest amount of insight into the correlation between the regions; e.g.the linking of Late Oligocene-
Middle Miocene sites east of Lakes Eyre and Frome. By contrast, the Pliocene sites show no evident
geographic clustering.

The apparent unity of the Late Oligocene-Middle Miocene faunas does accord with the idea that much of
the Quaternary and Recent fauna makes its first appearance in the Late Miocene and Pliocene, a major
tumover of the fauna having occurred immediately prior. Despite this, the component of the Late Oligocene-
Middle Miocene fauna that then carries through to the Quatemary and Recent is a remarkably high percentage
of the total compared to other continents (see Savage & Russell 1983 for non-Australian data).

It is not clear how much of the total interval of the Late Oligocene-Miocene these central and northern
Australian sites span. Attempts to establish such correlations have provided a range of results. Pollen dates
suggest Early to Middle Miocene while fossil bats suggest Early Miocene, a foraminiferan, Late Oligocene, a
radiometric date, Late Oligocene, and biogeographic considerations, Early Miocene.

The minimum age for these faunas and local faunas is established by the Beaumaris Local Fauna which
can be dated by marine macro- and micro-fossils as Mio-Pliocene.

Middle Pliocene - ? Early Pleistocene sites:

Floraville Local Fauna, p. 1046
Chinchilla Local Fauna, p. 1046
Quanbun Local Fauna, p. 1048
Bow Local Fauna, p. 1049

Krui River Local Fauna, p. 1050
Palankarinna Local Fauna, p. 1050
Kanunka Local Fauna, p. 1051
Talyawalka, p. 1051

Town Well Cave, p. 1051
Curramulka Local Fauna, p. 1052
Fisherman's Cliff Local Fauna, p. 1053

1014 - RICH, ET AL.

Bone Gulch Local Fauna, p. 1062
Dog Rocks Local Fauna, p. 1063
Coimadai Local Fauna, p. 1063
Smeaton, p. 1065
Morwell Local Fauna, p. 1065

Between the Late Pleistocene faunas dated by means of the 14c¢ technique and the Hamilton - Awe - Bluff
Downs local faunas dated as Early to Middle Pliocene by the Potassium-argon technique, there are a number
which are undated but fall somewhere in between. Unfortunately, there is not a single well-dated Late
Pliocene terrestrial mammal fauna known from Australia or Papua New Guinea. The largely undescribed
Nelson Bay Local Fauna (Flannery & Hann 1984) from near Portland, Victoria, is the single securely dated
Early Pleistocene assemblage from either Australia or Papua New Guinea (MacFadden et al. 1987). It is
associated with foraminifera indicative of Zone N22 on the Blow scale (Blow 1969). It is particularly
unfortunate that there are so few well-dated sites in this interval, because it was then that the final phases of
the change occurred from the relatively lush conditions that prevailed in central Australia during the greater
part of the Tertiary to the arid one of the modem day. In that interval, presumably, most of the arid-adapted
mammals appeared and became widespread.

The classification of marsupials we have followed is that of Aplin & Archer (1987).

EARLY CRETACEOUS

Lightning Ridge Local Fauna

Type locality: Opal field at Lightning Ridge, N.S.W. (291/2° S., 148° E.) (Archer et al. 1985).

Rock unit and age: Wallangulla Sandstone member of the Griman Creek Formation. The Griman Creek
Formation is regarded as Middle Albian on palynological evidence (Morgan 1984).
Faunai
OSTEICHTHYES
Dipnoi
Ceratodontidae
Ceratodus wollastoni Chapman 1914
Ceratodus sp.
Neoceratodontidae
Neoceratodus forsteri (Krefft 1870)
Teleostei
REPTILIA
Testudines
Plesiosauria
‘Crocodylus' selaslophensis (Etheridge, 1917)
Ornithischia
Fulgurotherium australe von Huene, 1932
Saurischia
Rapator ornitholestoides von Huene, 1932
MAMMALIA
Monotremata
?Ornithorhynchidae

Steropodon galmani Archer, Flannery, Ritchie & Molnar (1985)

Reference: Archer et al. (1985), Gaffney (1981), Molnar (1980), Molnar & Galton (1986), Rich,
Flannery & Archer (1989).

EARLY TERTIARY?

Tingamurra Local Fauna

Type locality: Near Boat Mountain, Queensland (261/6° S., 152° E.)

Rock unit and age: Fossils found in sediments associated with basalts. Basalts in the vicinity of the
fossil site have been equated with the Main Range basalts 40 km to the southwest, which have been dated at
22.1 myBP and others to the south which have been dated at 22.0-25.0 myBP (Gaffney & Bartholomai

1979). The primitive nature of the "polyprotodonts” found to date suggests an early Tertiary age for the site.
At this writing, a paper has been submitted to Nature justifying an age assignment of 54 myBP for the Tingamurra
Local Fauna (Godthelp, Archer & Hand, submitted).

AUSTRALIA'S MAMMALIAN RECORD - 1015

Fauna:
REPTILIA
Testudines
Trionychidae
Crocodilia

Squamata
AVES

MAMMALIA
Marsupialia
Reference: Gaffney (1981), Gaffney & Bartholomai (1979), Godthelp, Archer & Hand (submitted)
LATE OLIGOCENE-EARLY MIOCENE

Geilston Bay Local Fauna

Type locality: Geilston Bay on the north shore of the Derwent River across the estuary from Hobart,
Tasmania (42° 50'S, 147° 21'E.) (Tedford et al. 1975).

; Rock unit and age: Unnamed formation. Fossils found in an "... arenaceous clay, containing coarse
grit, and a few slightly rounded pebbles interbedded ...," within a travertine and beneath a basalt with a
minimum radiometric date of 22.4 + 0.5 myBP (Allport 1866, p. 74; Tedford et al. 1975).

Fauna:
MOLLUSCA
Gastropoda

"Bulinus" gunni
"Helix" tasmaniensis
MAMMALIA
Marsupialia
Dasyuromorphia
7Dasyuridae
Diprotodontia
Vombatiformes
Diprotodontidae or Palorchestidae
Phalangerida
Phalangeridae
Burramyidae

References: Ludbrook (1980), Tedford et al. (1975).

LATE OLIGOCENE-MIDDLE MIOCENE

Mammalian Faunal Zones in the Etadunna Formation,
Lake Eyre Basin

A system of five mammalian faunal zones has recently been proposed for the Etadunna Formation
exposed at Lakes Palankarinna, Kanunka, Pitikanta, and Ngapakaldi in South Australia (Woodburne ef al. in
prep.). These faunal zones are informally designated A-E from bottom to top.

Because the non-mammalian components of these zones have not as yet been distinguished (except in
the case of Zone D, e.g. Pledge 1984), a composite of the five zones is presented immediately below as the
Ngapakaldi Fauna as originally defined.

The age of the mammalian faunas of the Etadunna Formation has been variously estimated as Late
Oligocene to Early Miocene (Stirton, Tedford & Woodbume 1968) to Middle Miocene (Woodburne ef al.
1985; Rich et al. 1982). The estimate by Stirton, Tedford & Woodbume (1968) was made utilising
assumptions concerning the rate of evolution of the mammals. That reported in Woodbume et al. (1985)
relied on the correlation of fossil pollen recovered near the base of the Etadunna Formation with similar
palynomorphs recovered from Batesfordian to Balcombian marine deposits in Victoria and South Australia.
Truswell & Harris (1982) regarded pollen collected at Mammalon Hill at Lake Palankarinna to be mid-
Miocene, but with considerable uncertainty.

Additional evidence bearing on the question of the age of the Etadunna Formation has come from two
sources. Norrish & Pickering (1983) have published a rubidium/strontium radiometric determination of 25
myBP or Late Oligocene made by W. Compston of the Australian National University, Canberra. The sample
dated was collected from outcrops of the Etadunna Formation on Muloorina Station, some distance away from
all the published fossil mammal sites. From a borehole in the Etadunna Formation on the northwest side of
Lake Palankarinna have been recovered an abundant sample of the foraminifer Buliminoides sp., cf. B.

1016 - RICH, ET AL,

chattonensis. The occurrence of this foraminiferan suggests that the horizon of the Etadunna Formation thus
sampled is Late Oligocene (Lindsay 1987).

Flannery (1988) has pointed out that primitive phalangerids must have reached New Guinea before it was
separated from Australia by the Early Miocene. As undoubted phalangerids are unknown in the Etadunna
Formation and more primitive phalangeroids are known from there, it is likely that the faunas from them
must predate this time of separation. Flannery (1988) further points out that the presence of phalangerids in
the Late Oligocene Geilston Bay Local Fauna suggests it may be younger than the Etadunna Formation Faunal
Zones.

At the present time, all the evidence bearing on the age of the Etadunna Formation and its mammalian
fauna is tentative. So little is known about the history of terrestrial mammals in Australia that as long as
the final answer is somewhere in the present suggested range from the Late Oligocene to the Middle Miocene,
it will not fundamentally alter the picture of their late Cainozoic evolution. At most, the outcome will
expand or contract the chronology by a factor of two or fifteen million years.

When comparison is made with the amount of generic tumover between successive Mammalian Stage-
Ages in the North American sequence (Savage & Russell 1983, Woodburne 1987b), it appears that all the
Etadunna Faunal Zones plus the overlying Kutjamarpu Local Fauna could be readily encompassed within one
of the North American units. There is no readily observed pattem of generic replacement between these
Australian units (Fig. 67). Rather there appears to be a cohesiveness between them as compared with the
post-Middle Miocene assemblages suggesting that they represent a discrete, possibly quite brief, temporal
episode.

Ngapakaldi Fauna
(as originally defined by Stirton, Tedford & Miller 1961)

Type locality: East side of Lake Ngapakaldi, South Australia, 28° 17' S, 138° 17'E (Stirton, Tedford &
Miller 1961).

Referred localities: Lake Pitikanta, South Australia 28° 21'S, 138° 18'E. Lake Kanunka, South

Australia, 28° 23' S, 138° 18'E. Lake Palankarinna, South Australia, 28° 46'-47' S, 138° 24'E.
Rock unit and age: Etadunna Formation, Late Oligocene-Middle Miocene (Stirton, Tedford & Miller
1961).
Nonmammalian Fauna:
FORAMINIFERA
Triloculina tricarinata Ludbrook 1963
Elphididae
Elphidium advenum var. depressulum Ludbrook 1963
Miliolidae
Turmilinidae
Buliminoides sp. cf. B. chattonensis
MOLLUSCA
Gastropoda
Bulimulidae
Bothriembryon praecursor McMichael 1968
Camaenidae
Meracomelon lloydi McMichael 1968
ARTHROPODA
Ostracoda
OSTEICHTHYES
Dipnoi
Neoceratodontidae
Neoceratodus djelleh Kemp 1982
Neoceratodus gregoryi (White 1925)
Neoceratodus eyrensis (White 1925)
Neoceratodus nargun (Kemp 1983)
Neoceratodus sp. 1
Neoceratodus sp. 3
Neoceratodus sp. 4
Neoceratodus sp. 5
Teleostei
Siluriformes
Anidae
Genus and species unident.
Perciformes
Percichthyidae
Genus and species unident.

AUSTRALIA'S MAMMALIAN RECORD - 1017

AMPHIBIA
Hylidae (or Pelodryadidae)
Australobatrachus ilius Tyler 1976
Litoria sp., of. L. caerulea
Leptodactylidae
Limnodynastes archeri Tyler 1982
REPTILIA
Testudines
Chelidae
Emydura sp.
Meiolaniidae
Crocodilia
Squamata
Lacertilia
Varanidae
Scincidae
Egernia sp.
Ophidia
AVES
Pelecaniformes
Pelecanus tirarensis Miller 1966a
Phoenicopteriformes
Phoenicopteridae
Phoenicopterus novaehollandiae Miller 1963
Phoeniconotius eyrensis Miller 1963
Palaelodidae
Anseriformes
Anatidae
Falconiformes
Accipitridae
Gruiformes
Gmidae
Rallidae
Charadriiformes
Burhinidae
Columbiformes
Columbidae
Passeriformes

Comment: The non-mammalian taxa of the "classic Etadunna Fauna" is given above because as yet
representation of these taxa in Faunal Zones A-E has not been tabulated.

References: Archer (1976c 1982a), Archer & Bartholomai (1978), Archer, Plane & Pledge (1978,
1981), Estes (1984), Gaffney (1979, 1981), Ludbrook (1980), Stirton (1967a), Stirton et al. (1968), Tedford
et al. (1977), Tyler (1974, 1982), Woodburme & Tedford (1975).

Etadunna Formation Faunal Zone A [= ™"Wynyardiid" interval, includes Palankarinna South Local
Fauna
of Woodburne, Tedford, Archer & Pledge (1987)]

Type locality: West side of Lake Palankarinna, South Australia, 28° 47' S., 138° 24' E.
Rock unit and age: Members 3-5 of the Etadunna Formation exposed at Lake Palankarinna of Stirton,

Tedford & Miller (1961), Late Oligocene-Middle Miocene
Fauna:
MAMMALIA
Marsupialia
Diprotodontia
Vombatiformes
Phascolarctidae
Perikoala robustus Woodburme, Tedford, Archer & Pledge 1987
7Wynyardiidae
Muramura williamsi Pledge 1987
Phalangerida
Potoroidae
Potoroinae new genus
Pseudocheiridae
Paljara sp. A

1018 - RICH, ET AL.

References: Pledge (1987b), Woodburne (1986), Woodbume, Tedford, Archer & Pledge (1987).

Etadunna Formation Faunal Zone B
[= Ditjimanka Local Fauna of Woodburne (1986)]

Type locality: Tedford Locality, west side of Lake Palankarinna, South Australia, 28° 46' S., 138° 24°

E.
Rock unit and age: Members 6-7 of the Etadunna Formation at Lake Palankarinna of Stirton, Tedford &

Miller (1961), Late Oligocene-Middle Miocene (Woodbume et al. in prep).
Fauna:
MAMMALIA
Monotremata
Ornithorhynchidae
Obdurodon insignis Woodbume & Tedford 1975
Marsupialia
Dasyuromorphia
Dasyuridae
Ankotarinja tirarensis Archer 1976c
Ankotarinja sp. A
Keeuna woodburnei Archer 1976c
Peramelemorphia
Perameloidea
Perameloid sp. A
Diprotodontia
Vombatiformes
Phascolarctidae
Perikoala palankarinnica Stirton 1957a
Madakoala sp., cf. M. wellsi
Diprotodontidae
?Raemeotherium sp.

Tlariidae
Ilaria lawsoni Tedford & Woodbume 1987
Phalangerida
Miralinidae
Miralina doylei Woodbume, Pledge & Archer 1987
Miralina minor Woodbume, Pledge & Archer 1987

Ektopodontidae
Chunia illuminata Woodbume & Clemens 1986b

Pseudocheiridae
Pildra secundus Woodburne, Tedford & Archer 1987

Pilkipildridae
Pilkipildra taylorae Archer, Tedford & Rich 1987
Placentalia
Microchiroptera
7Rhinolophoidea

References: Archer (1976c, 1978c), Archer, Tedford & Rich (1987), Stirton (1957a), Tedford &
Woodbume (1987), Woodburne & Clemens (1986b), Woodbume, Pledge & Archer (1987), Woodburne &
Tedford (1975), Woodbume, Tedford & Archer (1987), Woodbume, Tedford, Archer & Pledge (1987).

Etadunna Formation Faunal Zone C [= Ngapakaldi Fauna]

Type locality: Ngapakaldi Quarry, east side of Lake Ngapakaldi, South Australia, 28° 17'S, 138° 17'E
(Stirton, Tedford & Miller 1961).
Referred localities: Lake Pitikanta, South Australia 28° 21'S, 138° 18'E. Lake Kanunka, South
Australia, 28° 23'S, 138° 18'E.
Rock unit and age: Etadunna Formation, Late Oligocene-Middle Miocene (Stirton, Tedford & Miller
1961).
Fauna:
MAMMALIA
Marsupialia
Dasyuromorphia

AUSTRALIA'S MAMMALIAN RECORD - 1019

Dasyuridae
New genus and species
Peramelemorphia
Perameloidea sp.
Diprotodontia
Vombatiformes
Palorchestidae
Ngapakaldia tedfordi Stirton 1967a
Ngapakaldia bonythoni Stirton 1967a
Pitikantia dailyi Stirton 1967a
Palorchestidae or Diprotodontidae
Thylacoleonidae
Priscileo pitikantensis Rauscher 1987
Phalangerida
Potoroidae
Purtia mosaicus Case 1984
Macropodidae
Nambaroo sp. A
Nambaroo sp. B
Macropodine gen. P sp. A

References: Campbell (1976; see Archer 1982a), Case (1984), Rauscher (1987), Rich & Rich (1987),
Stirton (1967a).

Etadunna Formation Faunal Zone D
[= Ngama Local Fauna of Pledge (1984)]

Type locality: Mammalon Hill, near the north end of the west side of Lake Palankarinna, South
Australia, 28° 41'S, 138° 24'E.

Referred locality: Lake Kanunka, South Australia.

Rock unit and age: Etadunna Formation, Late Oligocene-Middle Miocene. A depauperate pollen flora
immediately below site suggests a mid-Miocene age (Truswell & Harris 1982). Mammalian fauna suggests
placement between Etadunna Formation Faunal Zones A-C and the Kutjamarpu Local Fauna (Woodbume ef al.
in prep.). Ektopodon stirtoni of the Ngama Local Fauna is more advanced than Ektopodon sp., cf. E. stirtoni
from the Tarkarooloo Local Fauna and more primitive than Ektopodon serratus of the Kutjamarpu Local Fauna.
The pseudocheirids also are advanced over those from the Tarkarooloo Basin and from the stratigraphically
lower parts of the Etadunna Formation at Lake Palankarinna; one species shows affinity with a Kutjamarpu
taxon.

Fauna:
OSTEICHTHYES
Dipnoi
Neoceratodontidae
Neoceratodus sp. 4
Teleostei
Siluroidiformes
Percichthyidae
AMPHIBIA
Anura
Australobatrachus sp.
REPTILIA
Testudines
Chelidae
cf. spp. of Emydura
Meiolanidae
Meiolania sp.
Crocodilia
Gen. et sp. nov.

Scincidae
cf. spp. of Egernia
Ophidia
Boidae
AVES
Casuariiformes

1020 - RICH, ET AL.

Casuariidae

Anseriformes

Anatidae

Falconiformes

Accipitridae

Gruiformes

Rallidae

Charadriiformes

Burshinidae
Family undet.

Phoenicopteriformes

Phoenicopteridae

Columbiformes

MAMMALIA

Columbidae

Monotremata

Ornithorhynchidae
Obdurodon sp.

Marsupialia
Dasyuromorphia

Dasyuridae
of. spp. of Dasylurinja

Peramelemorphia

Perameloidea

Perameloid sp. B.

Diprotodontia

Vombatiformes

Phascolarctidae

Litokoala sp. of L. kanunkaensis
Diprotodontidae

Neohelos tirarensis Stirton 1967c
Palorchestidae

Ngapakaldia sp.
?Wynyardiidae

cf. spp. of Namilamadeta
Ilariidae?

Kuterintja ngama Pledge 1987c
Vombatidae

Phalangerida

Ektopodontidae

Ektopodon stirtoni Pledge 1986
Potoroidae

Purtia sp. A
Macropodidae

Nambaroo sp. B

Macropodine gen. P sp. B
Burramyidae

Burramys wakefieldi Pledge 1987e
Pseudocheiridae

Pildra magnus Pledge 1987d

Marlu sp., of. M. kutjamarpensis
Petauridae

References: Pledge (1984, 1986, 1987c-e), Truswell & Harris (1982).

Type locality:

Etadunna Formation Faunal Zone E

Theresa's Treasure (University of California Riverside locality RV-8506), Woodard

Promontory, west side of Lake Palankarinna, South Australia, 28° 46'S., 138° 24' E.
Referred locality: Kanunka North Local Fauna of Springer (1987), west side of Lake Kanunka, South

Australia, 28° 23' S., 138° 18' E.

Tentatively referred localities:

Etadunna Formation and overlain by the Tirari Formation.

Lungfish locality (RV-7233 and University of California, Berkeley
locality V-5766) and Lungfish North locality (RV 8456), west side of Lake Palankarinna, South Australia,
28° 46' S., 138° 24' E. These sites are in channels of the Etadunna Formation cut into lower strata of the

AUSTRALIA'S MAMMALIAN RECORD - 1021

Rock unit and age: Etadunna Formation, Late Oligocene-Middle Miocene (Stirton, Tedford & Miller
1961). On Lake Palankarinna, includes Member 9 of Stirton, Tedford & Miller (1961).

Fauna:

OSTEICHTHYES
Dipnoi
Neoceratodontidae
Neoceratodus nargun (Kemp 1983)
Neoceratodus sp. 1
Neoceratodus sp. 4
Neoceratodus sp. 5
MAMMALIA
Marsupialia
Dasyuromorphia
Dasyuridae
Keeuna sp. A
Peramelemorphia
Perameloidea
Perameloid sp. C.
Diprotodontia
Vombatiformes
Phascolarctidae
Litokoala kanunkaensis Springer 1987
Diprotodontidae
?Neohelos sp.
Palorchestidae
?Ngapakaldia sp.
Phalangerida
Potoroidae
Palaeopotorous sp. A
Macropodidae
Nambaroo saltavus Flannery & Rich 1986
Nambaroo sp. C.
?Bulungamaya sp.
Macropodine gen. P, sp. C
Macropodine gen. M, sp. A
Pseudocheiridae
Pildra sp. B

Reference: Springer (1987).

Pinpa Fauna

Type locality: West side of Lake Pinpa, South Australia, 31° 8'S, 140° 13'E.
Referred locality: Northeast bank of Billeroo Creek, South Australia, 31° 6'S, 140° 14'E. This is

Billeroo Creek Site 3 of Archer, Tedford & Rich (1987).
Rock unit and age: At top of unnamed member 1 of Namba Formation, Late Oligocene-Middle Miocene

(Callen & Tedford 1976).

Fauna:
OSTEICHTHYES
Dipnoi
Ceratodontidae
Neoceratodontidae
Neoceratodus djelleh Kemp 1982
Neoceratodus forsteri (Krefft 1870)
Neoceratodus eyrensis (White 1925)
Neoceratodus gregoryi (White 1925)
Neoceratodus nargun (Kemp 1983)
Neoceratodus sp. 1
Neoceratodus sp. 3
Neoceratodus sp. 4
Neoceratodus sp. 5
Teleostei
REPTILIA
Testudines

Chelidae
Emydura sp.

1022 - RICH, ET AL.

Meiolanidae
Crocodilia
AVES
Podicipediformes
Podicipedidae
Pelecaniformes
Pelecanidae
Pelecanus tirarensis Miller 1966a
Phalacrocoracidae
Anseriformes
Anatidae
Gruiformes
Rallidae
Charadriiformes
Burhinidae
Phoenicopteriformes
Passeriformes
MAMMALIA
Monotremata
Ornithorhynchidae
Obdurodon insignis Woodbume & Tedford 1975
Marsupialia
Diprotodontia
Vombatiformes
Phascolarctidae
Madakoala devisi Woodbume, Tedford, Archer & Pledge 1987
?Wynyardiidae
Muramura sp.
Ilariidae
Ilaria illumidens Tedford & Woodbume 1987 (=Vombatoidea genus B of
Tedford et al. 1977)
"Vombatoidea genus A' of Tedford et al. (1977)
Phalangerida
Miralinidae
Miralina sp., of. M. minor
Ektopodontidae
Chunia sp., cf. C. illuminata
Macropodidae or Potoroidae

Pseudocheiridae
Pildra antiquus Woodbume, Tedford & Archer 1987
Petauridae
Pilkipildridae
Pilkipildra handae Archer, Tedford & Rich 1987
Placentalia
Cetacea
Rhabdosteidae

References: Archer & Bartholomai (1978); Archer, Tedford & Rich (1987); Callen (1988), Callen &
Tedford (1976), Flannery & Rich (1986); Fordyce (1983); Gaffney (1981); Tedford & Woodbume (1987);
Tedford et al. (1977); Woodbume & Clemens (1986b); Woodbume, Tedford & Archer (1987); Woodbume,
Tedford, Archer & Pledge (1987).

Yanda Local Fauna

Type Locality: West side of Lake Yanda, South Australia, 31° 1/9 S., 140° 181/' E.

Rock unit and age: Namba Formation, Late Oligocene-Middle Miocene. Near the contact between the
two unnamed members of the Namba Formation. Tedford et al. (1977, fig. 2) show the fossiliferous unit at
Lake Yanda as below the contact between the two unnamed members placing it in unnamed member 1, and
regard the assemblage from there as part of the Pinpa Fauna. Investigations by two of us threw doubt upon
this physical correlation and consequently, the Yanda Local Fauna was recognized. Subsequent studies have
corroborated the original conclusion of Tedford et al. (1977). Although the Pinpa Fauna has mammalian
genera in common only with Etadunna Faunal Zones A and B, the Yanda Local Fauna not only shows
affinities with those units but also Etadunna Faunal Zone D in sharing the occurrence of Dasylurinja . For
this reason, the Yanda Local Fauna is retained as a unit distinct from the Pinpa Fauna.

AUSTRALIA'S MAMMALIAN RECORD - 1023

Fauna:
OSTEICHTHYES
Dipnoi
Neoceratodontidae
Neoceratodus djelleh Kemp 1982
Neoceratodus eyrensis (White 1925)
Neoceratodus gregoryi (White 1925)
Neoceratodus sp. 1
Neoceratodus sp. 4
Teleostei

Anseriformes
Anatidae
Phoenicopteriformes
Phoenicopteridae
Aves undet.
MAMMALIA
Marsupialia
Dasyuromorphia
Dasyuridae
Dasylurinja kokuminola Archer 1982b
Diprotodontia
Vombatiformes
Ilariidae
Ilaria sp.
Phalangerida
Miralinidae
Miralina sp., cf. M. minor
Pilkipildridae
Djilgaringa thompsonae Archer, Tedford & Rich 1987
Placentalia
Cetacea
Rhabdosteidae

References: Archer (1982a); Archer, Tedford & Rich (1987); Callen (1988); Fordyce (1983);
Woodbume, Pledge & Archer (1987).

Tarkarooloo Local Fauna

Locality: Tom O's Quarry, Lake Tarkarooloo, South Australia, 31° 81 )/y' S, 140° 61/3'B. (Rich &
Archer 1979).
Rock unit and age: Namba Formation, Late Oligocene-Middle Miocene. Near the contact between the
two unnamed members of the Namba Formation (Callen & Tedford 1976).
Fauna:
OSTEICHTHYES
Dipnoi
Neoceratodontidae
Neoceratodus gregoryi (White 1925)

Neoceratodus sp. 1
Neoceratodus sp. 5

Teleostei
REPTILIA
Testudines
Chelidae
Emydura sp
Crocodilia
Squamata
MAMMALIA
Marsupialia
Dasyuromorphia
Dasyuridae
Peramelemorphia

Perameloidea

1024 - RICH, ET AL.

Diprotodontia
Vombatiformes

Phascolarctidae
Madakoala devisi Woodbume, Tedford, Archer, & Pledge (1987)
Madakoala sp. 2

Palorchestidae
Negapakaldia tedfordi Stirton 1967a

Wynyardiidae
Namilamadeta snideri Rich & Archer 1979

Vombatidae

Phalangerida

Miralinidae
Miralina sp.

Ektopodontidae
Ektopodon sp., cf. E. stirtoni
Chunia omega Woodbume & Clemens 1986b

Potoroidae
Palaeopotorous priscus Flannery & Rich 1986
Gumardee sp.
Purtia sp.

Macropodidae
Nambaroo tarrinyeri Flannery & Rich 1986
Nambaroo saltavus Flannery & Rich 1986
Nambaroo novus Flannery & Rich 1986
Macropodine genus P

Pseudocheiridae
Pildra sp.

Pilkipildridae
Pilkipildra handae Archer, Tedford & Rich 1987

References: Archer, Tedford & Rich, (1987), Callen (1988), Flannery & Rich (1986), Flannery,
Tumbull, Rich & Lundelius (1987), Pledge (1986), Rich & Archer (1979), Rich & Rich (1982, 1987),
Woodbume & Clemens (1986b), Woodbume, Pledge & Archer (1987), Woodbume, Tedford, Archer & Pledge
(1987)

Wadikali Local Fauna

Type locality: West side of unnamed lake about 4.3 km east of Lake Tinko, South Australia, 31° 15'S,
140° 18' E.
Rock unit and age: Namba Formation, Late Oligocene-Middle Miocene
Fauna:
OSTEICHTHYES
Dipnoi
Neoceratodontidae

Neoceratodus sp. 3
MAMMALIA:
Marsupialia
Vombatiformes
Phascolarctidae
Madakoala sp.
Phalangerida
Miralinidae
Miralina sp., cf. M. minor
Pseudocheiridae
Marlu praecursor Woodbume, Tedford & Archer 1987

References: Woodbume, Pledge & Archer (1987), Woodbume, Tedford & Archer (1987), Woodbume,
Tedford, Archer & Pledge (1987)

AUSTRALIA'S MAMMALIAN RECORD - 1025
Ericmas Fauna

Type locality: Ericmas Quarry, Lake Namba, South Australia, 31° 12'S, 140° 14'E.
Referred localities: South Prospect B, Lake Namba, South Australia, 31° 14'S, 140° 14'E. Lake

Pinpa, South Australia, 31° 08'S, 140° 13'E.
Rock unit and age: At base of unnamed member 2 of the Namba Formation, Late Oligocene-Middle
Miocene (Callen & Tedford 1976).
Fauna:
OSTEICHTHYES
Dipnoi
Neoceratodontidae
Neoceratodus djelleh Kemp 1982
Neoceratodus eyrensis (White 1925)
Neoceratodus gregoryi (White 1925)
Neoceratodus sp. 1
Neoceratodus sp. 4
Teleostei

Anseriformes
Columbiformes
Columbidae
MAMMALIA
Monotremata
Ornithorhynchidae
Obdurodon insignis Woodbume & Tedford 1975
Marsupialia
Dasyuromorphia
Dasyuridae (1 or 2 species)
Diprotodontia
Vombatiformes
Phascolarctidae
Madakoala devisi Woodbume, Tedford, Archer & Pledge 1987
Madakoala wellsi Woodbume, Tedford, Archer & Pledge 1987
Diprotodontidae
Raemeotherium yatkolai Rich, Archer & Tedford 1978.
Phalangerida
Pseudocheiridae
Pildra antiquus Woodbume, Tedford & Archer 1987
Pildra secundus Woodbume, Tedford & Archer 1987
Petauridae
Placentalia
Cetacea
Rhabdosteidae

References: Archer & Bartholomai (1978), Callen (1988), Callen & Tedford (1976), Fordyce (1983),
Gaffney (1981), Rich et al. (1978), Tedford et al. (1977), Woodbume & Tedford (1975), Woodbume, Tedford
& Archer (1987), Woodbume, Tedford, Archer & Pledge (1987).

Kutjamarpu Local Fauna

Type Locality: Leaf Locality, east side of Lake Ngapakaldi, South Australia, (28° 17'S, 138° 17'E.)
(Stirton et al. 1967).
Rock unit and age: Wipajiri Formation, Late Oligocene-Middle Miocene (Stirton, Tedford & Woodburne
1967). The Wipajiri Formation is a channel-fill cut into the underlying Etadunna Formation.
Fauna:
OSTEICHTHYES
Dipnoi
Neoceratodontidae

Neoceratodus gregoryi (White 1925)
Neoceratodus djelleh Kemp 1982

1026 - RICH, ET AL.

Neoceratodus eyrensis (White 1925)
Neoceratodus sp. 3
Neoceratodus sp. 4
Neoceratodus sp. 5
Teleostei
REPTILIA
Testudines
Chelidae
Emydura sp.
Meiolanidae
Meiolania sp.
Crocodilia
Squamata
Lacertilia
Scincidae
Egernia sp.
Tiliqua sp.
Agamidae
AVES
Casuariiformes
Casuariidae
Dromaius gidju Patterson & Rich 1987
Dromornithiformes
Tromomithidae
Pelecaniformes
Pelecanidae
Pelecanus tirarensis Miller 1966a
Anseriformes
Anatidae
Charadriiformes
MAMMALIA
Marsupialia
Dasyuromorphia
Dasyuridae
Ankotarinja sp. A
Ankotarinja sp. B
Keeuna sp. A
Genus and sp. new
Peramelemorphia
Perameloidea
2 spp.
Diprotodontia
Vombatiformes
Phascolarctidae
Litokoala kutjamarpensis Stirton, Tedford & Woodbume 1967
Diprotodontidae
Neohelos tirarensis Stirton 1967c
Genus indet.
Vombatidae
Rhizophascolonus crowcrofti Striton, Tedford & Woodbume 1967
Thylacoleonidae
Wakaleo oldfieldi Clemens & Plane 1974
Phalangerida
Phalangeridae
Ektopodontidae
Ektopodon serratus Stirton, Tedford & Woodburme 1967
Potoroidae
Wakiewakie lawsoni Woodburne 1984
Bulungamaya sp. A
Bulungamaya sp. B
Macropodidae
Balbaroo sp. A
Balbaroo sp. B
Macropodine gen. P sp. C
Macropodine gen. W sp. A
Burramyidae
Burramyid sp. A-B
Pseudocheiridae
Pildra tertius Woodburne, Tedford & Archer 1987
Paljara tirarensae Woodbume, Tedford & Archer 1987

AUSTRALIA'S MAMMALIAN RECORD - 1027

Marlu kutjamarpensis Woodbume, Tedford & Archer 1987

References: Archer (1982a, b), Archer & Bartholomai (1978), Clemens & Plane (1974), Gaffney,
(1979, 1981), Godthelp, Archer & Plane (1989), Patterson & Rich (1987), Rich (1979), Stirton (1967c)
Tumbull & Lundelius (1970), Woodbume et al. (in prep.), Woodbume, Tedford & Archer (1987)

Kangaroo Well Local Fauna

Type Locality: Three kilometres northeast of Kangaroo Well, Deep Well Station, southem Northem

Territory, 24° 13'S., 134° 13'E. (Stirton et al. 1968).
Rock unit and age: Unnamed Formation, Late Oligocene-Middle Miocene (Stirton et al. 1968).
Flannery, Archer & Plane (1983) consider Balbaroo sp. from the Kangaroo Well Local Fauna to be more
primitive than the species of Balbaroo from Bullock Creek or Riversleigh System A.
Fauna:
MOLLUSCA
Gastropoda
Planorbidae
Physastra rodingae McMichael 1968
Camaenidae
Meracomelon lloydi McMichael 1968
ARTHROPODA
Ostracoda
OSTEICHTHYES
Teleostei
REPTILIA
Testudines
Crocodilia
AVES
Aves, undet. (?lost)
MAMMALIA
Marsupialia
Peramelemorphia
Perameloidea
Genus D
Diprotodontia
Phalangerida
Potoroidae
cf. Wakiewakie
Macropodidae
Balbaroo sp.

References: Flannery, Archer & Plane (1983), Gaffney (1981), Godthelp, Archer, Hand & Plane (1989),
Lloyd (1968).

Riversleigh District

Archer, Godthelp, Hand & Megirian (1989) recognize at least five significantly different intervals
among the Oligo-Miocene terrestrial mammal-bearing sites on Riversleigh Station, northwestem Queensland.
In ascending order these are (1) System A plus the D-Site Equivalents, (2) the Cave Assemblages [possibly
equivalent in age to (1)], (3) System B, (4) lower part of System C, and (5) upper part of System C. They
regard their report as a preliminary working hypothesis of the relationships between nearly one hundred
distinct sites that are known to occur on the limestone plateaus of Riversleigh Station. With this number of
sites, it is not feasible to give individual faunal lists. A representative site or sites from each interval is
given to convey the breadth of the fauna from each as it is known at the present time.

The mode of occurrence of the bulk of the numerous sites as isolated, bone-rich lenses commonly 3-5
metres in diameter with a maximum thickness of 0.5-1 metres is the major hindrance to the firm
establishment of the temporal relationships between them. The sites tend to be separated from one another
at distances of five to several hundred metres by homogencous limestone that except for the absence of bone
is similar to that of the fossil sites. The nearly uniform nature of the intervening limestone makes it all but
impossible to find inorganic markers to trace between sites. Apparently, these lenses represent pools
developed in the surrounding limestone that formed natural traps for the vertebrates found in them. These
pools were formed both in the open and within caves.

1028 - RICH, ET AL.

A more extensive body of water is hypothesized for the accumulation of many of the sites allocated to
System A. It was from such a facies that material was collected and reported on by Tedford (1967).

Locality: Riversleigh Station, northwestern Queensland (19° S., 1382/,° E.)

Riversleigh System A_ local faunas; e.g.
Riversleigh Local Fauna sensu stricto

Rock unit and age: Unnamed freshwater limestone (possibly equivalent to the Carl Creek Limestone).

Late Oligocene-Middle Miocene.
Faunai
MOLLUSCA
Gastropoda
Planorbidae
Physastra rodingae McMichael 1968
Camaenidae
Meracomelon lloydi McMichael 1968
OSTEICHTHYES
Dipnoi
Neoceratodontidae
Neoceratodus cf. N. gregoryi
Teleostei
AMPHIBIA
Anura
REPTILIA
Testudines
Squamata
Lacertilia
Ophidia
Pythonidae
Crocodilia
AVES
Dromornithiformes
Dromomithidae
Barawertornis tedfordi Rich 1979
Passeriformes
MAMMALIA
Marsupialia
Dasyuromorphia
Thylacinidae
Nimbacinus sp., cf. N. dicksoni
Dasyuridae
Genus et sp. nov.
Peramelemorphia
Perameloidea
Diprotodontia
Vombatiformes
Diprotodontidae
Bematherium angulum Tedford 1967b
Neohelos tirarensis Stirton 1967c
cf. Neohelos sp.
New zygomaturine genus (> 1)
New genus (> 1)
Palorchestidae
Ngapakaldia sp.
Palorchestes?
?Wynyardiidae
Genus indet.
Thylacoleonidae
Wakaleo sp., cf. W. oldfieldi
Phalangerida
Potoroidae
Wabularoo naughtoni Archer 1979
Bulungamaya delicata Flannery, Archer & Plane 1983
Gumardee pascuali Flannery, Archer & Plane 1983
Additional new genera of Balungamayinae
Macropodidae
Balbaroo gregoriensis Flannery, Archer & Plane 1983

AUSTRALIA'S MAMMALIAN RECORD - 1029

New genus (> 1) of Balbariinae
Macropodoidea

Galanaria tessellata Flannery, Archer & Plane 1983
Petauroidea

Gen. et sp. nov.

Comments: The original Riversleigh Local Fauna of Tedford (1967) and Stirton, Tedford & Woodbume

(1968) is part of System A.

The occurrence of species of Ngapakaldia, Neohelos, Wakaleo, and a wynyardiid in System A suggests it
correlates to one or more of the following units in South Australia: Faunal Zones A-C of the Etadunna
Formation, the Kutjamarpu and Tarkarooloo local faunas (Archer, Godthelp, Hand & Megirian 1989).

References: Archer (1979), Archer, Godthelp, Hand & Megirian (1989), Flannery, Archer & Plane

(1983), Tedford (1967)

Riversleigh District D-Site Equivalent to System A? local faunas; e.g.
Sticky Beak Local Fauna

Rock unit and age: Unnamed freshwater limestone (possibly equivalent to the Carl Creek Limestone).
Late Oligocene-Middle Miocene.

Faunai
REPTILIA
Testudines
Family indet.
Crocodilia
Crocodylidae
Genus indet.
AVES
Dromomithidae
Genus indet.
MAMMALIA
Marsupialia
Diprotodontia
Vombatiformes
Wynyardiidae
cf. Namilamadeta
Diprotodontidae
Neohelos sp. cf. N. tirarensis
Placentalia
Chiroptera

Riversleigh District Cave Assemblages local faunas; e.g.
Nooraleeba Local Fauna

Rock unit and age: Unnamed freshwater limestone. Late Oligocene-Middle Miocene.

Fauna:
MOLLUSCA
Gastropoda
OSTEICHTHYES
Teleostei
REPTILIA
Testudines
Crocodilia
AVES
Passeriformes
MAMMALIA
Marsupialia
Dasyurimorphia
Dasyuridae
Gen. et sp. nov.
Gen. indet.
Peramelomorphia
Perameloidea

Perameloid genus D
Diprotodontia

1030 - RICH, ET AL.

Phalangerida
Potoroidae
Two genera, indet.
Petauridae
Genus indet.
Placentalia
Chiroptera
Microchiroptera
Hipposideridae
Hipposideros (Brachipposideros) nooraleebus Sigé, Hand & Archer (1982)
Megadermatidae
Macroderma godthelpi Hand 1985

Comment: The bat Hipposideros (Brachipposideros) nooraleebus is considered to be a sister species of

the Burdigalian (Early Miocene) Hipposideros (Brachipposideros) aguilari from France (Sigé et al. (1982).
H. (B.) nooraleebus occurs in rocks which cut into limestones that yield a System A assemblage and is,

therefore, considered younger (Archer, Godthelp, Hand & Merigan 1989).
References: Archer (1982a), Hand (1985), Sigé, Hand & Archer (1982)

Riversleigh District System B_ local faunas; e.g.
Upper Site Local Fauna

Rock unit and age: Unnamed freshwater limestone. Late Oligocene-Middle Miocene.

Fauna:
MOLLUSCA
Gastropoda (3 spp.)
ARTHROPODA
Diplopoda (4 sp.)
Insecta
Hymenoptera
Formicidae (2 spp.)
Hemiptera
Cicadidae (1 sp.)
Coleoptera
Curculionidae (1 sp.)
Additional families? (2 spp.)
Crustacea
Isopoda (1 sp.)
OSTEICHTHYES
Teleostei (1 sp.)
AMPHIBIA
Anura
Leptodactylidae
Crinia sp.
Kyarranus (2 spp.)
Lechriodus intergerivus Tyler 1989
Limnodynastes (2 spp.)
Hylidae
Litoria (8 spp.)

Agamidae

Physignathus sp., cf. P. leseurii

Physignathus sp.

Genus and species new
Geckonidae

Genus and species new
Scincidae

Many species not yet determined
Varanidae

Genus and species indeterminate

Ophidia

Pythonidae

cf. Montypythonoides riversleighensis Smith & Plane 1985
Madtsoiidae

Genus and species new

AUSTRALIA'S MAMMALIAN RECORD

Typhlopidae
?Ramphotyphlops sp.
Elapidae
New genus? and species
Crocodilia
Crocodylidae
Genus? and species new; cf. Quinkana
AVES
Dromomithidae
Barawertornis tedfordi Rich 1979
Bullockornis sp., cf. B. planei Rich 1979
Casuariidae
Dromaius gidju Patterson & Rich 1987

Passeriformes
Menuridae
3 new genera and species
MAMMALIA
Marsupialia
Dasyuromorphia
Thylacinidae
Nimbacinus dicksoni Muirhead & Archer 1990
Thylacinus sp.
Dasyuridae
Genus and species new (> 3)
Genus? and species new
7Dasyuridae
Genus and species new
Peramelemorphia
New family 1
Genus and species new
Genus? and species new
Peroryctidae
Genus and species new
Genus? and species new (5)
Notoryctemorphia
Notoryctidae
Genus and species new
Diprotodontia
Vombatiformes
Phascolarctidae
Litokoala n. sp.
Genus? and species new
7New family
Genus and species new
Diprotodontidae
Neohelos tirarensis Stirton (1967)
cf. Neohelos sp. (> 1)
Genus and species new
Genus? and species new
7Wynyardiidae
Namilamadeta n. sp.
Tlariidae
Genus and species new
Thylacoleonidae
Wakaleo sp., cf. W. oldfieldi Clemens & Plane 1974
Genus and species new cf. Priscileo
New family
Genus and species new
Phalangerida
Phalangeridae
?Strigocuscus sp.
Trichosurus sp., cf. T. dicksoni
Potoroidae
Hypsiprymnodon n. sp.
Ekaltadeta sp., cf. E. ima Archer & Flannery 1985
cf. Wabularoo n. sp
of. Gumardee n. sp
Genus and species new
Genus? and species new (2)

- 1031

1032 - RICH, ET AL.

Wakiewakie lawsoni Woodbume 1984
Macropodidae

2Nambaroo n. spp. (2)
Burramyidae

Burramys n. sp.

Cercartetus n. sp.
Pseudocheiridae

Paljara n. sp.

Pildra n. sp.

cf. Pseudochirops spp. (3)
Petauridae?

New genus and new species
Acrobatidae

New genus? cf. Acrobates

Genus? and species new

Pilkipildridae
Djilgaringa sp.
New family
Genus ana species new (2)
New family?
Genus and species new
Yalkaparidontia
Yalkaparidontidae
Yalkaparidon coheni Archer, Hand & Godthelp 1988
?Marsupialia
Yingabalanaridae
Yingabalanara richardsoni Archer, Every, Godthelp, Hand & Scally 1990
Placentalia
Chiroptera
Megadermatidae
Macroderma sp.
Hipposideridae
Brachipposideros spp. (5)
Genus & sp. indet. (2)
Molossidae
Genus & sp. indet. (> 2)
Family?

Genus & sp. indet.

Comments: System B local faunas are most similar to the Tarkarooloo and Kutjamarpu local faunas of
South Australia in part because of the occurrence of Wakiewakie lawsoni, Wakaleo sp., of. W. oldfield,
Neohelos sp., cf. N. tirarensis, Dromaius gidgu, and species of Namilamadeta, Litokoala and Nambaroo.

Reference: Archer, Godthelp, Hand & Megirian (1989)

Riversleigh District lower System C local faunas; e.g.
Dwornamor Local Fauna

Rock unit and age: Unnamed freshwater limestone. Late Oligocene-Middle Miocene.

Faunai
MAMMALIA
Dasyuromorphia
Thylacinidae
Genus (> 1) and species (2)
Dasyuridae
Genus (> 1) and species (6)
Peramelemorphia
New family 1
Genus (> 1) and species (3)
Peroryctidae
Genus (> 1) and species (7)
Diprotodontia
Vombatiformes
Phascolarctidae
Litokoala n. sp.
Genus? and species new
Diprotodontidae
Neohelos sp., cf. N. tirarensis

AUSTRALIA'S MAMMALIAN RECORD - 1033

Nimbadon sp.
Palorchestidae
New genus and sp. (> 1)
Thylacoleonidae
Wakaleo sp., cf. W. oldfieldi
Phalangerida
Phalangeridae
Strigocuscus reidi Flannery & Archer 1987a
Trichosurus dicksoni Flannery & Archer 1987a
Potoroidae
Hypsiprymnodon bartholomaii Flannery & Archer 1987b
Ekaltadeta ima Archer & Flannery 1985
New genus (> 1) and n. sp. (> 5)
Macropodidae
New genus (> 1) and n. sp. (> 2)
Pseudocheiridae
Paljara n. sp.
Pildra n. sp.
cf. Pseudochirops spp. (3)
New genera and n. sp. (> 4)
?Pseudocheiridae
New subfamilies, genera, and species
Petauroidea
New genus (1) and n. sp. (> 2)
Burramyidae
Burramys n. sp.
Acrobatidae
Genus? and species new
Pilkipildridae
Djilgaringa gillespieae Archer, Tedford & Rich 1987
Yalkaparidontia
Yalkaparidontidae
Yalkaparidon jonesi Archer, Hand & Godthelp 1988
Placentalia
Chiroptera
Microchiroptera
Megadermatidae
Macroderma godthelpi Hand 1985
Macroderma? ("Dwomamor variant")
Hipposideridae
Brachipposideros spp. (4)

Comments: The local faunas in the lower part of System C are regarded as younger than the Kutjamarpu
and Tarkarooloo local faunas but older than the Bullock Creek Fauna on the basis of some of their
zygomaturines and thylacinids and their lack of wynyardiids, abundant and widespread elements in the System
B local faunas (Archer, Godthelp, Hand & Megirian 1989).

References: Archer & Flannery (1985), Archer, Godthelp, Hand & Megirian (1989), Archer, Hand &
Godthelp (1988), Flannery & Archer (1987a,b), Hand (1985, 1990)

Riversleigh District upper System C local faunas; e.g.
Henk's Hollow Local Fauna

Rock unit and age: Unnamed freshwater limestone. Late Oligocene-Middle Miocene.

Fauna:
OSTEICHTHYES
Dipnoi
Neoceratodontidae
Neoceratodus eyrensis (White 1925)
Neoceratodus gregoryi (White 1925)
Neoceratodus nargun (Kemp 1983)
Neoceratodus sp. 3
Neoceratodus sp. 5
Teleostei
AMPHIBIA
REPTILIA

Squamata

1034 - RICH, ET AL.

Lacertilia
Agamidae
Physignathus sp
Geckonidae
Genus and species indeterminate
Scincidae
Genus and species indeterminate
Ophidia
Pythonidae
Montypythonoides riversleighensis Smith & Plane 1985
Madtsoiidae
Genus and species new
Family indet.
Crocodilia
Crocodylidae
cf. Quinkana sp.

Marsupialia
Dasyuromorphia
Dasyuridae
Genus and species new (3)
Thylacinidae
Nimbacinus dicksoni Muirhead and Archer 1990
Peramelemorphia
New family
Genus and species new
Peroryctidae
Genus and species new (2)
Diprotodontia
Vombatiformes
Phascolarctidae
Genus and species new
Thylacoleonidae
Wakaleo sp.
Diprotodontidae
Neohelos sp.
Nimbadon sp.
Phalangerida
Phalangeridae
Trichosurus dicksoni Flannery & Archer 1987a
Strigocuscus sp.
Potoroidae
Ekaltadeta ima Archer & Flannery 1985
Genus and species new (3)
Macropodidae
Genus and species new (2)
Burramyidae
Burramys n. sp.
Pseudocheiridae
Pseudochirops sp.
Genus and species new
Acrobatidae
Genus and species new
Pilkipildridae
Genus and species new
Placentalia
Chiroptera
Megadermatidae
Genus & sp. indet. (2)
Hipposideridae
Genus & sp. indet. (4)

Comments: The younger, higher local faunas of System C may be closer to the Alcoota Local Fauna
than the Bullock Creek Fauna because of the presence of a pre-Kolopsis-like zygomaturine (Archer, Godthelp,
Hand & Megirian 1989).

AUSTRALIA'S MAMMALIAN RECORD - 1035

Bullock Creek Fauna

Type locality: 26 kilometres southeast of Camfield Station Homestead, north central Northern Territory

(17° 7'S., 131° 31-32’ E.).

Rock unit and age: Camfield beds, Late Oligocene-Middle Miocene. The type specimen of Neohelos
tirarensis was found in the Kutjamarpu Local Fauna. Neohelos sp., cf.N. tirarensis has been collected from
the Riversleigh Systems A, B, and C and Bullock Creek faunas. It is considered more primitive than
Zygomaturus keanei from Palankarinna Local Fauna, Zygomaturus gilli from the Beaumaris Local Fauna, and
Kolopsis torus from the Alcoota Local Fauna by Stirton, Tedford & Woodburne (1968). Clemens & Plane
(1974) conclude that the differences in stage-of-evolution of Wakaleo oldfieldi from the Kutjamarpu Local
Fauna and Wakaleo vanderleuri from the Bullock Creek Fauna suggest a younger age of the latter relative to
the former.

Fauna:
MOLLUSCA
Gastropoda
OSTEICHTHYES
Dipnoi
Neoceratodontidae
Neoceratodus sp. 1
Neoceratodus sp. 3
Neoceratodus sp. 5
REPTILIA
Testudines
Meiolanidae
Meiolania sp.
Squamata
Ophidia
Pythonidae
Morelia antiquus Smith & Plane 1985
Crocodilia
AVES
Casuariiformes
Casuariidae
Dromaius sp.
Dromornithiformes
Dromomithidae
Bullockornis planei Rich 1979
Bullockornis sp.
MAMMALIA
Marsupialia
Peramelemorphia
Perameloidea
Genus D
Dasyuromorphia
Thylacinidae
Nimbacinus dicksoni Muirhead & Archer 1990
Diprotodontia
Vombatiformes
Diprotodontidae
Neohelos sp., cf. N. tirarensis Stirton 1967c
Nimbadon sp.
Palorchestidae
Propalorchestes novaculacephalus Murray 1986
Thylacoleonidae
Wakaleo vanderleuri Clemens & Plane 1974
Phalangerida
Potoroidae
Genus undet.
Macropodidae
Balbaroo camfieldensis Flannery, Archer & Plane (1983)
cf. Dorcopsis

References: Megirian (1989), Patterson & Rich (1987), Plane & Gatehouse (1968), Rich (1979),
Flannery, Archer & Plane (1983); Murray, Wells & Plane (1987), Smith & Plane (1985).

1036 - RICH, ET AL.

EARLY MIOCENE
Wynyard

Locality: Fossil Bluff, Wynyard, Tasmania (40° 58.8'S, 145° 43.9'E.).

Rock unit and age: Fossil Bluff Sandstone. "Contains marine mega- and microfossils that compare best
with those characteristic of the Longfordian stage of Victoria whose planktonic foraminifera in turn compare
best with those of the Aquitanian (Early Miocene) of Europe (Quilty 1966)." (Stirton, Tedford &
Woodburne, 1968, p. 8).

Fauna:
MAMMALIA
Marsupialia
Diprotodontia
Vombatiformes
Wynyardiidae
Wynyardia bassiana Spencer 1901

MIDDLE MIOCENE

Batesford Quarry

Locality: Australia Portland Cement Ltd's Batesford Quarry, Victoria (38° 61/5'S., 144° 171 /y'B.).
Rock unit and age: Batesford Limestone. The exposures of the Batesford Limestone in the Batesford

Quarry were selected by Singleton (1941) as the type of the Batesfordian Stage. The Batesfordian is
considered to be Middle Miocene in age, between 15 and 16 myBP (Abele et al. 1988).

Fauna:
MAMMALIA
Marsupialia
Diprotodontia?

Comment: This record is based on a single, edentulous jaw fragment (Museum of Victoria, NMV
P164,999) of a marsupial about the size of Diprotodon optatum, yet is clearly quite distinct from that
species.

Canadian Lead

Locality: Canadian Lead. Beginning on the west bank of Cooyal Creek opposite Home Rule (formerly
Wyaldra), N.S.W. and extends westward approximately 3 km from that point (32° 25'S., 149° 341)5-
361/,' E.)

Rock unit and age: Auriferous clays and gravels filling both a palaeovalley cut into limestone and cave
systems originally opening into that valley to an average depth of 27 metres (Jones 1940). Across Cooyal
Creek, the Canadian Lead passes into the Home Rule Lead ([bid.). Neither is capped by basalts but a regional
study by Dulhunty (1964) concluded that all the auriferous gravels in the Gulgong Gold Fields predated basalts
with radiometric ages of 14.8 + 1.2 myBP and 13.8 + 1.1 myBP, Middle Miocene (Dulhunty 1971). A late

Early to Middle Miocene palynofloral has been reported from kaolin recovered from the Home Rule Lead
(McMinn 1981).

Fauna:
AVES
Dromornithiformes

Dromomithidae
MAMMALIA
Monotremata
Tachyglossidae
Zaglossus robusta (Dun 1895)
Marsupialia
Diprotodontia

References: Dulhunty (1964, 1971), Jones (1940), McMinn (1981), Murray (1978b), Rich (1979).

AUSTRALIA'S MAMMALIAN RECORD - 1037

LATE MIOCENE

Beaumaris Local Fauna

Type locality: Sea cliff exposures on Port Phillip Bay at Beaumaris, Victoria, 37° 59'S.,
145° 03' E.

Rock unit and age: Base of the Black Rock Sandstone. Singleton (1941) selected the Black Rock
Sandstone at Beaumaris as the type for his Cheltenhamian Stage. Based on its content of marine
macroinvertebrates, the Cheltenhamian is tentatively regarded as Mio-Pliocene by Abele et al. (1988), who

noted Ludbrook's opinion that the age of the stratotype of the stage has not been satisfactorily resolved
(Ludbrook 1973).

Fauna:
AVES
Sphenisciformes
Spheniscidae
Pseudaptenodytes macraei Simpson 1970d
?Pseudaptenodytes minor Simpson 1970d
Procellariiformes
Diomedeidae
Diomedea thyridata Wilkinson 1969
MAMMALIA
Marsupialia
Diprotodontia
Vombatiformes
Diprotodontidae
Zygomaturus gilli Stirton 1957b
Kolopsis sp.
Palorchestidae

Palorchestes sp.

References: Gill (1957b), Hall & Pritchard (1897), Simpson (1970d), Stirton (1957b), Stirton,
Woodbume & Plane (1967), Wilkins (1963), Wilkinson (1969), Woodbume (1969), Woods (1958, 1962).

Alcoota Local Fauna

Type locality: Seven kilometres southwest of Alcoota Station HS, central Norther Ternitory,
22° 52'S., 134° 27' E.

Rock unit and age: Waite Formation, Late Miocene.

"The thylacines and macropodids are more primitive than later descendants of their respective lineages,
these being known mainly from Pleistocene deposits. Palorchestes painei is more primitive than P. azael
(Pleistocene) and P. parvus (Chinchilla Local Fauna). The Alcoota species is closer, however, to those forms
than to the palorchestines of the Ngapakaldi fauna..... While the zygomaturine Plaisiodon centralis is more
advanced than Neohelos tirarensis (Kutjamarpu fauna), it apparently gave rise to no later forms. Kolopsis
torus is rather closely aligned in an ancestor-descendant relationship with N. tirarensis on the one hand and
with Zygomaturus gilli on the other. K. torus is also more primitive than K. rotundus (Awe fauna). The
Alcoota Local Fauna is thus post-Kutjamarpu-pre-Beaumaris in age.” (Stirton ef al. 1968).

The presence of Neohelos sp., cf. N. tirarensis in local faunas of Riversleigh Systems A, B, and C and
the Bullock Creek Fauna further supports the notion that like Kutjamarpu, these faunas precede Alcoota.

Fauna:
REPTILIA
Testudines
Crocodilia
Crocodylidae
Crocodylus sp.
cf. Pallimnarchus sp.
AVES
Casuariiformes
Casuariidae
Dromaiinae

1038 - RICH, ET AL.

Dromornithiformes
Dromomithidac
Dromornis stirtont Rich 1979
IIbandornis lawsoni Rich 1979
Ibandornis woodburnei Rich 1979
Anseriformes
Anatidac
Falconiformes
Accipitridac
Phoenicopteriformes
Phoenicopteridae (Rich, in prep.)
MAMMALIA
Marsupialia
Dasyuromorphia
Thylacinidac
Thylacinus potens Woodburne 1967b
Diprotodontia
Vombatiformes
Diprotodontidac
Kolopsis torus Woodbume 1967a
Plaisiodon centralis Woodburne 1967a
Pyramios alcootense Woodbume 1967a
Palorchestidac
Palorchestes painei Woodbume 1967b
Propalorchestes novaculocephalus Murray 1986
?Vombatidac
Thylacoleonidac
Wakaleo alcootaensis Archer & Rich 1982
Phalangerida
Macropodidac
Dorcopsoides fossilis Woodbume 1967b
Hadronomas puckridgi Woodburne 1967b
Another genus
Pseudocheiridac
Pseudochirops n.sp.

References: Archer (1982b), Archer & Bartholomai (1978), Archer & Rich (1982), Bartholomai
(1978a), Gaffney (1981), Lloyd (1968) Newsome & Rochow (1964), Patterson & Rich (1987), Rich (1979),
Woodbume (1967a, b).

EARLY PLIOCENE

Sunlands Local Fauna

Locality: Near the Sunlands Pumping Station, left bank of the River Murray, 8 km west of Waikerie,

South Australia, 34° 09'S., 1399 55" B.
Rock unit_and age: Lower part of the Loxton Sands, Early Pliocene (Kalimnan) on stratigraphic and
micropalacontological evidence (Lindsay 1965; Ludbrook 1961).

Kau na .

MOLLUSCA

BRYOZOA

CRUSTACEA

ECHINOIDEA

CHONDRICHTHYES

Elasmobranchi

Heterodontus sp., of. H. cainoxoicus
Odontaspis sp., of. O. acutissima
Isurus hastalis
Lamna sp., ef. L. cattica
of. Carcharodon megalodon
Orectolobus gippslandicus
Mustelus sp.
Carcharhinus sp., of. C. brachyurus
Galeorhinus sp., of. G. australis
Galeocerdo aduncus

AUSTRALIA'S MAMMALIAN RECORD - 1039

cf. Sphyrna sp.
Pristiophorus lanceolatus
cf. Myliobatis sp.
OSTEICHTHYES
Diodontidae
Diodon formosus
Monacanthidae sp. indet.
of. Labroidei
MAMMALIA
Marsupialia
Diprotodontia
Vombatiformes
Phascolarctidae
Phascolarctos maris Pledge 1987a
Diprotodontidae
cf. Zygomaturus
Phalangerida
Macropodidae
Dorcopsis sp.
Placentalia
Cetacea
Odontoceti
Delphinidae
cf. Steno
Mysticeti
Balaenidae indet.

References: Pledge (1985, 1987a).

Forsyth's Bank

Locality: West side of the Grange Bum approximately 8 kilometres west of Hamilton, Victoria
(37° 43' 42 + 03"S., 141° 56' 40 + 04" E.).
Rock unit and age: Grange Burn Formation, Early Pliocene. Ludbrook (1973) considers the molluscan
fauna from the Grange Burn Formation as indicative of the Kalimnan Stage.
Fauna:
MAMMALIA
Marsupialia
Diprotodontia
Phalangerida

Macropodidae
Protemnodon sp.

References: Abele et al. (1988), Boutakoff & Sprigg (1953), Gill (1957b), Ludbrook (1973), Stirton
(1957b), Tedford (1966a).

Hamilton Local Fauna

Type locality: South side of the Grange Bum about 100 metres downstream from a waterfall and
oximately 7 kilometres west of Hamilton, Victoria (37° 42' 58 + 03" S., 141° 57' 144 04" E.).

Rock unit and age: Unnamed lithic unit overlying the marine Grange Burn Formation and underlying a
basalt. The unnamed lithic unit which produced the fossil mammals is about 1.3 metres thick and is

interpreted as a fossil soil by Gill (1957b).
Both units bracketing the productive lens have been dated as Early Pliocene. Tumbull et al. (1965)

published a potassium-argon date of 4.35 + 0.1 myBP on the overlying basalt. Utilising the constants for
the potassium-argon technique recommended by Steiger & Jager (1978), McDougall (pers. comm. 1980) has
recalculated the date as 4.46+ 0.1 myBP. Ludbrook (1973) regards the molluscan fauna of the underlying

Grange Burn Formation as indicative of the Kalimnan Stage.
Faunai

MAMMALIA
Marsupialia
Dasyuromorphia

appr

1040 - RICH, ET AL.

Dasyuridae
Antechinus sp.
Peramelemorphia
Perameloidea
Diprotodontia
Vombatiformes
Palorchestidae
Palorchestes n. sp.
Diprotodontidae
New genus and species
Vombatidae
Genus and species indet.
Phalangerida
Phalangeridae
Trichosurus hamiltonensis Flannery, Tumbull, Rich & Lundelius 1987
Strigocuscus notialis Flannery, Tumbull, Rich & Lundelius 1987
Ektopodontidae
Darcius duggani Rich 1986
Potoroidae
Hypsiprymnodon sp.
Propleopus sp.
Milliyowi bunganditj Flannery et al. in press
Macropodidae
Dorcopsis wintercookorum Flannery et al. in press
of. Dendrolagus
Kurrabi pelchenorum Flannery et al. in press
Macropus (Notamacropus) sp.
Troposodon sp.
Simosthenurus sp.
Protemnodon sp.
Thylogale ignis
cf. Wallabia
Burramyidae
Burramys triradiatus Tumbull, Rich & Lundelius 1987c
Genus and species indet.
Pseudocheiridae
Pseudokoala erlita Tumbull & Lundelius 1970
Pseudocheirus marshalli Tumbull & Lundelius 1970
Pseudocheirus stirtoni Tumbull & Lundelius 1970
Petauridae
Petaurus sp., cf. P. australis
Petaurus sp., cf. P. norfolkensis
Placentalia
Chiroptera
Microchiroptera
Family indet.

References: Abele ef al. (1988), Archer (1982a), Boutakoff & Sprigg (1953), Flannery, Rich, Tumbull
& Lundelius (in prep.), Flannery, Tumbull, Rich & Lundelius “(1987); Gill (1957b), Ludbrook (1973), Rich
(1986)/ Ride (1964), Stirton (1957b), Turnbull & Lundelius (1970), Tumbull et al. (1965), Tumbull, Rich &
Lundelius (1987a-c).

Big Sink Local Fauna

Type locality: Southern wall of the Big Sink doline, Wellington Caves, New South Wales (32° 35' S.,

148° 59' E.) (Hand, Dawson & Augee 1988).

Rock unit and age: Big Sink Unit, the upper member of the Phosphate Mine Beds. Carbonate-cemented
osseous sandstones interbedded with thin layers of structureless mud. Early to Middle Pliocene age is
suggested by the presence of Pseudocheirus stirtoni and a unique peramelid otherwise known only from the
Hamilton Local Fauna, Protemnodon cf. P. devisi, which differs only slight from Protemnodon devisi from
Chinchilla Local Fauna, and Thylacoleo crassidentatus which occurs in the Bow and Bluff Downs local faunas
(Hand, Dawson & Augee 1988, Osborne 1983).

Fauna:
REPTILIA

Squamata
Lacertilia

AUSTRALIA'S MAMMALIAN RECORD - 1041

Scincidae
Tiliqua sp.
MAMMALIA
Marsupialia
Dasyuromorphia
Dasyuridae
Antechinus sp.
Thylacinidae
Thylacinus sp.
Peramelemorphia
New genus?
Diprotodontia
Vombatiformes
Thylacoleonidae
Thylacoleo crassidentatus Bartholomai 1962
Phalangerida
Macropodidae
Protemnodon cf. P. devisi
cf. Wallabia spp.
Burramyidae
Pseudocheiridae
Pseudocheirus stirtoni Tumbull & Lundelius 1970
Placentalia
Chiroptera
Megadermatidae
Macroderma koppa Hand, Dawson & Augee 1988 4
Rodentia
Muridae
New pseudomyine

References: Hand, Dawson & Augee (1988), Osbome (1983)

Bluff Downs Local Fauna

Type locality: Banks of Allingham Creek, Bluff Downs Station, north Queensland (19° 43'S.,

145° 36’ E.).
Rock unit and age: Allingham Formation. Sequence of terrigenous clays, silts, sands, calcareous sands,
and Chara limestones that appear to conformably underlie the Allensleigh "flow" of the Nulla Basalt (Archer

& Wade 1976) which is dated at 4.5 and 4 myBP at two sites 10 or more kilometres away from the fossil

vertebrate locality of Bluff Downs (Wyatt and Webb 1970).
Fauna:
ARTHROPODA
Crustacea
Unidentified gastrolith
OSTEICHTHYS
Teleostei
Unidentified spines and vertebrae
REPTILIA
Testudines, family indet.
Crocodilia
Crocodilidae
Crocodylus porosus
Squamata

Lacertilia
Agamidae
Near Amphibolurus spp.
Varanidae
Varanus sp.
cf. Megalania
Ophidia
Boidae
?Morelia sp.
Acrochordidae
AVES
Ciconiiformes

1042 - RICH, ET AL.

Ciconiidae
Ephippiorhynchus asiaticus (Latham, 1790)
Threskiornithidae
Threskiornis sp.
Anseriformes
Anatidae
Dendrocygna sp.
Cygnus sp.
Charadriiformes
Scolopacidae
Numenius sp.
MAMMALIA
Marsupialia
Dasyuromorphia
Dasyuridae
Planigale sp.
Peramelemorphia
Peramelidae
Perameles allinghamensis Archer in Archer & Wade 1976
Diprotodontia
Vombatiformes
?Phascolarctidae
Koobor jimbarratti Archer in Archer & Wade 1976
Diprotodontidae
Zygomaturus sp.
Euryzygoma sp.
Diprotodontinae, genus indet.
Vombatidae
Ramsayia lemleyi (Archer in Archer & Wade 1976)
Thylacoleonidae
Thylacoleo crassidentatus Bartholomai 1962
Phalangerida
Macropodidae
Sthenurus sp.
Troposodon minor (Owen 1877b)
Petrogale sp.
Protemnodon snewini Bartholomai 1978a
Macropus narada (Bartholomai) 1978a
Macropus dryas (de Vis 189Sa)
Osphranter pavana Bartholomai 19784
Placentalia
Rodentia
Muridae

References: Archer (in Archer & Wade 1976), Archer (1982a), Archer & Dawson (1982), Bartholomai
(1978a), Dawson (1981), Gaffney (1981), Molnar (1979), Wyatt & Webb (1970).

Rackham's Roost Local Fauna

Locality: Riversleigh Station, northwestem Queensland (19° S., 1382/4° E).
Rock unit and age: Cave deposit. Pliocene, probably Early Pliocene.

Fauna:
AMPHIBIA
Anura
REPTILIA
Squamata
AVES
MAMMALIA
Marsupialia
Dasyuromorphia
Dasyuridae
Planigale sp.
Sminthopsis sp.
New genus
Peramelomorphia
Diprotodontia

Phalangerida

AUSTRALIA'S MAMMALIAN RECORD - 1043

Potoroidae

Macropodidae
Protemnodon sp. cf. P. sneweni
Protemnodon sp. (> 1)
Macropus sp. (> 1)

Placentalia
Rodentia
Muridae
Zyzomys n. sp.
Pseudomys -n. sp.
One or more additional new genera, at least 11 additional new species
Chiroptera
Megadermatidae
Macroderma gigas
Hipposideridae
4 spp.
Vespertilionidae
4 spp.

Comments: The presence of Protemnodon sp., cf. P. sneweni as in the Bluff Downs Local Fauna,
together with the presence of abundant primitive rodents suggests that Rackham's Roost Local Fauna is of

Pliocene and probably Early Pliocene age.
References: Archer, Godthelp, Hand & Megirian (1989), Hand (1987), Godthelp (1987).

Tara Creek

Type Locality: Head of Tara Creek, north Queensland (19° 27'S., 145° 17' E.).
Rock unit and age: From unspecified sediments (possibly Allingham Formation) beneath the Nulla
Basalt (Bartholomai in Gaffney 1981), which is radiometrically dated as Pliocene (Wyatt & Webb 1970).
Fauna:
REPTILIA
Testudines
Chelodina sp.
Crocodilia
Crocodilus nathani
MAMMALIA
Marsupialia
Diprotodontia
Vombatiformes
Diprotodontidae

References: Gaffney (1981), Woods (1960a).

Lake Tyers

Locality: Second headland north of Lake Tyers House, on the Nowa Nowa arm of Lake Tyers, Victoria
(37° 49.5'S., 148° 7.5' E.). ;
Rock unit and age: Jemmy's Point Formation, Kalimnan Stage (Early Pliocene) on the basis of its
marine molluscan fauna (Abele et al. 1988).
Fauna:
MAMMALIA
Marsupialia
Diprotodontia
Phalangerida
Macropodidae

Protemnodon chinchillaensis Bartholomai 1973.

References: Flannery & Archer (1984), Plane (1972).

1044 - RICH, ET AL.
Bunga Creek

Locality: Southwest side of Bunga Creek road cutting on the Princes Highway, Victoria (37° 50' S.,
148° 00' E.). : A

Rock unit and age: Jemmy's Point Formation, Kalimnan Stage (Early Pliocene) on the basis of its
marine molluscan fauna (Abele et al. 1976).

Fauna:
MAMMALIA

Marsupialia

Reference: Warren (1965).

Great Buninyong Estate Mine

Locality: Found at a depth of 67 metres in an inclined shaft of the Great Buninyong Estate Mine, 10

kilometres southeast of Ballarat, Victoria (37° 39'S., 143° 53' E.) (Whitelaw 1899).

Rock unit and age: From unnamed deep lead, lacustrine sediments containing volcanic ejectamenta. The
deep lead is resting on Ordovician shale and is overlain by basalt. McDougall et al. (1966) and Aziz-Ur-
Rahman & McDougall (1972) have shown that nearby basalts are Pliocene in age, one date of
2.53 +.0.07 myBP coming from basalts 10 kilometres northeast of the mine, and another of
3.90 + 0.10 myBP from 12 kilometres northeast of the mine.

Fauna:
MAMMALIA
Marsupialia
Diprotodontia
Vombatiformes
Diprotodontidae
Phalangerida
Macropodidae
Macropus giganteus Shaw, 1790
Small macropodine

Comments: The apparent minimum age of this site is the oldest established radiometrically for
Macropus giganteus, one of the most common elements in Australian Quaternary terrestrial mammal
assemblages. Unfortunately, to obtain a basalt sample from directly above the fossil site today would only
be feasible by drilling from the surface, as flooding forced the closure of the Great Buninyong Estate Mine
shortly after the fossils were discovered.

References: de Vis (1899a), Flannery (1981), Hart (1899), Keble (1945), Rich (1976), Whitelaw
(1899).

Awe Fauna

[ype locality: Upper Watut River just south of its junction with Wiganda Creek, 6.1 kilometres

northeast of the Otibanda triangulation station, Morobe District, Papua New Guinea (7° 14.5' S., 146° 32.75'
E.) (Plane 1967a-b).

Referred localities: Upper Watut River, 14 sites including the Awe type locality, all within a radius of 5
kilometres of 7° 15' S., 146° 32' E. Sunshine localities, nine sites, 7° 04.5' S., 146° 36.6' E. Widubush
locality, 7° 5.6' S., 146° 37.57' E. Niba Gold locality, 7° 5.75' S., 146° 36.75' E. Unnamed locality, hel
7.67 S., 146° 37.83' E. Zoffman locality 7° 9.2' S., 146° 38.43' E. Koranga locality, 7° 19.2' S., 146°
43.25' E. (See Plane 1967a-b) for detailed listing of sites plus maps showing their location.)

Rock unit and age: Otibanda Formation, Late Pliocene. Three potassium argon dates have been
published for the Awe Fauna: 7.6 myBP (Evernden et al. 1964), 3.1-3.5 myBP (Page & McDougal 1972), and
3.3 + 0.1 and 2.5 + 0.1 myBP (Hoch & Holm 1986). Page & McDougal (1972) explained the greater age
recorded by Evernden et al. (1964) as owing to extraneous argon, a conclusion with which Hoch & Holm
(1986) are in agreement.

Faunai
MOLLUSCA

AUSTRALIA'S MAMMALIAN RECORD - 1045

Gastropoda
Lymnaea sp.
?Planispira sp.
Gabbia sp.
OSTEICHTHYS
Teleostei
REPTILIA
ata
Booidea
Crocodilia

Crocodylus sp., of. C. porosus
AVES
Casuariformes
Casuaridae
Casuarius sp.
MAMMALIA
Marsupialia
Dasyuromorphia
Thylacinidae
Thylacinus sp.
Dasyuridae
Myoictis sp.
Diprotodontia
Vombatiformes
Diprotodontidae
Nototherium watutense Anderson 1937
Kolopsis rotundus Plane 1967a
Kolopsoides cultridens Plane 1967a
Phalangerida
Macropodidae
Protemnodon otibandus Plane 1967b
Protemnodon buloloensis Plane 1967b
cf. Dorcopsis sp.
Watutia novaeguineae Flannery & Hoch 1989 (in Flannery, Hoch & Aplin 1989)
Placentalia
Rodentia
Muridae indet.

References: Anderson (1937), Archer (1982a), Dow (1961), Dow et al. (1974), Evernden et al. (1964),
Fisher (1944), Flannery, Hoch & Aplin (1989), Hoch & Holm (1986), (Page & McDougall (1972), Plane
(1967a-b 1972, 1976), Stirton, Woodburne & Plane (1967), Woods (1962).

Mogorafugwa

Type locality: Mogorafugwa (Swamp) about 10 km west of Koroba (5° 42' S., 142° 44' E.), Duna
Subdistrict, Southern Highlands, Papua New Guinea
Rock unit and age: Mudstone and clay (pers. comm. B.W. Houston to N. S. Pledge, 20/1/1972).
Species suggests equivalence with Awe Local Fauna.
Fauna:
MAMMALIA
Marsupialia
Diprotodontia
Phalangerida

Macropodidae
Protemnodon sp., cf. P. otibandus

LATE CAINOZOIC
Ian's Prospect

Locality: Billeroo Creek, Frome Downs Station, South Australia (31° 11' S., 140° 16' E).
i Uncertain, specimen found in loose sediments at the bottom of Billeroo Creek.
Presumably derived either from the Late Oligocene-Middle Miocene Namba Formation or the Pleistocene
Eurinilla Formation, both of which outcrop in the immediate vicinity. Fossil bones are common in the

1046 - RICH, ET AL.

Eurinilla Formation that forms the upper part of the banks of the stream at the site and none have been found
in the Namba Formation at this locality where it occurs near water level of the stream channel.
Unfortunately, on all occasions when this site has been visited, the Namba Formation has never been
completely dry. Therefore, it has not been possible to ascertain whether or not fossils occur there in the
Namba Formation.

Fauna:

MAMMALIA

Marsupialia
Dasyuromorphia

Dasyuridae
Wakamatha tasselli Archer & Rich 1979

References: Archer (1982a), Archer & Rich (1979).

MIDDLE PLIOCENE - EARLY PLEISTOCENE

Floraville Local Fauna

Type locality: South of Floraville Station HS and west of the Leichhardt River, Queensland (18° 17'S.,

139° 52' E.).

Rock unit and age: Unnamed surficial riverine sediments. Age is either Pliocene or Pleistocene, most
probably the latter. The local fauna so far lacks elements known to be restricted to one or the other epoch
further south other than Rattus species which so far have no known pre-Pleistocene record (Archer 1982b,
Godthelp in prep.).

Fauna:
CRUSTACEA
REPTILIA
Lacertilia
Varanidae
Varanus so.
Genus indet.
Ophidia
Testudines
Trionychidae
Chelidae
MAMMALIA
Marsupialia
Dasyuromorphia
Dasyuridae
Sminthopsis floravillensis Archer 1982b
Antechinus sp.
Dasyurus sp.
Peramelemorphia
Peramelidae
Diprotodontia
Vombatiformes
Diprotodontidae
Zygomaturus sp.
Genus undet.
Phalangerida
Macropodidae
Sthenurinae, genus undet.
Potoroidae
Potoroinae, genus undet.
Placentalia
Rodentia
Muridae
Rattus spp.
Pseudomys spp.

Chinchilla Local Fauna

['ype locality: Banks of the Condamine River near Chinchilla, Queensland (26° 48' S., 150° 41' E.).

AUSTRALIA'S MAMMALIAN RECORD - 1047

Rock unit and age: Chinchilla Sands. Early to Middle Pliocene on the basis of closer faunal

resemblance to the Early Pliocene Bluff Downs Local Fauna than to the Pleistocene fauna from the eastern
Darling Downs.

Many of the genera shared between the Bluff Downs and Chinchilla local faunas include closely related
species pairs. In most instances, the Chinchilla form appears to be somewhat more advanced than that of the
Bluff Downs one. This pattern repeated frequently across the faunal spectrum suggests that the Chinchilla
Local Fauna is somewhat younger than Bluff Downs rather than the differences being a result of the two fossil
samples having originated from different habitats.

On the other hand, the geographically much closer fauna from the Pleistocene eastern Darling Downs
differs more markedly from the Chinchilla Local Fauna. The following genera occur in the eastern Darling
Downs Fauna but not at Chinchilla or Bluff Downs: Diprotodon, Nototherium, Procoptodon, and Sarcophilus.
In addition, the Chinchilla and Bluff Downs local faunas share the following taxa which are unknown in the
easter Darling Downs fauna: Euryzygoma spp. and Macropus dryas.

Because the Bluff Downs Local Fauna at present appears to be Early Pliocene in age based on potassium-
argon analysis and the Chinchilla Local Fauna is younger but much closer in stage of evolution to it than to
the Pleistocene fauna from the eastern Darling Downs, an Early to Middle Pliocene age is suggested for the
Chinchilla Local Fauna.

Fauna:
OSTEICHTHYES
Dipnoi
Ceratodontidae
Ceratodus palmeri Krefft 1874b
Neoceratodontidae
Neoceratodus forsteri (Krefft 1870)
REPTILIA
Testudines
Trionychidae
Chelidae
Emydura sp.
Meiolanidae
Crocodilia
Crocodylidae
Pallimnarchus pollens DeVis 1907
AVES
Casuariiformes
Casuariidae
Dromaius novaehollandiae Latham 1790
Pelecaniformes
Pelecanidae
Pelecanus proavis de Vis, 1892.
Phalacrocoracidae
Microcarbo (Haliaetor) melanoleucos Vieillot 1817
Anseriformes
Anatidae
Anseranas cf. A. semipalmatus
cf. Cereopsis
Cygnus cf. C. atratus
Anas superciliosa Gmelin 1789
Aythya australis Eyton 1838
Biziura lobata Shaw 1796
Galliformes
Megapodiidae
Progura gallinacea (DeVis 1888)
Gruiformes
Rallidae
Fulica atra Linnaeus 1758
Charadriiformes
MAMMALIA
Marsupialia
Dasyuromorphia
Dasyuridae
Dasyurus dunmalli Bartholomai 1971b
Thylacinidae
Thylacinus cynocephalus Harris 1808

1048 - RICH, ET AL.

Diprotodontia
Vombatiformes

?Phascolarctidae
Koobor notabilis (de Vis 1889b)

Diprotodontidae
Euowenia grata (de Vis 1887)
Euryzygoma dunense de Vis 1887

Palorchestidae
Palorchestes parvus de Vis 1895a
cf. Palorchestes sp.

Vombatidae
2Vombatus prior de Vis 1883d

Thylacoleonidae
Thylacoleo crassidentatus Bartholomai 1962

Phalangerida

Phalangeridae
Genus indet.

Macropodidae
Troposodon n. sp.
Troposodon minor (Owen 1877»)
Sthenurus antiquus Bartholomai 1963
Sthenurus notabilis Bartholomai 1963
Protemnodon devisi Bartholomai 1973
Protemnodon chinchillaensis Bartholomai 1973
Wallabia indra de Vis 1895a
Macropus pan de Vis 1895a
Macropus dryas (de Vis 1895a)
Osphranter woodsi Bartholomai 1975
Prionotemnus palankarinnicus Stixton 1955

Placentalia
Rodentia

Muridae
Pseudomys vandycki Godthelp 1990
Pseudomys spp.

References: Archer (1977b 1982a), Archer & Dawson (1982), Archer & Wade (1976), Bartholomai
(1962, 1963, 1966, 1967, 1968, 1971b, 1973, 1975, 1976), Bartholomai & Woods (1976), Dawson
(1982a), Flannery & Archer (1983); Gaffney (1981), Gaffney & Bartholomai (1979), Godthelp (1990), Kemp
& Molnar (1981), Olson (1975, 1977), Patterson & Rich (1987), Woods (1956b, 1960a, 1962).

Quanbun Local Fauna

Type locality: Jubilee Dam (formerly Alligator Dam) about 15 km north of Quanbun Homestead,

Western Australia (18° 17' S., 125° 23' E.).

Rock unit and age: Unnamed light clay overlain by 1.5 metres of darker clay, which is in tum overlain
by conglomerate of variable thickness (Flannery 1984). Presence of Macropus pan suggests a Pliocene age
for the site as at Chinchilla. However, the presence of a large species of Protemnodon is a characteristic of
Pleistocene assemblages, not Chinchilla or other Pliocene sites.

Fauna:
REPTILIA
Crocodylia
Crocodylidae
MAMMALIA
Marsupialia
Diprotodontia
Vombatiformes
Diprotodontidae or Palorchestidae
Vombatidae
Phascolonus sp., cf. P. gigas
Phalangerida
Macropodidae
Macropus pan de Vis 1895a
Protemnodon sp., cf. P. roechus
cf. Osphranter

References: Flannery (1984), Glauert (1921b)

AUSTRALIA'S MAMMALIAN RECORD - 1049

Bow Local Fauna

Type locality: Roadcut exposures near Bow, N.S.W. on the Merriwa-Cassilis Road, (32° 8' S., 150° 9
E.).

Rock unit and age: Unnamed fluviatile sediments. Early to Middle Pliocene on the basis of correlation
with the Bluff Downs Local Fauna which is tentatively radiometrically dated at about 4 to 4.5 myBP.
Fauna:
BIVALVIA
Hyriinae
Hyridella sp., cf. H. australis
Crobiculidae
Corbicula sp.
GASTROPODA
Planorbidae
Physastra sp.
Bithyniidae
Gabbia sp., cf. G. australis

Monotremata
Ornithorhynchidae
Ornithorhynchus sp.
Marsupialia
Dasyuromorphia
Dasyuridae
Dasyurus dunmalli Bartholomai 1971b
Peramelemorphia
Peramelidae
Perameles sp.
Diprotodontia
Vombatiformes
Diprotodontidae
Palorchestidae
Palorchestes sp., cf. P. parvus de Vis 1895a
Vombatidae
Phascolonus sp.
Thylacoleonidae
Thylacoleo crassidentatus Bartholomai 1962
Thylacoleo sp., of. T. hilli Pledge 1977
Phalangerida
Phalangeridae
Genus indet.
Potoroidae
Propleopus sp.
Macropodidae
Simosthenurus sp.
Protemnodon chinchillensis Bartholomai 1973
cf. Protemnodon
Macropus dryas (DeVis 1895)
Macropus (Osphranter) pavana Bartholomai 1978a
Troposodon bowensis Flannery & Archer 1984
Troposodon spp.
cf. Dendrolagus spp. 1 and 2
Kurrabi mahoneyi Flannery & Archer 1984
Kurrabi merriwaensis Flannery & Archer 1984
Prionotemnus palankarinnicus Stirton 1955
Macropodinae indet. Types 1 and 2

References: Archer (1982a), Archer & Dawson (1982), Flannery & Archer (1984), Skilbeck (1980).

1050 - RICH, ET AL.

Krui River Local Fauna

Type locality: Roadcut exposures on the south side of the Krui River, 17 km northwest of Bow, N.S.W.

on the Merriwa-Cassilis Road, (32° 5' S., 150° 7' E.).
Rock unit and age: Unnamed coarse clastic sediments containing many basalt fragments that appear to

represent a river terrace.
Fauna:
REPTILIA
Crocodilia indet.
MAMMALIA
Marsupialia
Diprotodontia
Vombatiformes
Diprotodontidae
Phalangerida
Macropodidae
Protemnodon sp., cf. P. devisi
Troposodon sp., cf.T. minor
Troposodon sp., of. T. bowensis

Palankarinna Local Fauna

Type locality: West side of Lake Palankarinna, South Australia, (28° 46' S., 138° 24' E.).

Rock unit and age: Mampuwordu Sands. "The age assignment of this fauna is based chiefly on its
species of Zygomaturus, which seems to be older than typical Pleistocene forms, and later than Awe,
Beaumaris, or Alcoota zygomaturines” (Stirton et al. 1968, p. 16).

Dipnoi
Neoceratodontidae
Neoceratodus sp. 3
Teleostei
REPTILIA
Crocodilia
Crocodylus
Sebecosuchia or Pristichampsinae
AVES
Casuariformes
Casuariidae
Dromaius ocypus Miller 1963
Dromornithiformes
Dromomithidae
MAMMALIA
Marsupialia
Peramelemorphia
Thylacomyidae
Ischnodon australis Stirton 1955
Diprotodontia
Vombatiformes
Diprotodontidae
Zygomaturus keanei Stirton 1967b
Meniscolophus mawsoni Stirton 1955
Phalangerida
Macropodidae
Prionotemnus palankarinnicus Stirton 1955
Sthenurinae, genus indet.

References: Hecht & Archer (1977), Miller (1963), Molnar (1978b), Ride (1964), Stirton (1955,
1967b), Stirton et al. (1961), Stirton, Woodbume & Plane (1967), Rich (1979), Stirton, Tedford &
Woodbume (1967), Tedford, Williams & Wells (1986), Woods (1962).

AUSTRALIA'S MAMMALIAN RECORD - 1051

Kanunka Local Fauna

Type locality: West side of Lake Kanunka, South Australia (28° 23' S, 138° 18’ E).
Rock unit and age: Tirari Formation. The productive unit appears to be in the magnetically reversed

Matuyama Chron (Tedford, Williams & Wells 1986) just above the Matuyama/Gauss boundary (2.48 myBP)
and hence Late Pliocene in age.

Dipnoi
Ceratodontidae
Teleostei
AVES
Casuariiformes
Casuariidae
Dromaius novaehollandiae Latham, 1790
Pelecaniformes
Pelecanidae
Pelecanus cadimurka Rich and van Tets, 1981
Pelecanus conspicillatus Temminck, 1824
Anhingidae
Anhinga novaehollandiae Miller, 1966a
Phalacrocoracidae
Ciconiiformes
Ciconiidae
Ardeidae
Anseriformes
Anatidae
Falconiformes
Accipitridae
Gmuiformes
Gmidae
Grus sp.
Rallidae
Otididae
Charadriiformes
Family indet.
Phoenicopteriformes
Phoenicopteridae
Ocyplanus proeses de Vis , 1905 [=Phoeniconaias graciOlis Miller, 1963]
cf. Phoenicopterus ruber
Xenorhynchopsis minor de Vis, 1905
Passeriformes
MAMMALIA
Marsupialia
Dasyuromorphia
Dasyuridae
Diprotodontia
Vombatiformes
Diprotodontidae
Zygomaturus sp.
Diprotodon sp.
Vombatidae
Phascolonus sp.
Vombatus or Lasiorhinus
Phalangerida
Potoroidae
Bettongia sp.
Macropodidae
Lagorchestes sp.
Dendrolagus sp.
Kurrabi sp.
cf. Prionotemnus sp.
Troposodon kentii Campbell 1973
Troposodon sp., cf. T. minor
Protemnodon sp. cf. P. devisi

1052 - RICH, ET AL.

Protemnodon sp.
Osphranter sp., cf. O. woodsi
Macropus (Fissuridon) pearsoni (Bartholomai 1973)
Macropus (Notamacropus) sp.
Sthenurinae
Placentalia
Rodentia
Muridae

Talyawalka

Locality: Near (normally dry) White Water Lake, Talyawalka Anabranch of the Darling River, New
South Wales, 32° 25' S., 143° 18' E. From a water bore, at a depth of about 28 metres.
Rock unit and age: Unnamed fluvial sediments. Probably Pliocene or Pleistocene, based on species
correlation with the Plio-Pleistocene Chinchilla Local Fauna and the Pliocene Kanunka Local Fauna.
Fauna:
MAMMALIA
Marsupialia
Diprotodontia
Vombatiformes

Diprotodontidae
Euowenia grata (de Vis 1887)

Reference: Pledge (1989a)

Town Well Cave

Locality: Town Well Cave, Curramulka, South Australia. 34° 42'S, 137° 43' E.

Rock Unit and age: Uncertain. The specimen was found cemented to the wall of the cave at some
height above the floor, in an area where no other fossils were apparent. The cave is developed along deep
joint fissures in Cambrian limestone, and in places contains a typical Late Pleistocene fauna.

Comparisons with the Bow Local Fauna suggests a Mio-Pliocene age (Archer & Dawson 1982) for the
Town Well Cave assemblage.

Fauna:
MAMMALIA
Marsupialia
Diprotodontia
Vombatiformes

Thylacoleonidae
Thylacoleo hilli Pledge 1977

References: Archer & Bartholomai (1978).

Curramulka Local Fauna

Type locality: Corra Lynn Cave (5Y1), Curramulka, South Australia; 34° 43' S., 137° 43' E. About 3
km south of Town Well Cave.

Rock unit and age: Unnamed red silty cave fill. This occurs only in one small area of the extensive
labyrinthine cave. Deposit fills a fissure which may extend to the surface. Host rock is Early Cambrian
limestone (Crawford 1965).

The Town Well Cave Thylacoleo hilli may be of the same age. Comparison with other faunas indicates
a Mio-Pliocene age.

Fauna:

AMPHIBIA
Neobatrachus pictus Tyler 1988
REPTILIA
Testudines
Lacertilia
Madtsoiidae
Wonambi sp., cf. W. naracoortensis Smith 1976

AUSTRALIA'S MAMMALIAN RECORD - 1053

Elapidae indet.
Scincidae
Tiliqua sp.
Varanidae indet.
AVES
Dromornithiformes
cf. Ilbandornis sp.
MAMMALIA
Marsupialia
Dasyuromorphia
Thylacinidae
Thylacinus sp.
Dasyuridae
cf. Glaucodon
of. Phascogale
Diprotodontia
Vombatiformes
Phascolarctidae
Phascolarctos sp., cf. P. cinereus
Gen. et sp. nov., very large
Diprotodontidae
?Zygomaturus
Zygomaturine indet.
Palorchestidae
Palorchestes sp., cf. P. parvus
Vombatidae
Vombatus sp. indet
Phascolonus sp. indet.
Phascolomys sp., cf. P. medius
Thylacoleonidae
Thylacoleo sp. indet., small
Phalangerida
Potoroidae
Potorous sp.
Macropodidae
Protemnodon sp. indet.
Troposodon sp, cf. T. bluffensis
Macropodine small spp.
Pseudocheiridae
New species, very large
Petauridae
Petaurus sp.

References: Anon. (1985); Pledge (1987a, in prep.); Smith (1976); Tyler (1988).

Fisherman's Cliff Local Fauna

Type locality: Cliff developed on the right bank (north side) of the Murray River, New South Wales
(34° 7'S., 142° 39' E.).

Rock unit and age Mooma Formation (which may be a member of the Blanchetown Clay). Marshall
(1973) regarded this local fauna to be either Late Pliocene or Early Pleistocene. This age assignment was
accepted partly on the geological conclusions of Gill (1973b) and partly on the basis of a specimen of
Protemnodon sp. that was similar to Protemnodon devisi Bartholomai (1973). P. devisi is from the
Chinchilla Local Fauna of Queensland, then thought to be of Late Pliocene or Early Pleistocene age (see
above). With further analysis, Crabb (1977) regarded the Fisherman's Cliff Local Fauna to probably be Early
Pleistocene, and Crabb (1982) suggested that the Moorna Formation actually represented an intraformational
unit within the Blanchetown Clay. The Mooma Formation appears to occupy the same stratigraphic position
relative to the overlying Blanchetown Clay at Fisherman's Cliff as the basal member of the Blanchetown
Clay does to the overlying part of that formation at Chowilla (Woodbume ef al. 1985) and thus is in the
upper part of the Gauss normal polarity epoch (2.47-2.92 myBP) and Late Pliocene (Bowler 1980). Whitelaw
(in prep.) has corroborated this correlation by recognising the Karoonda Surface of Gill (1973b) both at
Chowilla and at Fisherman's Cliff and demonstrating that the magnetic polarity of that part of the Mooma
Formation containing the Fisherman's Cliff Local Fauna is in fact normal and comparable to that at Chowilla

described by An et al. (1986).

1054 - RICH, ET AL.

Fauna:
GASTROPODA
Basommatophora
Lymnaeidae
Lymnaea sp., of. L. tomentosa
OSTEICHTHYES
Dipnoi
Neoceratodontidae
Neoceratodus forsteri (Krefft 1870)
Neoceratodus gregoryi (White 1925)
Teleostei

Chelonia
Cheliidae
Emydura sp., cf. E. macquarrii
AVES
Casuariiformes
Dromaiinae
MAMMALIA
Marsupialia
Dasyuromorphia
Dasyuridae
Sarcophilus moornaensis Crabb 1982
Dasyuroides achilpatna Archer 1982b
Dasyurus hallucatus
Dasyurus sp.
Sminthopsis sp., cf. S. macroura
Antechinus sp.
Peramelemorphia
Peramelidae
Diprotodontia
Vombatiformes
Diprotodontidae
cf. Diprotodon
Vombatidae
Lasiorhinus sp.
Phalangerida
Potoroidae
Bettongia sp.
Macropodidae
Sthenurus sp.
Lagostrophus sp., cf. L. fasciatus
Petrogale sp.
Osphranter sp.
Protemnodon sp., cf. P. devisi
Placentalia
Rodentia
Muridae
Several genera and species, as yet undescribed.

Reference: Archer (1982a), Bowler (1980), Crabb (1977, 1982), Gill (1973b), Marshall (1973),
Woodbume et al. (1985).

Bone Gulch Local Fauna

Type locality: System of gullies on the right bank (west side) of the Murray River, New South Wales

(34° 7' S., 142° 37' E’. Part of the area is shown in pl. 6, fig. 2 of Gill (1973b).

Rock unit and age: Near the base of the Blanchetown Clay. On stratigraphic grounds, Firman (1965,
1966) regarded the Blanchetown Clay to be early Middle Pleistocene, Lawrence (1966) placed it in the Early
Pleistocene and Gill (1973b), in the Late Pliocene or Early Pleistocene. Marshall (1973) did not regard the
mammals to be of any utility in dating the Bone Gulch Local Fauna. The contact between the
palaeomagnetically reversed Matuyama Epoch and the normal Gauss Epoch which is dated at 2.47 myBP
(Berggren et al. 1985), Late Pliocene, is in the lower part of the Blanchetown Clay in the sections where
measurements have been taken including at Bone Gluch itself (Bowler 1980, An et al. 1986, Whitelaw in
prep.). Because the various identifiable horizons within the Blanchetown Clay bear a consistent relation to

AUSTRALIA'S MAMMALIAN RECORD - 1055

the palaeomagnetic boundaries, it is likely that the Bone Gulch Local Fauna, being in the lower part of the
formation, is Late Pliocene.

Teleostei
Dipnoi
Ceratodontidae
Ceratodus palmeri Krefft 1874b
Neoceratodontidae
Neoceratodus forsteri (Krefft 1870)
REPTILIA
Testudines
Chelidae
MAMMALIA
Marsupialia
Diprotodontia
Vombatiformes
Diprotodontidae
Thylacoleonidae
Thylacoleo sp.
Vombatidac
Phascolonus sp., cf. P. magnus
Phalangerida
Potoroidae
cf. Bettongia
Macropodidae
Sthenurus sp.
Macropodinae, genus indet.
Placentalia
Rodentia
Muridae

Dog Rocks Local Fauna

Locality: Floor of Australian Portland Cement Limited's Batesford Quarry (38° 6.5' S., 144° 17.5' E.).
Rock unit and age: Fissure-filling within the argillaceous Moorabool Viaduct Sand. Late Pliocene
(2.03-2.48 myBP) or possibly Early Pliocene (3.40-3.48 myBP).

Fauna:

OSTEICHTHYES
Teleostei, indet.
AMPHIBIA
Anura, indet.
REPTILIA
Squamata, indet.
AVES, indet.
MAMMALIA
Marsupialia
Dasyuromorphia
Dasyuridae
Antechinus sp.
Dasyurus sp.
Peramelomorphia
Peramelidae
Perameles sp.
Isoodon sp.
Diprotodontia
Vombatiformes
Diprotodontidae
Zygomaturus sp.
Vombatidac
Vombatus ursinus

1056 - RICH, ET AL.

Phascolonus sp.
Phalangerida

Phalangeridae
Phalanger sp.

Potoroidae
Potorous sp.
cf. Bettongia

Macropodidae
Macropus cf. M. giganteus
Macropus cf. M. fuliginosus
Macropus sp. A
Macropus sp. B
Sthenurus sp.
Protemnodon cf. P. anak
Protemnodon sp. A
Wallabia cf. W. bicolor
Wallapia sp.
Troposodon sp.

Pseudocheiridae
Pseudocheirus sp. A
Pseudocheirus sp. B

Placentalia
Rodentia

Muridae
Pseudomys sp. A
Pseudomys sp. B

Comments: The fossiliferous unit was overlain by a basalt flow of the Newer Volcanics. Three
kilometres to the south a magnetically reversed basalt flow with similar petrographic characteristics was dated
at 2.03 + 0.13 myBP in three different quarries (Aziz-ur-Rahman & McDougall, 1972, corrected with decay
constant of Faure, 1986, in Whitelaw, 1989).

A maximum age for the site is provided by the occurrence of Globorotalia crassiformis which has a first
appearance at 4.0 myBP (Whitelaw 1989). This species occurs in sediments of the Moorabool Viaduct Sand
stratigraphically below the fissures. The sediments are magnetically reversed (Whitelaw 1989), implying
that the age of the fissures within the interval between 1.9 and 4.0 myBP is either in the early Matuyama
Chron (2.03-2.48 myBP) or the late Gilbert Chron (3.40-4.88 myBP). The presence of a diversity of rodents
suggests the early Matuyama Chron is the more likely interval represented (Whitelaw 1989).

References: Aziz-ur-Rahman & McDougall (1972), Bowler (1963), Rich (1976), Whitelaw (1989).
Coimadai Local Fauna

Type locality: Alkemade's Quarry, about 10 km northnortheast of Bacchus Marsh, Victoria (37° 46' S.,
144° 26' E.).

Rock unit and age: Coimadai Limestone, early Pliocene. The fossils occur 2 to 3 metres beneath an
ashi which Coulson (1924) correlated with the basalt of the Bullengarook flow. The Bullengarook flow has
been radiometrically dated as 3.31 and 3.64 myBP (P. Roberts, pers. comm. to RHT, 1983, in Tumbull,
Lundelius & Tedford 1990) and has a reversed polarity, thus limiting the range to 3.40-3.64 myBP.

Fauna:
MAMMALIA
Marsupialia
Diprotodontia
Vombatiformes
Diprotodontidae
Euowenia sp.
Zygomaturus sp.
Vombatidae
Vombatus near V. hirsutus
Vombatus sp. ["Phascolomys parvus"]
Phalangerida
Macropodidae
Kurrabi sp.
Protemnodon sp.
Troposodon sp.

AUSTRALIA'S MAMMALIAN RECORD - 1057

cf. Macropus
References: Coulson (1924), DeVis (1897), Turnbull, Lundelius & Tedford (1990).

Smeaton

Type locality: Depth of 50 feet (15 metres) in a well in section 42, Parish of Smeaton, Victoria (37°

16' S., 143° 54' E.).

Rock unit and age: From sediments resting on basalts (Gill 1957b). Aziz-ur-Rahman & McDougall
(1972) reported a date of 2.1 myBP on a similar basalt from the West Berry Consols Mine No. 1 Bore, which
may be the same as the one below the fossiliferous sediment in the well which lies 4.2 kilometres to the
north. On the basis of the stage-of-evolution of Glaucodon ballaratensis, Stitton (1957b) considered it
possible that the specimen was Late Miocene or possibly older while noting that a Pliocene or younger age
was possible if the specimen were a relict form.

Fauna:
AVES
MAMMALIA
Marsupialia
Dasyuromorphia
Dasyuridae
Glaucodon ballaratensis Stirton 1957b
Diprotodontia
Phalangerida
Macropodidae

Comment: Tumbull, Lundelius and Tedford are currently analysing the bird and macropods from this

site.
References: Archer (1982a), Gill (1957b), Stirton (1957b), Turnbull & Lundelius (1967).

Morwell Local Fauna

Type locality: State Electricity Commission of Victoria, Morwell Open Cut Coal Mine, La Trobe

Valley, Victoria (38° 15' S., 146° 29' E.).

Rock unit and age: From unnamed lacustrine clays deposited in depressions at the top of the Morwell
1A Coal Seam. The depressions were formed by fires at the top of the coal seam (hence the mining term
“fireholes" for the sediments filling them). The bulk of the specimens were collected in Firehole no. 2 and a
few more in Firehole no. 3, neither of which any longer exist.

The underlying Morwell 1A Coal Seam is considered to be Late Oligocene to Early Miocene (Partridge
1971), the maximum age for the Morwell Local Fauna. Stratigraphically above the firehole deposits are the
Haunted Hills Gravels, which Jenkin (1966) considers to be Late Pliocene to perhaps Early Pleistocene.
Palynological studies of the firehole sediments by Kirshaw & Sluiter (unpub.) indicate an age of Early
Pliocene to mid-Pleistocene. Flannery (1981), on the basis of all available evidence, suggests that an age of
Late Pliocene to Early Pleistocene is the most likely one for the Morwell fauna.

Fauna:

ARTHROPODA
Ostracoda

Charadriiformes
Pedionomidae
Pedionomus sp., cf. P. torquatus
MAMMALIA
Marsupialia
Diprotodontia
Vombatiformes
Macropodidae
Macropus giganteus Shaw, 1790
Macropus mundjabus Flannery 1980
Protemnodon anak Owen 1874

1058 - RICH, ET AL.

References: Flannery (1980), Rich (1976), Rich & McEvey (1980).

APPENDIX II

LITERATURE REFERENCES TO THE FOSSIL TERRESTRIAL
MAMMALS OF AUSTRALIA

Where does one begin to find the palaeontological literature relating to mammalian genus X? This
question prompted the compiling of references to the genera of terrestrial Australian mammals.

The list is simply a directory with the names arranged in alphabetical order. Where a genus is not listed,
no reference to it in a palaeontological context is known to me. In cases where two genera have been
synonomised, reference is made under the junior synonym to the senior one. In a few instances where a
generic name is no longer used and the species within it have been allocated to two different genera, the
obsolescent name is still used with cross references to the two names currently accepted: e.g. Phascolomys,
Vombatus, and Lasiorhinus.

Virtually every genus published by 1983 and listed here is dealt with in its palaeontological context once
if not several times in the many chapters in Archer, M. & Clayton, G. (eds.) Vertebrate Zoogeography and
Evolution in Australasia, Hesperian Press, Carlisle. Page references to each genus in that volume may be
found in its index. This information is not duplicated here.

Finally, no claims to completeness or balance of treatment are made or intended. This listing is simply
a place to start to answer the question about where the literature can be found, not an exhaustive bibliography
on the topic.

Acrobates: Archer (1978d, 1981a); Archer & Bartholomai (1978); Drummond (1963); Gorter (1977); Hand, Dawson & Augee
(1988); Hope (1973a); Marshall (1981); Simpson (1945); Wakefield (1960a-b, 1963, 1964b, 1967a-b, 1972b).

Aepyprymnus: Abbie (1937); Archer (1978d, 1979, 1981a); Archer & Bartholomai (1978); Bartholomai (1972a, 1977);
Case (1984); Finlayson (1938); Flannery & Archer (1987c); Flannery, Archer & Plane (1983, 1984); Flannery &
Hope (1983); Flannery & Szalay (1982); Gill (1953b); Gillespie et al. (1978); Gillieson & Mountain (1983); Hill,
Playford & Woods, (1970); Hope (1973b); Hope & Wilkinson (1984); Jones (1931); Keast (1972);
Lydekker (1887); Marshall (1981); Pledge (1984, 1990); Ride (1971); Ryder (1974); Simpson (1945); Stirton,
Tedford & Woodburne (1968, 1984); Wakefield (1964b, 1967b, 1974).

Ankotarinja: Aplin & Archer (1987); Archer (1976c, 1981a, 1982a-b); Archer & Bartholomai (1978); Archer &
Rich (1979); Callen & Plane (1986); Marshall (1981); Murray (1984a); Woodbume (1986).

Antechinomys: Archer (1977c, 1981a, 1982a-b); Baynes (1984, 1987); Keast (1972); Lundelius & Turnbull (1989);
Marshall (1981); Morton & Baynes (1985); Smith (1977); Smith & Meilin (1982), Simpson (1945),

Antechinus: Aplin (1981); Archer (1978d, 1981c, 1982a); Archer & Bartholomai (1978); Archer & Baynes (1972); Archer
& Rich (1979); Balme, Merrilees & Porter (1978); Bowdler (1979); Burbidge & George (1978); Callen & Plane
(1986); Cook (1963a); Davison (1986); Drummond (1963); Gill (1968); Gillieson & Mountain (1983);
Gorter (1977); Hope (1973a-b, 1982); Hope & Wilkinson (1984); Hope et al. (1977); Kendrick & Porter (1973);
Lundelius (1957); Lundelius & Tumbull (1973, 1978, 1989); McNamara (1990); Marshall (1981); Merrilees (1979a,
1984); Morton & Baynes (1985); Mulvaney (1961); Murray & Goede (1977); Pledge (1990); Porter (1979);
Ride (1960); Roe (1971); Simpson (1945); Smith (1972); Van Dyck (1982); Wakefield (1960a-b, 1963, 1964a-b,
1967a-b, 1972 a-b); Wells, Moriarty & Williams (1984); White (1967); Whitelaw (1989); Woodburne (1986);
Woodburne, Campbell et al. (1986)

Archizonurus: Anderson (1933); de Vis (1889b); Mahony & Ride (1975).

Balbaroo: Flannery, Archer & Plane (1983); Flannery & Rich (1986).

Baringa: Flannery & Hann (1984); Flannery, Hoch & Aplin (1989); MacFadden et al. (1987).

Bematherium: Archer (1979, 1981a); Archer & Bartholomai (1978); Bartholomai (1972a); Flannery, Archer &
Plane (1983); Hand & Archer (1987); Mahoney & Ride (1975); Marshall (1981); Stirton, Tedford &
Woodbume (1968); Tedford (1967b).

Bettongia: Archer (1972, 1979, 1981a); Archer & Bartholomai (1978); Archer & Baynes (1972); Balme, Merrilees &
Porter (1978); Bartholomai (1972a, 1977); Baynes (1987); Bowdler (1979); Burbidge & George (1978);
Butler (1969); Case (1984, 1985); Drummond (1963), Finlayson (1938); Flannery & Archer (1987c); Flannery,
Archer & Plane (1983, 1984); Flannery & Szalay (1982); Gorecki et al. (1984); Hand & Archer (1987); Gill (1968);
Glauert (1926); Higgins & Petterd (1889); Hill, Playford & Woods (1970); Hope (1978, 1982); Hope &
Wilkinson (1984); Hope et al. (1977); Johnston (1887); Jones (1931); Keast (1972), Kendrick & Porter (1973);
Krefft (1874a); Longman (1924c); Lundelius (1957); Lundelius & Tumbull (1984, 1989); Marshall (1973, 1981);
Merrilees (1968, 1979a-b, 1984); Murray & Goede (1977); Pledge (1984, 1986, 1990); Porter (1979); Ride (1971);
Roe (1971); Ryder (1974), Simpson (1945); Smith (1971, 1972); Smith & Rogers (1981); Stirton, Tedford &
Woodburne (1968); Tedford (1966b); Tedford & Wells (1990); Tedford, Williams & Wells (1986); Thorne (1971);
Wakefield (1960a, 1963, 1964a-b, 1967b, 1972b); Wells, Moriarty & Williams (1984); Woodbume (1984, 1986);
Woodward (1914),

AUSTRALIA'S MAMMALIAN RECORD - 1059

Bohra: Dawson & Flannery (1985); Flannery & Szalay (1982); Quirk & Archer (1983).

Boriogale --- see Macropus

Brachalletes: Dawson & Flannery (1985); de Vis (1883e); Mahoney & Ride (1975); Marshall (1981); Simpson (1945).

Brachipposideros: Wand (1987); Quirk & Archer (1983); Sigé, Hand & Archer (1982).

Bulungamaya: Flannery, Archer & Plane (1983, 1984); Hoch & Holm (1986).

Burramys: Anderson (1933); Anon. (1966a-b); Archer (1981a); Archer & Bartholomai (1978); Broom (1895a, 1896a,
1898); Drummond (1963); Flannery & Hope (1983); Hope (1982); Imaizumi (1968); Keast (1972);
Longman (1924c); Mahoney & Ride (1975); Marshall (1981); Pledge (1981, 1987e); Ride (1956, 1960, 1964,
1971); Simpson (1933, 1945); Stirton, Tedford & Woodburne (1968); Turnbull (1968); Tumbull &
Lundelius (1970); Turnbull, Rich & Lundelius (1987a); Wakefield (1960a-b, 1967a, 1972a-b); Woodbume (1986).

Caloprymnus: Archer (1979, 1981a); Case (1984); Finlayson (1938); Flannery (1983); Flannery & Archer (1987c);
Flannery, Archer & Plane (1984); Flannery & Szalay (1982); Hope (1978); Lundelius (1957); Lundelius & Tumbull
(1984, 1989); Marshall (1981); Merrilees (1968); Simpson (1945); Tedford (1966b),; Woodbume (1984).

Canis: Allen (1972); Anon. (1871); Archer (1981a); Archer & Brayshaw (1978); Baynes (1984, 1987); de Vis (1883d,
1899b); Gill (1953a); Glauert (1926); Godwin (1980); Jones (1921); Krefft (1867); McCoy (1862);
Merrilees (1968, 1979a-b, 1984); Milham & Thompson (1976); Simpson (1945); Smith (1972); Smyth (1869);
Wakefield (1967b); White (1967).

Cercartetus (=Dromicia, Eudromicia): Archer (198la); Archer & Bartholomai (1978); Archer & Baynes (1972); Balme,
Merrilees & Porter (1978); Baynes (1987); Bowdler (1979); Broom (1896a); Cook (1963a); Drummond (1963);
Gill (1965, 1968); Gillieson & Mountain (1983); Godwin (1980); Gorter (1977); Hope (1973a, 1982);
Hope et al. (1977); Keast (1972); Lundelius (1957); Lundelius & Turnbull (1982, 1989); Marshall (1981);
Merrilees (1968, 1984); Murray & Goede (1977); Pledge (1974, 1990); Porter (1979); Ride (1960);
Simpson (1945); Smith (1971); Stirton, Tedford & Woodburne (1968); Tedford et al. (1975); Turnbull &
Schram (1973); Wakefield (1960a-b, 1963, 1964b, 1967a-b, 1972b); Wells, Moriarty & Williams (1984);
Woodbume, Campbell, Rich & Pledge (1986).

Chaeropus: Archer (198la, 1982b); Baynes (1984, 1987); Keast (1972); Lundelius & Tumbull (1981, 1989);
Marshall (1981); Merrilees (1967b, 1968, 1979a); Morton & Baynes (1985); Simpson (1945); Smith (1977);
Tedford (1966b); Wakefield (1964a).

Chalinolobus: Baynes (1987); Simpson (1945); Wakefield (1963, 1967a, 1972b).

Chunia: Callen & Plane (1986); Hand & Archer (1987); Pledge (1986); Rich (1986); Woodburne (1986, 1987a);
Woodbume, Campbell, Rich & Pledge (1986); Woodbume (1986); Woodbume, Campbell (1986); Woodburne &
Clemens (1986a-c).

Conilurus: Archer (1981a); Baynes (1987); Drummond (1963); Hope (1973a); Hope & Wilkinson (1984); Morton &
Baynes (1985); Pledge (1990); Simpson (1945); Wakefield (1960b, 1963, 1964a-b, 1967a-b, 1972b); Wells,
Moriarty & Williams (1984).

Dactylonax: Gillieson & Mountain (1983).

Dactylopsila: Pledge (1986).

Darcius: Rich (1986); Woodbume (1986, 1987a); Woodbume & Clemens (1986b-c); Woodbume, Campbell, Rich &
Pledge (1986)

Dasycercus: Abbie (1937); Archer (1981a, 1982a-b); Archer & Rich (1979); Baynes (1982, 1984, 1987); Burbidge &
George (1978); Crabb (1982); Hope (1978); Keast (1972); Lundelius (1957); Lundelius & Tumbull (1978, 1989);
Marshall (1973, 1981); Morton & Baynes (1985); Simpson (1945); Smith (1977); Smith & Medlin (1982);
Tedford (1966b); Thome (1971); Wakefield (1964a).

Dasykaluta: Archer (1982a-b); Van Dyck (1982).

Dasylurinja ; Archer (1982a-b).

Dasyurinus --- see Dasyurus

Dasyuroides: Archer (198la, 1982a-b); Archer & Rich (1979); Crabb (1982); Lundelius & Turnbull (1978, 1989);
Marshall (1973, 1981); Morton & Baynes (1985); Simpson (1945); Smith & Medlin (1982).

Dasyurus (= Dasyurops, Dasyurinus): Abbie (1937); Anderson (1914); Aplin (1981); Archer (1978d, 1981a, 1982a-b);
Archer & Bartholomai (1978); Archer & Baynes (1972); Archer & Rich (1979); Balme, Merrilees & Porter (1978);
Bartholomai (1971b, 1977); Baynes (1987); Bensley (1901); Bowdler (1979); Butler (1969); Crabb (1982);
Dawson (1982a); Drummond (1963); Flannery & Archer (1984); Gill (1953a, 1968); Gillieson & Mountain (1983);
Glauert (1926); Goede & Murray (1979); Gorter (1977); Hill, Playford & Woods (1970); Hope (1973a-b, 1978;
1982); Hope et al. (1977); Howchin (1930); Keast (1972); Kendrick & Porter (1973); Krefft (1865); Lundelius &
Turnbull (1978, 1989); Lydekker (1887); McCoy (1862); McNamara (1990); Mahoney (1964); Mahoney &
Ride (1975); Marshall (1973, 1981); Marshall & Hope (1973); Merrilees (1967b, 1968, 1979a, 1984); Morton &
Baynes (1985); Murray & Goede (1977); Murray, Goede & Bada (1980); Owen (1843a, 1859c); Pledge (1974, 1980,
1990); Porter (1979); Ryder (1974); Smith (1972); Smith & Medlin (1982); Spencer & Kershaw (1910);
Simpson (1945); Tedford (1966b); Tedford, Williams & Wells (1986); Thorne (1971); Wakefield (1960a, 1964a-b,
1967a-b, 1972b, 1974); Wells, Moriarty & Williams (1984); Woodward, (1914).

Dendrolagus: Abbie (1937); Archer (1979, 1981a); Bartholomai (1972a); Clarke (1878); Dawson & Flannery (1985);
Flannery (1983); Flannery & Archer (1984); Flannery & Hann (1984); Flannery, Hoch & Aplin (1989); Flannery,
Mountain & Aplin (1982); Flannery & Szalay (1982); Gillieson & Mountain (1983); Marshall (1981);
Owen (1877b); Ride (1971); Simpson (1945); Woodbume (1986).

Diarcodon --- see Diprotodon

Dinotherium: Owen (1843b); Simpson (1945).

Diprotodon (=Diarcedon): Abel (1912); Aplin (1987); Anderson, C (1923, 1924a-b, 1925, 1926, 1933); Anderson,
W. (1888, 1890); Anon. (1916, 1924); Aplin & Archer (1987); Archer (1977a, 1981la, 1982a); Archer &
Bartholomai (1978); Archer & Wade (1976); Barrett (1955); Bartholomai (1972a); Beddard (1902); Bennett (1872,
1875, 1876, 1878); Bensley (1903); Browne (1945); Case (1985); Clarke (1878); Daintree (1872); David (1916); de

1060 - RICH, ET AL.

Vis (1883a-b, 1887a, 1888a-b, 1889a, 1891a, 1895b, 1900, 1907); Dennant & Kitson (1903); Dugan (1981);
Dulhunty, Flannery & Mahoney (1984); Dun (1892, 1893, 1894, 1895, 1900); Duncan (1884, 1885); Etheridge
(1878, 1890, 1891, 1892, 1894a-c, 1897, 1918); Flannery & Gott (1984); Flannery & Hann (1984); Flannery &
Hope (1983); Flannery & Szalay (1982); Fletcher (1954), Flower & Lydekker (1891); Forbes (1894); Gill (1953a-
b, 1955a-b, 1957a-b, 1962, 1963, 1965, 1978), Gill & Banks (1956); Gillespie ef al. (1978); Glauert (1912a-b,
1921a, 1926, 1948); Gorecki ef al. (1984); Gorter & Nicoll (1978); Gould (1863); Gregory (1910, 1912); Hand,
Dawson & Augee (1988); Hardman (1884); Harlon (1846); Harper (1945); Hill, Playford & Woods (1970);
Hobson (1844, 1845a-b, 1846a-b, 1848); Hochstetter (1859); Hope (1973b, 1978, 1982); Hope, Dare-Edwards &
Melntyre (1983); Horton (1978, 1979, 1980, 1981, 1984); Horton & Connah (1981); Horton & Wright (1981);
Howchin (1891, 1918, 1930); Huxley (1862, 1899); Jack (1879); Jack & Etheridge (1892); Johns &
Ludbrook (1963); Keast (1968, 1972); Keble (1945); Kirk (1981); Krefft (1865, 1870, 1871la-b, 1872, 1873,
1874a, 1875a); Lester et al. (1988); Lindsay (1966); Longman (1916, 1921, 1924a-d, 1925, 1926, 1927, 1929,
1935); Lydekker (1887, 1888, 1896a-b); McCoy (1861, 1862, 1865a, 1866, 1867, 1874a, 1875, 1876a-b, 1877);
MacFadden et al. (1987); McIntyre & Hope (1978); McNamara (1990); Mahoney & Ride (1975); Marcus (1976).
Marshall (1973, 1981); Merrilees (1968, 1969, 1979); Mitchell (1831, 1838, 1862); Murray (1984b); Murray &
Goede (1977); Nicholson & Lydekker (1889); Noetling 1911, 1912); Owen (1838, 1839, 1840, 1845, 1859a-c,
1870, 187la-b, 1872a, 1876, 1877a-b, 1882a-b, 1883a, 1884); Palmer (1904); Pilling & Waterman (1970);
Pledge (1973, 1974, 1981, 1990); Pritchard (1899), Pulleine (1927); Quirk & Archer (1983); Ramsay (1881,
1892); Rich, P. (1981); Rich, P. & van Tets (1985); Rich, T. (1976, 1981, 1986); Ride (1964, 1966, 1970,
1971); Schmidt (1885); Scott (1910, 1912, 1915b); Scott & Lord (192la-d, 1923a), Simpson (1930, 1945);
Stephenson (1963, 1964); Stetson (1933); Stirling (1893, 1894, 1900a-b, 1901, 1907a-b, 1913); Stirling &
Zeitz (1896, 1899a, 1900); Stirton (1953, 1954, 1955, 1957b); Stirton, Tedford & Miller (1961); Stirton, Tedford
& Woodbume (1968); Stirton, Woodburne & Plane (1967); Sussmilch (1922); Tate (1948); Tedford (1955, 1966b,
1973); Tedford & Wells (1990); Tedford, Williams & Wells (1986); Tindale (1957); Troughton (1959); Vanderwal &
Fullagar (1989); Walcott (1920); Waterhouse (1846); Wells(1978); White & O'Connell (1979); Whitley (1966);
Wilkinson (1972); Winge (1923, 1941), Woods, J.E.T. (1860); Woods, J.T. (1956a, 1960a, 1962); Woodward,
A.S. (1907); Woodward, B.H. (1909, 1910, 1914); Wright (1986).

Djilgaringa: Archer, Tedford & Rich (1987)

Dorcopsis: Archer (1979, 1981a); Archer & Barthollomai (1978); Bartholomai (1972a); Clarke (1878); Dawson &
Flannery (1985); Flannery (1983); Flannery & Archer (1983); Flannery, Archer & Plane (1983); Flannery &
Hann (1984); Flannery, Hoch & Aplin (1989); Flannery, Mountain & Aplin (1982); Flannery & Szalay (1982);
Hoch & Holm (1986); Marshall (1981); Owen (18776); Plane (1967b); Simpson (1945); Stirton, Tedford &
Woodbume (1968); Tumbull & Lundelius (1970); Woodbume (1967b, 1984, 1986); Woodburne, Campbell, Rich &
Pledge (1986).

Dorcopsoides: Archer (1981a); Archer & Bartholomai (1978), Bartholomai (1972a); Dawson & Flannery (1985);
Flannery (1983); Flannery, Archer & Plane (1983); Flannery, Hoch & Aplin (1989); Keast (1972); Mahoney &
Ride (1975); Marshall (1981); Murray (1984a); Ride (1971); Stirton, Tedford & Woodburne (1968);
Woodburne (1967b).

Dorcopsulus: Archer (1981a); Archer & Bartholomai (1978); Bartholomai (1972a); Dawson & Flannery (1985);
Flannery (1983); Flannery, Archer & Plane (1983); Flannery & Hann (1984); Flannery, Mountain & Aplin (1982)
Gillieson & Mountain (1983); Marshall (1981).

Dromicia --- see Cercartetus

Echidna (see also Zaglossus & Tachyglossus): Archer (1981a); Broom (1896a); de Vis (1885a); Dun (1895); Flower &
Lydekker (1891); Glauert (1910b, 1914, 1926); Howchin (1930); Krefft (1868); Lydekker (1887, 1896a); Mahoney
& Ride (1975); Owen (1884, 1887b, 1889); Simpson (1945); Woodward, B. (1910).

Echymipera: Archer (1981a), Archer & Bartholomai (1978); Callen & Plane (1986); Gillieson & Mountain (1983);
Keast (1972); Marshall (1981); Simpson (1945); Stirton, Tedford & Woodbume (1968).

Ekaltadeta: Archer & Flannery (1985).

Ektopodon: Archer (1981a); Archer & Bartholomai (1978); Callen & Plane (1986); Clemens (1977); Mahoney &
Ride (1975); Marshall (1981); Pledge (1984, 1986); Rich (1986); Rich & van Tets (1985); Stirton, Tedford &
Woodburne (1967, 1968); Woodburne (1986, 1987a); Woodburne, Campbell et al. (1986); Woodbume &
Clemens (1986a-c); Woodbume & Tedford (1975).

Eptesicus: Simpson (1945); Wakefield (1967a, 1972b).

Eudromicia --- see Cercartetus

Euowenia (=Owenia): Archer (1981a); Archer & Bartholomai (1978); Archer & Wade (1976); Bartholomai (1972a); de
Vis (1887, 1888b, 1891a, 1895b, 1907); Dun (1894), Etheridge (1918); Hand & Archer (1987); Hill, Playford &
Woods (1970); Horton (1984); Keast (1972); Longman (1916, 1921); McNamara (1990); Mahoney & Ride (1975);
Marshall (1981); Murray (1984a-b); Pledge (1989a); Scott & Lord (1921a-c); Simpson (1945); Stirton, Tedford &
Woodburne (1968); Stirton, Woodbume & Plane (1967); Tedford, Williams & Wells (1986); Tumbull, Lundelius &
Tedford (1990).

Euryzygoma: Anderson (1933); Aplin (1987); Archer (1977a, 1981a); Archer & Bartholomai (1978); Archer &
Flannery (1985); Archer & Wade (1976); Bartholomai (1972a); Fletcher (1951); Hill, Playford & Woods (1970);
Horton (1984); Howchin (1930); Keast (1972); Longman (1921, 1924b-c, 1926, 1927, 1929, 1935); Mahoney &
Ride (1975); Marshall (1981); Murray (19842); Quirk & Archer (1983); Rich, T. (1981), Scott (1927); Scott &
Lord (1922a); Simpson (1945); Stirton, Tedford & Woodbume (1968); Stirton, Woodbume & Plane (1967); Tedford
& Wells (1990); Tedford, Williams & Wells (1986).

Fissuridon: Archer (1981a); Archer & Wade (1976); Bartholomai (1973); Dawson & Flannery (1985); Flannery &
Archer (1982); Horton (1984); Marshall (1981); Murray (1984b); Sanson (1978); Tedford, Williams &
Wells (1986).

Galanaria: Flannery, Archer & Plane (1983).

AUSTRALIA'S MAMMALIAN RECORD - 1061

Glaucodon: Archer (1981a, 1982a-b); Archer & Bartholomai (1978); Crabb (1982); Keast (1972); Mahoney &
Ride (1975); Marshall (1973, 1981); Murray (1984a); Quirk & Archer (1983); Ride (1964); Stirton (1957b).

Gumardee: Flannery, Archer & Plane (1983, 1984); Flannery & Rich (1986); Hoch & Holm (1986); Woodbume (1984).

Gymnobelideus (=Palaeopetaurus): Archer (1981a); Broom (1895b, 1896a); Drummond (1963); Hope (1973a, 1982);
Mahoney & Ride (1975); Marshall (1981); Ride (1960); Simpson (1945); Tumbull, Rich & Lundelius (1987a);
Wakefield (1960a-b, 1967a, 1972a-b)

Gyomys: Drummond (1963); Ride (1960); Simpson (1945); Wakefield (1960b, 1963, 1964»).

Hadronomas: Archer (1979, 1981a); Archer & Barthoholomai (1978); Bartholomai (1972a); Dawson & Flannery (1985);
Flannery (1983); Flannery, Archer & Plane (1983); Flannery, Hoch & Aplin (1989); Mahoney & Ride (1975);
Marshall (1981); Ride (1971); Stirton, Tedford & Woodbume (1968); Woodbume (1967b).

Halmaturotherium (=Halmatutherium): Dawson & Flannery (1985); Krefft (1874); Mahoney & Ride (1975).

Halmaturus --- see Macropus.

Halmatutherium --- see Halmaturotherium

Hipposideros (Brachipposideros): Flannery, Archer & Plane (1983); Hand (1987); Lester & Hand (1987); Rich &
van Tets (1985); Sigé, Hand, & Archer (1982)

Homo: Allen (1972); Marshall (1973); Merrilees (1968); Simpson (1945); Thome (1981); White & O'Connell (1979).

Hulitherium: Flannery & Plane (1986); Hand & Archer (1987)

Hydromys: Archer (1981a); Archer & Baynes (1972); Baynes (1987); Bowdler (1979); Hope (1973a);
Hope et al. (1977); Marshall (1973); Merrilees (1979a, 1984); Morton & Baynes (1985); Murray & Goede (1977);
Pledge (1990); Porter (1979), Ryder (1974); Simpson (1945); Tedford (1966b); Wakefield (1964a-b, 1967a-b,
1972b); White (1967).

Hyomys: Gillieson & Mountain (1983)

Hypsiprymnodon: Archer (1979, 1981a, 1982b); Archer & Bartholomai (1978); Archer, Bartholomai & Marshall (1978);
Archer, Hand & Godthelp (1988); Bartholomai (1972a); Bensley (1901); Broom (1896a); Case (1984);
de Vis (1888c); Finlayson (1938); Flannery (1983); Flannery & Archer (1987b); Flannery, Archer & Plane (1983,
1984); Flannery & Rich (1986); Flannery & Szalay (1982); Keast (1972); Marshall (1981); Pledge (1981, 1984);
Ride (1971); Simpson (1945); Stirton, Tedford & Woodburne (1968); Woodburne (1984, 1986);
Woodburne et al. (1986).

Hypsiprymnus --- see Potorous.

Naria: Pledge (1987c); Tedford & Woodbume (1987)

Ischnodon: Archer (1981a); Archer & Bartholomai (1978); Mahoney & Ride (1975); Marshall (1981); Ride (1964),
Stirton (1955); Stirton, Tedford & Woodbume (1968); Tedford, Williams & Wells (1986).

Isoodon (=Thylacis): Aplin (1981); Aplin & Archer (1987); Archer (1972, 1978d, 1981a); Archer & Bartholomai (1978);
Archer & Baynes (1972); Archer & Brayshaw (1978); Balme, Merrilees & Porter (1978); Bartholomai (1977);
Baynes (1982, 1987); Bowdler (1979); Burbidge & George (1978); Butler (1969); Callen & Plane (1986);
Case (1985); Cook (1963a); Drummond (1963); Gill (1968); Glauert (1926); Godwin (1980); Gorter (1977);
Keast (1972); Kendrick & Porter (1973); Hope (1973a-b, 1978, 1982); Hope & Wilkinson (1984);
Hope et al. (1977); Lundelius & Turnbull (1981, 1989); McNamara (1990); Marshall (1973, 1981);
Merrilees (1967b, 1968, 1969, 1979a-b, 1984); Morton & Baynes (1985); Murray & Goede (1977); Pledge (1974,
1990); Porter (1979); Roe (1971); Ryder (1974); Simpson (1945); Smith (1972, 1977); Tedford (1966b);
Wakefield (1960a-b, 1963, 1964a-b, 1967a-b, 1972b, 1974); White (1967); Wells, Moriarty & Williams (1984);
Whitelaw (1989); Woodward, B. (1914).

Kangurus --- see Lagostrophus

Keeuna: Archer (1976c, 1981a, 1982a-b); Archer & Bartholomai (1978); Callen & Plane (1986); Marshall (1981);
Woodbume (1986).

Koalemus: Anderson (1933); Bartholomai (1968); Chapman (1934); de Vis (1889b); Mahoney & Ride (1975);
Marshall (1981).

Kolopsis: Archer (1981a); Archer & Bartholomai (1978); Archer & Wade (1976); Bartholomai (1972a); Clemens &
Plane (1974); Flannery & Plane (1986); Flannery, Mountain & Aplin (1982); Hoch & Holm (1986); Mahoney &
Ride (1975); Marshall (1981); Plane (1967a-b); Rich, Archer & Tedford (1978); Stirton, Tedford &
Woodbume (1968); Stirton, Woodbume & Plane (1967); Tedford, Williams & Wells (1986); Woodburne (1967a-b,
1969).

Kolopsoides: Archer (1981); Archer & Bartholomai(1978); Archer & Wade (1976); Bartholomai (1972a); Flannery &
Plane (1986); Flannery, Mountain & Aplin (1982); Mahoney & Ride (1975); Marshall (1981); Plane (1967a-b);
Stirton, Tedford & Woodbume (1968); Stirton, Woodbume & Plane (1967), Woodburne (1969).

Koobor: Aplin & Archer (1987); Archer (1977b, 1981a); Archer & Bartholoomai (1978); Archer & Wade (1976);
Marshall (1981); Pledge (1987a,c); Springer (1987); Tedford & Woodburne (1987); Woodbume, Tedford, Archer &
Pledge (1987).

Kurrabi: i & Archer (1984); Flannery, Hoch & Aplin (1989); Tumbull, Lundelius & Tedford (1990).

Kuterintja: Aplin & Archer (1987); Pledge (1987c)

Lagorchestes: Archer (1981a); Balme, Merrilees & Porter (1978); Bartholomai (1972a); Baynes (1982, 1984, 1987);
Burbidge & George (1978); Clarke (1878); Dawson & Flannery (1985); Flannery (1983); Flannery & Hann (1984);
Flannery, Hoch & Aplin (1989); Flannery & Szalay (1982); Gorecki ef al. (1984); Hope (1978);
Hope et al. (1977); Huxley (1862); Lundelius & Turnbull (1989); Marshall (1973, 1981); Merrilees (1968, 1984);
Pledge (1990); Simpson (1945); Tedford (1966b); Tedford, Williams & Wells (1986); Wakefield (1964a-b, 1967b);
Wells, Moriarty & Williams (1984); White (1967).

Lagostrophus: Archer (1981a); Baynes (1982); Dawson & Flannery (1985); Flannery (1983), Flannery & Archer (1983);
Flannery, Archer & Plane (1984); Lundelius & Tumbull (1989); Marshall (1973, 1981); Merrilees (1968);
Simpson (1945); Wakefield (1964a).

1062 - RICH, ET AL.

Lasiorhinus (see also Phascolomys): Archer (1981a); Archer & Bartholomai (1978); Baynes (1987); Dawson (1981,
1983a-b); Flannery & Hope (1983); Gorter (1977); Hope (1978); Hope & Wilkinson (1984); Hope et al. (1977);
Horton (1984); Keast (1972); Leach (1977); Lundelius & Turnbull (1982, 1989); Marshall (1973, 1981);
Merrilees (1967a, 1968); Pledge (1990); Simpson (1945); Tedford (1966b); Wakefield (1964a).

Leggadina --- see Pseudomys

Leporillus: Baynes (1982, 1984, 1987); Hope (1978); Kendrick & Porter (1973); Lundelius (1957, 1964);
Marshall (1973); Merrilees (1979a); Morton & Baynes (1985); Simpson (1945); Smith (1977); Tedford (1966b);
Thorne (1971).

Leptosiagon --- see Macropus

Litokoala: Aplin & Archer (1987); Archer (1977b, 1981a), Archer & Bartholomai (1978); Callen & Plane (1986);
Mahoney & Ride (1975); Marshall (1981); Springer (1987); Stirton, Tedford & Woodburne (1967, 1968);
Woodbume, Campbell et al. (1986); Woodbume, Tedford, Archer & Pledge (1987).

Lyroderma Hand (1987).

Macroderma: Archer (1981a); Archer & Brayshaw (1978); Hand (1985, 1987); Hand & Archer (1987); Hand, Dawson &
Augee (1988); Lundelius (1957); Merrilees (1979a, 1984); Molnar et al, (1984); Simpson (1945); White (1967).

Macropus (=Boriogale, Halmaturus, Leptosiagon, Megaleia, Osphranter, Phascolagus); Abbie (1937); Allen (1972);
Anderson, C. (1926, 1929a, 1932); Anderson, W. (1889, 1914); Anon, (1916); Aplin (1981); Archer (1972,
1978d, 1981d); Archer & Bartholomai (1978); Archer & Brayshaw (1978); Archer & Wade (1976); Balme, Merrilees
& Porter (1978); Bartholomai (1963, 1966, 1971a, 1972a, 1975, 1977, 1978a); Baynes (1984, 1987);
Bennett (1878); Bowdler (1979); Broom (1896a, 1935); Butler (1969); Case (1985); Clarke (1878);
Cook (1963a); Dawson & Flannery (1985); de Vis (1883a,c-e, 1895a, 1897, 1899a-b, 1907); Downie &
White (1978); Dun (1893, 1900); Finlayson (1948, 1949); Flannery (1980, 1984); Flannery & Archer (1982,
1984); Flannery & Gott (1984); Flannery & Hann (1984); Flannery, Hoch & Aplin (1989); Flannery &
Hope (1983); Flannery & Szalay (1982); Flower (1867); Gill (1953a-b, 1954a, 1957b, 1965, 1968, 1978);
Gillespie et al. (1978); Glauert (1912b, 1921b, 1926); Godwin (1980); Goede & Murray (1979); Gorecki et al.
(1984); Gorter (1977); Hand, Dawson & Augee (1988); Higgins & Petterd (1889); Hill, Playford & Woods (1970);
Hope (1973a-b, 1978, 1982); Hope & Wilkinson (1984); Hope et al. (1977); Horton (1978, 1980, 1984); Horton
& Connah (1981); Horton & Murray (1980); Howchin (1930), Huxley (1862); Johnston (1887); Jones (1931);
Keast (1972); Kendrick & Porter (1973); Kreffft (1865, 1867, 1870, 1875b); Leach (1977); Longman (1924a-b,
1926, 1927); Lydekker (1887, 1891, 1895, 1896a); Lundelius & Tumbull (1989); McCoy (1862, 1874b, 1879);
McIntyre & Hope (1978); MacFadden et al. (1987); Mahoney & Ride (1975); Marcus (1976); Marshall (1973,
1974, 1981); Marshall & Corruccini (1978); Merrilees (1968, 1979a-b, 1984); Murray (1984b); Murray &
Goede (1977); Murray, Goede & Bada (1980); Owen (1843a-b, 1859c, 1870, 1871b, 1873, 1874a-b, 1876, 1877,
1882b, 1883b-c); Pledge (1974, 1980a-b, 1981, 1990); Porter (1979); Ride (1960, 1964, 1971); Ryder (1974);
Scott (1905); Scott & Lord (1922a, 1924a); Simpson (1945); Spencer & Kershaw (1910); Stirling &
Zietz (1899a); Stixton, Tedford & Woodburne (1968); Tate (1948); Tedford (1966b); Tedford & Wells (1990);
Tedford, Williams & Wells (1986); Thome (1971); Turnbull, Lundelius & Tedford (1990); Vanderwal & Fullagar
(1989); Wakefield (1964a-b, 1967b, 1974); Walcott (1920); Wells (1978); Wells, Moriarty & Williams (1984);
White 1967); White & O'Connell (1979); Whitelaw (1989); Wilkinson,C.S. (1892); Wilkinson, H.E. (1972);
Woodward (1914); Wright (1986).

Macrotis (=Peragale, Thalacomys, Thylacomys): Abbie (1937); Archer (1981a); Archer & Bartholomai (1978);
Baynes (1987); Glauert (1926); Keast (1972); Lundelius (1957); Lundelius & Turnbull (1981, 1989);
Lydekker (1887); Marshall (1973, 1981); Merrilees (1967b, 1968); Morton & Baynes (1985); Pledge (1984);
Simpson (1945); Tedford (1967a); Tedford & Wells (1990); Tedford, Williams & Wells (1986); Woodward,
B. (1914).

Madakoala: Aplin & Archer (1987); Tedford & Woodbume (1987); Springer (1987); Woodbume, Tedford, Archer &
Pledge (1987)

Mallomys: Gillieson & Mountain (1983)

Marlu: Pledge (1987d); Woodbume, Tedford & Archer (1987)

Mastacomys: Bowdler (1979); Drummond (1963); Gill (1968); Gillespie et al. (1978); Godwin (1980); Goede &
Murray (1979); Hope (1973a-b, 1982); Hope et al. (1977); Mahoney & Ride (1975); Marshall (1974);
Mulvaney (1961); Murray & Geode (1977); Murray, Goede & Bada (1980); Pledge (1990); Ride (1960);
Simpson (1945); Thomas (1922); Wakefield (1960a, 1963, 1964b, 1967a-b, 1972a-b); Wells, Moriarty &
Williams (1984).

Megaleia --- see Macropus.

Melomys: Baynes (1987); Godthelp (1990); Morton & Baynes (1985); Ryder (1974); Simpson (1945); White (1967);
Wakefield (1967a, 1974).

Meniscolophus: Archer (1981a); Archer & Bartholomai (1978); Bartholomai (1972a). Mahoney & Ride (1975);
Marshall (1981); Stirton (1955, 19667b); Stirton, Tedford & Woodbume (1968); Stirton, Woodbume &
Plane (1967); Tedford, Williams & Wells (1986); Woodbume (1967b, 1969).

Mesembriomys: Archer & Brayshaw (1978); Baynes (1987); Kendrick & Porter (1973); Morton & Baynes (1985);
Simpson (1945); White (1967).

Microperoryctes: Archer (1981a, 1982a); Marshall (1981).

Miniopterus: Drummond (1963); Simpson (1945); Wakefield (1967a, 1972b).

Miralina: Tedford & Woodbume (1987); Woodbume, Pledge & Archer (1987)

Muramura: Aplin & Archer (1987); Pledge (1987b)

Murexia: Archer (1981a, 1982a); Archer & Bartholomai (1978). Marshall (1981).

Mus: Baynes (1987); Drummond (1963); Higgins & Petterd (1889); Johnston (1887); Lundelius (1964);
Simpson (1945); Smith (1977); Spencer & Kershaw (1910).

Mylodon --- see Thylacoleo.

AUSTRALIA'S MAMMALIAN RECORD - 1063

Myoictis: Archer (1981a, 1982a); Archer & Bartholomai (1978); Archer & Rich (1979); Hoch & Holm (1986);
Marshall (1981).

Myotis: Drummond (1963); Simpson (1945).

Myrmecobius: Abbie (1937); Archer (198la, 1982 a-b); Bensley (1901); Hofer (1952); Keast (1972); Krefft (1874a);
Lundelius & Turnbull (1978, 1989); Marshall (1981); Morton & Baynes (1985); Ride (1964); Simpson (1945);
Tedford (1966b); Wakefield (1964).

Nambaroo: Flannery & Rich (1986)

Namilamadeta: Aplin & Archer (1987); Archer, Hand & Godthelp (1988); Hand & Archer (1987); Lester et al. (1988);
Marshall (1981); Pledge (1987b); Rich & Archer (1979); Tedford & Woodburne (1987); Woodburne (1986)
Woodburne, Campbell et al. (1986).

Neohelos: Aplin (1987); Archer (1981a); Archer & Bartholomai (1978); Bartholomai (1972a); Callen & Plane (1986);
Clemens & Plane (1974); Flannery, Archer, & Plane (1982); Jupp et al. (1989); Mahoney & Ride (1975);
Marshall (1981); Murray (1984a); Pledge (1984); Quirk & Archer (1983); Rich, Archer & Tedford (1978);
Stirton (1967c); Stirton, Tedford & Woodbume (1968); Stirton, Woodburne & Plane (1967); Woodburne (1967b,
1986); Woodburne, Campbell et al. (1986).

Neophascogale: Archer (1981a, 1982a); Marshall (1981).

Nimbacinus: Muirhead & Archer (1990).

Ningaui: Morton & Baynes (1985)

Ngapakaldia: Aplin & Archer (1987); Archer (1981a), Archer & Bartholomai (1978); Bartholomai (1972a); Callen &
Plane (1986); Hand & Archer (1987); Hoch & Holm (1986); Lester et al. (1988); Mahoney & Ride (1975);
Marshall (1981); Murray (1984a); Pledge (1984); Rich, T. (1981); Rich, Archer & Tedford (1978); Rich &
Rich (1987); Rich & van Tets (1985); Stirton (1967a); Stirton, Tedford & Woodbume (1968); Stirton, Woodburne
& Plane (1967); Tedford et al. (1975, 1977); Waters & Savage (1969); Woodbume (1967b, 1986); Woodburne,
Campbell et al. (1986); Woodbume & Tedford (1975).

Notelephas: Longman (1916); Mahoney & Ride (1975); Owen (1882c, 1883a).

Notomys: Archer (1972, 1981a); Baynes (1982, 1984, 1987); Hope (1978); Kendrick & Porter (1973); Lundelius (1964);
Marshall (1973); Merrilees (1979a, 1984); Morton & Baynes (1985); Simpson (1945); Smith (1977);
Tedford (1966b); Tedford & Wells (1990); Thorne (1971); Wakefield (1964a).

Notoryctes: Abbie (1937); Anderson (1925); Archer (1981a); Archer & Bartholomai (1978); Bensley (1901);
Marshall (1981); Morton & Baynes (1985); Simpson (1945).

Nototherium (see also Zygomaturus): Anderson, C. (1924a-b, 1933, 1937); Anderson, W. (1914); Anon. (1916);
Archer (1981a); Archer & Bartholomai (1978), Archer & Wade (1976); Bartholomai (1972a, 1977); Bennett (1875,
1876); Case (1985); Clarke (1878); Cook (1963a); de Vis (1883a-b, 1887a-b, 1888a-b, 1889a, 1891a, 1895b,
1899a, 1907); Dun (1892, 1893, 1894); Etheridge (11918); Flannery, Mountain & Aplin (1982); Flower &
Lydekker (1891); Gill (1953c, 1978); Gill & Banks (1956); Glauert (1912a-b, 1921a,1926); Guérin &
Faure (1987); Guérin, Winslow, Piboule & Faure (1981); Hill, Playford & Woods (1970); Hope (1973b);
Horton (1980, 1984); Horton & Murray (1980); Howchin (1930); Huxley (1862); Keast (1972); Keble (1945);
Krefft (1865, 1866, 1870, 1872, 1874a, 1875b); Lester, Boyde, Gilkeson & Archer (1987); Longman (1916, 1921,
1924c, 1926); Lydekker (1887, 1890, 1896a); McCoy (1862, 1865b); Mahoney & Ride (1975); Marshall (1981);
Murray (1984b); Murray & Goede (1977); Noetling (1912); Owen (1859a-c, 1866, 1870, 1871a-b, 1872a, 1874 a-b,
1876, 1877a-b, 1880, 1882a-b, 1884, 1889); Plane (1967a-b); Pledge (1973); Ride (1971); Scott (1912, 1915a-b,
1917, 1927); Scott & Harrison (1911); Scouw & Lord (1921a-d, 1922a-b, 1923a, 1924a-d, 1925a-b, 1926);
Simpson (1945); Stirling (1900b); Stirton, Tedford & Woodburne (1968); Stirton, Woodbume & Plane (1967);
Tedford (1966b); Tedford & Wells (1990); Tedford, Williams & Wells (1986); Whitley (1966); Woodbume (1967b);
Woodward, B. (1909, 1910, 1914).

Nyctophilus: Archer & Baynes (1972); Baynes (1987); Cook (1963a); Gill (1968); Pledge (1990); Simpson (1945);
Wakefield (1963, 1967a, 1972b).

Obdurodon: Archer (1981a); Archer & Bartholomai (1978); Archer, Every, Godthelp, Hand & Scally (1990); Archer, Plane
& Pledge (1978); Callen & Plane (1986); Lester & Archer (1986); Lester, Boyde, Gilkeson & Archer (1987);
Murray (1984a); Quirk & Archer (1983); Woodburne (1986); Woodbume, Campbell et al. (1986); Woodburne &
Clemens (1986b-c); Woodbume & Tedford (1975).

Onychogalea: Archer (1981la); Bartholomai (1972a); Baynes (1987); Dawson & Flannery (1985); Flannery &
Hann (1984); Flannery, Hoch & Aplin (1989); Flannery & Szalay (1982); Hope (1978); Jones (1931); Lundelius &
Tumbull (1989); Marshall (1973, 1981); Merrilees (1968, 1979a, 1984); Simpson (1945); Tedford (1966b); Tedford
& Wells (1990); Wakefield (1964a-b, 1967b).

Ornithorhynchus: Archer (1981a); Archer & Bartholomai (1978); Archer, Plane & Pledge (1978); de Vis (1885a-b);
Dun (1895); Mahoney & Ride (1975); Murray (1978b); Simpson (1945); Woodbume & Clemens (1986c);
Woodbume & Tedford (1975).

Ornoryctes --- see Peroryctes.

Osphranter --- see Macropus.

Owenia --- see Euowenia.

Pachysiagon --- see Procoptodon.

Palaeopotorous: Flannery & Rich (1986)

Palaeopteraurus -— sce Gymnobelideus.

Paljara: Aplin & Archer (1987); Woodburne, Tedford & Archer (1987)

Palorchestes: Anderson (1924a-b, 1933); Aplin & Archer (1987); Archer (1981a); Archer & Bartholomai (1978); Archer
& Wade (1976); Banks, Colhoun & van de Geer (1976); Bartholomai (1962, 1963, 1972a, 1977, 1978b);
Cudmore (1926); de Vis (1883a,c,e, 1895a); Dun (1893, 1894); Flannery (1983); Flannery & Archer (1984);
Flannery & Gott (1984); Flannery & Hann (1984); Gill (1953b-c); Glauert (1912b, 1926); Hall & Pritchard (1897),
Hill, Playford & Woods (1970); Hope (1973b, 1982); Horton (1980, 1984); Howchin (1930); Keast (1972);

1064 - RICH, ET AL.

Lester et al. (1988); Longman (1924c); Lydekker (1887, 1895, 1896a); MacFadden et al. (1987); Mahoney &
Ride (1975); Marshall (1981); Molnar (1978a); Murray (1978a, 1984a-b); Murray & Goede (1977); Owen (1873,
1874a, 1876, 1877a, 1879, 1880); Quirk & Archer (1983); Ramsay (1886a,c); Raven (1929); Raven &
Gregory (1946); Rich, P. & van Tets (1985); Rich, T. (1981); Ride (1971); Scott (1916, 1917); Scou &
Lord (1921c, 1925b); Simpson (1945); Stirton, Tedford & Woodburne (1968); Stirton, Woodbure & Plane (1967);
Tate (1948); Tedford (1966b); Tedford, Williams & Wells (1986); Turnbull & Lundelius (1970); Wells (1978);
Wells, Morisrty & Williams (1984); White & O'Connell (1979); Woodburne (1967a-b); Woodburne &
Clemens (1986c); Woods (1958).

Parantechinus: Archer (1982a); Keast (1972); Lundelius (1957); Lundelius & Tumbull (1978); Marshall (1981); Stirton,
Tedford & Woodbume (1968); Van Dyck (1982).

Peradorcas: Archer (1981a); Flannery (1983); Flannery & Szalay (1982); Marshall (1981); Simpson (1945).

Peragale --- see Macrotis.

Perameles: Abbie (1937); Archer (1972, 1978d, 1981a); Archer & Bartholomai (1978); Archer & Wade (1976); Balme,
Merrilees & Porter (1978); Bartholomai (1977); Baynes (1982, 1987); Bensley (1901); Bowdler (1979);
Broom (1896a); Burbidge & George (1978); Callen & Plane (1986); Case (1985); Drummond (1963), Flannery &
Hope (1983); Freedman & Joffe (1967); Gill (1968); Glauert (1926); Gorter (1977); Hill, Playford &
Woods (1970); Hope (1973a-b, 1978, 1982); Hope & Wilkinson (1984); Hope et al. (1977); Howchin (1930);
Keast (1972); Kendrick & Porter (1973); Lundelius (1957); Lundelius & Tumbull (1981, 1989); Lydekker (1887);
Mahoney & Ride (1975); Marshall (1973, 1974, 1981); Morton & Baynes (1985); Merrilees (1967b, 1968, 1979a,
1984); Murray & Goede (1977); Murray, Goede & Bada (1980); Owen (1877a) Partridge & Thome (1963);
Pledge (1974, 1990); Porter (1979); Ride (1960); Ryder (1974), Simpson (1945), Smith (1972); Tedford (1966b);
Thome (1971); Wakefield (1960a-b, 1963, 1964a-b, 1967a-b, 1972b, 1974); Wells, Moriarty & Williams (1984);
Whitelaw (1989); Woodbume (1986); Woodward (1914).

Perikoala: Aplin & Archer (1987); Archer (1981a); Archer & Bartholomai (1978); Callen & Plane (1986); Keast (1972);
Mahoney & Ride (1975); Marshall (1981); Springer (1987); Stirton (1957a); Stirton, Tedford & Woodbume (1967,
1968); Tedford & Woodburne (1987); Tedford et al. (1977); Woodburne (1986); Woodburne,
Campbell et al. (1986); Woodbume & Clemens (1986b); Woodburne, Tedford, Archer & Pledge (1987).

Peroryctes (=Ornoryctes): Archer (1981a, 1982b); Archer & Bartholomai (1978); Callen & Plane (1986); Gillieson &
Mountain (1983); Keast (1972); Marshall (1981); Simpson (1945); Stirton, Tedford & Woodbume (1968).

Petauroides --- see Schinobates.

Petaurus: Abbie (1937); Aplin (1981); Archer (1978d, 1981a); Broom (1895b, 1896a-b); Drummond (1963); Gillieson &
Movntain (1983); Hope (1973a); Jones (1931); Marshall (1981); Pledge (1990); Ride (1960); Ryder (1974);
Simpson (1945); Smith (1971); Turnbull, Rich & Lundelius (1987a); Wakefield (1960a-b, 1963, 1964b, 1967a-b,
1972a-b, 1974); Wells, Moriarty & Williams (1984); White (1967), Woodbume (1986).

Petramops: Hand 1990.

Petrogale: Aplin (1981); Archer (1981a); Archer & Brayshaw (1978); Bartholomai (1972a, 1978a); Baynes (1982, 1984,
1987); Clarke (1878); Dawson & Flannery (1985); Flannery (1980, 1983); Flannery, Hoch & Aplin (1989);
Flannery & Szalay (1982); Hope (1973a); Jones (1931); Kendrick & Porter (1973); Krefft (1875b);
Longman (19244); Lundelius & Tumbull (1989); Marshall (1973, 1981); Merrilees (1968, 1979b, 1984);
Ride (1971); Simpson (1945); Wakefield (1964b, 1967a, 1972b, 1974); White (1967).

Petropseudes -— see Pseudocheirus.

Phalanger (=Phalangista): Archer (1981a); Archer & Bartholomai (1978); de Vis (1888c); Downie & White (1978);
Flannery, Archer & Plane (1983); Flannery & Rich (1986); Gillieson & Mountain (1983); Higgins &
Petterd (1889); Hooijer (1950, 1952); Johnston (1887); Jones (1931); Keast (1972); Krefft (1867); Mahoney &
Ride (1975); Marshall (1981); Owen (1877a); Pledge (1984, 1986); Simpson (1945); Stirton (1957b); Stirton,
Tedford & Woodburne (1968); Tedford et al. (1975); Turnbull & Lundelius (1970); Whitelaw (1989);
Woodbume (1986); Woodbume & Clemens (1986b).

Phalangista --- see Phalanger.

Phascogale: Archer (1978d, 1981a, 1982a); Archer & Baynes (1972); Balme, Merrilees & Porter (1978); Baynes (1982,
1987); Bensley (1901); Broom (1896a); Drummond (1963).; Gorter (1977); Hope (1973a, 1982); Kendrick &
Porter (1973); Longman (1924d); Lundelius (1957); Lundilius & Turnbull (1973, 1975, 1978, 1989);
Marshall (1981); Merrilees (1979a, 1984); Morton & Baynes (1985); Owen (1871a); Pledge (1974, 1990);
Porter (1979); Ride (1960); Simpson (1945); Smith & Medline (1982); Spencer & Kershaw (1910);
Tedford (1966b); Wakefield (1960a-b, 1964b, 1967a-b, 1972b).

Phascolagus -— see Macropus.

Phascolarctos: Abbie (1937); Anderson (1932); Aplin (19981); Archer (1972, 1977b, 1981a); Archer &
Bartholomai (1978); Balme, Merrilees & Porter (1978); Bartholomai (1968, 1977); Case (1985); Chapman (1934);
de Vis (1889b); Frechkop 1965); Glauert (1910b, 1926); Hill, Playford & Woods (1970); Hope (1973a, 1982);
Keast (1972); Longman (1921); Lundelius & Turnbull (1982, 1989); Mahoney & Ride (1975); Marshall (1973,
1981); Merrilees (1968, 1969, 1979a, 1984); Milham & Thompson (1976); Murray (1984b); Owen (1871a-b,
1882a, 1883b); Pledge (1974, 1981, 1986, 1987a, 1990); Rich (1986); Rich & Archer (1979); Rich, Archer &
Tedford (1978); Scott & Lord (1922a, 1925a); Simpson (1945); Tedford (1966b); Tedford & Wells (1990);
Wakefield (1967b); Wells, Moriarty & Williams (1984); Woodward (1910, 1914).

Phascolomis -— sce Vombatus.

Phascolomys (see also Lasiorhinus & Vombatus): Abbie (1937); Anderson, C. (1924a); Anderson, W. (1914);
Anon. (191b); Bensley (1903); Dawson (1981, 1983a-b); de Vis (1883b, 1886, 1891b, 1897, 1899b, 1907);
Frechkop (1930); Gill (1972); Glauert (1910b, 1912b, 1914, 192l1a, 1926); Higgins & Petterd (1889);
Howchin (1930); Johnston (1887); Krefft (1865, 1870); Longman (1917, 1921, 1926b-d, 1926); Lydekker (1887,
1890); McCoy (1862, 1874b, 1879, 1882); Mahoney & Ride (1975); Merrilees (1967a, 1969); Murie (1866);
Murray (1984b); Owen (1839, 1840, 1871b, 1872a-d, 1874a, 1877a, 1882a-b, 1886, 1887a); Ramsay (1886a-b);

AUSTRALIA'S MAMMALIAN RECORD - 1065

Scott (1917); Scott & Lord (1921b, 1922a); Simpson (1945); Spencer & Kershaw (1910); Stirling (1893, 1900b);
Stirling & Zietz (1899a-b); Wakefield (1964b); Walcott (1920); Wilkinson (1892); Woodward (1910, 1914).
Phascolonus (= Sceparmodon): Aplin & Archer (1987); Archer (198la); Archer & Bartholomai (1978), Archer &
Wade (1976); Bartholomai (1977); Dawson (1981, 1983a); de Vis (1891la-c, 1893a); Dun (1892, 1894);
Flannery (1984); Flannery & Archer (1984); Flannery & Hope (1983); Glauvert (1912b, 1921b, 1926); Gorter &
Nicoll (1978); Hill, Playford & Woods (1970)., Hope (1973b, 1978); Hope & Wilkinson (1984); Horton (1978,
1984); Horton & Connah (1981); Keast (1972); Leach (1977); Longman (1924b-c, 1926); Lydekker (1887, 1890,
1896a); McNamara (1990); Marcus (1976); Mahoney & Ride (1975); Marshall (1973, 1981); Merrilees (1968);
Murray (1984a-b); Murray & Goede (1977); Owen (1877a, 1884, 1889); Partridge & Thome (1963); Quirk &
Archer (1983); Ramsay (1881, 1886b); Ride (1967, 1971); Scott (1915b, 1917); Scott & Lord (1921c, 1925b);
Simpson (1945); Stephenson (1964); Stirling (1900b); Stirling & Zietz (1899b); Tedford, Williams &

Wells (1986); Wells (1978); Whitelaw (1989); Woodward, B.H. (1909).

Phascolosorex: Archer (1981a, 1982a); Marshall (1981).

Pildra: Pledge (1987d); Tumbull, Rich & Lundelius (1987b); Woodbume, Tedford & Archer (1987)

Pilkiptldra: Archer, Tedford & Rich (1987)

Pipistrellus: Simpson (1945); Wakefield (1967a, 1972b).

Pitikantia: Archer (198la); Archer & Bartholomai (1978); Bartholomai (1972a); Callen & Plane (1986); Mahoney &
Ride (1975); Marshall (1981); Rich & Rich (1987); Stirton (1967a); Stirton, Tedford & Woodbume (1967); Stirton,
Woodbume & Plane (1967); Woodburne (1967b, 1986); Woodburne, Campbell et al. (1986).

Plaisiodom: Archer (1981a); Archer & Bartholomai (1978); Flannery & Plane (1986); Hand & Archer (1987); Mahoney &
Ride (1975); Marshall (1981); Murray (1984a); Rich, Archer & Tedford (1978); Stirton, Tedford &
Woodbume (1968); Stirton, Woodbume & Plane (1967); Woodburne (1967a-b, 1969).

Planigale: Archer (1976a, 1981a, 1982a-b)., Archer & Bartholomai (1978); Archer & Rich (1979); Lundelius &
Turnbull (1973, 1989); Marshall (1981); Morton & Baynes (1985); Simpson (1945); Smith & Medlin (1982).

Plectodon --- see Thylacoleo.

Pogonomys: Godthelp (1990).

Potorous (=Hypsiprymnus): Abbie (1937); Aplin (1981); Archer (1972, 1979, 1981a); Archer & Baynes (1972);
Balme (1980); Balme, Merrilees & Porter (1978); Bartholomai (1972a, 1977); Bowdler (1979); Broom (1895,
1896a-b); Butler & Merrilees (1971); Case (1985); Clarke (1878); Cook (1963a); Drummond (1963);
Finlayson (1938); Flannery, Archer & Plane (1983, 1984); Flannery & Szalay (1982); Gill (1968); Glauert (1926);
Godwin (1980); Gorter (1977); Hill, Playford & Woods (1970); Hope (1973a-b, 1982); Hope et al. (1977);
Johnston (1887); Kendrick & Porter (1973); Krefft (1865); Lundelius (1957); Lundelius & Tumbull (1984, 1989);
Mahoney (1964); Mahoney & Ride (1975); Marshall (1981); Merrilees (1968, 1979a-b, 1984); Murray &
Goede (1977); Pledge (1974, 1990); Porter (1979); Ride (1960, 1971); Ryder (1974); Simpson (1945);
Smith (1971, 1972); Spencer & Kershaw (1910); Wakefield (1960a-b, 1963, 1964a-b, 1967a-b, 1972b, 1974);
Wells, Moriarty & Williams (1984); Whitelaw (1989); Woodbume (1984).

Prionotemnus: Archer (1981a); Archer & Bartholomai (1978); Bartholomai (1975); Dawson & Flannery (1980, 1985);
Flannery & Archer (1982); Flannery, Hoch & Aplin (1989); Flannery, Mountain & Aplin (1982); Flannery &
Szalay (1982); Hill, Playford & Woods (1970); Mahoney & Ride (1975); Marshall (1981); Sanson (1978);
Stirton (1955); Stirton, Tedford & Woodbume (1968); Tedford, Williams & Wells (1986); Woodbume et al. (1986).

Priscaleo: Aplin & Archer (1987); Murray, Wells & Plane (1987); Rauscher (1987)

Prochaerus --- see Thylacoleo.

Procoptodon (=Pachysiagon): Anderson (1932, 1933); Archer (1978d, 1981a); Archer & Bartholomai (1978); Archer &
Wade (1976); Bartholomai (1963,1970, 1972a, 1977); Bennett (1878); Case (1985); Clarke (1878); Dawson &
Flannery (1985); de Vis (1883c, 1899b); Dun (1893, 1900); Flannery (1983); Flannery & Archer (1983); Flannery
& Hope (1983); Flannery & Szalay (1982); Gill (1953a-b, 1965); Glauert (1910a, 1912b); Gorecki et al. (1984);
Hand & Archer (1987); Hand, Dawson & Augee (1988); Hill, Playford & Woods (1970); Hope (1978); Hope,
Dare-Edwards & MclIntyre (1983); Horton (1978, 1980, 1984); Horton & Connah (1981); Howchin (1930);
Keast (1972); Leach (1977); Longman (1924c),; Lydekker (1887, 1890. 1891, 1895. 1896a); McCoy (1879);
McIntyre & Hope (1978); Mahoney & Ride (1975); Marshall (1973, 1976, 1981); Merrilees (1968); Merrilees &
Ride (1965); Murray (1984a-b); Owen (1873, 1874b, 1876, 1877a-b, 1880); Quirk & Archer (1983); Pledge (1973,
1990); Ride (1959a, 1971); Sanson (1978); Scott (1906, 1916); Scott & Lord (1921c); Simpson (1945); Stirton &
Marcus (1966); Tate (1948); Tedford (1966b); Tedford & Wells (1990); Tedford, Williams & Wells (1986);
Wells (1975, 1978); Wells, Moriarty & Williams (1984); White & O'Connell (1979); Wright (1986).

Propalorchestes: Murray (1986).

Propleopus (=Triclis): Anderson (1933); Archer (1979, 1981a); Archer & Bartholomai (1978); Archer, Bartholomai &
Marshall (1978); Archer & Flannery (1985); Bartholomai (1972a-b); Case (1984, 1985); de Vis (1888c);
Flannery (1983); Flannery & Archer (1984); Flannery, Archer & Plane (1984); Flannery & Szalay (1982);
Gillespie et al. (1978); Hand & Archer (1987); Hope (1978); Horton (1984); Horton & Connah (1981); Hill,
Playford & Woods (1970); Keast (1972); Longman (1924b-c); Lydekker (1896a); Mahoney & Ride (1975);
Marshall (1981); Merrilees (1968); Murray (1984a-b); Murray, Wells & Plane (1987); Pledge (1981, 1990);
Raven (1929); Rich & van Tets (1985); Ride (1971); Simpson (1945); Stirton, Tedford & Woodburne (1968);
Tedford (1966b); Tedford, Williams & Wells (1986); Woodbure (1984); Woods (1960b).

Protemnodon: Archer (1978d, 1981a); Archer & Bartholomai (1963, 1978); Archer & Wade (1976); Balme, Merrilees &
Porter (1978); Bartholomai (1972a, 1973, 1977, 1978a); Case (1985); Cook (19634); Dawson & Flannery (1985);
de Vis (1883, 1888b); Dun (1893); Finlayson (1948); Flannery (1980, 81983, 1984); Flannery & Archer (1983,
1984); Flannery, Archer & Plane (1984); Flannery & Gott (1984); Flannery & Hann (1984); Flannery, Hoch & Aplin
(1989); Flannery & Hope (1983), Flannery, Mountain & Aplin (1982); Flannery & Szalay (1982);
Gillespie et al. (1978); Gillieson & Mountain (1983); Glauert (1910a); Goede & Murray (1979); Gorecki et al.
(1984); Gorter & Nicoll (1978); Hand, Dawson & Augee (1988); Hill, Playford & Woods (1970); Hoch &

1066 - RICH, ET AL.

Holm (1986); Hope (1973b, 1978, 1982); Hope & Wilkinson (1984); Horton (1978, 1980), Horton (1984); Horton
& Connah (1981); Horton & Wright (1981); Howchin (1930); Krefft (1875b); Leach (1977); Lester et al. (1988);
Lundelius & Tumbull (1989); Lydekker (1896a); Mahoney & Ride (1975); MacFadden et al. (1987); Marcus (1976);
Marshall (1973, 1974, 1981); Merrilees (1968, 19792, 1984); Milham & Thompson (1976); Mountain (1981);
Murray (19782, 1984a-b); Murray & Goede (1977); Owen (1874a-b, 1876, 1877a-b, 1887a); Plane (1965, 1967b,
1971); Pledge (1974, 1990); Raven & Gregory (1946); Ride (1962a, 1971); Ryder (1974), Sanson (1978); Scott &
Lord (1921c, 1924a); Simpson (1945); Stirton, Tedford & Woodbume (1968); Tedford (1966b); Tedford & Wells
(1990); Tedford, Williams & Wells (1986); Turnbull, Lundelius & Tedford (1990); Vanderwal & Fullagar (1989);
Wakefield (1964b, 1967b); Wells, Moriarty & Williams (1984); White & O'Connell (1979); Whitelaw (1989);
Wilkinson, C.S. (1892); Wilkinson, H.E. (1972); Wright (1986).

Pseudantechinus: Archer (1982a-b); Marshall (1981); Van Dyck (1982).

Pseudocheirus (=Petropseudes, Pseudocheirops): Aplin (1981); Archer (1977b, 1978d, 1981a); Archer &
Bartholomai (1978); Archer & Baynes (1972); Archer, Hand & Godthelp (1988); Baynes (1987); Bowdler (1979);
Broom (1896a); Butler (1969); Callen & Plane (1986); Case (1985); de Vis (1889b); Drummond (1963); Flannery
& Hann (1984); Flannery & Rich (1986); Gill (1968); Gillieson & Mountain (1983); Godwin (1980);
Glauert (1926); Goede & Murray (1979); Hope (1973a-b, 1982); Keast (1972); Lundelius (1957); Lundelius &
Turnbull (1982, 1989); Lydekker (1887); MacFadden et al. (1987); Mahoney & Ride (1975); Marshall (1981);
Merrilees (1968, 1979a-b, 1984); Mulvaney (1961); Murray & Goede (1977); Pledge (1974, 1990); Porter (1979);
Rich, Archer & Tedford (1978); Ride (1960); Roe (1971); Ryder (1974); Simpson (1945); Smith (1971); Spencer &
Kershaw (1910); Stirton, Tedford & Woodbume (1968); Tumbull & Lundelius (1970); Tumbull, Rich &
Lundelius (1987b); Wakefield (1960a-b, 1963, 1964a-b, 1967a-b, 1972b, 1974); Wells, Moriarty &
Williams (1984); White (1967); Whitelaw (1989); Woodburne (1986); Woodburne ef al. (1986).

Pseudochirops (see also Pseudocheirus): Archer, Hand & Godthelp (1988)

Pseudokoala: Archer (1981a); Archer & Bartholomai (1978); Marshall (1981); Pledge (1987d); Tumbull &
Lundelius (1970); Tumbull, Rich & Lundelius (1987b).

Pseudomys (=Leggadina, Thetomys): Aplin (1981); Archer (1972, 1981a); Archer & Baynes (1972); Baynes (1982, 1984,
1987); Bowdler (1979); Drummond (1963); Gill (1968); Godthelp (1990); Godwin (1980); Hand & Archer (1987);
Hope (1973a-b, 1978, 1982); Hope & Wilkinson (1984); Hope et al. (1977); Kendrick & Porter (1973);
Lundelius (1957, 1964); McNamara (1990); Marshall (1973, 1974); Merrilees (1979a, 1984); Morton &
Baynes (1985); Murray & Goede (1977); Pledge (1990); Porter (1979); Roe (1971); Ride (1960); Simpson (1945);
Smith (1977); Tedford & Wells (1990); Thorne (1971); Wakefield (1960b, 1963, 1964a-b, 1967a-b, 1972a-b);
Wells, Moriarty & Williams (1984).Whitelaw (1989).

Pteropus: Aplin (1981); Archer & Brayshaw (1978); Simpson (1945); Wakefield (1974); White (1967).

Purtia: Callen & Plane (1986); Case (1984); Flannery & Archer (1987c); Flannery, Archer & Plane (1984);
Woodbume (1986).

Pyramios: Aplin (1987); Archer (1981a); Archer & Bartholomai (1978); Bartholomai (1972a); Mahoney & Ride (1975);
Marshall (1981); Murray (1984a); Stirton, Tedford & Woodbume (1968); Stirton, Woodburne,.& Plane (1967);
Woodbume (1967a-b, 1969).

Raemeotherium: Callen & Plane (1986); Flannery & Plane (1986); Marshall (1981); Rich, Archer & Tedford (1978); Rich
& Rich (1987); Woodbume (1986).

Ramsayia: Dawson (1981, 1983a); Flannery & Hope (1983); Hope & Wilkinson (1984); Horton (1984); Murray (1984b).

Rattus: Allen (1972); Aplin (1981); Archer (1981a); Archer & Bartholomai (1978); Archer & Baynes (1972);
Baynes (1982, 1984, 1987); Bowdler (1979); Cook (1963a); Downie & White (1978); Drummond (1963);
Gill (1968); Godthelp (1990); Godwin (1980); Hope (1973a-b, 1978, 1982); Hope & Wilkinson (1984);
Hope ef al. (1977); Kendrick & Porter (1973); Lundelius (1964); McNamara (1990); Marshall (1973);
Merrilees (1979a, 1984); Morton & Baynes (1985); Mulvaney (1961); Murray & Goede (1977), Pledge (1974,
1990); Porter (1979); Simpson (1945); Smith (1977); Tedford (1966b); Tedford, Williams & Wells (1986);
Wakefield (1960b, 1963, 1964a-b, 1967a-b, 1972a-b, 1974); Wells, Moriarty & Williams (1984); White (1967).

Rhinolophus: Archer (1981a); Drummond (1963); Simpson (1945); Wakefield (1967a, 1972b).

Rhinoycteris: Sigé, Hand & Archer (1982).

Rhizophascolonus: Archer (1981a); Archer & Bartholomai (1978); Callen & Plane (1986); Hope & Wilkinson (1984);
Mahoney & Ride (1975); Marshall (1981); Stirton, Tedford & Woodburne (1967, 1968); Woodburne,
Campbell et al. (1986).

Rhinomeles: Archer (1981a, 1982a).

Saccolaimus Hand (1987).

Sarcophilus: Abbie (1937); Anderson, C. (1924a); Anon, (1871); Archer (1978d, 1981a, 1982a-b); Archer &
Bartholomai (1978); Archer & Baynes (1972); Archer & Wade (1976); Bartholomai (1977); Bartholomai &
Marshall (1973); Baynes (1987); Bowdler (1974, 1979); Butler (1969); Calaby & White (1967); Cook (1963a-b);
Crabb (1982); Dawson (1982a); de Vis (1883d, 1893b); Douglas, Kendrick & Merrilees (1966); Flannery &
Gott (1984); Flannery & Hope (1983); Flood (1973); Gill (1953a, 1965, 1968); Gillespie et al. (1978);
Glauert (1912b, 1914, 1926); Goede & Murray (1979); Hill, Playford & Woods (1970); Hope (1973a-b, 1978,
1982); Hope, Dare-Edwards & McIntyre (1983); Hope & Wilkinson (1984); Hope et al. (1977); Horton (1980,
1984); Horton & Connah (1981); Horton & Wright (1981); Howchin (1930); Keast (1972), Krefft (1865);
Longman (1924c-d),; Lundelius (1966); Lundelius & Tumbull (1978, 1989); Lydekker (1887, 1896a).,
McCoy (1882); MacIntosh (1971); McIntyre & Hope (1978). Mahoney & Ride (1975); Marshall (1973, 1974,
1981); Marshall & Corruccini (1977); Merrilees (1967b, 1968, 1969, 1979a-b, 1984); Milham &
Thompson (1976); Murray (1984b); Murray & Goede (1977); Owen (1859c, 1871a, 1877a); Pledge (1974, 1990);
Porter (1979); Ride (1964); Simpson (1945); Smith (1972); Sobbe (1990); Tedford (1966b); Tedford, Williams &
Wells (1986); Thorne (1971); Tedford & Wells (1990); Wakefield (1964a-b, 1967a-b, 1972b); Walcott (1920);
Wells, Moriarty & Williams (1984); White (1967), Wilkinson (1978).

AUSTRALIA'S MAMMALIAN RECORD - 1067

Satanellus: Archer (1981a, 1982a); Simpson (1945); White (1967).

Schinobates (=Petauroides): Aplin (1981); Archer (198la); Archer & Bartholomai (1978); Bensley (1901);
Drummond (1963); Hand, Dawson & Augee (1988); Hope (1982); Jones (1931); Marshall (1981); Ryder (1974);
Simpson (1945); Tumbull, Rich & Lundelius (1987b); Wakefield (1960a-b, 1967a, 1972a-b, 1974).

Schizodon -— see Thylacoleo.

Scoteinus: Simpson (1945); Wakefield (1963, 1967a, 1972a-b, 1974).

Setonix: Archer (1972, 1979, 1981a); Archer & Baynes (1972); Balme, Merrilees & Porter (1978); Bartholomai (1972a);
Butler (1969); Cook (1963a); Dawson & Flannery (1985); Flannery (1983); Flannery & Hann (1984); Flannery &
Rich (1986); Flannery & Szalay (1982); Glauert (1926); Marshall (1981); Merrilees (1968, 1979a-b, 1984);
Porter (1979); Ride (1971); Simpson (1945); Tedford (1966b).

Simoprosopus --- see Zygomaturus.

Simosthenurus: Flannery (1983); Flannery & Archer (1983, 1984); Flannery & Gott (1984); Flannery & Hope (1983);
Flannery & Pledge (1987); Lundelius & Tumbull (1989); Mahoney & Ride (1975); Marcus (1976); Marshall (1981);
Murray (1984a); Pledge (1980b, 1981, 1990); Rich & van Tets (1985); Tedford (1966a); Tedford & Wells (1990);
Wells (1978),

Sminthopsis: Abbie (1937); Archer (1978d, 1981a, 1982a-b); Archer & Baynes (1972); Archer & Rich (1979); Balme,
Merrilees & Porter (1978)., Baynes (1982, 1984, 1987); Bensley (1901); Burbidge & George (1978);
Drummond (1963); Gill (1968); Gorter (1977); Hand & Archer (1987); Hope (1973a, 1982); Hope &
Wilkinson (1984); Hope et al. (1977); Kendrick & Porter (1973); Lundelius (1957); Lundelius & Turnbull (1975);
Marshall (1981); Merrilees (1979a, 1984); Morton & Baynes (1985); Murray & Goede (1977); Pledge (1974,
1990); Porter (1979); Roe (1971); Simpson (1945); Smith (1972, 1977); Smith & Medline (1982);
Tedford (1966b); Thome (1971); Wakefield (1963, 1964a-b, 1967a-b, 1972b); Wells, Moriarty & Williams (1984).

Steropodon: Archer, Every, Godthelp, Hand & Scally (1990); Archer, Flannery, Ritchie & Molnar (1985); Hand &
Archer (1987); Kielan-Jaworowska, Crompton & Jenkins (1987), Rich, Flannery & Archer (1989).

Sthenomerus: de Vis (1883a, 1888b, 1891a); Mahoney & Ride (1975); Marshall (1981); Scott & Lord (1921a,c);
Simpson (1945).

Sthenurus: Anderson (1932, 1933); Archer (1972, 1978d, 1981a); Archer & Bartholomai (1978); Balme, Merrilees &
Porter (1978); Bartholomai (1963, 1966, 1972a, 1977, 1978a); Case (1985); Clarke (1878); Cook (1963a);
Cudmore (1926); Dawson & Flannery (1985); de Vis (1883c,¢, 1899a-b); Dulhunty, Flannery & Mahoney (1984);
Dun (1893); Flannery (1983); Flannery & Archer (1983); Flannery & Gott (1984); Flannery & Hope (1983);
Flannery & Szalay (1982); Flood (1973); Gill (1953c); Gillespie et al. (1978); Glauert (1909, 1910a-b, 1912b,
1926); Goede & Murray (1979); Gorecki ef al. (1984); Hand, Dawson & Augee (1988); Hill, Playford &
Woods (1970); Hope (1973a-b, 1978, 1982); Hope, Dare-Edwards & McIntyre (1983); Hope & Wilkinson (1984);
Hope et al. (1977); Horton (1980, 1984); Horton & Connah (1981); Horton & Wright (1981); Howchin (1930);
Keast (1972); Longman (1924c, 1926); Lundelius (1963); Lundelius & Tumbull (1989); Lydekker (1887, 1895,
1896a); Mahoney & Ride (1975); McCoy (1879); Marcus (1976); Marshall (1973, 1981); Merrilees (1965, 1967c,
1968, 1969, 1979a-b, 1984); Milham & Thompson (1976); Murray (1978a, 1984a-b); Murray & Goede (1977);
Owen (1874a-b, 1876, 1877a-b, 1887a); Pledge (1974, 1980b, 1981, 1990); Raven (1929); Raven &
Gregory (1946); Ride (1971); Sanson (1978); Scott (1917); Scott & Lord (1925b); Simpson (1945);
Stirton (1957b); Stirton, Tedford & Woodbume (1968); Tate (1948); Tedford (1966a-b); Tedford & Wells (1990);
Tedford, Williams & Wells (1986); Vanderwal & Fullagar (1989); Wakefield (1967b, 1972); Wells (1975, 1978);
Wells, Horton & Rogers (1982); Wells, Moriarty & Williams (1984); Wells & Murray (1979); White &
O'Connell (1979); Whitelaw (1989); Woodward, B. (1909, 1910, 1914); Wright (1986).

Strigocuscus: Archer, Hand & Godthelp (1988); Flannery & Archer (1987a); Flannery, Archer & Maynes (1987); Flannery,
Tumbull, Rich & Lundelius (1987).

Sus: Allen (1972); Pledge (1980b); Simpson (1945).

Synaptodon: de Vis (1895a); Dawson & Flannery (1985); Mahoney & Ride (1975); Marshall (1975); Simpson (1945).

Tachyglossus (see also Echidna): Archer (1981a); Baynes (1987); Gill (1968); Glauert (1910b, 1914); Hope (1973b,
1982); Merrilees (1979a, 1984); Murray (1978b, 1984a); Murray & Goede (1977); Pledge (1974); Ride (1960);
Ryder (1974); Spencer & Kershaw (1910); Tedford (1966b); Wakefield (1974); Woodward, B. (1914).

Tadarida: Simpson (1945); Wakefield (1967a, 1972b).

Tarsipes: Archer (1972, 1981a); Balme, Merrilees & Porter (1978); Marshall (1981); Merrilees (1979a); Porter (1979);
Simpson (1945).

Thalacomys --- sec Macrotis.

Thetomys --- see Pseudomys.

Thylacinus: Abbie (1937); Anderson (1924a, 1925, 1929b); Anon. (1871); Archer (1971, 1974, 1978d, 1981a, 1982a-b);
Archer & Bartholomai (1978); Archer & Dawson (1982); Balme, Merrilees & Porter (1978); Bartholomai (1977);
Baynes (1987); Bennett (1876); Bensley (1901, 1903); Bowdler (1974, 1979); Broom (1896a); Clemens (1977);
Cook (1963a-b); de Vis (1893b, 1899b, 1900); Etheridge (1892); Flannery & Hann (1984); Flannery &
Hope (1983); Flannery, Mountain & Aplin (1982); Flood (1973); Gill (1965, 1968); Gillespie et al. (1978);
Gillieson & Mountain (1983); Glauert (1914, 1921a, 1926); Gorter (1977); Graves (1958); Gregory (1929); Hand,
Dawson & Augee (1988); Hill, Playford & Woods (1970), Hoch & Holm (1986); Hope (1973b, 1978, 1982);
Horton (1980); Horton & Wright (1981); Howchin (1930); Howlett (1960); Keast (1972); Krefft (1865, 1867);
Kendrick & Porter (1973); Longman (1921, 1924c-d); Lundelius (1966); Lundelius & Tumbull (1978, 1989);
Lydekker (1887, 1896a); MacFadden et al. (1987); Mahoney & Ride (1975); Marcus (1976); Marshall (1973, 1974,
1977b, 1981); Merrilees (1967b, 1968, 1979a-b, 1984); Milham & Thompson (1976); Muirhead & Archer (1990);
Murray (1984a); Murray & Goede (1977); Owen (1871a, 1877a, 1883c); Partridge & Thome (1963); Plane (1976);
Pledge (1973, 1980b, 1981, 1990); Quirk & Archer (1983); Ride (1959b, 1960, 1964); Simpson (1945);
Smith (1972); Stirton, Tedford & Woodburne (1968); Tate (1947); Tedford (1966b); Wan Deusen (1963);
Wakefield (1964a-b, 1967b, 1972b); Wells, Moriarty & Williams (1984); Woodbume (1967b).

1068 - RICH, ET AL.

Thylacis --- see Isoodon.

Thylacoleo (=Mylodon, Plectodon, Prochaerus, Schizodon, Thylacopardus): Abbie (1941); Aplin (1987); Anderson,

C. (1924a-b, 1929a-b, 1933); Anderson, W. (1888); Anon. (1887, 1916); Aplin & Archer (1987); Archer (198 1a,
1982b); Archer & Bartholomai (1978); Archer & Dawson (1982); Archer & Flannery (1985); Archer & Rich (1982);
Archer & Wade (1976); Bartholomai (1962, 1963, 1977); Bensley (1903); Broom (1898); Case (1985);
Clarke (1878); Clemens & Plane (1974); Daily (1960); de Vis (1883d, 1886, 1888b, 1899a-b, 1900);
Drummond (1963); Etheridge (1918); Finch (1971, 1982); Finch & Freedom (1982); Flannery & Archer (1984);
Flannery & Gott (1984); Flannery & Hope (1983); Flower (1868); Gill (1953a-b, 1954c, 1957c, 1963, 1965,
1973a, 1978); Glauert (1912b, 1926); Gregory (1929); Hand, Dawson & Augee (1988); Hope (1973b, 1978, 1982);
Hope & Wilkinson (1984); Horton (1979, 1980, 1984); Horton & Connah (1981); Horton & Wright (1981);
Howchin (1930); Keast (1972); Krefft (1866, 1870, 1872, 1874a, 1875b); Lane & Richards (1963); Leach (1977);
Lindsay (1966); Longman (1916, 1924b-d, 1926); Lundelius (1966); Lundelius & Turnbull (1978, 1989);
Lydekker (1887, 1895, 1896a); McCoy (1865b, 1876a, 1879); Mahoney & Ride (1975); Marcus (1976);
Marshall (1973, 1974, 1981); Megirian (1986); Merrilees (1967b, 1968, 1969, 1979a, 1984); Murray (1978a,
1984a-b); Murray & Goede (1977); Murray, Wells & Plane (1987); Owen (1859c, 1866, 1870, 1871a, 1872a,
1874a, 1877a-b, 1883a-d, 1884, 1887a-b, 1889); Quirk & Archer (1983); Partridge & Thorne (1963); Pledge (1973,
1977, 1981, 1990); Rauscher (1987); Rich & van Tets (1985); Scott & Lord (1921b-c, 1922b, 1924b);
Simpson (1945); Smith (1972); Stirling (1900b); Stirton, Tedford & Woodbume (1968); Tedford (1966b); Tedford
& Wells (1990); Tedford, Williams & Wells (1986); Vanderwal & Fullagar (1989); Wakefield (1967b, 1972a);
Wells, (1975); Wells, Horton & Rogers (1982); Wells, Moriarty & Williams (1984); Wells & Nichol (1977); White
& O'Connell (1979); Wilkinson (1972); Woods (1956b); Wright (1986).

Thylacomys --- see Macrotis.

Thylacopardus --- see Thylacoleo.

Thylogale: Aplin (1981); Archer (1981a); Archer & Wade (1976); Bartholomai (1972a, 1977); Bowdler (1979); Dawson
& Flannery (1985); Downie & White (1978); Flannery (1983); Flannery & Hann (1984); Flannery, Hoch & Aplin
(1989); Flannery, Mountain & Aplin (1982); Flannery & Szalay (1982); Gill (1968); Gillespie et al. (1978);
Gillieson & Mountain (1983); Godwin (1980); Goede & Murray (1979); Gorter (1977); Hope (1973b); Horton &
Murray (1980); Marshall (1981); Mulvaney (1961); Murray & Goede (1977); Murray, Goede & Bada (1980);
Ride (1971); Ryder (1974); Simpson (1945); Stirton, Tedford & Woodbume (1968); Turnbull & Lundelius (1970);
Wakefield (1964a-b, 1967b, 1974); Woodburne (1986).

Trichosurus: Abbie (1937); Aplin (1981); Archer (1981a); Archer & Bartholomai (1978); Archer & Baynes (1972); Archer
& Brayshaw (1978); Archer & Rich (1982); Baynes (1984, 1987); Bensley (1901); Bowdler (1979); Broom (1895,
1296a); Case (1985); Drummond (1963); Flannery & Archer (1987a); Flannery, Archer & Maynes (1987); Flannery,
Archer & Plane (1983); Flannery & Rich (1986); Flannery, Tumbull, Rich & Lundelius (1987); Gill (1953c, 1957b,
1968); Glauert (1926); Godwin (1980); Gorecki et al, (1984); Hope (1973a-b, 1978); Hope & Wilkinson (1984);
Hope et al. (1977); Horton & Connah (1981); Jones (931); Keast (1972); Kendrick & Porter (1973);
Longman (1924b); Lundelius & Tumbull (1982, 1989); Marshall (1981); Merrilees (1968, 1979a-b, 1984);
Mulvaney (1961); Murray & Goede (1977); Pledge (1974, 1980b, 1986, 1990); Porter (1979); Rich &
Archer (1979); Roe (1971); Ryder (1974); Scott & Lord (1925a); Simpson (1945); Stirton, Tedford &
Woodbume (1968); Tedford (1966b); Tedford & Wells (1990); Tedford, Williams & Wells (1986); Turnbull &
Lundelius (1970); Wakefield (1960a, 1964a-b, 1967a-b, 1972a-b, 1974); White (1967); Woodburme (1986);
Woodburme & Clemens (1986b)

Triclis --- see Propleopus.

Troposodon: Archer (1981a); Archer & Bartholomai (1978); Bartholomai (1967, 1972a, 1978a); Campbell (1973);
Dawson & Flannery (1985); Flannery (1983); Flannery & Archer (1982, 1983, 1984); Flannery & Szalay (1982);
Hill, Playford & Woods (1970); Mahoney & Ride (1975); Marshall (1981); Murray (1984); Quirk & Archer (1983);
Sanson (1978); Tedford, Williams & Wells (1986); Tumbull, Lundelius & Tedford (1990); Tedford & Wells (1990);
Whitelaw (1989); Woodburne et al, (1986).

Uromys: Gillieson & Mountain (1983); Godthelp (1990).

Vombatus (=Phascolomis) (see also Phascolonus): Aplin (1981); Archer (1972, 1978d, 1981a); Archer &
Bartholomai (1978); Balme, Merrilees & Porter (1978); Bartholomai (1977); Bowdler (1979); Case (1985);
Dawson (1983a-b); Flannery & Hope (1983); Gill (1953a-c, 1972); Gillespie et al. (1978); Goede &
Murray (1979); Hope (1973a-b, 1982); Hope & Wilkinson (1984); Horton & Connah (1981); Keast (1972);
Marcus (1976); Marshall (1973, 1974, 1981); Merrilees (1967a, 1968, 1979a, 1984); Murray (1984b); Pledge
(1990); Scott & Lord (1925a); Simpson (1945); Tedford (1966b); Tedford, Williams & Wells (1986); Tumbull,
Lundelius & Tedford (1990); Wakefield (1967b); Wells, Moriarty & Williams (1984); Whitelaw (1989);
Wilkinson (1978).

Wabularoo: Archer (1979, 1981a); Flannery, Archer & Plane (1983, 1984); Flannery & Rich (1986); Marshall (1981);
Quirk & Archer (1983); Pledge (1984); Woodburne (1984).

Wakaleo: Aplin (1987); Aplin & Archer (1987); Archer (1981a); Archer & Bartholomai (1978); Archer & Dawson (1982);
Archer & Rich (1982); Callen & Plane (1986); Clemens & Plane (1974); Finch (1982); Finch & Freedman (1982);
Flannery, Archer & Plane (1983); Hand & Archer (1987); Marshall (1981); Megirian (1986); Murray (1984a);
Murray, Wells & Plane (1987); Rauscher (1987); Woodburne (1986); Woodbume, Campbell et al. (1986).

Wakamatha: Archer (1981a, 1982a); Archer & Rich (1979); Marshall (1981).

Wakiewakie: Archer, Hand & Godthelp (1988); Callen & Plane (1986); Case (1984); Flannery & Archer (1987c);
Flannery, Archer & Plane (1984); Godthelp, Archer, Hand & Plane (1989);Woodburne (1984, 1986)

Wallabia: Aplin (1981); Archer (1981a); Archer & Bartholomai (1978); Archer & Brayshaw (1978); Bartholomai (1972a,
1976); Case (1985); Flannery & Archer (1983); Flannery & Hann (1984); Flannery, Hoch & Aplin (1989); Flannery
& Szalay (1982); Gill (1968); Gorecki ef al. (1984); Gorter (1977); Hand, Dawson & Augee (1988); Hope (1973a,

AUSTRALIA'S MAMMALIAN RECORD - 1069

1982); Horton (1984); Marshall (1973, 1981); Merrilees (1968, 1979a); Murray (1984b); Pledge (1990);
Ride (1971); Simpson (1945); Stirton, Tedford & Woodburne (1968); Tedford (1966b); | Tedford, Williams &
Wells (1986); Wakefield (1967b, 1974); Wells, Moriarty & Williams (1984); White (1967); Whitelaw (1989);
Woodbume (1984).

Warendja: Aplin & Archer (1987); Dawson (1983b); Flannery & Pledge (1987); Hope & Wilkinson (1984)

Watutia: Flannery, Hoch & Aplin (1989).

Wynyardia: Abbie (1937, 1941); Anderson (1925, 1933); Aplin (1987); Aplin & Archer (1987); Archer (1981a); Archer
& Bartholomai (1978); Bensley (1903); Chapman (1941); Clemens (1977); Gill (1953c, 1954b, 1957b);
Gregory (1929); Haight & Murray (1981); Hofer (1952);. Keast (1972); Jones (1931), Longman (1921, 1924c,
1926); Mahoney & Ride (1975); Marshall (1981), Murray (1984a); Pledge (1987b); Rich & Archer (1979); Rich &
van Tets (1985); Ride (1964); Sera (1942); Simpson (1945); Spencer (1901); Stirton, Tedford &
Woodburne (1968); Tedford et al. (1977); Wood Jones (1931); Woods (1962).

Wyulda: Flannery, Archer & Plane (1983).

Xeromys: Morton & Baynes (1985)

Yalkaparidon: Archer, Hand & Godthelp (1988); Hand & Archer (1987); Lester, Archer, Gilkeson, & Rich (1988).

Yingabalanara: Archer, Every, Godthelp, Hand & Scally (1990).

Zaglossus (see also Echidna): Archer (1981a); Archer & Bartholomai (1978); Flannery & Hann (1984); Gillieson &
Mountain (1983); Glauert (1914, 1926); Hope (1973b); Horton (1984); Howchin (1930); Keast (1972); MacFadden
et al. (1987); Mahoney & Ride (1975); Merrilees (1979a, 1984); Murray (1976, 1978a-b, 1984a-b); Murray &
Goede (1977); Quirk & Archer (1983); Pledge (1980a, 1990); Scott & Lord (1922c, 1924a,c); Simpson (1945);
Tedford (1966b); Wells, Moriarty & Williams (1984).

Zygomaturus (=Simoprosopus) (see also Nototherium): Anderson, C. (1933); Anon. (1916); Archer (1978d, 1981a); Archer
& Bartholomai (1978); Archer & Wade (1976); Bartholomai (1972a); Bertrand (1986); Clemens & Plane (1974); de
Vis (1888a-b, 1889a, 1891a, 1895b, 1907); Dun (1894); Flannery & Gott (1984); Flannery & Hann (1984);
Flannery & Hope (1983); Flannery & Plane (1986); Glauert (1912a); Goede & Murray (1979); Guérin & Faure
(1987); Guérin, Winslow, Piboule & Faure (1981); Hardjasasmita (1985); Hill, Playford & Woods (1970);
Hope (1973b, 1982); Hope & Wilkinson (1984); Horton (1984); Huxley (1862); Keast (1972); Krefft (1872,
1874a); Lester et al. (1988); Longman (1916, 1921); MacFadden et al. (1987); Mahoney & Ride (1975);
Marcus (1976); Marshall (1973, 1974, 1981); Merrilees (1968, 1979a, 1984); Murray (1978a, 1984b); Murray &
Goede (1977); Owen (1859a-b 1882a); Pledge (1974, 1990): Quirk & Archer (1983); Rich, Fortelius, Rich &
Hooijer (1987); Ride (1971); Scott & Lord (1921la-c); Simpson (1945); Stirton (1967b); Stirton, Tedford &
Woodbume (1968); Tedford & Wells (1990); Tedford, Williams & Wells (1986); Stirton, Woodbume & Plane (1967);
Wakefield (1967b); Tumbull, Lundelius & Tedford (1990); Wells (1975, 1978); Wells, Moriarty & Williams (1984);
White & O'Connell (1979); Whitelaw (1989); Whitley (1966); Woodburne (1967b, 1969); Wright (1986); Wyrwoll
& Dortch (1978).

Zyzomys: Hand & Archer (1987); Kendrick & Porter (1973); Morton & Baynes (1985); Simpson (1945); White (1967).

PLATES

Plate 1. A, /laria illumidens, occlusal view of right 1]-M3. x 1.5; B, Djilgaringa gillespiei, a
pilkilpildrid, occlusal view of right P3-Ms5, x 10. Drawn by Airi Bennett.

Plate 2. Upper dentitions of mega- and microchiropterans. Nyctymene major (megachiropteran): A, upper
dentition; B, lower dentition; x 4. Rhinolophus ferrum-equinum (microchiropteran): C, upper dentition;
D, lower dentition; x 9.6. [After Miller 1907].

1070 - RICH, ET AL.

PLATE 1

PLATE 2

CHAPTER 24

THE PLEISTOCENE
MEGAFAUNA OF
AUSTRALIA

Peter Murray!
PAL OUICHOR A jo Picsvesisenmietiatseceistaidaonks 1072 Family Thylacoleonidae............ 1107
The Megafauna Concept.............:eeceeeee 1073 Family Potoroidae...............06+ 1111
What is Megafauna...............cceeee 1073 Family Macropodidae............... 1111
Late Pleistocene Australian Pleistocene Faunas.............cccccceeeeeeeees 1125
Mee af aiittissicccaceieescor seve nsvenaes 1073 Palaecobiology............:scccescseeceeceeeenees 1128
How Big Were They?............csccescees 1074 Defining a Megafauna Community ..1128
Estimation of Body Size................. 1074 The Significance of Body Weights ... 1129
Relative Megafauna...................6666 1075 Pattern of Speciation..................05 1131
Non-Mammalian Megafauna........... 1076 Dwarfing.............ccsseccsssceescceseeeees 1131
Evolution, Morphology and Systematics 1077 Predator Diversity ...............ceeeeeeees 1133
PVE ONY 0 iiectaye ese lankaaackinceb neds 1077 Behaviour and Intelligence.............. 1133
Morphology and Systematics.......... 1083 Locomotor Behaviout............... 1133
Species of Australian Pleistocene Brain SiZe...........ceceeceee eee eeee ees 1137
ME DAEAUIA, vncevecesspscicerstevsoevcns 1088 Intelligence ............ccceeeeee seco ees 1138
Order Monotremata.............cesseeeee 1088 Defensive Structures and
Family Tachyglossidae ............. 1088 Morphological
Order Dasyuromorphia .................. 1090 Differentiation .................5066 1140
Family Dasyuridae..................- 1090 Reprodtnsta ne «cs cgiexcpsa'voceeargonienes gen 1141
Order Diprotodontia................:.006 1094 Concluding Remarks...............:cceeeseeees 1141
Family Phascolarctidae............. 1994. SUMMMALy’s,.001..0.05-covecessesccdecceuvensop eee 1143
Family Vombatidae................. 1096. References oo ocisducitaiaes ee ieetieaetee betes 1144
Family Diprotodontidae ............ TOSS Appendix t ..0.cccscepessseeseressderesemeareegtes 1150
Family Palorchestidae............... LTOG: Plates BA's co.3. cseeeeee senna dees tates ease ensss 1159

a

1 Spencer & Gillen Museum, GP.O. Box 2109, Alice Springs, Northern Territory 0871, Australia.

1072 - MURRAY

INTRODUCTION

This chapter is primarily intended to serve as an introduction to the morphology and
systematics of the larger extinct mammals of the Late Pleistocene - the marsupials and
monotremes often referred to as the Australian "megafauna". It briefly summarizes the Tertiary
record and includes a few technical notes for those who wish to pursue the systematic aspects
of Quaternary mammalogy. It is aimed primarily at Quaternary specialists whose principle
interest is other than marsupial palaeontology and vertebrate palaeontologists who require a
summary of the somewhat disparate literature. Because the element of controversy is often
ignored or sometimes is not known to cross-disciplinary researchers, I have attempted to
delineate a few of these areas,

Large vertebrates other than mammals are important elements of the megafauna and
although they are also included in the chapters on birds and reptiles in this book, a
consideration of the community palaeontology of the Late Pleistocene of Australia cannot be
made without reference to them. While excellent textbooks on Quaternary mammals are
available for North America (Kurten & Anderson 1980) and Europe (Kurten 1968) no
comparable contemporary work exists for Australia. Consequently, Richard Owen's rare (1877)
two volume work has long filled this gap, and tired though it is, it is unlikely that any modern
mode of publication will ever surpass the large, detailed and wonderfully rendered lithographic
illustrations it contains, nor has Owen's text been entirely drained of its insights.

A consideration of the Australian megafauna would be incomplete without a discussion of
the cause or causes of their extinction. Because space is limited in a book chapter, I have
discussed the problem in terms of the palaeobiological information presented here rather than
attempt a comprehensive review. This should be read as a particular argument. Numerous
reviews of the subject have been published, for example, Horton (1979, 1980), Hope (1978)
and Merrillees (1968a, b). A knowledge of the current systematics of the taxa is considered
essential as is some basic anatomical information, necessary to interpret the text and
illustrations. For further reading on the history of the subject, marsupial anatomy and
systematics, I recommend Archer & Clayton (1984) which complements this and other
chapters of Vertebrate Palaeontology of Australia (this volume).

In its broadest and most convenient sense, the idea of a Pleistocene "megafauna" denotes a
conspicuous element of a Quaternary mammalian community dominated by large animals, of
which many genera and species became extinct sometime prior to the Holocene. In part, the
notion of a megafauna is a product of contrast with the Holocene faunas, which appear to be
dominated by smaller genera and species. In Australia, at least, there is also an apparent
gradual trend in increased body size from the late Palaeogene, culminating in the Late
Pleistocene. However, the impression that Pleistocene mammals were exceptionally gigantic
is not an entirely accurate or an especially useful frame of reference for understanding this
marked transition from Quatemary to modern faunas. The magnitude of body size differences
are more relative than absolute, although the factor of absolute body size is obviously
important and the categorization of a megafauna, albeit unruly, is valid enough. Thus, while
this chapter emphasizes the morphology of the larger vertebrates of the Australian Pleistocene,
the underlying theme is an attempt to understand the significance of large body size and certain
morphological specializations in the Quaternary extinction of Australian marsupials.

Numerous papers have been written on the subject of Pleistocene extinctions in Australia
nominating various causal agencies - hunting and ecological disruption by aboriginal man,
climatic change, epidemic diseases and even psychological stress (Gill 1955a, b, Jones 1968,
Merrilees 1968a,b, Balme et al. 1978, Gillespie et al. 1978, Hope 1978, Main 1978, Archer et
al. 1980, Martin 1984). However splendidly detailed these arguments are, their explanatory
powers are no better than the implicit understanding of the biology of the animals in question.

PLEISTOCENE MEGAFAUNA - 1073

Ironically, the most magnificent species of the Australian megafauna, Diprotodon spp. and

Zygomaturus spp. are among the least understood and not entirely due to a lack of effort, for

as Stirton (1955) observed, Diprotodon is the most-often mentioned genus in the literature.
Data tables discussed in this article are to be found in Appendix I, at the end of the text.

THE MEGAFAUNA CONCEPT

WHAT IS THE MEGAFAUNA?

The term megafauna is nothing more than a convenient reference applied primarily to
the large, extinct vertebrates of the Pleistocene. It has also been applied to invertebrate fauna
in a more specific usage by marine ecologists. I prefer to use the term operationally,
redefining the term to suit the circumstances of its usage. The flexible megafauna concept is
used in three different ways in this chapter. In the “absolute” sense, megafauna refers to
Australian mammals weighing 40 kg or more in contrast to the "relative" usage in which an
extinct Pleistocene species attained a significantly larger size than a closely allied form, even
though the relatively large animal may have weighed less than the designated absolute
criterion, A third usage is based on the concept of scaling (Schmidt-Neilsen 1984) in which
the body sizes of a community of animals are considered in an ecophysiological context. This
approach attempts to compare the body sizes attained by continental and island faunas by
examining the many factors that control animal body size.

Even though the notion of megafauna differs greatly among scientists, it is generally agreed
that North America, South America, Eurasia, Africa, Australia and some islands each supported
a megafauna community and that a major extinction phenomenon resulted in the extirpation of
the larger taxa at various times in the late Quaternary or Recent, leaving faunas with
significantly fewer large-bodied vertebrates (Martin & Wright 1967, Martin & Klein 1984).

Australia was one of the most severely affected of the continental faunas in that no
mammal greater than about 60 kg in weight survived among the living fauna encountered by
the Europeans. More than 40 species of megafauna became extinct in Australia, although the
majority of these are not found in every site (Martin & Murray 1983, Murray 1984). The
fossil taxa differ from one site to another due to the agencies of preservation, nature of the
community sampled at the site, geographic differences and time period. In the southeastern and
southwestern portions of Australia there is a fairly typical megafaunal assemblage that provides
4 convenient introduction to the constituents of the Late Pleistocene megafaunal community.

LATE PLEISTOCENE AUSTRALIAN MEGAFAUNA

A typical Pleistocene mammalian community from the temperate portions of Australia
about fifty-thousand years ago consisted of the genera Zaglossus (Tachyglossidae), long-beaked
echidnas: the Marsupial Lion Thylacoleo (Thylacoleonidae); giant wombats Phascolonus and
Ramsayia (Vombatidae); Palorchestes, the Marsupial Tapir (Palorchestidae); Diprotodon
(Diprotodontidae); Zygomaturus, (Diprotodontidae); Nototherium (Diprotodontidae); the giant
potorooid Propleopus (Potoroidae); giant wallabies, Protemnodon (Macropodidae); short-faced
kangaroos Simosthenurus, Sthenurus and Procoptodon (Macropodidae) and some exceptionally

1074 - MURRAY

Figure I. Reconstructions of Australian megafauna drawn to scale: 1, Sarcophilus laniarius, 15 kg; 2,
Zaglossus hacketti,, 20 kg; 3, Zaglossus ramsayi, 10 kg; 4, Megalania prisca, 1,000 kg; 5, Zygomaturus
trilobus, 450 kg; 6, Diprotodon optatum, 1150 kg; 7, Thylacoleo carnifex, 45 kg; 8, Palorchestes azeal 300
kg; 9, Phascolonus gigas, 150 kg; 10, Procoptodon goliah, 120 kg; 11, Macropus titan, 85 kg; 12,
Protemnodon anak, 50 kg; 13, Simosthenurus occidentalis, 50 kg; 14, Sthenurus atlas, 50 kg; 15, Propleopus
oscillans, 45 kg; (see text). Estimated weights are the average for each species; Thylacinus cynocephalus
(unnumbered) serves as scale).

large species or subspecific morphs of the living genus Macropus (Macropodidae) (Fig. 1).
Large Pleistocene forms of other extant genera include the giant Tasmanian devil, Sarcophilus
laniarius (Dasyuridae) a large swamp wallaby, Wallabia vishnu (=bicolor) and, for example,
local morphs of the Rednecked or Rufous Wallaby Macropus rufogriseus (Macropodidae). In
some localities, certain extant species, especially macropodine kangaroos, are the dominant
element in the fauna, with the megafauna in the background. In such assemblages, the very
large species (>100 kg) tend to be comparatively uncommon and incomplete, represented by
isolated teeth and robust postcranial fragments. There are many notable exceptions that relate
io the taphonomy, the type of community sampled and the geological age of each site
(Douglas et al. 1966, Behrensmeyer & Hill 1980, Brain 1980, Horton & Murray 1980). For
example, many individuals of Diprotodon have been recovered from Lake Callabonna in South
Australia (Tedford 1973). The majority of these factors are explained in detail in the chapter on
taphonomy (by Baird in this volume). Typical megafaunal assemblages, best known from
cave sediments, present in the following approximate categories of relative abundance:
tachyglossid monotremes, uncommon; large dasyurids, common; thylacinids, uncommon;
thylacoleonids, uncommon; giant vombatids, uncommon; palorchestids, very rare; large
diprotodontids, uncommon; giant potorooids, rare; giant wallabies, common; sthenurine
macropodid kangaroos, common; large macropodine kangaroo morphs or subspecies, common;
extant medium-sized macropodine species, very common. Many Pleistocene cave deposits
contain an excellent representation of smaller, extant mammalian species, which give a more
accurate indication of the type of communitics than the extinct megafaunal elements associated
with them.

HOW BIG WERE THEY?
ESTIMATION OF BODY SIZE.

In order to define megafauna in the absolute sense - on the basis of body mass, it is
necessary to estimate the body weight of each species, particularly those forms thought to be
near the designated rubicon. Most continental faunas, those of North America, Eurasia and
Africa in particular, retain many closely related, although somewhat smaller living forms,

PLEISTOCENE MEGAFAUNA - 1075

which can serve as a basis for estimation of body weight of similar extinct species. The
extinct Cave Lion, Panthera spelaeus, is very similar in form, but larger than, the living
African Lion, P. leo. A satisfactory estimation of the weight of a Cave Lion can be
extrapolated directly from a regression of African Lion proportions. In the case of Australia,
there are many extinct marsupial species which have no closely related living counterparts.
However, it is known that the average relationship of body mass to body length for all living
mammals is P=0.025 L3 (Jerison 1973). By omitting small mammals from the regression
(those under 5 kg) a slightly heavier average of P=0.035 L3 with an identical slope closely
approximates the weights of living male marsupial species given in Strahan (1983) and means
for the range of body weights of larger mammals given by Walker (1968). Single significant
figure estimations of Australian extinct marsupial body weights extrapolated from these
regressions are considered to be average adult male body weights from which, where there is
evidence of marked sexual dimorphism, weight of much smaller females can be extrapolated.

A general observation is that the Australian Pleistocene megafauna had fewer species of
very large animals than North America, Eurasia or Africa and that no Australian marsupial
species attained the truly gigantic proportions of the largest placental mammals (proboscidians,
edentates, hippopotamids, rhinocerotids) of those continents. The largest Australian marsupial,
Diprotodon optatum, probably weighed about 1 metric ton, equivalent to a Javan rhino,
although there is a great deal of variation in body size in these diprotodontids and a maximum
weight of up to 2 metric tons might have been attained by some individuals. Zygomaturine
and nototheriine diprotodontids, although massive, barrel-shaped mammals probably weighed
about one-half of a metric ton. The giant wombat, Phascolonus gigas, probably weighed less
than 200 kg; the giant kangaroos up to 150 kg, about three times heavier than the average
males of the largest living species of Macropus. Although the magnitude of estimated weights
of the Australian megafauna are not unusual in comparison to other continental faunas, the
contrasts between the surviving larger Australian marsupial body sizes and those attained by
the extinct Pleistocene forms are rather remarkable. The ecological and evolutionary
significance of these and other observations on body size will be considered further on.

RELATIVE MEGAFAUNA.

Certain smaller-bodied genera had species or morphs that also became extinct during the
Pleistocene. Among these are the giant Tasmanian Devil, the long-beaked echidnas and large-
bodied subspecies of living macropodid genera, sometimes considered separate species on the
basis of larger body size alone. These forms ranged from 20% to as much as 50% heavier than

1076 - MURRAY

vombatids _ large living macropodids

3044 living marsupials “

™ | @ living placentals

4 extinct .
marsupials Sthenurus

Macropus

; = 4)"
Simosthenurus, Sthenurus e
aad

T A
Thylacoleo, Propleopus) a Ae ae

2.0

1.0

Head-body length (m)

10 25 50 100

Figure 2. Method of weight estimation based on W = PL?- A regression line based entirely on living
mammals > 5 kg results in a predictive coefficient of P = 0.035; snout - rump lengths of restored
megafaunal species are placed in relation to the regression line with + 10% adjustment on either side, made
on the basis of living analogs and skeletal proportions of the fossils. (The regression was plotted
independently of the megafauna, admittedly an educated guess, but at least educated!).

their living counterparts (Marshall & Corrucini 1978) and are considered a part of the
megafauna extinction phenomenon regardless of their absolute size or their persistence
("dwarfing") after the Pleistocene as a smaller morph. The relative megafauna includes
vertebrates that weighed from about 10-15 kg (Zaglossus, Sarcophilus) to as much as 100 kg
in the case of Macropus titan, the Giant Grey Kangaroo and Macropus cooperi, the Giant Euro.

NON-MAMMALIAN MEGAFAUNA.

The estimated weights of the avian and reptilian megafauna species are arrived at in the
same manner as those for mammals. The raw figures obtained from the body lengths of these
forms are not entirely satisfactory because of the proportional elongation of the body in reptiles
(especially snakes) and birds show a foreshortened affect which results in an underestimation of
the weight of the giant flightless species. The giant varanid, Megalania prisca, may have
attained a nose to tail length of over 6 m (Fig. 2). By analogy with similarly proportioned
Crocodylus porosus, Megalania could have attained weights approaching a metric ton. Living
komodo dragons (Varanus komodoensis) grow to just under 3 m in length, and although they
are very impressive when viewed in their habitat, such specimens weigh no more than 40-50
kg (Auffenberg 1980). Perhaps an adjusted, but still prodigious, estimate of half to one-third
of the calculated weight of Megalania should be kept in mind. Miolania, the homed turtle, of
which several species are known, probably weighed between 50-200 kg. Giant flightless birds
of the family Dromornithidae are among the largest known birds. Some Miocene species
(Dromornis stirtoni) have massive limb bones that supported a short, deep, obviously heavy

PLEISTOCENE MEGAFAUNA - 1077

suids ursids equids cervids camelids bovids erhing

tapirids e 208
e A
N, a) |
© «eo? e
© ee F
2 a/P zygomaturines, nototheriines
| A Palorchestes

; Phascolonus

todon

200 500 1000 Weight (kg)

body and weights of over 300 kg have been postulated (Rich & Balouet 1984). Genyornis
newtoni was not as massive as some of the Tertiary species and may have weighed under 200
kg, but perhaps it was significantly more massive than the calculated >100 kg (Ostrich-sized).
Pleistocene crocodylids are poorly known and are assumed to follow proportions similar to
those of living Australian Crocodylus species. Quinkana appears to have been a small
freshwater crocodile attaining lengths of under 3 m and an estimated weight of 50 kg.
Pallimnarchus and perhaps Baru were closer to the proportions of C. porosus (Molnar 1982, P.
Willis, pers. comm.).

Obtaining accurate estimations of the body weights of the extinct megafauna is obviously
very difficult, but the implications of body size in ecology and physiology may prove to be
important for understanding the factors underlying Pleistocene extinction in Australia (Main
1978). An algorithmic basis of estimation such as W = PL3 (weight is proportional to length
cubed) is an improvement over guessing or the use of unspecified criteria such as "20% larger
than..." where it is not made clear whether a linear or an exponential relationship is implied.

EVOLUTION, MORPHOLOGY AND SYSTEMATICS

PHYLOGENY

The evolution of Pleistocene vertebrate communities and the phylogeny of its constituents
are essential to understanding the phenomenon of Pleistocene extinction from a biological
point of view. Until recently this broader perspective played a lesser role because few Tertiary
faunas had been described and because of a focus on interdisciplinary approaches emphasizing
prehistory, geomorphology, palaeoclimatology and other Quaternary specializations.

Early to Late Miocene faunas from South Australia, the Northern Territory and Queensland
are sufficiently well known to allow some speculations on the evolution of the distinctive
Pleistocene communities (Stirton et al. 1967, Archer & Bartholomai 1978, Archer

1078 - MURRAY

1984,Woodburne et al. 1985). Fossil quarries in the dry lake beds of the Simpson Desert of
South Australia contain some of the earliest representatives of the most conspicuous
megafauna taxa, the Diprotodontidae. This family, which contains the largest of all of the
marsupial species (Diprotodon), is represented by comparatively small and very generalized
Miocene genera that appear to have begun to differentiate into their respective subfamilial
groups by the end of the Oligocene. The genera Ngapakaldia and Pitikantia share features with
the marsupial tapirs (Propalorchestes and Palorchestes) as well as with the zygomaturines. An

Figure 3. Proposed phylogeny of the marsupial tapirs (Palorchestidae) A, Propalorchestes
novaculacephalus (mid-Miocene); B, Palorchestes painei (Late Miocene); C, Palorchestes azeal (Late
Pleistocene). Note gradual increase in size over approximately twelve million years.

PLEISTOCENE MEGAFAUNA - 1079

Early to mid-Miocene Diprotodon ancestor is unknown, although it probably shared a
generalized ancestor with the zygomaturines. Primitive zygomaturine or zygomaturine-like
forms, such as Nimbadon from the Carl Creek Limestone of the Northern Territory and
Bematherium from an unnamed formation at Riversleigh, Queensland represent continuations
of plesiomorphic diprotodontid lineages that radiated earlier in the Palaeogene. A small,
primitive marsupial tapir-like form (Propalorchestes) (Fig. 3) was already present in the late
part of the mid-Miocene, as were advanced zygomaturines such as Neohelos tirarensis, which
were probably ancestral to the Plio-Pliestocene genus Zygomaturus (Fig. 4). These small
(approximately sheep-sized) forerunners to the Pleistocene megafauna appear to have lived in
tropical rainforest habitats in association with many species of phalangeroid possums that
inhabited trees and scrub emarginating, crocodile-infested streams and lakes, Potorooid-like
kangaroos were the dominant macropodoids. Some primitive forms retained the second
premolar rather than losing it after the eruption of the large blade-like third premolar. This
strange radiation included the remote relatives of the Pleistocene giant musk-rat kangaroo,
Propleopus, among which also arose Ektadelta, a lineage of extinct carnivorous kangaroos.
Few of these early Neogene mammals attained large body sizes, among the largest perhaps
being the cow-sized /laria (Ilariidae). Neohelos tirarensis shows a gradual trend of body size
increase from the presumbly earlier Tirari Local Fauna of South Australia to the older D Site
faunas at Riversleigh, Queensland. In the later mid-Miocene, represented by the Bullock
Creek Local Fauna in the Northern Territory, Neohelos tirarensis attains much larger sizes
anticipating the earliest marsupial megafauna known in Australia from the Late Miocene at
Alcoota, Northern Territory.

As in the Pleistocene faunas, larger mammalian carnivorous species are few. The two mid-
Miocene genera of marsupial lions are also small. Priscileo was no larger than a cuscus, and
Wakaleo was about the size of a dingo (Fig. 5). The thylacinids were represented by a form no
larger than a fox. The largest carnivores were freshwater crocodiles, such as Baru, which may
have been more terrestrial than the living saltwater crocodile. The 12 million year old Bullock
Creek Local Fauna may represent the last phase of continuous wet tropics in the interior of the
continent. By the Late Miocene, perhaps 4-5 million years later, there is evidence of episodic
and severe droughts in central Australia where the fossil remains of at least five genera of large
diprotodontids appear to have succumbed to waterhole tethering. This may have been a direct
consequence of climatic changes due to the gradual northward drift of the continent, which by
Late Miocene or Early Pliocene times had reached a position near where it lies today. The
Alcoota Local Fauna contains many more very large species than any preceding fauna, certainly
rivalling, if not surpassing, the Pleistocene megafauna in terms of the number of mammalian
genera weighing over about 300 kg. Plaisiodon centralis was probably as large as the
Pleistocene Zygomaturus trilobus (Fig. 4), and the diprotodontine Pyramios alcootaensis, an
early relative of Diprotodon, was nearly as large (Fig. 6). The Alcoota palorchestid,
Palorchestes painei was considerably larger than the mid-Miocene marsupial tapir
Propalorchestes. Interesting is the appearance at that time of the first macropodid kangaroos,
including the genera Protemnodon and Sthenurus. Dorcopsoides and Hadronomus, two
common Alcoota kangaroo genera, appear to be very primitive macropodid kangaroos having
features transitional with potoroids. Large carnivores continue to be rare in the Late Miocene.
Thylacinus potens was slightly larger than the Recent species T. cynocephalus. The marsupial
lion Wakaleo alcootaensis was slightly larger than W. vanderleuert from the Bullock Creek
Local Fauna. At least three species of dromornithid birds were present, among them the huge
Dromornis stirtoni. Large freshwater crocodiles were common, and the giant varanid,
Megalania is represented. The significance of the Alcoota assemblage is that many of the
features of the Pleistocene megafauna are clearly recognizable some eight million years earlier,
presumably for similar reasons. The community structure consisting of few mammalian
carnivores and several genera of large-bodied bulk feeding browsers in the company of giant

1080 - MURRAY

A

Figure 4. Phylogeny of zygomaturine diprotodontids: A, Neohelos tirarensis, (mid-Miocene;, B, Kolopsis
torus, Late Miocene; C, Zygomaturus trilobus, Late Pleistocene, cranial outlines suggest a simple
anisometric transform progressing from low and elongated to deep and deflected in the cranial base.

giant birds, crocodiles and the giant varanid Megalania was established long before the
Pleistocene, in response to the reduction of forest cover and concommitant increase in savanna
grassland. The Pleistocene megafauna appears to be a culmination of this trend in which the
number of large browsing specialists becomes reduced to a few very large genera and species

accompanied by radiations of grazing forms (macropodids) that are only beginning to appear in
the Late Miocene.

PLEISTOCENE MEGAFAUNA - 1081

Figure 5. Phylogeny of marsupial lions (Thylacoleonidae): A, Priscileo pitikantensis, (late-mid-Miocene);
Wakaleo vanderleueri, (Late Miocene);Thylacoleo carnifex, (Late Pleistocene). The family shows a general
increase in size through the later Tertiary. (After Owen 1877, Rauscher 1987, Murray ef al. 1987).

The mid- to Late Pliocene is represented in several localities. Perhaps the most complete of
these assemblages is the Chinchilla Local Fauna of Darling Downs Queensland. This fauna
contains many forms transitional between the Alcoota Local Fauna and the characteristic
Pleistocene megafauna. A remarkable feature of the Chinchilla is the appearance of highly
specialized, gigantic diprotodontids such as Euryzygoma and Zygomaturus and a proliferation of
the macropodid kangaroo genera and species that were faintly indicated in the background of the
Alcoota assemblage. A large marsupial lion, Thylacoleo crassidens, was about the same size
as but slightly more primitive than the Pleistocene marsupial lion, Thylacoleo carnifex. The
Chinchilla palorchestid, Palorchestes parvus, is larger than its Late Miocene predecessor,
Palorchestes painei, but smaller than Palorchestes azeal, the typical Pleistocene form.
Chinchilla probably represents an open riverine community surrounded by wooded grasslands.
Nearly all of the genera known from the Pleistocene were already present at Chinchilla, with
the transition to the typical Pleistocene fauna being principally one of species succession. In
fact, the faunal transition from the Late Pliocene to the Early to mid-Pleistocene is only

1082 - MURRAY

scarcely evident. Certain localities in South Australia are thought to represent Early
Pleistocene faunas because they contain some characteristic Pliocene species along with

Figure 6. Phylogeny of diprotodontine diprotodontids: A, Pyramios alcootense (Late Miocene);
Euryzygoma dunenese, (early -Middle Pliocene); C, Diprotodon sp., (Pleistocene). (Woodburne 1967, after
Archer & Bartholomai 1978, Archer 1984).

some typical Pleistocene forms (Williams 1980). It is unlikely that a faunal definition of the
Plio-Pleistocene boundary of the sort employed by palaeontologists in Eurasia and Africa will
prove satisfactory in Australia, principally because the Australian fauna is confined
biogeographically.

The available information seems to indicate that the distinctive Late Pleistocene megafauna
of Australia is largely an expression of a general trend of aridification of the continent that

PLEISTOCENE MEGAFAUNA - 1083

becomes evident in the Late Miocene. Until the later part of the mid-Miocene, the mammalian
fauna contained few large, and absolutely no gigantic, species, possibly because they were
adapted to practically continuous temperate and tropical rainforest habitats. The increase in
body size among diprotodontids and early macropodids was apparently in reponse to more open
habitats, presumably in association with the invasion of savanna grasslands. Initially, these
habitats recruited a large number of specialist genera of the megafauna type, perhaps in relation
to rich ecotonal environments. As this deteriorated into greater habitat uniformity, the number
of large genera was reduced in favour of fewer generalized, bulk feeding genera that underwent
selection for even greater body size in response to the lower per unit nutritional quality of the
browse. Simultaneously, as the habitats favoured grazing and distance efficient locomotion,
the macropodid kangaroos radiated rapidly in the Late Miocene and Early Pliocene. Because of
their locomotor efficiency, specialist macropodid browsers, such as sthenurine kangaroos filled
some of the vacant niches of the waning diprotodontid specialist browsers. Even the highly
specialized, selective feeding browser Palorchestes shows an interesting trend towards increased
body size and higher crowned molars over a period of about twelve million years.

The palaeontological record implies that the Pleistocene megafauna was highly evolved and
finely adjusted to the conditions under which it was living in the Late Pleistocene and that the
habitat changes which shaped this community were for the most part gradual until about forty-
thousand years ago. Hope (1982) has already provided an excellent review of the development
of aridity in relation to the Australian Cainozoic faunas.

MORPHOLOGY AND SYSTEMATICS.

The extreme diversity of the megafauna assemblage presents an unwieldy collection for
morphological and systematic treatment. Consequently, only the broader aspects of the topic
will be introduced. Additional systematic and morphological data are presented with the
introduction to individual taxa.

Readers should refer to the systematic accounts elsewhere (e.g. Marshall 1984) and in other
chapters of this book in order to supplement the piecemeal quality of megafauna systematics.
The classification used here is abridged from Aplin & Archer (1987).

CLASS MAMMALIA
SUBCLASS THERIA
ORDER MONOTREMATA
SUPERLEGION Incertae Sedis
Family Tachyglossidae
SUPERCOHORT MARSUPIALIA

ORDER DAS YUROMORPHIA
Family Thylacinidae
Family Dasyuridae

ORDER DIPROTODONTIA

SUBORDER VOMBATIFORMES
INFRAORDER PHASCOLARCTOMORPHIA
Family Phascolarctidae
INFRAORDER VOMBATOMORPHIA
Family Diprotodontidae*
Family Palorchestidae*
Family Vombatidae
Family Thylacoleonidae*
SUBORDER PHALANGERIDA
SUPERFAMILY MACROPODOIDEA

Family Potoroidae

1084 - MURRAY

Family Macropodidae
*extinct forms

Fossil and Recent morphological evidence indicates that the monotremes are closer to the
therian mammals than was generally believed. The morphology of the molar teeth of the Early
Cretaceous ormithorhynchid monotreme Steropodon are incipiently tribosphenic, although
highly specialized (Archer e7 al 1985, Kielan-Jaworoska et al. 1987). Re-evaluation of the
cranial morphology by Kemp (1983) suggests that the anterior lamina of the periotic of
monotremes is homologous with the alisphenoid of therian mammals.

Pleistocene and Recent monotremes are highly specialized egg-laying mammals that retain
numerous primitive skeletal features characteristic of theriomorph cynodont reptiles. Their
basic morphology is also very similar to that of primitive triconodont prototherian mammals
such as morganucodonts. Miocene platypuses retain a permanent dentition throughout their
lives. In Pleistocene and Recent ornithorhynchids, the dentition is suppressed early in life and
replaced by cornified grinding pads. They also show a trend of body size reduction after the
Miocene. Ornithorhynchids are poorly represented in Pleistocene assemblages and are not
included with the megafauna.

Tachyglossids (echidnas) are reasonably well represented as Quaternary fossils, and some of
them were large enough to be included among the megafauna assemblage (Murray 1978a-c).
Their postcranial elements are robust and compact, the humerus and femur are broad and flat
with poorly differentiated articular surfaces and faint indications of epiphyseal growth. The
vertebrae are comparatively undifferentiated from front to back, and the fibula retains a peculiar
flabelliform process. The pectoral girdle retains a separate coracoid, procoracoid or interclavicle
and an independent episternum. As in cynodont reptiles, the anterior fossa of the scapula is
not present. The distinctive tachyglossid cranium is elongated and narrow with a large,
smooth, thin-walled braincase. The rostrum is an elongated tubular construction terminating
in an oval narial aperture. Echidnas are totally edentulous, processing their insect prey with a
tongue-palatal grinding complex. The inferolateral wall of the braincase is composed of the
anterior process or lamina of the periotic rather than the lateral or ascending wing of the
alisphenoid bone. The sutures of the cranial lamina are obliterated early in life. The palate is
elongated and terminates in a pair of "echidna pterygoids" homologous with the reptilian
epipterygoids or the pterygoid hamulus of typical therian mammals. Monotremes totally lack
a bony tympanic bulla and meatal canal, which is entirely cartilagenous.

The majority of australidelphian marsupial taxa are small mammals; less than half of the
twenty-seven known living and extinct families contain species over 10 kg body weight. Older

Figure 7. Osteological characters of monotremes and marsupials: A, cranium and dentary of tachyglossid
monotreme, Tachyglossus aculeatus; B, humerus and femur of tachyglossid monotreme, Tachyglossus
aculeatus; C, monotreme pectoral girdle, (Tachyglossus); [Cf, coracoid foramen, Cl, clavicle; Co, coracoid;
Ec, epicoracoid; Es, episternum; Gf, glenoid fossa; Ps, presternum); D, Polyprotodont marsupial, Dasyurus
viverrinus, brackets show incisors and premolars for comparison with E, diprotodont, Trichosurus vulpecula;
F, left dentary viewed from behind, showing inflection of angle [A]; G, macropodid humerus showing
epicondylar foramen, often present in marsupials; H, right foot of Vombatus ursinus, showing fused digits I
and III (symphalangy); I, innominate bones (pelvis) of a kangaroo, showing epipubic bones [Ep]. (After
Jones 1968; Marshall 1984).

PLEISTOCENE MEGAFAUNA - 1085

1086 - MURRAY

classifications divided the Australian marsupials into two groups, those with more than two
lower incisors, the polyprotodonts and those with only two lower incisors, the diprotodonts
(Fig. 7). The diprotodont marsupials also have a fusion of pedal digits II and II, whereas, with
the exception of the bandicoots (Perameloidea), all other polyprotodonts lack this
specialization.

Very few Australian polyprotodont marsupials qualify as megafauna. Two families of the
Dasyuroidea, the Thylacinidae and the Dasyuridae, contain a single genus each of medium to
large-sized carnivores. Dasyuroid marsupials share many similarities with American
didelphoids. Their molars are of the basic tribosphenic type with a series of well-developed,
labial stylar cusps, the canine is large and a post-incisor diastema is not present. The incisors
are small and peg-like, and more than one incisor is present in each quadrant of the upper and
lower jaws. Characteristic of plesiomorphic marsupials, they possess 4 molars, an inflected
angle of the dentary, an alisphenoid tympanic process accommodating the middle ear and with
the notable exception of thylacinids, epipubic bones, Although thylacinids have been separate
from dasyurids since before the Middle Miocene, their dentitions are similar, reduction of
stylar cusps in the thylacinids being the primary distinction dentally (Archer 1982),
Thylacinids have converged with placental canids and South American borhyaenid marsupials,
while dasyurids show relatively little differentiation from a basic didelphimorphian condition.

The Pleistocene thylacinid Thylacinus cynocephalus has a superficially wolf-like cranium
and long, slender dentaries in contrast to the dasyurid Sarcophilus laniarius, which has a short,
low, broad triangular cranium and comparatively deep, short dentaries with concomitant loss of
the first premolar from the tooth row. The postcranial skeleton of. Thylacinus is moderately
convergent with canids in having long, slender limbs and narrow girdles (Keast 1982).

Diprotodontian marsupials are divided into two suborders, the Phalangerida and the
Vombauformes, Macropodoids (kangaroos) are the only group that attain very large body sizes
among the phalangeridans. The majority of large Australian marsupials belong to the
Vombatiformes, which includes the marsupial lions, diprotodontids, marsupial tapirs, wombats
and koalas. Both subordinal groups appear to be very ancient and have probably coevolved in a
complex manner. The current classification (Aplin & Archer 1987) strongly dichotomizes
these suborders, although many basic, ostensibly symapomorphic (shared-derived or advanced)
similaritics pervade among certain vombatimorph families and the macropodoid-phalangerid
lineages of the Phalangerida. The megafaunal diprotodontians are characterized by varying
degrees of bilophodonty in their molars (with the exception of thylacoleonids), the presence of
a long post-incisive diastema, reduced or absent canines, reduction or loss of the first and
second premolars in conjunction with hypertrophy of the third premolar and of the lower
central incisor pair. The phalangeroid-like thylacoleonids are united with the vombatimorphs
by their basicranial morphology, in which the middle ear, like that of wombats and
diprotodontids is surrounded by processes of the squamosal rather than the alisphenoid.
Differences in the postcranial skeletons of these groups are dominated by the contrasts between
the predominately quadrupedal plantigrady of the vombatimorphs and the bipedal saltatory
(hopping) locomotion of the kangaroos.

The systematics and morphology of other Australian vertebrate megafauna taxa are
discussed in detail elsewhere in this book. These include large marine placental mammals
(pinnepeds and cetaceans), reptiles, principally crocodylids, varanid lizards, tortoise-like
miolanids and large birds, predominantly casuariids, dromornithids and megapodiids.

Figure 8. Reconstructions of: A, Zaglossus hacketti and B, Zaglossus ramsayi compared with C,
Tachyglossus aculeatus. Cranium of: D, Z. hacketti, restored from Zaglossus robusta; E, Zaglossus ramsayi; F,
and G, Zaglossus bruijnii,

PLEISTOCENE MEGAFAUNA - 1087

1088 - MURRAY
SPECIES OF AUSTRALIAN PLEISTOCENE MEGAFAUNA

Order Monotremata Bonaparte 1838
Family Tachyglossidae Gill 1872
Genus Zaglossus Gill 1872

Long-beaked Echidnas (Figs 7-10; Table 1). Two or possibly three species of
Zaglossus occur in Australian Pleistocene deposits. Remains of Z. ramsayi (Owen 1844), a
species similar to the living New Guinea form Z. bruijnii, have been identified in many
localities in south and eastern Australia. The postcranial elements are distinguished from

Figure 9. Late Pleistocene tachyglossid cranial material from Mammoth Cave, Western Australia; A,
Tachyglossus aculeatus, B,-E, Zaglossus ramsayi.

PLEISTOCENE MEGAFAUNA - 1089

E F

0 5cm li

Figure 10. Selected Late Pleistocene tachyglossid postcranial elements from Mammoth Cave, Western

Australia: A, innominate Zaglossus hacketti; B, Zaglossus ramsayi; C, Tachyglossus aculeatus, D, Scapula,
Zaglossus hacketti; E, Zaglossus ramsayi; F, Tachyglossus aculeatus . G, Humerus (restored) Zaglossus

hacketti; H, Zaglossus ramsayt; I, Tachyglossus aculeatus; J, Femur, Zaglossus hacketti; K, Zaglossus ramsayi
L, Tachyglossus aculeatus; M, Tibia, Zaglossus hacketti, N, Zaglossus ramsayi; O, Tachyglossus aculeatus.

1090 - MURRAY

Tachyglossus aculeatus (Illiger 1811) by their much larger size and certain proportional
differences in the limb segments. The cranium has an elongated, decurved rostrum like that of
Zaglossus bruijnii, although it appears to be somewhat shorter, broader and straighter (Murray
1978a; Fig. 9). A much larger tachyglossid Z. hacketti (Glauert 1914), known only from
Mammoth and nearby Labyrinth caves in southwestern Australia, differs from other Zaglossus
species in some particulars of its postcranial skeleton (Fig. 10). There are no remains of
the

cranium. Its limb segment ratios are more similar to T. aculeatus than to Z. bruijnii or Z.
ramsayi. A cranial fragment and a humerus of an exceptionally large fossil echidna, Z. robusta
(Dun 1895), thought to be Middle Miocene in age (T.H. Rich eg al., this volume) may have
some connection with the west Australian "giant echidna". In Pleistocene deposits,
Tachyglossus fossils are less abundant than those of Zaglossus. Z. bruijnii has been identified
in Pleistocene deposits in New Guinea. The youngest age for Zaglossus in Australia is 13,000
BP from Main Cave, Montagu, Tasmania. This date was determined using aspartic acid and
electron spin resonance techniques (Goede & Bada 1985). Goede & Bada pointed out that
reworking of older bone was a possibility to explain such a young date. Murray & Chaloupka
(1984) identified a prehistoric Zaglossus painting in the Arnhem Land escarpment.

Order Dasyuromorphia Gill 1872
Family Dasyuridae (Goldfuss 1820) Waterhouse 1838
Genus Sarcophilus Geoffroy and Cuvier 1837

Tasmanian Devils (Figs 11-12; Table 2). The "Giant" Tasmanian Devil, Sarcophilus
laniarius (Owen), was a common element in Late Pleistocene cave deposits in southern South
Australia, southern Victoria, eastern New South Wales and southeastern Queensland. S.
laniarius is essentially a statistically defined species, as there are no particular morphological
features other than significantly larger size and associated allometry that serves to distinguish
them from the “average-sized" present-day Tasmanian devils that survive today on the island of
Tasmania and those that recently became extinct during the late Holocene on the Australian
mainland (Ride 1964, Dawson 1982a; Figs 11-12; Table 2). Gill (1953) observed that S.
laniarius specimens were from 15% to 50% larger than the average Tasmanian Devil. Dawson
(1982a) found that S. /aniarius dentitions were on the average about 14% larger than those of
S. harrisii. Werdelin (1987) estimated the large S. laniarius to be 16% larger than the living
devil. Exceptionally small Tasmanian devil morphs were also present in the Late Pleistocene
and early Holocene. Some temporal overlap is indicated among the large, medium and small
devils (Dawson 1982). Devils are easily distinguished from other dasyurids by their small,
crowded, blunt-crowned premolars, low-crowned M2 and stout canines in addition to their larger
size. Small devils such as the Nelson, Laura and Padypadiy specimens (Calaby & White 1967,
Horton 1977, Werdelin 1987) are not much larger than the bowlingi subspecies of Dasyurus
maculatus from King Island in the Bass Strait (Hope 1973). After about 10,000 yBP, Devil
populations from deposits in the southeastern states were about the same size as the living
Tasmanian Devil (S. laniarius Werdelin), with a small form persisting until the late Holocene
in the tropical north (Calaby & White 1967), Dawson (1982a) suggested a 23,000 yBP
Clogg's Cave !4C date as terminal for S. laniarius dawsoni Werdelin. Werdelin (1987) argues
that S. laniarius has priority over harrisii and that the living Tasmanian Devil is a subspecies
of §. laniarius (S. laniarius harrisii).

Figure Il. Late Pleistocene devils and thylacine: A, Sarcophilus laniarius laniarius, B, Sarcophilus
laniarius dixoni; C, Sarcophilus laniarius (small morph); D, humerus of Sarcophilus harrisii compared with
Late Pleistocene; E, Sarcophilus laniarius laniarius; F, Skull of Sarcophilus harrisit, (Tasmanian devil
compared with Thylacinus cynocephalus (Tasmanian Wolf). (After Owen 1877; Merrilees & Porter 1979 and
Merdelin 1987).

PLEISTOCENE MEGAFAUNA - 1091

1092 - MURRAY

I have retained the older generic and specific nomenclature throughout the text in order to avoid
the inevitable confusion that this causes, if not accompanied by lengthy synonomies and
discussions, although I agree with Werdelin's diagnosis.

Prehistoric Tasmanian Devils figure prominently in the rock art of the Arnhem Land
escarpment (Calaby & White 1967; Murray & Chaloupka 1984).

Order Dasyuromorphia (Gill,1872)
Family Thylacinidae Bonaparte, 1838
Genus Thylacinus Temminck 1827

\N 0 20cm
ae |

Figure 12. Skeletal elements of Tasmania devil, Sarcophilus : A, living Tasmanian devil skeleton; B, C,
foot [Ca, calcaneus; As, astragalus; Cu, Cuboid; Na, navicular, V, metarsal V; I, metatarsal I, En,
entocuneiform] D, innominate; E, Femur, F, scapular; G, humerus; (D-G after Merrilees & Porter 1979).

Tasmanian "Wolf" (Figs 11-13; Table 3). The once ubiquitous pouched "wolf"
Thylacinus cynocephalus (Temminck 1827) is now confined to Tasmania where it is
exceedingly rare or recently extinct. Thylacines survived on the mainland until at least 3,300
years ago (Partridge 1967) and into the Holocene in New Guinea (Van Deusen 1963, Plane
1967). Thylacines are included here as a "surviving" megafaunal element (Figs 11, 13; Table

PLEISTOCENE MEGAFAUNA - 1093

3). Pleistocene thylacines were once considered to be two separate species, T. spelaeus and T.
rostralis. These have since been demonstrated to be no different than T. cynocephalus (Ride
1964, Dawson 1982). Thylacine remains are less abundant in fossil assemblages than those of
Sarcophilus. Worn, isolated molars of Thylacinus are rather similar in size and shape to those
of S. laniarius but can be readily distinguished by their proportionally longer, narrower
shearing crest (metacrista), reduced protocones and more lingual position of the (also reduced)
stylar cusp D, which gives the buccal, occlusal outline a more concave appearance. The lower
molars of Sarcophilus are broader buccolingually and M4-5 are absolutely much smaller than
those of thylacines. Occasional confusion of isolated Thylacinus and Sarcophilus lower molars
can have annoying consequences; for example, the Thylacinus major dentary in plate V, Fig. 8
of Owen (1877) was drawn with the last three molars of a Sarcophilus laniarius, presumably
forming the basis for Owen's new specific designation (Stephenson i963). Thylacines have an
additional premolar, and these are comparatively high-crowned and uncrowded. The thylacine
was a favourite subject of prehistoric aboriginal artists in Arnhem Land and the Kimberleys
(Murray & Chaloupka 1984).

Figure 13. Skeletal elements of pouched wolf, Thylacinus : A, recent pouched wolf skeleton; B,
innominate; C, scapula; D, femur, E, humerus. (After Merrilees & Porter 1979, Archer 1984).

1094 - MURRAY

Order Diprotodontia Owen 1866
Family Phascolarctidae Owen 1839
Genus Phascolarctos De Blainville 1816

Koalas (Fig. 14; Pl. 1). Phascolarctids, koalas, are not a prominent element in Late
Pleistocene fossil assemblages, presumably because they are highly arboreal (Fig. 14). Their
distribution has changed markedly since 35,000 years ago when their range extended to Western
Australia (Merrilees 1968b). The Pleistocene species Phascolarctos stirtoni (Bartholomai
1968) was about one third again larger than the living species P. cinereus according to
Bartholomai (1968), but differs little otherwise.

Figure 14. Skeletal elements of recent Koala, Phascolarctos cinereus : A, cranium and dentary; B,
skeleton; C, scapula; D, humerus; E, innominate; F, femur. (After, Merrilees & Porter 1979, Archer 1984).

Koalas are an endangered species at present. Their populations are waning due to habitat
disturbances and epidemic venereal Chlamydia infection. Their highly specialized diet, arboreal
adaptations and various morphological peculiarities (Aplin & Archer 1987, Haight & Nelson
1987) indicate that koalas have been evolving independently from other diprotodontians for an
extremely long period of time. Aplin & Archer (1987) suggest that phascolarctids represent
the sole survivor of an ancient radiation that mirror the ancestors of the vombatomorphian
suborder of diprotodontians. Distinctive selenodont molars of this lineage are the basis of
Archer's (1976) hypothesis that selenodonty preceded development of bilophodonty in diproto-

PLEISTOCENE MEGAFAUNA - 1095

Figure 15. Pleistocene wombat fossils: A, dentary of Vombatus hacketti, B, dentary of Warendja
wakefieldei,; C, dentary of Phascolomys medius; D, cranium of living Tasmanian Vombatus ursinus compared
with; E, Vombatus hacketti; F, dentary of Phascolomys magnus (=Ramsayia curvirostris), G, entary a
Phascolonus gigas, H, palatal fragment of Phascolonus medius; lines show interdiastemal palate; I, ventr;
and lateral aspects of diastemal palate of Ramsayia curvirostris; J, Phascolonus gigas.

1096 - MURRAY

dontan molar evolution and that this condition is only slightly removed from that of
perameloids (bandicoots) (Archer 1984). The Tertiary fossil record documents a gradual
reduction of the number of phascolarctid genera and species down to a single (of late
precariously) existing form. Morphological, serological and karyotypological evidence
indicates that the vombatids are the closest living relatives of the phascolarctids (Aplin &
Archer 1987).
Order Diprotodontia Owen 1866
Family Vombatidae Burnett 1830

Genus Phascolonus Owen 1872

Genus Phascolomys Owen 1873

Genus Ramsayia Tate 1951

Genus Warendja Hope and Wilkinson 1982

Genus Lasiorhinus Gray 1863

Genus Vombatus Geoffroy 1803

Wombats (Figs 15-17; Table 4). The largest vombatid, Phascolonus gigas (Owen 1872)
was between 1.6-1.8 m long and probably weighed about 200 kg. Marshall (1984) noted that
these wombats attained the size of acow. The postcranial morphology of P. gigas is similar
to Vombatus ursinus, including its forelimb morphology, which reflects an ability to dig by
scratching like its smaller relatives. The broad, flat-topped skull of Phascolonus gigas is about
twice as long as that of Vombatus, and its dentaries are about four times as massive (Figs 15-
17; Pl. 1). Phascolonus's body was elongated, wide and low-slung, and it would have stood
less than a metre high at the shoulder. Its distinctive broad, flat incisor teeth suggest that it
may have diverged from other wombats at an early stage of wombat evolution, but not before
the characteristic fossorial features were present. Another very large wombat, Ramsayia
curvirostris (Tate 1951), known only from incomplete material, was a narrow-incisored species
like those of the living wombats, Lasiorhinus and Vombatus. Ramsayia was smaller than
Phascolonus but certainly attained a prodigious size. Dawson (1981) placed the remaining two
large wombats, Phascolomys medius (=Lasiorhinus medius) and Phascolomys magnus in the
genus Ramsayia. Retaining the old, probably invalid, genus for descriptive purposes,
Phascolomys magnus dentaries are a little more than twice the size of those of V. ursinus.
The dentaries of Phascolomys medius are only about a quarter to one-third larger than
Vombatus or Lasiorhinus. The remaining Pleistocene wombat species were not particularly
large forms but became extinct along with the megafauna species (Figs 15, 16; Tables 4-5) and
are of interest.

Warendja wakefieldi (Hope & Wilkinson 1982) is a plesiomorphic wombat dentary with a
relatively straight as opposed to rocker-like, inferior border of the horizontal ramus, a straighter
cheek tooth row and an unspecialized ascending ramus that distinguishes the species from all
known wombats younger than the Miocene. Vombatus hacketti, known only from Western
Australia, was similar in size to the living V. ursinus but shows some proportional differences
as does Lasiorhinus angustidens, an extinct P. medius-sized Hairy-nosed wombat. The living
Northern Hairy-nosed wombat, Lasiorhinus kreffti, now confined to a small range in western
Queensland, is embarking on a similar fate.

Specific identification of the larger fossil wombats is made difficult by the often poor
quality of specimens, a lack of adequate samples and overlapping size ranges. The peculiarities
of wombat dental morphology makes them easy to distinguish from all other living and extinct
marsupial species because, with the exception of the Miocene Rhizophascolonus crowcrofti,
their molars are open-rooted and divided into two simple-crowned moieties, each having a thin,
prominent enamel perimeter and an exposed cementum interior. Phascolonus gigas is
distinguished from Ramsayia spp. by the shape of its diastemal palate, which is broad
anteriorly and narrow posteriorly between its third premolars. Ramsayia's diastemal palate is
broad posteriorly and narrow anteriorly, and the diastemal palate of Phascolomys medius is

PLEISTOCENE MEGAFAUNA - 1097

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Dentitions of larger Pleistocene Vombatids; A, Phascolonus gigas maxillary and mandibular
D, Ramsayia curvirostris, E, upper incisor of

dentitions; B, Phascolomys magnus, C, Phascolomys medius;
Phascolonus gigas; F, upper incisor of Ramsayia curvirostris. (After Owen 1877, Stirling 1913, Marcus 1976).

Figure 16.

1098 - MURRAY

constricted in the middle and broad both anteriorly and posteriorly (Dawson 1981).
Phascolomys medius differs from Vombatus ursinus in having a deeper dentary, in which the
anterior edge of the masseteric fossa lacks a definite ridge and the masseteric foramen is absent.
The molars of Phascolonus medius are larger and have rounded lobes separated by U-shaped
valleys on the lingual side (Marcus 1976). Phascolomys magnus (=Ramsayia curvirostris) has
a deep, thin horizontal ramus, large but not remotely as massive as that of Phascolonus gigas.
At the level of M3 Phascolonus magnus's dentary is about 50 mm high. In comparison with
Phascolomys medius, the occlusal outlines of the molar moieties are more diamond-shaped.
The length of the cheek tooth row of Phascolonus magnus is on the order of 60-65 mm. The
dentaries of Phascolonus gigas are extremely short and deep with the inferior border of the
horizontal ramus describing a tight-radius rocker from which the incisor alveolus protrudes
horizontally beyond a massive torus. At the level of M3.4, the dentaries are from 75-85 mm
deep. The molars decrease in length posteriorly. The molar moieties are separated by a
shallow, parabolic concavity lingually. The buccal valley is a deep, obliquely-oriented open V-
shaped indentation. The lobes of the molar moieties are more rounded than in Phascolomys
magnus but narrower than in Ramsayia curvirostris. An important discriminating feature noted
by Marcus (1976) is the pseudolophodont character of the molar crowns when viewed from the
labial side, resulting from wearing a transverse ridge on each lobe. The occlusal surfaces of the
molars of R. curvirostris are worn flat in contrast to the undulating surfaces of Phascolonus.
The upper incisors of Phascolonus gigas are diagnostic for the genus. These are extraordinarily
broad, flat, thin, gently curved, chisel-like teeth that resemble the upper incisors of
Diprotodon. Although R. curvirostris molars overlap in size with those of Phascolonus, the
lophs are more rounded and the median valleys are acutely V-shaped. The teeth are broad
relative to their length and the dentine extends over the alveolar margins. Ramsayia's upper
incisors are much narrower, relatively thicker and have a much lighter radius of curvature.
Individual elements of the postcranial skeleton of Phascolonus are illustrated in Stirling
(1913). The distal limb segments are exceptionally short and broad, and the femur, although
considerably smaller, is strikingly similar to that of Diprotodon (Fig. 17).

Order Diprotodontia Owen 1866
Family Diprotodontidae Gill 1872
Subfamily Zygomaturinae Stirton, Woodburne and Plane 1967
Genus Zygomaturus Owen 1859

Marsupial "Rhinoceros" (Figs 18-22; Pls 2,3; Table 5). The definitive Pleistocene
zygomaturine, Zygomaturus trilobus (Macleay 1858) was the largest and most highly derived
member of this once diverse subfamily of diprotodontids. A smaller, less specialized Early
Pleistocene species, Zygomaturus keani (Stirton et al. 1967) is known from the Palankarinna
Local Fauna of South Australia. The systematics of Late Pleistocene Zygomaturus species
need revision for Z. trilobus is either highly variable, or there was more than one species. Our
understanding of the genus is also bedeviled by a confusion with the genus Nototherium,
stemming from a series of mixed-up assignments by Owen (1873). Although subsequently
rectified by Woods (1968), specimens of Zygomaturus in museum catologues may still bear
the names Nototherium mitchelli, N. victoriae and N. tasmanicum.

For convenience, I will treat the familiar barrel-shaped, bluff-browed Pleistocene giant,
Zygomaturus, as a single species, Z. trilobus. "Pygmy" zygomaturines are also known from
poorly sampled Late Pleistocene Pureni Local Fauna in New Guinea (Woodburne et al. 1985),
and it would come as no great surprise if similar forms were eventually found in the sparsely
sampled Pleistocene deposits in the tropical north of Australia. Z. trilobus was probably the
second largest marsupial species. It attained a length of just under 2.5 m and a shoulder height

PLEISTOCENE MEGAFAUNA - 1099

50cm

Figure 17. Reconstruction and postcranial elements of Phascolonus gigas. A, reconstructed skeleton
based on: B, scapula; C, humerus; D, ulna; E, femur; F, tibia; G, radius; H, metatarsal V of Vombatus
ursinus compared with; I, Phascolonus gigas and; J, Diprotodon australis, K, fibula. (After Stirling

1913).

1100 - MURRAY

of about 1 m. It probably weighed between 300-500 kg. The massive cranium of Z. trilobus
is characterized by highly angled, prow-like nasal bones terminating in a pair of lateral crests or
bosses that have been variously interpreted as horn-like structures or rhinarial callosities
(Murray 1978c). The small orbits are also surmounted by roughened tuberosities on the frontal
crests, suggesting an incipient development of a titanothere-like craniofacial armament. The
widely flared and exceptionally deep zygomatic arches terminate anteriorly in long, stout
zygomatic processes. In contrast to the wide nasal complex, the rostrum narrows precipitously
in the diastemal region. The central incisors, both upper and lower sets, are recumbent and
tusk-like with labially divergent crowns. The upper third premolar is a distinctive 5-cusped
molariform tooth, unique to this group of diprotodontids (Fig. 18). The postcranial skeleton
of Z, trilobus is completely known except for some missing elements of the feet (Fig. 22).
The cervical vertebral centra are moderately compressed with the neural spines diminishing in
length from C2 to C7. The thoracic centra are broad and deep with low, stout neural spines
and robust, subround to flattish oval ribs having progressively less curvature and higher
articulations from front to back. The bodies of the lumbar vertebrae are markedly constricted
and wide, with short, stout transverse processes and low, broad posteriorly directed neural
spines. The innominates are exceptionally broad and flat with laterally flared ilia. The scapula
is long and narrow with a rugose tuberosity on its vertebral border. The scapular spine is
exceptionally prominent and terminates in a powerful, strap-like acromion process. The
anterior limb segments are subequal and relatively longer than the markedly unequal hind limb
segments. The humerus is relatively long, broad and flattish-oval in section. The deltoid
crests are moderately well-developed suggesting, in conjunction with its stout claws, that
Zygomaturus used its forelimbs for more than locomotion, although its short olecranon
process indicates that it was not a powerful scratch-digger. The femur is much longer than the
tibia and fibula, but slightly shorter than the humerus. The head of the femur is little deflected
from the flattish diaphysis, which is expanded distolaterally into a massive trochanter. The
fibula is stout and bears an articular facet for the lateral condyle of the femur.

The armatured skeletal restorations of these beasts look rather uncomfortable, and I suspect
that they are not quite right. Zygomaturus has many elements in its skeleton (long acromion
process, scapula shape, vertebral proportions, limb segment ratios, highly modified tarsal
elements and innominate morphology) that are reminiscent of the least specialized American
giant ground sloths. Regardless of their probable appearance, the zygomaturines comprise one
of the best known evolutionary sequences in Australia, and they will undoubtedly prove
valuable in later Tertiary biochronology.

Zygomaturus trilobus is in a complementary distributional relationship with Diprotodon in
many parts of the continent. They appear to overlap in ecotonal situations but are mutually
exclusive in habitat extremes, with Diprotodon preferring the semiarid plains of the interior and
Zygomaturus favouring the wooded, hilly coastal regions (Glauert 1910, Gill & Banks 1956,
Hope 1982). Circumstantial evidence of the association of aboriginal man with Z. trilobus
comes in the form of a fascinating artifact, an upper third premolar mounted in spinifex resin
given to Kim Akerman by Kimberley Aborigines (Akerman 1973). It may have been
fossilized already, as isolated Zygomaturus teeth have been found eroding from stream deposits
in the nearby Victoria River district of the Northern Territory. Archer et al. (1980) felt that the
Zygomaturus trilobus bones in Mammoth Cave showed evidence of butchering by aboriginals.
A small, as yet formally unnamed zygomaturine ("Danitherium") (Fig. 20) from a cave in the
the Baliem Valley of Irian Jaya is associated with artifacts and a number of longbones showing
butchering cuts (Phillip Walker, pers. comm.).

PLEISTOCENE MEGAFAUNA - 1101

Figure 18. Pleistocene Diprotodontidae: A, Diprotodon optatum; B, Zygomaturus ?trilobus (tasmanicum);
[Compare B, with Fig. 19 ]; C, buccal views of upper and lower P3's of Diprotodons; D, occlusal views of
upper and lower P3's, of Diprotodon; E, occlusal views of upper and lower P3's of Zygomaturus trilobus;
Pa, paracone; Me, metacone; Pr, protocone; Ps, parastyle; hy, hypocone]; F, G, M? of Diprotodon
and Zygomaturus for comparison. ( After Stirton ef al. 1967, Owen 1877).

1102 - MURRAY

Order Diprotodontia Owen 1866
Family Diprotodontidae Gill 1872
Subfamily Diprotodontinae Stirton, Woodburne and Plane 1967
Genus Diprotodon Owen 1838
Genus Nototherium Owen 1845

Diprotodons and Nototheres (Figs 18-22; Pl. 3). With the exception of Pyramios
alcootense (Woodburne 1967), diprotodontine marsupials are principally a Pliocene radiation.
Although Pyramios is a relatively plesiomorphic diprotodontine, Diprotodon appears to have
developed from a later Euryzygoma-like form. The genus is present in Late Pliocene Kanunka
and Fisherman's Cliff local faunas (Woodburne et al. 1985) and throughout the Pleistocene.
The systematics of Pleistocene Diprotodon species is at least as problematic as that of
Zygomaturus. The principal species is D. australis (Owen 1843) sometimes equated with D.
optatum (Owen 1838) (Fig. 22; Pl. 3). A smalier species was described by Huxley (1862) as
Diprotodon minor and yet another species as D. longiceps (McCoy 1876). Each one of these
designations is problematic, and the genus is crying out for a revision. Principally because
there are two sizes of diprotodons, I have used D. optatum and D. minor merely to facilitate a
description, as this is not the place to open a formal discussion of systematics.
Diprotodontines are distinguished from zygomaturines on the basis of their simpler P? crown
morphology, narrower mid-valley between the anterior and posterior moieties of the molars and
a tendency to have a marked obliquity of the crescentic cementum exposures of the lophs. The
P3 of Diprotodon when worn, forms a characteristic horseshoe-shaped ridge with the opening
on the labial side between the closely approximated paracone and metacone (Fig. 18). The
molars are also higher crowned, narrower relative to length and absolutely larger than those of
Zygomaturus (Fig. 19; Table 6).

Both species of Diprotodon preferred the drier, open expanses of the interior of Australia,
although D. minor is less common, Diprotodon sp. is present in all regional faunas except for
Tasmania (King Island excepted) and the extreme southwest (Keble 1945), Many Diprotodon
finds have been made in the old central lake basins, such as Lake Callabonna (Tedford 1973).
D. optatum, the largest marsupial known, stood 1.6-1.8 m at the shoulder and attained a length
of 2.75-3.4 m. They probably weighed from just under one, up to two metric tons.
Diprotodon had a long, narrow head, moderately long neck and a compact body with fairly long
proximal and short distal limb segments terminating in small, apparently heavily padded feet,
the carpals being characterized by a heel-like enlargement of the pisiform, The pes is also
highly modified and although somewhat wombat-like with a short, laterally expanded
metatarsal V, the greatly broadened plantar surface of the astragalus, the massive hook-like
calcaneus and enlarged, elongated navicular are very distinctive (Figs 21,22), Digits II-III are
syndactylous, and all digits are reduced. Diprotodons were graviportal animals in having their
weight transmitted to pillar-like limbs in the manner of elephants. The olecranon process of
the ulna is deflected outward to allow full extension of the forelimb as is characteristic of this
type of locomotory suspension. Diprotodon's nasal region is similar to that of Zygomaturus
in having a high, flange-like premaxillary septum, presumably for the attachment of powerful
levator labii muscles, and the nasals are somewhat retracted suggesting a high degree of facial
mobility in the region. A deep fossa is developed in the maxilla, presumably to accommodate
powerful maxillolabialis muscles. This muscular complex apparently complemented the
extremely long procumbant, spatulate upper and nearly horizontal, chisel-like lower incisors in
a powerful cropping mechanism (Archer & Bartholomai 1978).

Diprotodon minor was originally described from teeth found at Darling Downs (de Vis
1888). Subsequent finds come from a wide area of southeastern Australia. The dentary, upper

PLEISTOCENE MEGAFAUNA - 1103

Figure 19. Dentitions of Pleistocene “diprotodontoids": A, upper and lower cheek teeth of Diprotodon,
B, ibid. lower teeth in side view; C, upper and lower cheek teeth of Zygomaturus trilobus; D, buccal view
of upper and lower cheek teeth of Zygomaturus (note higher crowns and narrower mid-valleys in

Diprotodon);, E, upper and lower cheek teeth of Palorchestes azeal inset: unwom P3: note also worn,

obscured crown morphology of Zygomaturus p3: F, incisor of Palorchestes azeal [Me, metacone Pa,

paracone; Pr, protocone, Hid, hypolophid; Med, metaconid; prd, protoconid]. (After Owen 1877,
Woods 1958, Banks et al. 1976, Archer 1984).

1104 - MURRAY

incisors, assorted molars and isolated postcranial elements suggest that Diprotodon minor was
about one-third smaller than D. optatum.

Figure 20. Comparison of Pleistocene "nototheriine" and zygomaturine dentaries; A, Nototherium
inerme, B, undescribed "pygmy" zygmaturine from New Guinea; note lophid orientation. (After Woods 1968;
photo of zygomaturine, courtesy of Philip Walker).

The conspicuous and durable bones and teeth of Diprotodon make it the most likely
megafaunal species to occur in post-Pleistocene sediments, because they would tend to survive
reworking and because of the tendency for prehistoric aboriginal people to "souvenir" them.
This is not to deny the possibility that Diprotodon survived longer than any other megafaunal
species, but is mentioned as an alternative explanation as to why this genus, more often than
any other large extinct form, is sometimes associated with terminal Pleistocene and early

PLEISTOCENE MEGAFAUNA - 1105

Figure 21. Comparison of diprotodontid postcranial elements: A, scapula of Diprotodon optatum, B,
Zygomaturus trilobus (tasmanicum), C, humerus of Diprotodon optatum, D, Zygomaturus trilobus; E, femur
of Zygomaturus trilobus (tasmanicum) arrow points to "Owen's line"; F, Diprotodon optatum, G,
innominate of Diprotodon optatum, H, Zygomaturus, I - J, foot of Diprotodon [Na, navicular, V;
metatarsal I; En, entocuneiform; As, astragalus]. (After Owen 1877; Merrilees & Porter 1979; Archer

1984).

1106 - MURRAY

Holocene sediments. Diprotodon has also been "recognized" from time to time in aboriginal
rock art, the most recent discovery being from Laura, north Queensland.

Nototherium inerme (Owen 1845) is a poorly known, medium-sized Pleistocene notothere
also requiring revision. Judging from its dentaries, it was slightly smaller than Zygomaturus
trilobus. The dentary is less robust and the ascending ramus is lower and narrower than in
Zygomaturus. Also, the more closely approximated lophids are obliquely crescentic, the
symphysis is longer and narrower and the P3 is substantially smaller. Undoubted Nototherium
fossils appear to be confined to certain Queensland local faunas.

Order Diprotodontia Owen 1866
Family Palorchestidae (Tate 1948) Archer and Bartholomai 1978
Genus Palorchestes Owen 1873

Marsupial "Tapirs" (Figs 3, 19; Table 6). Palorchestine palorchestids are rare fossils
throughout their long Tertiary and Quaternary record. The earliest palorchestid is recorded from
the Late Oligocene Geilston Bay Local Fauna (Tedford et al. 1975). Palorchestines with an
apparently complete suite of Palorchestes autapomorphies are present in the medial Miocene
Bullock Creek Local Fauna in the form of lamb-sized Propalorchestes novaculacephalus
(Murray 1986). As is often the case with highly apomorphic forms, palorchestines retain
many equally plesiomorphic features in the cranial base suggesting a close relationship with
Ngapakaldia. Among these features are the retention of a large squamosal epitympanic
fenestra and a squamosal dorsal process of the tympanic cavity, but in Propalorchestes, it is
floored by a large ventral alisphenoid tympanic wing that may have been closed-off posteriorly,
in contrast to more derived diprotodontids and later palorchestids which have lost the
alisphenoid wing. In comparison with living diprotodontians, Propalorchestes resembles both
kangaroos and wombats about equally, depending upon how much weight the various isometric
distortions of the cranial base structures are given to the interpretation. This technical
indulgence is no mere digression, because on the basis of its dentition, Palorchestes azeal was
originally thought to be a giant kangaroo (Owen 1877, Tate 1948, Raven & Gregory 1948)
and even after Woods (1958) recognized the diprotodontid affinity of palorchestids, Archer was
willing to entertain the notion that palorchestids might have some special relationship with
macropodoids after all (Archer 1984). At present, they are snuggled among the Vombatiformes
(Aplin & Archer 1987).

A single species Palorchestes azeal (Owen 1873) is associated with the Late Pleistocene
megafauna. Perhaps, significantly, the two earlicr species known (P. painei, Late Miocene; P.
parvus, Early to mid-Pliocene) do not appear to overlap temporally and show a gradual trend of
increased body size and molar crown height and complexity through time. The dentition of P.
azeal is distinguished by the presence of well-developed fore and mid links on the molars and a
simple, 3-cusped upper third premolar (Fig. 19). The cranium is remarkable for its drastically
reduced and retracted nasals, which in conjunction with conspicuous fossae for large
nasomaxillolabialis muscles and narrow, protracted rostrum Iead to the conclusion that ithada
trunk at least as well developed as that of placental tapirs (Bartholomai 1978; Figs 1,3). The
dentaries are slender and kangaroo-like in having a sinuous profile to the inferior border. The
long, narrow deeply-grooved mandibular symphysis is associated with a slender protrusible
tongue like that of giraffes. Its broad, spatulate lower incisors lie almost parallel to its
strongly developed diastemal crests, indicating their probable use in stripping leaves or
cropping clumped vegetation. Details of its postcranial skeleton await publication. Archer
(1984) makes note of its powerful forelimbs equipped with large, laterally compressed claws.
What little is known of these large animals, which probably weighed 350-400 kg, suggests
that they may have been the approximate Australian community equivalents of certain
American giant ground sloths.

PLEISTOCENE MEGAFAUNA - 1107

The fossil remains of Palorchestes azeal date from >30,000 to >50,000 yBP. A good
association of P.azeal material with wood fragments at Pulbeena Swamp, northwestern
Tasmania yielded a 14C determination of about 54,000 yBP (Banks et al. 1976). Murray &
Chaloupka (1984) described a spectacular rock painting in the Arnhem Land escarpment region
that resembles some reconstructions of Palorchestes (P1. 1).

0 im
T— 1 of pf os) ST) ade f

Figure 22. Restorations of A, Diprotodon optatum and B, Zygomaturus trilobus.

Order Diprotodontia Owen 1866
Family Thylacoleonidae Gill 1872
Genus Thylacoleo Owen 1858

Marsupial "Lions" (Figs 23, 24; Pls 5, 6; Table 7). Marsupial lions were predaceous
and scavenging marsupials convergent with placental cats, analogous with and no less
remarkable than the convergence of the South American borhyaenid Thylacosmilus with the

1108 - MURRAY

Figure 23. Marsupial lion, Thylacoleo carnifex : A, drawing of a restoration by Rod Wells, Flinders
University; B, restoration of manus, note large, hooded pollical claw, C, lingual view of dental apparatus;
note reduced canines and premolars; hypertrophied, shearing P3 complex and robust incisors. (After Owen
1877, Wells & Nichol 1977).

PLEISTOCENE MEGAFAUNA - 1109

placental sabre-toothed "tigers". Thylacoleonids are represented in the Middle Miocene by two
genera, Priscileo (Rauscher 1987) and Wakaleo (Clemens & Plane 1974). Compared to the
leopard-sized Pleistocene Thylacoleo carnifex (Owen 1858), these are small forms ranging from
the size of a large possum (Priscileo) to kelpie or dingo-sized (Wakaleo). Both Priscileo and
Wakaleo show incipient enlargement of the sectorial third premolar to form a shearing complex
analogous to the placental carnassial set and reduction of the molar row. They also differ
sufficiently by the mid-Miocene to separate them into two distinct subfamilies, the
Thylacoleoninae and Wakaleoninae (Murray, Wells & Plane 1987). Although the general
trends of thylacoleonid dental evolution are well documented by these forms (Archer & Rich
1982), interpretations of the morphological details of the cranial base will ultimately determine
their relationship to other diprotodontians. This is important, because for many years T.
carnifex was considered to be a gigantic phalangeroid on the basis of its laniary incisors,
sectorial premolars, simple molars and other, phenetic similarities (Broom 1898). Details of
the cranial base of Thylacoleo say otherwise. Picking up from observations by Winge (1941)
and Woods (1958), Aplin recognized that Thylacoleo is decidely more wombat-like than
phalangeroid-like and accordingly has placed it among the Vombatiformes (Aplin & Archer
1987). Consequently, the tabloid headlines will now read "giant killer wombat” instead of
"giant killer possum". The striking similarity of the cranial base of Thylacoleo to Vombatus
and the additional synapomorphic feature of frontal-squamosal contact almost closes the case.
Almost, because the very phalangerid-like W. vanderleueri possesses a well-developed
alisphenoid ventral tympanic process and lacks frontal-squamosal contact on the cranial wall.
Thus, at least some of the major potential vombatimorph synapomorphies may merely be
convergences (Murray, Wells & Plane 1987), and it seems that the relationship between the
wakaleonines and the thylacoleonines is a distant one.

The extreme variability in body size of Thylacoleo carnifex (Archer & Dawson 1982)
makes it difficult to determine a typical weight for the species. The New England specimen,
about 1.25 m long from nasals to sacrum, produces a calculated weight of 68 kg. Some large
crania indicate that live weights of >100 kg were likely. The major morphological and
adaptive features of Thylacoleo carnifex are well known (Woods 1956, Wells & Nichol 1977,
Horton & Wright 1981, Finch 1982, Finch & Freedman 1982, Wells et al. 1982). T. carnifex
was a powerfully built animal with generalized, if not slightly long limb proportions, not far
removed, in fact, from a placental lion or leopard (Fig. 23; Pl. 6; Table 8). The pollex was
equipped with a large, hooded claw and stout, curved claws were present on the remaining
manual digits, proportioned like a phalangerid. There is no indication that they were
digitigrade like felids. The opposable, large-clawed pollex and their apparent association with
woodland habitats has led to the suggestion that they were semi-arboreal (Pledge 1977).
Although there is general agreement that Thylacoleo was carnivorous, its capability as a
predator has been cast in doubt, most recently by Horton & Wright (1981), who portray it as a
scavenger. This conclusion seems unlikely in a community with so few other predators upon
which a large mammalian scavenger must depend for regular kills. A large and powerful beast
equipped with a beak-like piercing incisor complex, Thylacoleo was probably an efficient and
formidable predator.

Thylacoleo carnifex has widespread occurrence in Late Pleistocene deposits in Australia and
Tasmania, but is unknown from New Guinea. It was among the longer-surviving Pleistocene
megafauna, occurring in assemblages such as that at Lanceficld, Victoria. Murray &
Chaloupka (1984) illustrated some Thylacoleo-like rock paintings from the Arnhem Land
escarpment region of the Northern Territory.

Figure 24. Restoration A, and postcranial skeleton of Thylacoleo carnifex : B, scapula; C, humerus; D,
radius; E, ulna; F, innominate; G, femur; H, fibula; I, tibia. (After Finch 1971; Wells & Pledge 1983;
measurements courtesy of Dirk Megirian, Northem Territory Museum and R. Wells Flinders University).

1110 - MURRAY

PLEISTOCENE MEGAFAUNA - I111

Order Diprotodontia Owen 1866
Family Potoroidae (Gray 1821) Pearson 1950
Subfamily Hypsiprymnodontinae Pearson 1950 or
Subfamily Propleopinae Archer and Flannery 1985
Genus Propleopus De Vis 1888

Giant Rat-Kangaroos (Figs 25-27 Table 8). Omnivorous and perhaps carnivorous rat-
kangaroos have been present since the mid-Miocene (Archer 1984). If for no other reason, they
are potentially important to palaeoecologists as possible members of the apparently
depauperate large predatory carnivore guild in Australian megafaunal communities. The notion
of predatory potoroid kangaroos was created by both Archer (1984) and Flannery (1984) on
reflection of the habits of living potoroids and their possession of large, serrated shearing
premolars, and I can think of no reasonable argument to oppose the idea. Propleopus oscillans
(de Vis 1888) was about the size of an adult male living Grey Kangaroo (Macropus giganteus);
a fair estimate of its live weight is between 40 and 60 kg. Propleopus are rare fossils
compared to other large kangaroo species, which lends Eltonian support to the likelihood of
their carnivory. For this reason, they are incompletely known, especially postcranially.
Pledge (1981) described a right humerus, and there is an undescribed nearly complete cranium
from Victoria Cave (Rod Wells, pers. comm.) (Fig. 25). Otherwise, the species is recognized
by its large, distinctive serrated third premolars and its low-crowned, simple bunolophodont
molars. A second species, P. chillagoensis (Archer & Bartholomai 1978) is of doubtful
Pleistocene age. Another, definitely Pleistocene species from Wellington Caves, New South
Wales awaits formal description.

Potoroid kangaroos differ from macropodid kangaroos in a number of important features,
among these are the large, extensive masseteric canal and squamosal-frontal contact on the
lateral braincase wall. Propleopines differ from their diminutive hypsiprymnodontine relatives
in having a larger and more anteriorly positioned metaconid on M2 and in having separate
anterior ends of the masseteric and dental canals. Archer & Flannery's (1985) diagnosis of this
subfamily division is probably based on structures under allometric control.

So little is known about the postcranial skeleton of Propleopus that we can only speculate
on its locomotor behaviour by way of its close relationship to the living musk-rat kangaroo,
Hypsiprymnodon moschatus. Hypsiprymnodon is the most primitive of the living kangaroos
in retaining the hallux, although it also expresses the major macropodoid synapomorphies of
the foot, which include a stepped articulation of the calcaneo-cuboid joint and elaborations of
the tibia-astragalar and astragalo-calcaneal joints in relation to bounding locomotion.
Potoroids are typically quadrupedal bounders, but are capable of short bursts of bipedal saltation
(Buchmann & Guiler 1974).

Order Diprotodontia Owen 1866
Family Macropodidae Gray 1821

Subfamily Sthenurinae Glauert 1926
Genus Sthenurus Owen 1838
Genus Procoptodon Owen 1845
Genus Troposodon Campbell 1973

Subfamily Macropodinae (Pearson 1950)
Genus Bohra Flannery and Szalay 1982
Genus Protemnodon Owen 1874
Genus Wallabia Desmarest 1803
Genus Macropus Shaw 1789

1112 - MURRAY

Kangaroos and Wallabies (Figs 25-35; Pls 7, 8; Tables 9-14). The identification of
Late Pleistocene kangaroos has become a specialist field because of the diversity and phyletic
complexity of the group. Stirton (1963), Tedford (1966), Bartholomai (1972,1973, 1975) and
Flannery (1984) have clarified many aspects of macropodid relationships that have been in a
confused state since the time of Owen's monograph (1877) and the pioneering work of de Vis
(1888). Perhaps the only advantage in writing a chapter devoted to the large extinct
Pleistocene marsupials is that the established macropodid subfamilies are not quite as difficult
to define synchronously as they are working back through time, and the systematically
troublesome smaller extant kangaroo species can be ignored. The macropodid kangaroos
represent the largest radiation of marsupials still in existence, even though the diversity of the
family was greatly diminished by the beginning of the Holocene. From the genus Macropus
alone, at least eight species became extinct by about 15,000 years ago. M. titan and M.
ferragus were the giants among the "true" (macropodine) kangaroos (Table 9). M. ferragus, a
giant Euro, may have stood 2.5 m tall and weighed over 150 kg. M. titan, an almost equally
large Late Pleistocene kangaroo, was closely related to the living Grey Kangaroo (M.
giganteus) (Tedford 1967). Marshall & Coruccini (1978) concluded that many of these "giant"
forms represent large morphs of "dwarfed" extant species. M. siva, for example, appears to be
a large racial form of the living M. agilis and several, perhaps synonomous species (M.
cooperi and M. altus) may represent larger Pleistocene conspecifics of M. (Osphranter)
robustus (Marshall 1973, Main 1978). Bartholomai (1973) remarked that M. birdselli differs
from M. titan only in the length of the diastema, a highly variable character. Obviously there
is some synonomy due primarily to the fragmentary nature of the material. A large number of
Pleistocene species of Macropus are found only in the Darling Downs and Wellington caves
local faunas, including M. rama (similar in size and morphology to the Grey Kangaroo) and M.
gouldi, (Figs 15-16 of Pl. 23 in Owen 1874; the type has been lost), the distinctive M.
piltonensis with unique dental characters and M. thor, which closely resembles the living M.
parryi (Bartholomai 1975). A "giant" tree kangaroo, Bohra paulae, identified from postcranial
elements in Wellington Caves deposits, may have weighed about 50 kg (Flannery & Szalay
1982). The more common and, therefore, important Late Pleistocene macropodines (Macropus
spp.) can be subdivided into three subgenera: 1) the Osphranter group (Reds and Euros) having
an inflated rostrum, narrow dentaries with a conspicuous subalveolar excavation, a posterior
groove in the lower molars, upper molars with a weak forelink and a U-shaped occlusal profile
of the upper incisors; 2) the Prionotemnus group, (e.g. Rufous and Toolache wallabies) having
a narrow rostrum and dentaries deepened below the anterior cheek tooth row and 3) the
Macropus group (Grey kangaroos) with the upper incisors forming a V-shaped occlusal outline
and strong forelinks on the upper molars (Bartholomai 1973).

The genus Protemnodon ("giant wallabies") (Table 10) contains at least three Australian and
one New Guinea species (Table 11). P. anak was about the size of a Grey kangaroo; the New
Guinea species P. otibandus was slightly smaller. Protemnodon brehus and P. roechus were
usually much larger, but considerable size overlap occurs among all known Protemnodon
species, and it is an untrustworthy criterion for separating them. At least some of the
Protemnodon species were compactly built kangaroos with powerful upper limbs, short, broad
feet and elongated skulls lacking the characteristic downgrowth of the snout found in modern
Macropus species (Bartholomai 1973). Postcranial elements collected by Merrilees (1973)
from Lake Tandou indicate that although the cranial and body length of P. brehus equals that of
Macropus fuliginosus, its femur and tibia are both shorter. Wallabia bicolor, the living

Figure 25. Pleistocene macropodoid dentaries: A, Propleopus oscillans; B, Protemnodon anak, C, Sthenurus
andersoni; D, Sthenurus atlas; E, Macropus coopert; F, Macropus ferragus; G, Macropus titan; H, Troposodon
minor; I, Sthenurus oreas; J, Sthenurus pales; K, Procoptodon pusio; L, Macropus birdselli. (After Owen 1877,
Tedford 1967, Bartholomai 1973, Marshall 1973),

PLEISTOCENE MEGAFAUNA - 1113

1114 - MURRAY

Swamp Wallaby has been tentatively linked to this group, although serological evidence and
the fact that they will hybridize with M. agilis weighs heavily against this (Bartholomai 1975,
Archer 1984).

The Sthenurinae (Tables 11-13) comprise a large subfamily of entirely extinct, browsing
kangaroos, characterized by short, deep skulls and extreme reduction of metatarsal V to a small
splint (Tables 12-15). These are medium to large kangaroos that can be further divided into
two distinct subgenera: a short-faced (brachycephalic) group Sthenurus (Simosthenurus) and a
long-faced (dolichocephalic) S. (Sthenurus) group. They can also be divided into long-footed
and short-footed forms with equally satisfying results (Bartholomai 1963; Merrilees 1965a,
1968a, Tedford 1966, Wells & Murray 1979). Pledge (1980a) recommends that the subgenera
be given full generic rank. Sthenurines have highly distinctive molar teeth characterized by
low crowns, sharp, straight lophs and poorly developed midlinks. The premolars, in contrast
to the long, blade-like sectorial pair in Protemnodon, are typically broad and worn flat on the
occlusal surface. Sanson (1976) considers the dolichocephalic sthenurines to represent a
parallel trend with macropodines toward grazing.

Procoptodon goliah (P\. 7; Table 14) was probably the largest kangaroo species, standing
over 2,5 m tall and weighing perhaps, 200 kg. The size of individuals from different localities
varies on a Cline from west (Lake Menindee, where they are largest) to east (Darling Downs
where they are smallest) (Marcus 1976). Currently placed among the sthenurines on the basis
of numerous synapomorphies of the feet and cranium, Sanson (1976) argues that these may be
parallel features and places them within the Macropodinae. He also considers them to be
grazers rather than browsers. The massive, squarish crania of Procoptodon spp., having
distinctly straight, parallel molar rows and peg-like, reduced upper incisors, are matched to
short, deep dentaries with high ascending rami, short diastems, elaborate, broad, high-crowned
molars, crushing third premolars and small, lanceolate-crowned incisors. P. goliah had long
forelimbs that may have been used to pull down branches to reach the foliage (Tedford 1967).
Alternatively, they may reflect an habitually quadrupedal locomotor pattern associated with
grazing. There are currently four Pleistocene Procoptodon species, P. goliah, P. rapha, P.
pusio and P. texasensis, given in order of descending body size and complexity of dentition
(Owen 1874, Tedford 1967, Marcus 1976, McIntyre & Hope 1978, Sanson et al. 1980). An
uncommon Pleistocene genus, Troposodon, is considered to be a sthenurine on the basis of its
molar morphology (Flannery 1984).

A comprehensive element by element key to the identification of each Pleistocene fossil
kangaroo species is not possible within the confines of this chapter. This chapter, instead,
best serves the interest of the reader as a guide to where and equally important, how to look for
the answer. Essential works on Pleistocene kangaroos are Stirton (1963), Bartholomai (1973,
1975), Tedford (1966, 1967) and Marcus (1976), Macropodid skeletal material is highly
distinctive and every element bears the stamp of the specialized locomotor adaptations unique
to the group. However, the fossil record is selective, and certain elements are more often
preserved than others. Of these elements, certain structures are more informative than others.
The commonest and most informative kangaroo skeletal structures are their teeth and the
cuboid, calcaneus and metatarsal IV elements of the foot. Given the above limitations, the
following discussion concentrates on these.

Figure 26. Pleistocene macropodid appendicular elements: A, humerus of Macropus sp.; B, Sthenurus sp.;
C, Procoptodon goliah, D, Propleopus oscillans; E, femur of Macropus titan; F, Simosthenurus sp.; G,
Procoptodon goliah; H, ulna of Protemnodon otibandus; I, Simosthenurus sp.; J, Procoptodon goliah; K,
tibia of Macropus titan; L, Protemnodon; M, Simosthenurus; N, Procoptodon goliah. (After Tedford 1967,
Plane 1967, Merrilees & Porter 1979, Pledge 1980a, 1981).

PLEISTOCENE MEGAFAUNA - 1115

1116 - MURRAY

Macropus molars are long, narrow and high-crowned with thick enamel and well-developed
midlinks, The molars erupt successively and drift mesially throughout the lifetime of an
individual. The third premolars are sectorial, usually possessing a posterolingual cuspule or
thickening and the cheek-tooth rows are distinctly arched. In addition to being larger than M.
titan, the P3 of M. ferragus has a detached or angled posterior lobe and the previously
mentioned Osphranter characters. M. cooperi was smaller than M. ferragus but larger than the
living Euro, and is further distinguished from M. robustus by its larger P? and stronger
forelinks on the upper molars and from M. ferragus by its lower molar crowns and straighter
disposition of the longitudinal crest of the P3 (Bartholomai 1975).

The metatarsal IV's of both M. titan and M. ferragus are enormous compared to those of M.
giganteus or M. robustus. Very large, typically Macropus metatarsal IV's are almost certain to
belong to one or the other of these species. The metatarsal IV of M. ferragus is similar to M.
fuliginosus proximally and not as elongated anteroposteriorly as M. giganteus. The proximal
plantar surface is relatively larger than in any living species, and the plantar crest is relatively
stronger than in any species of Macropus, living or fossil (Tedford 1967).

Large Protemnodon species are distinguished from large Macropus species by their long,
robust, deeply rooted blade-like permanent premolars, absence of a forelink on the upper molars
and the presence of a strong anterior ridge from the paracone to the anterior cingulum. The
immediate impression given by Protemnodon dentitions is a greater symmetry of the lophs and
lophids, especially when worn, and the smooth, broad concave surfaces of the backs of the
lophs and the front of the lophids, a feature related to the less steep ascent of the lophs or
lophids from the median valley. The cheek tooth rows are straight rather than arched, as in
Macropus, with less anteromedial torsion of the horizontal ramus. In general, there is less
curvature of the occlusal surfaces of the tooth rows and the wear gradient is less "twisted."
Protemnodon spp. lower incisors are broadly U-shaped, approaching in form those of
Palorchestes, with which this genus has been confused in the past (Woods 1958).
Protemnodon anak differs from P. brehus in its smaller size and narrower P? with a more
concave labial surface. The P3 of P. brehus is more flexed than in that P. anak, and the lower
molars have a less well-developed posterior cingulum (Bartholomai 1973). Protemnodon
roechus differs from the other two species in lacking prominent vertical ridges on the lower
permanent premolar, and the upper is crescent-shaped in occlusal view with oblique, rather than
vertical transecting ridges. The crown bases of the lower molars are swollen, and the upper
molars usually have a small tubercle at the lingual extremity of the median valley.

The construction and orientation of the calcaneum and astragalus of Protemnodon are
basically similar to Macropus, although relatively more massive (Stirton 1963, Tedford 1967).
The astragalus is distinguished from Macropus by its massive head and relatively narrower
width across the trochlea. The calcaneum is relatively shorter and broader than in Macropus,
and both the peroneal tubercle and the sustentaculum tali are more prominent and broader. The
cuboid is very similar to Macropus and should be distinguished by specimen comparison. The
metatarsal IV is extremely short and broad with a posteriorly elongated plantar facet and
relatively larger facets for the fifth metatarsal. The plantar crest is sharper than in Macropus,
and the second and third metatarsal grooves are more deeply excavated.

The Sthenurinae are distinguished from other macropodids by their often omamented, low
molar crowns with nearly straight, thinly enamelled lophs, low, weak midlinks, short, deep
dentaries, nearly straight tooth rows and broad, swollen permanent premolars. The dolicho-
cephalic sthenurines usually have more distinct, higher midlinks and less ornamentation

Figure 27. Occlusal views of cheek dentitions of macropodids (upper below): A, Propleopus oscillans; B,
Troposodon minor, C, Protemnodon anak, D, Protemnodon brehus; D, Protemnodon roechus; F, Macropus
ti.an; G, Osphranter cooperi; H, Macropus ferragus: K, Procoptodon pusio. (After Tedford 1967, Bartholomai
1973, Marshall 1973, Bartholomai 1975).

PLEISTOCENE MEGAFAUNA - 1117

aD

&
BUDE

1118 - MURRAY

Figure 28. Restored macropodid skulls: A, Protemnodon anak; B, Protemnodon roechus; C, Macropus
titan; D, Sthenurus gilli; E, Protemnodon anak, F, Macropus titan; G, Procoptodon goliah; H,

Procoptodon rapha. D-F, occlusal view of lower incisors and symphyseal morphology. (After Tedford 1967,
Bartholomai 1973 in Owen).

of the molar crowns than the brachycephalic species. The short-faced Simosthenurus subgenus
is Characterized by a "macrodont" and a "microdont" species complex. The macrodont complex
is composed of three dentally similar species §. occidentalis, S. orientalis and S. brownei. If
entire dentaries are present, S. brownei can be distinguished from the other two macrodont
species by its much deeper horizontal ramus posterior to M4, The lower permanent premolar
of S. brownei is shorter and has finer crenulations than S. occidentalis (Merrilees 1968a). S.
vrientalis differs from S. occidentalis in its larger size, extension of the mandibular symphysis
posterior to P3 and its larger P3 relative to the molars (Tedford 1966). The crania of these

PLEISTOCENE MEGAFAUNA - 1119

species are extremely broad across the zygomatic arches and have remarkably short, narrow
rostra, although S. brownei has a much broader narial aperture than the others.

The microdont Simosthenurus species are S. gilli and S. maddocki. S. maddocki is similar
in mandibular form and dentition to S. gilli, but it appears to have been a much larger and very
different animal. The permanent lower premolar of §. maddocki is much narrower relative to
length than S. gilli, and the dentary is readily identified by its possession of an exceptionally
long, deep and slightly obliquely oriented subalveolar sulcus (buccinator sulcus) that
commences at the base of P3 and extends posteriorly to between Mz and M3_ The lower
incisors are narrow as in S. gilli but more procumbant. S. gilli is the smallest of the
sthenurine kangaroos. It can be distinguished from S$. maddocki by its sparser and coarser
molar ornamentation and its relatively longer, narrower cranium in addition to the previously
mentioned characters (Wells & Murray 1979),

Figure 29. Comparison of sthenurine crania: A, brachycephalic sthenurine Sthenurus maddocki; B,
compared with dolichocephalic sthenurine, Sthenurus atlas. (After Pledge 1980b, Pledge 1989a).

Virtually complete skeletons of Simosthenurus have been recovered by Neville Pledge and a
cave diving team from a drowned cave near Tantanoola, South Australia (Pledge 1980a).
These unique specimens show that there are many important differences between sthenurine and
macropodine postcranial skeletons. Sthenurines have more robust femora with a relatively

1120 - MURRAY

greater diameter and stronger curvature of the diaphysis. The tibiae are also robust and
relatively shorter than in Macropus spp. The scapulae are shorter and much broader, with a
stouter, longer acromial process and relative reduction of the supraspinatus fossa. The
innominates differ from Macropus and Protemnodon in having broader ilia, a short pubic ramus
and a more acute upward flexion of the ischium. These features indicate that there were some
fundamental locomotor and postural differences between sthenurines and macropodines.

D

Figure 30. Outline drawings of the lateral aspect of skulls of brachycephalic sthenurines (Simosthenurus):
A, Sthenurus occidentalis; B, Sthenurus maddocki; C, Sthenurus orientalis; D, Sthenurus gilli, E, Sthenurus
brownei. Nasal aperture shapes compared. F, Sthenurus occidentalis, G, Sthenurus brownei; H, Sthenurus
maddocki. Symphyseal outlines of dentaries compared: I, Sthenurus occidentalis, J, Sthenurus maddocki;
K, Sthenurus gilli. (After Pledge 1980a).

PLEISTOCENE MEGAFAUNA - 1121

B 0 50cm
Sen |-eepemnens |

A, Sthenurus occidentalis, B, Procoptodon

Figure 31. Restorations of short-faced browsing kangaroos:
goliah.

1122 - MURRAY

alk
DS

Figure 32. Occlusal views of upper and lower check dentitions of Sthenurine kangaroos (uppers below):
A, Sthenurus brownei; B, Sthenurus oreas, C, Sthenurus gilli; D, Sthenurus maddocki; E, Sthenurus pales,
F, Sthenurus occidentalis, G, Sthenurus orientalis, H, Sthenurus atlas; I, Sthenurus andersoni; J, Sthenurus
tindalei. (After Merrilees 1965, Tedford 1966, Merrilees 1967, Wells & Murray 1979).

PLEISTOCENE MEGAFAUNA - 1123

.

-

Figure 33. Pleistocene macropodoid limb girdle elements: A, scapula of Macropus sp.; B, Sthenurus
occidentalis; C, Procoptodon goliah; D, innominate of Macropus sp.; E, Sthenurus sp.; F, Procoptodon
goliah. (After Tedford 1967, Merrilees & Porter 1979, Pledge 1980).

Two other, short-faced sthenurines include the gigantic S. pales and the S. occidentalis-
sized S. oreas. Sthenurus pales is conspicuously larger than any other Simosthenurus species,
and is also distinguished by possessing higher crowned cheek teeth, particularly the p3- Due to
its large size, it may become confused with the larger long-faced sthenurines, but it can be
readily differentiated from them on the basis of dental morphology. The plesiomorphic
Sthenurus oreas was about the same size as S. occidentalis, distinguished by its shallower
dentary and narrower and smaller P3 relative to the molars, in conjunction with the simplicity
and coarseness of the ornamentation. It shares, with long-faced sthenurines, relatively high
molar crowns, stronger links and coarse, simple ornamentation that is considered structurally
intermediate between the Sthenurus subgenera and perhaps the genus Procoptodon (Tedford

B, Macropus titan;
[Ca, calcaneus;

cuneiform and entocuneiform, IV, metatarsal IV].

1124 - MURRAY

1966). In some respects it resembles the smallest of the long-faced forms Sthenurus atlas, but
no more so, perhaps, than Simosthenurus maddocki; hence some caution in raising their
subgeneric status is justified. The named species of long-faced sthenurines overlap in morph-

Figure 34. Foot structure (restored) of Pliocene and Pleistocene macropodids: A, Protemnodon otibandus;
C, Macropus ospranter sp.; D, Simosthenurus sp.; E, Procoptodon goliah; F,

As, astragulus; V, metatarsal V; Me, mesectocuneiform = fused ectocuneiform, meso-
(Plane 1967, Tedford 1967, Marshall 1973, Owen 1977).

PLEISTOCENE MEGAFAUNA - 1125

ology and size. Sthenurus tindalei and an even larger unnamed species (R. Wells, pers.
comm.) are the largest sthenurines, rivalling Procoptodon goliah and Macropus ferragus
(Tedford 1966). Similar to S. atlas is S. andersoni, which has higher crowned upper molars

and a narrow, more elongated P3_ Because S. tindalei is unlikely to be confused with any form
other than S. pales, it can be distinguished from the latter by its slightly smaller, lower
eee P3 that bears a continuous lingual crest around the anterior end of the tooth (Tedford

The genus Procoptodon is the morphological extreme of the brachycephalic sthenurines in
having massive, short, broad, high skulls reminiscent of the now extinct New World
glyptodonts. Procoptodon material is instantly recognizable from its massiveness, its
distinctive, large lophodont molars with high complex links, its short, broad premolars and its
diminutive incisors. Procoptodon rapha and P. pusio separate out reasonably well on standard
molar crown dimensions (Marcus 1976; Table 14). P. pusio and P. texasensis have the
simplest molar crown morphologies and smallest dimensions of the Pleistocene genera. The
yak me Procoptodon texasensis is the most plesiomorphic species of the genus (Archer

Troposodon is a rare Pleistocene form that survives as a remnant of the Pliocene kangaroo
radiations. It may be related to the sthenurines (Flannery 1984), and the two species of
Pleistocene Troposodon (T. kenti and T. minor) are, thus, appropriately mentioned in this
context. These are medium-sized kangaroos with strongly tapered dentaries, low-crowned
molars and low, simple blade-like permanent premolars. The upper premolars are distinctively
L-shaped due to the off-set of a prominent distolingual cuspule.

The pes elements of the sthenurines are as diagnostic of the group as their dentition. The
calcaneum, astagalus, cuboid and metarsal IV of Procoptodon and Sthenurus (Simosthenurus)
are very similar and readily distinguished from those of large Macropus. The calcaneum is
proportionally broader, and the tuber calcaneus wider than in Macropus. The plantar surface
tapers more rapidly posteriorly, becoming continuous with the posterior margin of the cuboid
facet. The sustentaculum tali extends further anteriorly and is broader than in Macropus, the
tarsal groove describing a deep, narrow sulcus along its base. Compared to Macropus, the
cuboid facet is more expanded distally, and the anterior portion of the facet is not as strongly
stepped. The anterior calcaneal facets of the cuboid reflect the shallowly-stepped articulation;
the astragalar facet is relatively larger, and the plantar process is not set off laterally by a deep
groove as seen in typical Macropus (Tedford 1967). The metatarsal IV is relatively short and
broad, although less compact than that of Protemnodon. The distal articulation is wide, with a
low keel restricted to the ventral surface. The dorsal surface is somewhat flattened distally,
deepening proximally to a sub-triangular section. The plantar surface is large and bears a
conspicuous facet for the plantar sesamoid. A deep, short, roughend plantar crest extends about
one-third of the length of the bone. The sulci for metatarsals II and III are small and ventrally
disposed. The fifth metatarsal facet is short and tapers distally, often retaining a remnant of the
splint-like vestige that is partially ankylosed to the shaft of the fourth. Interestingly, the
dolichocephalic sthenurines differ from the brachycephalic forms in being more Macropus-like
in precisely those characters that distinguish the group as a subfamily (Tedford 1966).

PLEISTOCENE FAUNAS

The Australian Pleistocene regional faunas closely correspond with the modern
zoogeographic sub-regions, presumably because the major drainage systems and precipitation
regimes have not drastically changed since the late Tertiary. While it is tempting to redefine
these regions by superimposing palacontologist's names over the existing drainage pattern

1126 - MURRAY

nomenclature, I have mercifully opted instead for a bland geographic nomenclature (Figs 35,
36; Table 15),

PLEISTOCENE MODERN (Whitley 1959)
1) Northern Leichardtian

2) Northeastern Jardinean/Krefftian

3) Eastern Lessonian

4) Southeastern 2 =

5) Tasmanian Tobinian

6) Central Mitchellian/Sturtian

7) Southwestern Vilaminghian/Greyian

IRE ag sag

P-RORR aR

10 11

”

$444 —

47 49

Figure 35. Australian Late Pleistocene megafauna reconstructions; Key: 1, Palorchestes azeal: 2,
Zygomaturus trilobus; 3, Diprotodon optatum,; 4,D. minor; 5, Nototherlum inerme,; 6, Thylacoleo
carnifex; 7, Phascolonus gigus; 8, Ramsayia curvirostris; 9, Phascolomys magnus; 10, P. medius; 11,

Vombatus hacketti; 12, Phascolarctos cinereus/stirtoni, 13, Propleopus oscillans; 14, Procoptodon goliah;
15, P. rapha; 16, P. pusio; 17, Sthenurus brownei, 18, S. maddock; 19, S. occidentalis; 20, S. orientalis;

21, S. gilli; 22, S. atlas; 23,8. tindalei; 24, S. pales; 25, S. oreas; 26, S. andersoni; 271, Troposodon

minor, 28, Wallabia indra; 29, Protemnodon brehus; 30, P. anak, 31, P. roechus; 32, Macropus ferragus;

33, M. titan; 34,M. siva; 35, M. cf. Siganteus, 36,M. rama; 37,M. thor; 38, M. piltonensis; 39, M.

eouldi: 40, M. stirtoni; 41, Bohra paulae; 42, Sarcophilus spp.; 43, Zaglossus hacketti; 44, Z. ramsayi;

45, Aboriginal man for scale; 46, Flamingo; 47, Progura naracoortensis; 48, P. gallinacea; 49,

dromomithid: 50, Megalania prisca; 51, Wanambi naracoortensis; 52, Meiolania spp.

PLEISTOCENE MEGAFAUNA - 1127

Each regional fauna is characterized by the presence or absence of a suite of megafauna
species.

The Northern regional fauna is poorly known, but appears to be somewhat depauperate, as
the modern monsoon-effected faunas are today. It is known from poorly sampled local faunas
at Riversleigh, Queensland; Victoria R., Northern Territory; and the Kimberleys, Western
Australia. The habitat was probably monsoonal savanna-woodland as it is today.
Protemnodon brehus, Zygomaturus trilobus and Diprotodon sp. have been identified as well as
a small Tasmanian devil morph.

ROP

NORTHEASTERN

>

ad h
Tacha

CENTRAL S
Rais =“

EASTERN

a

A_

wd

SOUTHWESTERN

YALL TASMANIAN

Figure 36. Regional Late Pleistocene faunas; see Fig. 35 for key to species.

The Northeastern regional fauna was the most species-rich. The large number of local
faunas sample a wider time-range than other regions, except perhaps, for the Central region.
The habitat was principally open grassland with patchy scrub and riparian woodland. This
regional fauna contains all the known species of Procoptodon and many species of Macropus
not recorded elsewhere (Longman 1924, Bartholomai 1975).

The Eastern regional fauna is the most widespread and, therefore, the most typical
Pleistocene community. It was continuous with a broadly ecotonal and geographically
transitional Southeastern regional fauna, Tasmanian regional fauna and Southwestern regional
fauna. This was a complex ecotone of shrubby sclerophyll forest, savanna woodland, dry

1128 - MURRAY

grassland and heath occurring locally as vegetational mosaics in relation to a complex
topography (Frank 1972, Marcus 1976, Sanson et al. 1980. The Tasmanian and Southwestern
regions can be viewed as representing isolated extremes of the Eastern biome. The eastern
fauna contains fewer species of Procoptodon and Macropus but more species of Sthenurus
(Simosthenurus). Although Phascolonus gigas is not present, other vombatids (Phascolomys
spp., Ramsayia spp.) are well represented. The Southeastern regional fauna is distinguished by
the presence of Sthenurus maddocki and Sthenurus gilli, which are rare or absent in the eastern
region. It also contains Sthenurus brownei, a species also common to the Southwestern
regional fauna, but absent from Tasmania, Victoria and New South Wales (Gill 1957,
Merrilees 1965, 1968, Flood 1973, Erry & Flannery 1978, Wells et al. 1984). The Tasmanian
regional fauna is distinguished by its lack of many common genera and species, among them
Diprotodon (known however, from King island, Bass Strait) Propleopus, Phascolonus spp.,
Nototherium and Procoptodon. A single specimen of S. gilli was identified from the
Montague Local Fauna, otherwise dominated by the ubiquitous S. occidentalis. On the other
hand, the Tasmanian megafauna shows no obvious signs of morphological endemicity, in
contrast to the Southwestern region. Eastern Tasmania was predominantly lightly wooded
grassland and western Tasmania was predominantly grassland in the north grading to forested
mosaic grasslands to the south (Colhoun 1975, Colhoun et al. 1977, 1979, Goede et al. 1978,
Goede & Murray 1979, Murray et al. 1980).

The Southwestern regional fauna contains several endemics, including a large tachyglossid
and a wombat. Its sthenurine fauna is restricted to two species, and many other macropodines
characteristic of the adjacent faunas are absent. The region was clearly isolated to some extent
during the Late Pleistocene, presumably by arid land to the north and the Nullarbor Plain to the
east. A complex late Quaternary ecological succession has been documented for the Cape
Naturaliste region (Dortch & Merrilees 1973, Balme et al. 1978, Merrilees 1979, Porter 1979).
The Pleistocene faunal material from local caves Suggests that a woodland-heath-forest
association was present throughout the Pleistocene. The shrub communities were replaced by
forest about 20,000 years ago.

The Central region exemplifies an extensive semiarid region characterized by open woodland
with light mulga scrub and grass cover, with riparian woodland communities along perennial
watercourses and shallow lakes, transitional to monsoonal savanna woodland to the north and
grading into open shortgrass plains and saltbush steppe to the northwest and northeast. This is
an enormous region comprised of large and small lakes and the western Murray-Darling
drainage in South Australia, New South Wales and Victoria. The region contained many
characteristic open country species which include giant Osphranter spp. large dolichocephalic
sthenurines (S. tindalei), Procoptodon goliah, Phascolonus gigas and Diprotodon spp.

With the exception of the Tasmanian and Southwestern regions, a number of non-
mammalian megafauna are important members of these Late Pleistocene communities. The
Northeastern and Eastern regional faunas contain the Giant monitor, Megalania prisca, Horned
turtles, Miolania spp. The large python, Wonambi naracoortensis, and the dromornithid,
Genyornis newtoni, are represented in the Central and Southeastern regional faunas.

PALAEOBIOLOGY

DEFINING A MEGAFAUNA COMMUNITY

Ideally, complete, individual communities based on information from local faunas should
form the basis of such an investigation (cf. Tedford 1967, Balme et al. 1978). For the

PLEISTOCENE MEGAFAUNA - 1129

purposes of this introduction, a synthetic community, based on the large vertebrates common
to most regional faunas must suffice, thus reflecting the introductory notes of this chapter in
which a "typical" Australian megafaunal community was constructed. This brief and greatly
oversimplified modelling exercise attempts to synthesize inferences from body weight
estimations, functional anatomy, systematics and behaviour into a construct against which the
various causal factors of Pleistocene extinction might be examined.

Figure 37. Scaled skeletal restorations of Australian megafauna to provide a visual impression of
probable weight relationships. The human skeleton represents 177 cm stature and body mass of 80 kg.
Skeletal robusticity is correlated with increased body mass on the order of Y = 0.061 x 1.09 (Schmidt-
Neilsen 1984): A, Megalania prisca redrawn with slight modifications from T. Rich & B. Hall, (Aust. Nat.
Hist. 19, 1979; B, Simosthenurus occidentalis drawn from a photograph in R. Wells & N. Pledge Chap.
16: Vertebrate Fossils, Natural History of the Southeast, R. Soc. So. Aust., 1983; C, Phascolonus gigas,

restoration based on proportions in E. Stirling, Mem. R. Soc. So. Aust.1 (4) 1913; D, Zygomaturus sp.,

drawn from a photograph of specimen in the Tasmanian Museum, Hobart; E, Procoptodon goliah restoration
based on proportions in R. Tedford, Univ. Calif. Publs. geol. 64, 1967; F, Genyornis newtoni after P. V.
Rich & E. M. Thompson 1982; G, Diprotodon optatum, drawn from photograph of Australian Museum
specimen, Sydney; H, Macropus titan, based on proportions in R. Owen, Extinct Mammals of Australia,

1877; photograph in R. Wells & N. Pledge in Chap. 15: Vertebrate Fossils, Natural History of the
Southeast, R. Soc. So. Aust., 1983; I, Thylacoleo carnifex (restoration based on a photograph in R. Wells
& N. Pledge in Chap. 16: Vertebrate Fossils, Natural History of the Southeast, R. Soc. S. Aust., 1983).

THE SIGNIFICANCE OF BODY WEIGHTS

The estimated body weights of inferred Australian ecological equivalents with those of
North America, where the megafauna is well documented, makes an interesting comparison.
Australia is defined as a connected land mass, which includes the islands of Tasmania and New
Guinea. North America is the contiguous land area of Canada, the United States and Mexico to
the Tropic of Cancer. In spite of the considerable difference in land area (Australia: 7.7 X 106

1130 - MURRAY

km2; North America: 24 X 106 km2), both continents supported a relatively similar number of
megafaunal species that attained at least 40 kg body weight.

Figure 38. Scaled skeletal restorations of examples of Late Pleistocene North American megafauna with
human skeleton (height = 177 cm) for comparison: A, Castoroides, giant beaver, redrawn, Romer, Vertebrate
Paleontology Univ. Chicago Press, 1967; B, Mammut americanus, Mastodon, based on proportions given for
a Benton County Missouri U.S.A. Specimen recorded in A Guide to the Fossil Mammals and Birds in the
British Museum of Natural history, 1909, skeleton redrawn from A. Romer, Vertebrate Paleontology, 1967;
C, Bison sp., example of a Late Pleistocene bison, (redrawn from A, Romer, Vertebrate Paleontology, 1967);
D, Panthera atrox, American Lion, based on data in Anderson (1984) and Anderson, pers. comm.; E,
Eremotherium mirabil, Giant ground sloth, drawn from a photograph of a specimen from the Daytona Beach
Bone Bed on display in the Daytona Beach Museum of Arts and Sciences, Daytona Beach Florida U.S.A.; F,
Nothriotheriops shastense, Shasta ground sloth (redrawn from A. Romer, Vertebrate Paleontology, 1967).

The pooled estimate of the mean body mass of the Australian mammalian megafauna,
constituting 35 species >40 kg is 196 kg. The North American megafauna, constituting 44
species >40 kg is 595.4 kg, approximately three times heavier than that of Australia (Figs 37-
39; Tables 16-19). In North America, there is a 27% survival rate of post-Pleistocene
mammals >40 kg compared to only 15% in Australia. If it can be assumed that the long-term,
overall productivity of a continent has some relationship to its area, then the factor of three
difference between land area and the body mass of the megafaunal species might serve as an

PLEISTOCENE MEGAFAUNA - 1131

| empirical estimator for relative scaling of body sizes in terms of community structure. These

estimations are extremely crude and are intended to demonstrate no more than an approximate
order of magnitude. Because there is no real demarcation between the extinct megafauna and
large living mammals (>40 kg) they are included as "surviving megafauna". The Australian
surviving megafauna (>40 kg) comprise six species with a pooled mean adult male weight of
55 kg. North American survivors include sixteen species (>40 kg) having a pooled mean
weight of 295 kg. If the arbitrary megafauna weight definition is divided by three (=13.3 kg)
the lowered Australian rubicon includes a total of 13 Australian surviving megafauna with a
mean pooled weight of 30.9 kg, which exactly equals the 27% survivorship of North America.
Coincidental? Probably, given the nature of the assumptions and imprecision of the
estimations. However, when the entire Australian megafauna is adjusted by this factor, its
species much more closely approximate the trophic structure of Pleistocene North America,
both qualitatively and quantitatively (Table 20). This strongly suggests that the absolute
weight equivalents of North American mammals should not be used as populational analogues.
Large Australian marsupials such as Diprotodon were probably the trophic and populational
equivalents of proboscidians rather than rhinos; zygomaturines were the populational
equivalents of rhinos, not American bison.

PATTERN OF SPECIATION

Prolific Pleistocene speciation in Australia can be inferred by comparison with North
America. There were 19 genera and 35 species of Australian megafauna compared to 42 genera
and about 44 species for North America. The Rancholabrean megafauna was predominatly
monogeneric, while in Australia a small number of megafaunal genera had speciated very
prolifically and finely, implying very narrow niche insinuations (Fig. 39). The tendency for
high speciation among the Australian megafauna supports the proposition that high
diversification of the community is a strategy for attaining maximum control of the
comparatively smaller amount of energy available to it, this being accomplished at the expense
of maintaining high population densities. In other words, population densities of large
herbivores in Australia were comparatively low in order to accommodate an energy-extraction
strategy of species-richness within an oligotrophic ecosystem. As pointed out, it appears
likely that the largest Australian marsupials were bound by population restrictions analogous
to the largest of placental herbivores (mammoths, giant sloths) rather than populations attained
by placentals of approximately the same actual weight. Consequently, populations of the
larger herbivorous marsupials were generally lower than would be attributed to them on the
basis of absolute weights of placental analogues.

DWARFING

The greater the extent of speciation, the more bound-up and secure the available energy. An
important aspect of this speciation is the evolution of a range of body sizes adjusted to the
productivity limitations of the habitat. In a general sense, body size in mammals is a function
of the rate of energy turnover in the system. Large mammals store energy longer than small
ones, contributing to stability. This is the principle behind the high degree of speciation in
Australia in comparison with North America and provides a likely explanation for the great
discrepancy in the mean weight of the surviving megafauna in Australia (30.9 kg) compared to
North America (295 kg).

The dwarfing phenomenon is documented in certain lineages of Late Pleistocene
macropodids. The biological function of dwarfing is to increase the local population density of

1132 - MURRAY
a species. According to Dameth (1981) change in body size offers no direct energetic

advantages in the relationship of population density to body mass where DR = W-
0.75

North America Australia

15

N genera and species

10

15-35

40-100
100-350
350-1500
1500-3000
35-115
115-500
500-1000

Wt. kg.

Figure 39, Paired histograms of numbers of genera and species of extinct megafauna grouped by weight
categories, demonstrating similarity in the distribution of body weights through both North American and
Australian communities. The mean body weight of the North American megafauna is approximately three
limes greater than that of Australia’s. Note the discrepancy in numbers of species (although some of the
North American species are disputed or unassigned) but similarity in the distribution of genera by weight.

PLEISTOCENE MEGAFAUNA - 1133

(D=population density, R=metabolic requirements and W=body mass). Because there is no
direct energetic advantage gained by the reduction of body size, a gradual reduction in habitat
productivity due to climatic change should result in a decline in the local population density
rather than a reduction inthe body size. Late Pleistocene dwarfing appears instead, to be
associated with an acceleration of the energetic turnover in the habitat due to perturbations for
which accelerated sexual maturity and increased reproductive rates and perhaps opportunistic
reproductive patterns are more suitably adaptive. Because body-size is an important adjunct to
many feeding adaptations, dwarfing is not a possibility open to all species (Fig. 40). Certain
specialized browsers cannot continue in their characteristic niches because their large bodies are
an essential physical and physiological adjunct to the adaptation. The grazing macropodine
kangaroos however, were able to scale-down without shifting their adaptive zone, although
some species exclusion and accommodative downward sliding of the already suitably small
kangaroos is an expected and, indeed, observable phenomenon (Marshall & Corrucini 1978,
Horton 1980, 1984).

PREDATOR DIVERSITY

From the foregoing, it may be inferred that the populations of the Australian megafauna
were too sparse to support a high diversity of large mammalian carnivores. Thylacoleo
carnifex is the only large "felid" equivalent and Thylacinus cynocephalus is the only proxy
large "canid". Even with the addition of Propleopus oscillans to the predator guild, the poor
recruitment of large carnivores must be a reflection of the limited biomass and low standing
crop of large herbivorous species. Such a system would provide an optimum support base for
carnivore-scavengers such as Megalania prisca and crocodiles. The lack of highly specialized
scavenging birds (e.g. vultures) and large, hyena-like mammalian scavengers (Australian avian
scavengers are largely opportunistic) may relate to the low mortality rate. In the absence of a
high biomass with nightly predator kills (Serengeti-type) a lower biomass of megamarsupials
suffering periodic catastrophic mortality (drought) would be better suited to low metabolic
reptilian scavengers.

BEHAVIOUR AND INTELLIGENCE
Locomotor Behaviour

The thick-set browsing kangaroos (Sthenurinae) differ anatomically from the equally large
but more lightly built macropodine kangaroos. The distinguishing functional features relate to
the mechanism of the thigh. In Simosthenurus the semimembranosus muscles have a more
acute line of action across the hip joint than in macropodines, created by the upward sweep of
the ischial tuberosity of the pelvis (Fig. 41). The femur in Simosthenurus is also relatively
longer, and with regard to the retraction of the femur, the gluteus medius is longer, and
provided with a much broader attachment to the ilium. Simosthenurus, therefore, had a more
powerful and faster retracting mechanism of the hip than Macropus spp. However, its lower
limb segment was relatively shorter, resulting in a "lower-geared" system. It appears that the
locomotion of Simosthenurus was optimally suited for short bursts of rapid bounding. Small
macropodines show conditions somewhat similar to Simosthenurus. This is a predictable
relationship because their strategy for escape from predators is a short sprint for cover. The
peculiar limb proportions of Simosthenurus suggest that it could have been habitually
quadrupedal during slow locomotion. Procoptodon, being very large and long-limbed, may
have been considerably faster than Simosthenurus but perhaps less capable of sustaining a

1134 - MURRAY

sprint unless it was able to unload heat very efficiently. Its hindlimb was also much more
"high-geared," and thus it was probably a slow starter.

Figure 40. Dwarfing phenomenon in Australian Late Pleistocene megafauna: A, Macropus titan; B,
Macropus cooperi compared with Macropus robustus,; C, Sarcophilus laniarius with S. harrissii; D, Wallabia
indra with W. bicolor; E, Phascolarctos stirtoni with P. cinereus; F, Macropus siva with M. agilis. (After

Murray 1984a).

Protemnodon exhibits another unique macropodid locomotor system. It also had a "low-
geared" hind limb with rebust proportions of the femur, tibia and fibula. Its foot was broader
and shorter than that of any other macropodid except for tree kangaroos. The out-lever to in-
lever proportions of the foot of P. otibandus is 1:3.8 compared with between 1:4.3 to 1:4.7 of
other macropodines. The stocky hindlimb array and the short, broad feet of Protemnodon

PLEISTOCENE MEGAFAUNA - 1135

suggest that it may have required a great deal of energy to initiate a rapid locomotor mode and
to sustain it for any appreciable distance. This assertion depends upon the degree of elastic
recoil of the achilles tendon, and the assumption of course, that Protemnodon locomotion was
similar to that of Macropus.

Figure 41. Hind limbs of: A, Macropus giganteus; B, Simosthenurus occidentalis; and C, Thylogale
billardieri, not to scale; size adjusted to tibia length.

The diprotodontids were plantigrade, graviportal species. The short lever of the olecranon
process in both Zygomaturus and Diprotodon indicate that the line of action of the medial head
of the triceps was inserted close to the joint, yielding little torque, but fast retraction of the
forearm. In common with placental graviportal herbivores, the olecranon was deflected
posteriorly and the ulna and radius were both large and free for weight-bearing. The shoulder
musculature was highly developed. The scapulae of both genera have large acromion processes
and wing-like extensions of the axillary border producing powerful but restricted movements in
relation to the stabilization of the upper segment of the forelimb (Fig. 42). The hind
limbs featured “high-geared" retraction of the knee joint as inferred from the short ischial
tuberosities (Fig. 43). The broad ilia reflect short, massive gluteus medius muscles. In-lever
to out-lever proportions of the feet indicate weak leverage, characteristic of graviportal beasts.
The femur of Diprotodon is relatively long in proportion to the inferior segment of the limb.
This may have yielded a greater velocity of retraction of its hind limb. These beasts were
probably capable of a sustained, smooth and efficient, but comparatively slow, maximum-

speed gait.

1136 - MURRAY

The limbs of the giant wombat, Phascolonus gigas, are structurally similar to its nearest
living relative, Vombatus ursinus. The olecranon process forms a long and superiorly directed
lever arm characteristic of scratch-digging fossors. Both the scapular acromion and the deltoid
tuberosity of the humerus attest to large, well-developed deltoid muscles. The morphology of
the giant wombat's scapula is in marked contrast to that of Diprotodon in having a long,
straight axillary border, like that of the living wombat. The hind-limb of the giant wombat is
also similar to that of the living forms in having a comparatively long and upward-flexed
ischial tuberosity. This effectively lengthens the hamstrings while simultaneously keeping
their line of action close to the hip joint. Both the manus and the pes have comparatively
longer in-levers than those in the diprotodontids. Thus, it appears that P. gigas was
anatomically suited for digging, although its size would make it the largest tunnelling
mammal ever known. Its locomotor capabilities would have been similar to the living
wombats, which are effective short-distance sprinters with speeds up to 40 km having been
recorded.

A

Figure 42. Comparison of forelimbs of: A, Vombatus ursinus; B, Zygomaturus trilobus; C, Diprotodon
optatum; D, Phascolonus gigas; not to scale, size adjusted to humerus length.

Of the known Australian marsupial carnivores, only the thylacine Thylacinus
cynocephalus, may have been capable of sustained cursorial pursuit predation of larger animals.
Even in this case, the thylacine's hunting prowess would have been severely limited without
the social component of pack hunting. We lack direct evidence of pack hunting in thylacines,
but rock paintings from Arnhem Land, Northern Territory depict human beings surrounded by
thylacine packs, suggesting the possibility that this dog-like marsupial may have been a social
hunter over part of its extensive range. Keast (1982) discusses the cursorial adaptations of the
thylacine suggesting that it was not as cursorial as the placental wolf. Werdelin (pers. comm.)
argues against the cursorial pursuit mode of hunting by Thylacinus, based on functional
grounds, and instead, favours an ambush mode of hunting similar to that of foxes.

The marsupial lion, Thylacoleo carnifex, has an overall lion-like visage, but lacks many of
the cursorial specializations of the large placental cats. In comparison with the lion, the limbs

PLEISTOCENE MEGAFAUNA - 1137

are remarkably similar, but the inferior limb segments of the marsupial lion are relatively

heavier (Figs 44, 45). The fibula, which is reduced to a thin splint in the lion is robust in
Thylacoleo. The forelimbs bear the greatest similarity, possibly due to their similar roles in
predation. The greatest differences are in the structure of the manus and pes. The lion is
highly digitigrade, with elongated metacarpals and metatarsals, In Thylacoleo the metacarpals
are broad and short, and, in particular, the metatarsals are much shorter and the entire foot is
relatively smaller. The in-lever (calcaneus) of Thylacoleos's foot is relatively much shorter
than that of the lion. The propulsive thrust of the hindlimb of Thylacoleo was probably
greatly inferior to that of the lion. The marsupial lion was, therefore, principally an ambush
hunter with a limited pursuit capability.

Figure 43. Comparison of the hind limbs of: A, Vombatus ursinus, B, Zygomaturus trilobus; C,
Diprotodon optatum, D, Phascolonus gigas; not to scale, size adjusted to femur length.

Brain Size

The estimation of brain size relative to body size depends on the determination of a satisfactory
allometric adjustment that allows comparison of the enormous range of mammalian body
forms. Jerison (1973) argues that increased brain size in mammals may be at least partly
related to a positive feedback relationship between predators and prey, concluding that the actual
functions that developed from this interaction was the perceptual equipment necessary for
flexible response. Given the apparently low level of predator-prey interaction among
Australian marsupials in combination with their relatively small-brained didelphoid ancestors,
the rate of encephalization would be predictably lower in Australidelphians. Using Jerison’s
(1973) allometric formula EQ=E /0,12P2/3 (where EQ = encephalization quotient; E = brain
weight; P = body weight), the average Australian marsupial brain (Encephalization quotient) is
about two-thirds that of the average placental. Of course, included among the placental sample
are the exceptionally large primate brains, and both Jerison's (1973) allometric formula and
Crile & Quiring's (1940) data base have certain flaws. These calculations, however, seem to
support the direct anatomical observations, that is, marsupials have relatively smaller brains
than placentals (Fig. 46; Tables 21,22).

1138 - MURRAY

Figure 44. Comparison of the forelimb structure of: A, placental lion, Panthera leo, and B, Thylacoleo
carnifex.

Intelligence

Some possible differences in intelligence between placentals and marsupials appear to be
reflected in learning and discrimination tests. In a series of simple discriminations, the Red
Kangaroo performs favourably with that of a house mouse (Mus musculus) but was generally
inferior to many other eutherians tested (Neuman 1961). Munn (1964) concludes that
Australian marsupials are capable of only moderate improvement on discrimination tests.
Kirkby (1977) found that the discriminatory powers of the Brushtail Possum (Trichosurus
vulpecula) was greatly inferior to that of placentals. Tindale-Biscoe (1973) suggests that one
possible explanation for the poor australidelphian discriminatory responses is slower neural
responses associated with a lower metabolism. Both the brain-size calculations and intelligence

PLEISTOCENE MEGAFAUNA - 1139

intelligence testing methods have been severely critici i
y criticized but appear to be supporte
phylogenetic considerations (Lillegraven et al. 1987). ~ Dah aa

Figure 45. Comparison of the hind limb structure of: A, placental lion Panthera leo, and Thylacoleo
carnifex.

While these studies suggest that there are important differences in encephalization and
discriminatory behaviour under certain circumstances between marsupials and placentals, their
actual adaptive significance is not well understood. Without a doubt, the neural adaptations of
marsupials are as refined relative to their particular adaptive zones as those of placentals. The
difference in marsupial and placental mental capability only becomes relevant in the context of
entirely novel conditions, such as the introduction of placental predators.

1140 - MURRAY

0 5cm

Figure 46. Comparison of the endocranial cavities of : A, Sarcophilus harrisii (Tasmanian devil) and B,
a young wolverine (Gulo luscius); the skulls have equal-sized visceral components but differ markedly in the
size of the neurocranium.

Defensive Structures and Morphological Differentiation

Australian marsupials lack highly developed defensive and offensive structures like horns,
antlers and tusks, characteristic of many herbivorous placentals. Of the Late Pleistocene
species, only Zygomaturus trilobus has any indication of the development of defensive horns
or knobs on its head. A Pliocene genus, Euryzygoma dunense, had elongated masseteric
processes resembling the laterally projecting zygomatic processes of American Oligocene
entelodonts. Many giant marsupial species had large claws, and certain large diprotodontians
had large incisor teeth that could have been used defensively, but for the most part, these appear
to be primarily associated with feeding. Kangaroos are moderately effective defensively in
being able to strike out with their huge hind feet while balancing on their tail. This is a
proven strategy for a single predator but would not have the required mobility to keep several

PLEISTOCENE MEGAFAUNA - 1141

predators from eventually reaching it from behind, and particularly vulnerable is the tail itself,
that when apprehended would completely unbalance the kangaroo's defensive capabilities.

Associated with the lack of elaborate defensive specializations is a general morphological
uniformity or paucity of major adaptive zone level of differentiation in the Marsupialia in
general, in comparison to the placentals. There are no flying marsupials or marsupial whales,
seals or otters, but there are adaptations that represent rudimentary steps along some of these
evolutionary pathways. The pattern of differentiation is quantitatively similar to that of
placentals, but qualitatively, the differentiation is generally less radical. Thus, the extensive
speciation of the sthenurine kangaroos is composed of very similar forms. Whatever the cause
of Late Pleistocene extinction, the factors effecting one of the dozen or so species, for example,
the genus Sthenurus, appear to have been equally detrimental to all of them.

Reproduction

This is the most fundamental element in understanding the nature of marsupial speciation
and ultimately of Pleistocene extinction. All that has been previously mentioned about
speciation, body weights and population sizes of the Australian megafauna can be reformulated
and explained in terms of the marsupial pattern of reproduction. The uniquely marsupial
reproductive pattern is basic to their comparative lack of differentiation (morphological
conservatism) and more importantly, in terms of extinction phenomena, their relative lack of
chromosomal (karyotypic) diversity (Lillegraven et al. 1987). According to Lillegraven et al.
(1987), the comparative lack of genotypic differentiation and lower encephalization in
marsupials are related to the degree of connection between energetics and reproduction. In
marsupials, fecundity, gestation period and postnatal growth rates are largely independent of
body mass and metabolic rates. Eutherians have metabolically-linked reproductive pattems that
involve rapid and prolonged intra-uterine development which appears to be correlated with
relatively larger brain-size and complexity, longevity, gregariousness and chromosomal
diversity.

On the less theoretical side, there is less extreme sexual dimorphism among the extinct
megafaunal marsupial genera, whereas the larger surviving megafauna (macropodine kangaroos)
are highly dimorphic. Although this point requires more detailed investigation, it has
important implications in relation to energetics and reproductive rate as obvious advantages are
accrued in the shorter period of sexual immaturity resulting in increased reproductive turnover.
Extreme sexual dimorphism is only possible in forms that have less investment in their body
size for obtaining and processing food. If extreme sexual dimorphism were an option for
population-stressed diprotodontids and short-faced browsing kangaroos, it would have been
present. As with dwarfing, their potential for dimorphism was constrained by the close
relationship of a particular body size to their respective feeding specializations.

CONCLUDING REMARKS

Understanding the interaction between man and the environment and the effect of
environmental changes on biological communities is one of the most important of the
contemporary scientific issues confronting us today. The Pleistocene extinction problem
undoubtedly has some bearing on this. Unfortunately, the compelling attraction of certain
“causes” have found their way into scientific issues, where they offer little or no insight. The
evidence that marsupials may not have been as intelligent or that their mode of reproduction
places significant constraints on their diversification might offend and the notion that
aboriginal hunters may not have been conservation-minded will be obstinately denied by those
committed to the idea. Climatic change is a conveniently neutral causal factor that can
extinguish a megafauna without any emotive connotations.

1142 - MURRAY

In the first place, without being able to determine from scientific evidence, any specific
causal mechanism, the agency or agencies responsible for extinction remain aloof. In order to
implicate human activities, there must be some evidence of their association with the
megafauna. This we have in both circumstantial (temporal overlap) (Merrilees 1968b, Dortch
& Merrilees 1973, Bowler 1976, Chaloupka 1977, Hope et al. 1977, Murray & Chaloupka
1984) and direct (artifact associations) forms (McIntyre & Hope 1978, Archer et al. 1980). If
climatic extremes, particularly drought, is responsible, then there must be supporting evidence
also. This evidence is present from sites about 25,000 years old at Lancefield (Gillespie et al.
1978) and Lake Menindee (Tedford 1967). The evidence does not seem to support a "Bow-
wave" or "Blitzkreig" effect as proposed by Martin (1984), but this does not rule out the
possibility that aboriginal man was involved in the megafauna extinctions (Jones 1968, 1979).
The arrival of Homo sapiens on the Australian continent is the only new element in the
evolution of Australian communities since the invasion of murid rodents in the Pliocene.
Episodic droughts severe enough to result in ecological tethering such as proposed for
Lancefield (Horton 1978) have been occurring in Australia since the Late Miocene (Woodbume
1967; Murray, in prep.). These droughts did not result in the extirpation of the megafauna
then, and there seem to be no compelling reasons why drought alone should have become any
more effective eight million years later.

The foregoing palaeobiological arguments suggest that the Australian megafauna was, if
anything, extremely well adjusted to the drying conditions of the later Tertiary and that many
of their specialized locomotor and feeding mechanisms, the increased body size of bulk feeding
browsers, the trophic and population structure of their communities and the pattern of
speciation also made them highly vulnerable to almost any introduced perturbation, particularly
that of predator pressure. The precise nature of these ecological disturbances is not known.
The use of fire by aboriginal man has been implicated (Merrilees 1968b), but it would have the
same drawback as the idea of the Bow-wave effect of hunting that if it were the primary agency
of extinction, effective extirpation would be almost immediate. However, if the population of
the megafaunal species was relatively low and if they were comparatively slow-reproducing,
those specialists among which body size was an important aspect of their feeding adaptation
were unable to respond to higher population demands by decreasing their body size were more
at risk to predator pressure than others. It is, therefore, not difficult to imagine that a gradual
attrition of these populations by hunting could eventually lead to their extinction. Of the
various preferred scenarios, this would seem to account for the differential nature of the
extinction pattern better than any drought or fire ecology argument,

If extinction by population attrition through hunting were the case, we would expect to find
archaeological sites with some associated megafauna and palaeontological assemblages that
reflect a gradually diminishing megafauna element from 50,000 yBP to the Holocene. If
drought were the primary causal agency of extinction, then it must be proven that the
potentially enormous refugia in Australia were severely and equally affected, presumably to the
extent that no animal over 40 kg was able to survive, The late persistence of megafauna in the
very Late Pleistocene or early Holocene could be taken as a clear refutation of the drought
hypothesis. If the firing of the bush by aboriginal people was responsible, its pattern should
not differ from that of the "Bow-wave" Overkill model, for if the megafauna made an initial
adaptation to it, as twenty thousand years of overlap with aboriginal man in Australia seems to
imply, there is no direct support for the idea. However, it is possible that the necessarily
detrimental ecological modifications of the environment took some twenty thousand years to
fully establish, and the final energetic circumstances could not be endured by the megafauna.

More succinctly, the cause of megafaunal extinction was probably due to a combination of
all the above agencies, but without the influence of aboriginal man, the megafauna would have
survived until the arrival of the Europeans.

PLEISTOCENE MEGAFAUNA - 1143

SUMMARY

The study of Late Pleistocene megafauna in Australia was initiated by the British anatomist
Richard Owen in the mid-19th Century. His monumental two-volume work (1877) is a tribute
to the brilliance of the old anatomist, as is still stands as the single most complete and
valuable reference on the subject. Since the 1950's, the palaeontological and geochronological
background to the evolution of the Australian Pleistocene megafauna has been substantially
enriched by new discoveries. Subsequent systematic investigations have also clarified the
taxonomic relationships of some groups, although much remains to be done. At the present
time, the major mammalian constituents of the Australian Late Pleistocene community are
identified with sufficient precision that more detailed studies of Pleistocene communities and
broader comparisons can be anticipated. Of the monotremes, there are two genera and three
species of large echidnas. The dasyurids have one large-bodied genus, Sarcophilus, with one
relative megafaunal subspecies, S. laniarius dawsoni, The thylacines are reduced to a single
Pleistocene species, Thylacinus cynocephalus. Koalas are represented by a slightly larger
Pleistocene morph, Phascolarctos stirtoni. The large wombats are still in a somewhat
ambiguous state. It appears that there is a single, very large form, Phascolonus gigas, and a
medium-sized Ramsayia-group composed of one or two species. A third large-bodied genus,
Phascolomys, contains a single species, P. medius, that was about half again larger than the
living Lasiorhinus or Vombatus.

The marsupial tapirs, Palorchestes spp., have been reduced to a single Late Pleistocene
species, P. azeal. The presence of P. parvus at Strathdownie Cave may be a misidentification,
or, perhaps, not all of the Strathdownie material is Late Pleistocene in age. These are rare and
poorly represented fossils about which little is known. They are among the largest marsupials
which include the marsupial rhinoceros, Zygomaturus trilobus, and the marsupial proboscidian
analogue, Diprotodon spp. There may be two distinct Pleistocene species of Diprotodon. A
single "nototheriine" Nototherium inerme, survived into the Late Pleistocene. "Pygmy"
zygomaturines have been recorded from Late Pleistocene cave localities in New Guinea.

The marsupial lions, although highly variable in size, appear to represent a single species,
Thylacoleo carnifex. They represent the largest Australian mammalian carnivores. The
suggestion of a scavenger status for the species in unsupportable due to the absence of any
other primary carnivore to make kills. Because the carnivore recruitment in megafaunal
communities appears to be remarkably low, it has been proposed that the giant rat-kangaroos,
Propleopus, may have been at least an opportunistic predator. The macropodoid megafaunal
complex was an enormous radiation, of which the genera Macropus and Sthenurus each contain
at least a dozen species. Procoptodon goliah was the largest Late Pleistocene kangaroo,
attaining an estimated weight of between 150 and 200 kg.

In terms of absolute weights, the Australian megafauna were comparatively small-sized
compared to the North American, Eurasian and African megafaunas. The mean body weight of
the Australian megafauna over 40 kg is about one-third that of the North American megafauna.
The pattern of speciation of the Australian megafauna also differs from that of North America
in that there were few genera and many finely differentiated species in Australia compared to the
predominantly monospecific but large number of genera in North America. The Australian
marsupials evolved few specialized defensive and offensive structures and appear to have been
less fleet than the placental cursorial specialists. Anatomical and behavioural evidence suggest
that the marsupial megafauna may not have evolved as high a flexibility of response to
predators as the larger herbivorous placentals. The Australian megafauna had been adapting to
cyclic drought for at least 8 million years. Many of their morphological specializations
related to the marginal conditions of a semiarid climate and pedological senility. They appear

1144 - MURRAY

to have developed a high level of fitness for local and regional droughts and are unlikely to
have entirely succumbed to the singular conditions of the Late Pleistocene. Therefore, in
combination with the inference that the Australian megafaunal populations were comparatively
low, the cause of extinction may be attributed to gradual populational attrition by human
predation.

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APPENDIX I
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Table 1. Mean dimensions (mm) of selected skeletal elements of three tachyglossid species from Late Pleistocene deposits
in Mammoth Cave, Westem Australia.

Tachyglossus cf. T. aculeatus Zaglossus ramsayi Zaglossus hackett

Condylobasal

Length of skull - 165.0 -
Length of Beak - 89.3 =
Length of Humerus 54.0 64.8 -
Length of Radius - 76.0 100.0
Length of Femur 57.0 72.0 126.0
Length of Tibia - 87.7 112.0

Table 2. Mean dimensions (mm) of the cheek teeth of giant devil, Sarcophilus laniarius (data from Dawson 1982a).
p> Ps) M2 Mp) M3 Mz” M4 Mg Ms Mg

Length 6.7
6.1

0 12.3 9.9 13.7 11.6 14.
Width 3 1

10.0 6.9 11.0 7.2

Table 3. Mean dimensions (mm) of the cheek teeth of Late Pleistocene thylacines, Thylacinus cynocephalus, from
Wellington caves (data from Dawson 1982b).

Table 4. Mean dimensions (mm) of lower molars of vombatid, Phascolomys medius, compared with Vombatus ursinus (data
from Marcus 1976).

Phascolomys medius Vombatus ursinus
Depth of Mandible
at M2 - M3 43.9 32.4
M2 Length 14.3 11.6

M2 Anterior Width 8.1 6.1

M2 Posterior Width

M3 Length
M3 Anterior Width
M3Posterior Width

Mg Length
Mg Anterior Width
M4Posterior Width

Ms Length
Ms Anterior Width
Ms Posterior Width

M2 - Mg Length

PLEISTOCENE MEGAFAUNA - 1151

Table 5. Mean dimensions (mm) of upper cheek teeth of Zygomaturus trilobus and Diprotodon optatum (data from Owen

1877).

p> Length
P? Width

M2 Length
M2 Anterior Width
M2 Posterior Width

mM? Length
M? Anterior Width
M?> Posterior Width

m4 Length
m4 Anterior Width
M4 Posterior Width

M> Length
M> Anterior Width
M> Posterior Width

Zygomaturus trilobus

28.7
24.5

31.5
29.8
31.2

37.6
35.5
33.5

44.3
38.5
35.0

43.8
37.6
29.2

Diprotodon optatum
24.6
21.5

38.4
32.3
35.1

47.5
40.3
41.1

57.5
45.6
43.6

54.2
44.9
37.8

Table 6. Mean dimensions (mm) of upper cheek teeth of Late Pleistocene palorchestid, Palorchestes azeal, compared with

Pliocene (?Pleistocene) Palorchestes parvus (data drom Woods 1958).
Palorchestes azeal

p> Length
p>? Width

M2 Length
M? Width

mM? Length
M? Width

m4 Length
M4 Width

M> Length
M> Width

19.5
18.5

26.3
21.9

26.8
23.2

27.0
22.6

27.7
22.0

Palorchestes parvus
16.8
14.5

19.6
15.5

20.1
16.3

21.0
16.6

23.7
16.6

1152 - MURRAY

S606

Table 7. Mean dimensions (mm) of Thylacoleo carnifex cheek teeth (data from Archer & Dawson 1982).

P3 p3 M2 M2 P3+Mz P>+ M3
Length 39.2 49.8 14:2 14.2 51.8 47.2
Width 14.0 14.7 10.0 9.3 . :

—aeG=—uaeR60o020o773°0
SSS

Table 8. Mean dimensions (mm) of lower cheek dentitions of the giant rat-kangaroo, Propleopus oscillans (data from
Pledge 1980a).
Propleopus oscillans

P3 Length 14.2
P3 Width 9.5
M2 Length 9.7
M2 Anterior Width 9.2
M2 Posterior Width 9.0
M3 Length 10.9
M3 Anterior Width 9.9
M3 Posterior Width 10.2
Mg Length 11.4
Mg Anterior Width 10.5
M4 Posterior Width 9.8
Ms Length 11.0
Ms Anterior Width 9.7
Ms Posterior Width 8.5

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Table 9. Mean dimensions (mm) of lower cheek teeth of large Late Pleistocene Macropus species (data from Bartholomai).

Macropus titan Macropus ferragus Macropus cooperi

P3 Length 7.4 9.0

P3 Width 4.0 4.0 3.0
M2 Length 13.0 - -
M2Width 7.6 8.3 >
M3 Length 14.5 16.9 13.2
M3 Width 9.0 9.7 ‘
Mg Length 16.5 18.8 15.3
Mg Width 9.7 11.1 -
Ms Length 17.5 20.8 17.4
Ms Width 10.3 11.4 9.0

Table 10. Mean dimensions (mm) of lower check teeth of Late Pleistocene Protemnodon species (data from Bartholomai
1973).

Protemnodon anak Protemnodon brehus Protemnodon roechus
P3 Length 16.5 16.7 16.5! 17.7 20.0! 18.8
P3 Width $7 5.4 6.4 7.1 7.3 7.0
M2 Length 10.5 12.0 13.4 13.5 13.1 13.3
M2 Width 7.5 Ay | - 9.9 - 9.4
M3 Length 13.4 13.9 15.3 16.3 16.2 16.1

M3 Width 953 9.3 10.6 11.5 - 11.3

PLEISTOCENE MEGAFAUNA - 1153

M4 Length 16.2 15.5 17.9 18.2 18.9 18.5
Mg Width 10.5 10.4 12.1 12.6 12.4 12.5
Ms Length 17.2 16.5 19.7 19.2 20.0 19.7
Ms Width - 10.8 12.3 12.7 12.3 12.6
1 holotypes

Table 11. Mean dimensions (mm) of lower cheek teeth of occidentalis group of Simosthenurus (leaf-eating kangaroos)
(data from Merrilees 1965, Tedford 1966).

S. occidentalis S. brownei S. orientalis
P3Length 16.7 14.4 17.0
P3 Anterior Width 7.8 9.61 8.1
P3Posterior Width 9.9 - 10.0
M>) Length 12.1 10.2 13.1
M2 Anterior Width 9.0 9.1} 10.3
M? Posterior Width 9.4 - 10.6
M3 Length 12.8 11.1 14.3
M3 Anterior Width 10.1 9.91 11.6
M3 Posterior Width 10.1 - 11.9
Mg Length 13.1 11.6 15.6
Mg Anterior Width 10.4 10.2! 12.5
Mg Posterior Width 10.4 - 12.6
Ms Length 12.1 11.3 14.6
Ms Anterior Width 10.3 10.3! 12.7
Ms Posterior Width 9.5 - 12.1

1 maximum width
———————————————=—========—

Table 12. Mean dimensions (mm) of lower cheek teeth of microdont leaf-eating kangaroo, Simosthenurus gilli and S.
maddocki (data from Merrilees 1965, Wells & Murray 1979)).

Sthenurus maddocki Sthenurus gill
P3 Length 15.9 14.4
P3 Maximum Width 8.2 8.6
M2 Length 10.3 8.8
M2 Maximum Width 8.6 8.0
M3 Length 11.1 9.7
M3 Maximum Width 9.3 8.5
Mg Length 12.0 10.1
Mg Maximum Width 10.2 9.2
Ms Length 11.7 9.7
Ms Maximum Width 10.2 9.3

————
i eee
Table 13. Mean dimensions (mm) of lower cheek teeth of dolichocephalic sthenurines, Sthenurus (data from Tedford
1966).

S. tindalei S. atlas S. andersoni S. oreas
P3 Length 18.7 16.8 14.5 14.2
P3 Anterior Width 8.2 6.5 6.1 6.9
P3 Posterior Width 9.7 8.2 FA 8.5

M2 Length - 12.3 11.1 13.3

1154 - MURRAY

M2 Anterior Width - 9.2 8.8 9.9
M2 Posterior Width < 9.5 8.9 10.2
M3 Length - 13.6 12.7 14.8
M3 Anterior Width - 10.5 9.7 11.0
M3 Posterior Width 13.6 10.4 10.0 11.2
Mg Length 17.0 14.8 13.9 16.4
Mg Anterior Width - - 11.0 12.2
Mg Posterior Width - 11.4 10.7 12.2
Ms Length 16.4 14.0 12.6 -

Ms Anterior Width 13.6 11.9 11.1 -

Ms Posterlor Width 11.4 10.5 9.6 -

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Table 14. Mean dimensions (mm) of lower check teeth of Procoptodon species (data from Marcus 1976).

P. goliah P. rapha P. pusio
P3 Length 14.1 15.2 10.0
P3 Anterior Width 8.6 7.9 5.6
P3 Posterior Width 11.0 11.0 6.6
M2 Length 18.3 17.9 13.7
M2 Anterior Width 13.9 13.1 10.1
M2 Posterior Width 14.7 13.3 10.4
M3 Length 22.2 21.8 16.5
M3 Anterior Width 16.9 15.2 11.8
M3 Posterior Width 16.5 14.9 11.7
Mg Length 24.7 23.5 18.5
Mg Anterior Width 18.5 16.2 12.7
M4Posterior Width 17.7 15.8 12.6
Ms Length 24.9 23.3 18.0
Ms Anterior Width 18.0 15.3 12.7
Ms Posterior Width 16.7 14.9 11.8

eee SSS SS

Table 15. Check list of Pleistocene occurrences of Australian mammalian megafaunal species by state.
Qid. N.S.W Vic. Tas. S.A. W.A.
Tachyglossidae
Zaglossus ramsayi Xx x xX ».4 Xx
Zaglossus hacketti
Dasyuridae
Sarcophilus laniarius Xx Xx Xx Xx xX ?
Thylacoleonidae
Thylacoleo carnifex x Xx XxX Xx X
Thylacoleo sp.
Vombatidae
Phascolonus gigas x
Ramsayia curvirostris
Phascolomys magnus
Phascolomys medius
Vombatus hacketti ».4
Phascolarctidae
Phascolarctos stirtoni Xx
Palorchestidae
Palorchestes azeal Xx
Palorchestes parvus
Diprotodontidae

x
x

~_
x x

x mK mM
xx

~<
~ >
x
x
x

Diprotodon minor x x 4 4
Diprotodon optatum X xX xX x X ?
Zygomaturus trilobus X X xX ? X xX
Zygomaturus ?tasmanicum 4

X xX

Nototherium inerme

PLEISTOCENE MEGAFAUNA - 1155

Macropodidae
Propleopus oscillans
Wallabia vishnu
Protemnodon anak
Protemnodon brehus
Protemnodon roechus
Troposodon minor
Macropus siva
Macropus rama
Macropus gouldi
Macropus piltonensis
Macropus thor
Macropus birdselli
Macropus titan
Macropus ferragus
Macropus stirtoni
Macropus pearsoni
Sthenurus andersoni
Sthenurus atlas
Sthenurus oreas
Sthenurus tindalei
Sthenurus pales
Sthenurus occidentalis
Sthenurus brownei
Sthenurus gilli
Sthenurus orientalis X
Sthenurus maddocki
Procoptodon goliah Xx
Procoptodon rapha x

x
x

~ MRK KM
~~ KM OM
ta
~~ OM
>

mK M
bas
~*~

mm

~_

> DS OM DE Od OK KOK OK OK OK OK OK OK OO OM
)

mx KK OM

mK OK OK
*
pa Et idl i i i ia!
KK

Procoptodon pusio
Procoptodon texasensis

OOOO

Table 16. Data base from which the estimation of body mass of megafauna was derived; < indicates underestimation of
upper limit of body mass range; cranial and dentary lengths can be used as a crude estimator of body length; (~ Body
length = 4.5 x 5.0 x cranial length; ~ Body length = 7.0 x dentary length, see Jerison, 1973).

Range of Supplementary Estimators Range Estimated

Head-Body (mm)(example) Mass (kg)

Length (M) Cranial Dentary 0.025 L3 - 0.035 L3

Length Length
Sarcophilus laniarius 0.7-0.8 140 129 8.6-17.9
Propleopus oscillans 0.9-1.1 187 150 18.2-46.5
Sthenurus atlas 1.0-1.1 180 180 25.0-46.5
Sthenurus occidentalis 1.0-1.15 185 140 25.0-53.2
Sthenurus orientalis 1.0-1.2 190 140 25.0-60.0
Sthenurus gilli 0.8-0.9 163 115 12.8-25.5
Sthenurus maddocki 1.0-1.2 190 140 25.0-60.0
Procoptodon pusio 1.1-1.25 - 140 33.0-68.0<
Procoptodon rapha 1.15-1.3 235 165 38.0-76.9<
Procoptodon goliah 1.2-1.6 244 180 43.2-143.3
Protemnodon anak 1.1-1.3 230 155 33.0-76.9
Protemnodon brehus 1.2-1.5 250 155 43.2-118.0
Protemnodon roechus 1.3-1.5 260 190 54.9-118.0
Macropus titan 1.3-1.5 265 170 54.9-118.0
Thylacoleo carnifex 1.02-1.25 230 135 26.5-68.0
Phascolonus gigas 1.5-1.8 370 280 84.4-204.1
Ramsayia magna 1.0-1.2 - 160 25.0-60.0
Phascolonus medius 0.875-1.0 - 140 16.7-35.0
Palorchestes azeal 2.0 384 - <280
Nototherium inerme 2.0 - 370 <425
Zygomaturus trilobus 2.15-2.4 480 345 248.5-483
Diprotodon optatum 2.9-3.2 680 450 609.0-1146.9
Megalania prisca 3.0-3.6 - - 675.0-1632.3
1.5 - - 84.0-118.0

Genyornis newtoni

ao.

1156 - MURRAY

—— ——___.___ ee

Table 17. Single significant figure body weight estimations of extinct Australian megafauna species; line indicates
arbitrary 40 kg cutoff for comparison with North American mega fauna. These estimations are derived primarily from the
data base in Table 17, with slight upward or downward adjustments based on analogies with living species and proportional
peculiarities, offered as “average” body mass estimations. Obviously this is not a precise measure; it is an attempt to
present an approximate figure for comparison with other continental Pleistocene megafauna based on an objective

methodology.

Genus & Species Est. Male Genus & Species Est. Male
Weight (kg) Weight (kg)

Zaglosus ramsayi 10 Macropus stirtont 65
Sarcophilus laniarius 15 Macropus birdselli 70
Zaglossus hacketti 20 Protemnodon brehus 75
Sthenurus gilli 25 Procoptodon pusio 75
Warendja wakefteldi 30 Protemnodon roechus 85
Phascolonus medius 36 Macropus ferragus 85
Vombatus hacketti 35 Macropus titan 85
Troposodon minor 45 Procoptodon rapha 90
Propleopus oscillans 45 Sthenurus pales 90
Bohra paulae 45 Sthenurus tindalei 90
Thylacoleo carnifex 45 Genyornis newtoni 100
Sthenurus atlas 50 Procoptodon goliah 120
Sthenurus anderson 50 Phascolonus gigas 150
Sthenurus occidentalis 50 Palorchestes parvus 250
Protemnodon anak 50 Palorchestes azeal 300
Sthenurus maddocki 55 Zygomaturus ?tasmanicum 450
Sthenurus oreas 55 Nototherium inerme 450
Sthenurus brownei 60 Zygomaturus trilobus 500
Sthenurus orientalis 60 Diprotodon minor 850
Macropus pearsoni 60 Megalania prisca 1000
Ramsayia magna 60 Diprotodon optatum 1150

eee

EE...

Table 18. North American megafauna body mass estimates expressed as a single significant figure for comparison with
Australia's megafauna. The majority of these species are thought to be Rancholabrean equivalent (Late Pleistocene).

Genus & Species Est. Male Genus & Species Est. Male
Weight (kg) Weight (kg)

Stockoceros sp. 1 40 Navahoceros 225
Stockoceros sp. 2 40 Panthera atrox 235
Mylohyus 40 Glyptotherium floridanus 280
Hydrochoerus 45 Holmesina 280
Saiga 50 Bootherium 300
Dasypus bellus 60 Tapirus veroensis 330
Tetrameryx 60 Equus sp. 350
Acinonyx 65 Megalonyx jeffersoni 370
Miracinonyx 65 Symbos cavifrons 400
Canis dirus 65 Euceratherium collinum 450
Neochoerus 80 Arctodus simus 720
Hemiachenia 100 Cervalces sp. 850
Palaeolama 100 Bison priscus 850
Blastoceros 145 Alces latifrons 1000
Ovis 150 Bison latifrons 1000
Casteroides 150 Glossatherium sp. 1100
Tremarctos 150 Camelops hesternus 1100
Platygonus 150 Titanotylops sp. 1500
Nothrotheriops shastense 180 Eremotherium mirabil 2500
Sangamona fugitiva 180 Mammut americanus 3000
Smilodon californicus 225 Mammuthus primigenius 3000
Homotherium simum 225 Mammuthus jeffersoni 4000

ee _____

PLEISTOCENE MEGAFAUNA - 1157

SSO

Table 19. Indices and coefficients derived from comparisons of estimated body masses (mean) and continental areas from
which the following observations can be recognized: 1, the pooled body mass of the Australian megafauna was on the order
of one-third that of the North American; 2, based on the arbitrary equal to or greater than 40 kg megafauna definition

( Martin's line"). The North American "surviving megafauna" is about 5 times greater in average body mass than that of
Australia; 3, adjusting the definition of megafauna in Australia by a factor of 0.3, the number of “surviving megafauna” in
Australia becomes equivalent to that of North America (27%). This suggests that the correct magnitude of trophic

comparisons of species by body weight should include Australian species considerably smaller than 40 kg body mass.

1. Mean Body Mass of Extinct Megafauna
Number of Species Mean Body Mass Coefficient
Mean N.A./Mean Aust.
North America 44 595.4 kg
3.04
Australia 35 196.0

UNE EE

2. Mean Body Mass of "Surviving" Megafauna >40 kg.

Number of Species Mean Body Mass % Survival Coefficient
North America 16 295 kg 21%
5.4
Australia 6 55 kg 15%

i

3. Mean Body Mass of "Surviving" Megafauna Using a Factor of 3 to Derive Relative Weight
Categories.

Number of Species Mean Body Mass % Survival Coefficient
North America 16 295 27%
9.5
Australia 13 30.9 271%

4. Megafauna Weight/Area Index

Number of Species Mean Body Mass Continent Area Index
North America 44 595.4 kg 4.4 x 10° km? 0.41
Australia 35 196.0 kg 7.7 x 10° km? 0.39

ee ____..__e

ES

Table 20. Examples of Australian ecological (Eltonian) analogs classified by relative equivalent categories of body mass
adjusted by a factor of 0.3.

NORTH AMERICA AUSTRALIA

40 - 100 kg 15 - 35 kg

Canis Sarcophilus

Dasypus Zaglossus

Noechoerus Phascolonus

Hydrochoerus Vombatus
TT

>100 - 350 kg >35 - 115 kg

Platygonus Propleopus

Panthera Thylacoleo

Castoroides Ramsayia

Palaeolama Troposodon, Sthenurus

Nothrotheriops Simosthenurus

Sangamona, Navahoceras Protemnodon

cervids, bovids (grazers) Macropus

Glyptotherium, Holmesina Phascolonus

nnn

>350 - 1500 kg >115_ - 500 kg
Procoptodon

Megalonyx (folivorous browser)
Tapirus (large browser) Palorchestes

1158 - MURRAY

Zygomaturus
Nototherium

Glossatherium

a ee aE

>1500 - 3000 kg >500-1000 kg
Camelops, Titanotylops (grazer - browsers) Diprotodon
Eremotherium, Mammut (browsers) Diprotodon

EE... eee

——————

Table 21. Encephalization quotients for some living Australian marsupials for comparison with placentals (EQ formula
from Jerison, 1973). Sample mean EQ for Australian marsupials = .57. (Data from Haight & Nelson 1987).

Common Name Scientific Name Body Mass Brain EQ=
P (kg) Mass EO x 0.12p 2/3
E(g)
Marsupial Mouse Antechinus swainsonit .054 95 55
Brown Bandicoot Isoodon obesulus 925 4.95 .43
Native Cat Dasyurus viverrinus 1.320 6.10 -42
Tiger Cat Dasyurus maculatus 3.000 10.40 -42
Tasmanian Devil Sarcophilus harrisii 5.100 13.50 -38
Sugar Glider Petaurus breviceps 0.116 2.95 1.03
Brush-Tailed Possum Trichosurus vulpecula 3.150 13.00 50
Spotted Cuscus Phalanger maculatus 4.000 17.10 57
Common Wombat Vombatus ursinus 16.100 48.10 .63
Rat Kangaroo Potorous tridactylus 1.460 12.00 -78
Pademelon Thylogale billardierii 4.900 22.50 .65
Grey Kangaroo Macropus giganteus 34.530 61.40 48

Table 22. Encephalization quotients of placental mammals (EQ formula from Jerison 1973). Sample mean EQ for
placentals = 0.95. (Data from Crile & Quiring (1940).

Common Name Scientific Name Body Mass Brain EQ=
P (kg) Mass E/0 x 0.12p 2/3
E(g)
Racoon Procyon lotor 4.38 41.0 1.28
Coatimundi Nasua narica 6.25 44.2 1.08
Brown Bear Ursus arctos 197.0 407.0 1.00
Dog Canis familiaris 12.47 81.5 1.20
Wolf Canis lupus 22.68 119.0 1.24
Hyena Crocuta crocuta 43.50 168.0 1.30
Jaguar Felis onca 34.47 147.0 1.16
Leopard Felis pardus 48.00 135.0 85
Lion Felis leo 124.00 229.2 -77
Hare Lepus arcticus 1.90 14.4 -78
Agouti Dasyprocta punctata 3.63 21.8 78
Beaver Castor canadensis 5.83 29.5 .76
Norway Rat Rattus norvegicus .20 1.6 39
Capybara Hydrochoerus isthimus 27.70 52.2 48
Tapir Tapirus indicus 201.00 265.0 64
Rhinoceros Rhinoceros bicornis 763.00 666.0 66
Horse Equus caballus 461.70 618.0 .86
Hippopotamus Hippopotamus amphibius 1351.00 720.0 -49
Deer Odocoileus virginianus 61.50 209.0 1.12
Bush Buck Tragelaphus scriptus 34.50 139.0 1.09
Giraffe Giraffa camelopardalis 529.00 680.0 -87
Hyrax Procavia capensis 3.50 19.2 69
Elephant Loxodonta africanus 6700.00 5700.0 1.34

Vervet Monkey Cercopithecus aethiops 4.00 61.5 2.05

PLEISTOCENE MEGAFAUNA - 1159

PLATES

Plate 1. Drawing of the skull of the giant wombat, Phascolonus gigas. (After a photograph in Pledge
1980b).

Plate 2. Drawing of the skull of Zygomaturus trilobus. (After Owen 1877).

Plate 3. Drawing of the skull of Diprotodon optatum, from a specimen in the Australian Museum, Sydney.
Plate 4. Amhem Land rock art and artist's reconstruction of the Marsupial Tapir, Palorchestes.

Plate 5. Drawing of skull of Thylacoleo carnifex. (After Owen 1877).

Plate 6. Steps in the reconstruction of the now extinct Thylacoleo: A, muscle overlay; B, final
reconstruction.

Plates 7, 8. Reconstruction of Procoptodon; drawing of skull of Procoptodon goliah. (Tedford 1967).

1160 - MURRAY PLATE 1

PLATE 3 PLEISTOCENE MEGAFAUNA - 1161

PLATE 4

1162 - MURRAY PLATE §

PLATE 6

PLATE 7 PLEISTOCENE MEGAFAUNA - 1163

PLATE 8

1164 - MURRAY

Palorchestes azael and Palorchestes parvus, a large and a small Palorchestes, were amongst the
most bizarre of marsupials in Australia. The large form was as large as a bull and may have
ripped off the bark of trees in order to feed. It has extremely powerful forearms, massive claws

and a cranial morphology that suggests it may have had a trunk. (From Rich & van Tets 1985,
with permission of The Museum of Victoria).

CHAPTER 25

THE AUSTRALASIAN
MARINE VERTEBRATE
RECORD AND ITS
CLIMATIC AND
GEOGRAPHIC
IMPLICATIONS

R. Ewan Fordyce!

IMtrOMUCHON: «02.2.5 seccsvceenvsenvcag sepecies 1166 BitdSs, ce¢siifed eilediscetentest eyes. eee 1175
Procedures and Problems............. 1166 Summary of the Stratigraphic
Climates, Geography and Record of Penguins.......... 1175

Biogeography..............seseeeees 1167 Interpretation of the Penguin
Climate and Its Influence on REGCOTO) craisinreeestoectzennyie' 1176
Marine Vertebrates........... 1167 Marine Mammals ...............+.+806 1178
Geography and Its Influence on SULOMIA « cvcisegesistagdesosseestoneses 1178
Marine Vertebrates........... 1170 Seals: the Stratigraphic Record
Biogeography and and Its Interpretations....... 1178
Zoogeography..............066 1170 Cetacea: the Stratigraphic
The Australian Marine Vertebrate Record and Its
Record and Its Implications.....1172 Interpretation............. 1179
Marine Reptiles:s.s,c.ccenesssseceeeses’ 1172 The Stratigraphic Record... 1179
The Australian Cretaceous Interpretation of the
ReCord supe ha tehsess big oes te 1172 RECOM. ...decsascerereosse 1183
The New Zealand Cretaceous References ...icssecevecseiseevedestegerecessess 1185
RECOM cette ss sone Sete 1173
Interpretation of the Australian
CIetaCCOUS........ceceeeeeeeee 1174
The Australasian Tertiary
Recordhs 22. eo. Re eee 1175

nnn EEE
1 Department of Geology, University of Otago, Dunedin, New Zealand.

1166 - FORDYCE
INTRODUCTION

The distribution of animals, including marine vertebrates, is controlled markedly by
geography (that is, the distribution of land and sea), and climate. Why study the spatial
distribution of fossils? Range disjunction, or allopatry, nearly always precedes evolutionary
differentiation. Therefore, our knowledge of the factors that influence the distribution of living
animals, together with palaeogeography and palaeoclimatology, may allow us to interpret the
interaction between palaeobiogeography and major evolutionary change (e.g. adaptive
radiation, extinction). The aim here is to consider the fossil record of some Australasian
marine vertebrates and its climatic and geographic implications.

This article consists of three sections:

1. An outline of procedures and problems encountered in interpreting the relationship
between fossils, palaeoclimates, and palacogeography.

2. A review of the theoretical ways in which changing geography and climate could affect
marine vertebrates. Useful general references for this section are Valentine (1973), Frakes
(1979), Cocks (1981), Forey (1981).

3. A summary and analysis of the Australasian marine vertebrate record, which covers the
Cretaceous and Tertiary of Australia and New Zealand. The earlier record is known too poorly
to warrant inclusion here. Fish are not discussed, as they are covered elsewhere in this volume
(in chapters by Long, Turner, A. Kemp and N. Kemp, this volume). A more detailed
discussion of New Zealand fossil vertebrates (Fordyce 1982a and this volume) is also included
in this volume. This section is complemented by a useful selection of articles in a symposium
volume edited by Ballance (1980).

This chapter is only slightly changed over that of Fordyce (1982a) and has been reproduced
with permission of the editors.

PROCEDURES AND PROBLEMS

Theoretically, one could analyse the palaeoclimatic and palaeogeographic implications of
fossil marine vertebrates as follows:

1. Establish that the taxon under study is monophyletic (that is, descended from a single
common ancestor); or, if a whole fauna is studied, that the fauna is adequately defined. This
goes hand in hand with the assumption that the fossil record of the study taxon is reasonably
complete,

2. Assume, on the basis of knowledge of living representatives of the taxon under study,
that the extant animals possessed a range of responses to climate and geography similar to
those of their living relatives. Alternatively, if there are no close living relatives, one would
have to determine the animals’ possible interactions with their physical environment by
analogy with unrelated animals that possess similar body structures.

3. Given these assumptions, one could then look at patterns of geographic and stratigraphic
change in morphology, diversity and distribution of the taxon under study. Changing patterns
might indicate changes in palaeoclimate and palaeogeography, and could support (or be
supported by) inferences about paleoclimate and paleogeography, obtained by other methods
(for example, stable isotopes and plate tectonics).

4. Finally, but very rarely, could come the most exciting part of this type of study: to
wa a causal relationship between biotic events and palaoclimatic and palaeogeographic
change.

Alternatively, one can work the other way, by proposing a causal relationship between
biotic and geological events (4, above), then testing this hypothesis by working through points

THE AUSTRALASIAN MARINE VERTEBRATE RECORD - 1167

1-3 above. In fact, this is probably the way most people work, whether consciously or
unconsciously.

Although this procedure appears simple enough, it is actually fraught with problems. The
systematic hurdle is the first and often most difficult to overcome. Different systematists
always have different ideas as to what constitute either monophyletic taxa (both in theory and
practice; see for example, Cracraft & Eldredge, 1979) or faunas. At least for the group I am
currently working on (Cetacea - whales, dolphins, and porpoises), most of the type-specimens
described anywhere more than about twenty years ago are based on specimens so incomplete as
to defy reasonable diagnosis. This reflects the fact that well-preserved specimens, which are
vital for good taxonomy, are uncommon (cetacean species in the past, as in the present, have
small populations relative to other animals). It is rare that any fossil cetacean species can be
demonstrated unequivocally to occur at two or more separate localities. The same probably
applies to most fossil vertebrates. In fact, the incompleteness of the record is a universal
problem in palaeontology: the presence of a fossil provides positive evidence, but the absence
of a fossil does not necessarily constitute negative evidence. The functional morphology of
species and analysis of adaptations to different physical environments (2, above) are not
considered here. (For reading on this topic, see, for example, Hildebrand 1974 and Raup &
Stanley 1978).

Accurate stratigraphic correlation is essential in determining contemporaneity of fossils in
different localities before patterns of biological change be established and correlated with
accurately dated changes in the physical environment, However, precise correlation is rarely
achieved. For example, the Oligocene was a time of major oceanic climate change, and,
therefore, perhaps a major time of change for marine vertebrates, yet biostratigraphic problems
prevent accurate correlation between Australasia and the type-localities of the Oligocene in
Europe. Other problems arise: could the biostratigraphic indicators have been influenced by
the palaeoenvironmental changes that they are being used to study? There is always a danger of
circular reasoning. Again, the absence of fossils in an area can be interpreted in more than one
way.

We can never be absolutely certain about causal relationships between biotic and physical
events, Many supposedly linked events are probably just coincidental, and many supposedly
unrelated events could be causally linked. In a few cases, synchroneity of change (whether
evolution or extinction) in a multitude of taxa could indicate a single causal event, for example,
the extinction of large reptiles at the end of the Cretaceous (Alvarez et al. 1980, Hsu 1980,
discussed below), and the invasion of North American terrestrial mammals into South America
in the latest Pliocene, when the Panamanian land bridge was established (Marshall et al. 1979),
Usually, however, the fossil record of a group and its relation to the changing physical
environment can be interpreted in radically different ways, for example, in the case of marine
mammals and Oligocene climates discussed by Lipps & Mitchell (1976) and Fordyce (1980b),
and in the general way that fossils used to counter the notion of continental drift before the late
1960's were used subsequently as evidence that drift had occurred. And, in most cases, no
causal relationship can be established at all. I could continue to discuss these problems, but I
think that it is appropriate to conclude this section, and to preface the next, with a quote from
Pielou (1981): "an explanation for any given disjunction [whether it reflects paleogeography or
palaeoclimate] is more likely to be obtained by common-sense than by abstract theorizing."

CLIMATES, GEOGRAPHY AND BIOGEOGRAPHY

CLIMATE AND ITS INFLUENCE ON MARINE VERTEBRATES

What is climate? Temperature and light constitute probably the most important features of
the marine climate. Other phenomena, such as wind, salinity, water density, currents, tides and

1168 - FORDYCE

waves, also could be regarded as part of the marine climate, but their influence on observable
patterns among fossil marine vertebrates generally is known too poorly to warrant discussion
here. All features of the marine climate ultimately are controlled by solar energy, and all
interrelate in complex feedback loops with all other physical features of the world (for example,
terrestrial climates, geography; see Fig. 1). For reviews of past climates, see Frakes (1979),
and Frakes & Rich (this volume).

<— o ridge
GLOBAL orogeny =
HORA,
iia ki rate
CHANGING pitiendate r Ne
GEOGRAPHY Events 5 iu
Sh jae

<

Physical

Pa Effects

change in es
variance or

dispersal events

Barriers Dispersal Routes

faunal separation «——————_——»_ faunal mixing

faunal divergence «—_________» faunal convergence

allopatric speciation «—________» sympatric speciation

ecological convergence «> ecological divergence,
competition, extinction

Ecological
oer Effects on

direct consequences

General Biotic Changes «—___—_—_ Regression Transgression
Y
phyletic size eliminate <———— habitat —————_»_ increase
K-r strategies smaller ~—————_ faunal province -» larger
diversity less ~—________ shallow shelves -» more
|______» radiation & extinction isolated ~—____. deep basins ——» united
increase? ~—_—— salinity —————» decrease?
less {stratigraphic | more
complete ~ \ record j > complete

(erosion)

enhanced extinctions ~ bias
& initial appearances

Figure 1. Environmental factors affecting marine vertebrates.

A uniformitarian approach is used generally in interpreting the effects of climate on fossil
marine vertebrates (although this does not preclude catastrophic explanations). Simply stated,
modem climates vary, modern climates affect animals, and animals respond to climates, and
these observations help us to interpret the fossil record and its relationship to past climates.
Below, I consider this in more detail.

Climate varies, both spatially and temporally, on an immense scale. This article will not
consider past climatic variation on the small scale, for example, the diurnal or seasonal changes
that often are recorded as growth layers in the fossil hard tissues of one individual. Larger
changes in the past, such as variations over tens of kilometres or over tens of years are only

THE AUSTRALASIAN MARINE VERTEBRATE RECORD - 1169

really important in order for us to get some idea of what is (or was) normal on a geologically
meaningful scale. The changes that are important when considering longer-term changes in
biological evolution are the changes over time of the conspicuous modern latitudinal (polar to
equatorial) gradients.

Variation in the marine climate is caused by two inter-relating factors: firstly, solar
changes, which we can measure today but really can not infer for the past, and, secondly,
geological (and extraterrestrial) changes which influence the manner in which solar energy
reaches the sea, and which we believe we can infer from the past. Below are some examples of
factors that may cause climate variation (see Frakes 1979 for further discussion).

1. Polar ice. The amount of polar ice, which affects global heat budgets, reflects cold and
available precipitation. Polar cooling probably results from thermal isolation of the poles, é.g.
by palaeooceanic changes such as the development of the Circum Antarctic Current, and the
relative distribution of land to sea.

2. Distribution of land to sea. Local effects are relatively well known, but global effects
are less certain. Changes in continental arrangement may result in the formation of
"supercontinents", with very "continental" climates on land and in surrounding seas (i.e. hot
summers, cold winters). Fragmentation of supercontinents may result in smaller land areas
with more “oceanic" climates (i.e. warm summers, mild to cool winters). Continental
movements have allowed circulation changes to develop, e.g. the separation of the southern
continents from Antarctica allowed establishment of the Circum Antarctic Current (Kennett
1977), and the formation of the Panama Isthmus may have resulted in circulation changes in
the Atlantic, which led to formation of the Arctic ice cap (Arthur 1979: 1482). The
distribution of land to sea can also reflect eustatic, or global, sea-level changes, which in turn
may be caused by changes in the volumes of polar ice and mid-oceanic ridges. More localised
sea-level changes can be caused by minor changes in the earth's radius (Morner 1981) and by
local tectonics.

3. Volcanism, The amount of volcanic dust in suspension in the atmosphere will
influence the amount of energy reaching the earth (see also 7).

4. Carbon dioxide. This is normally distributed in equilibrium between air and sea.
Anything that changes this distribution, e.g. changes in carbonate pathways or in the
biological production of CO, will affect global temperatures, for CO2 absorbs infrared
radiation that otherwise is reflected from earth. Increased atmospheric CO? will cause increased
surface temperature - the "greenhouse effect".

5. Magnetic reversals, In fact, there is no positive evidence of the correlation of reversals
and climate change, and some biological events which were thought previously to be linked
with reversals via climate change now appear not to have been (¢.g. Plotnick 1980).

6. Geothermal events. The level of activity of geothermal fields in deep sea rift zones
could significantly affect local marine climates (Macdonald & Luyendyk 1981), although it is
unlikely that such events would have widespread effects.

7. Extraterrestrial events. Any event which, like volcanism, produced particles that
blocked out solar radiation could influence the amount of solar energy reaching the earth's
surface. Asteroid and cometary impact (Alvarez et al. 1980, Hsu 1980) and tectite belts
(O'Keefe 1980) may constitute such events. Poisoning, following asteroid impact (Hsu 1980)
also could lead to climate changes via its influence on the plants involved in the CO? pathway.

8. Milankovitch parameters. Changes in earth orbit (Frakes 1979: 9) will affect the
availability of solar radiation.

What are the results of climate change? As regards global geography, the amount of global
ice dictates the amount of shallow marine shelf available via eustatic effects. Note, however,
that there is a feedback mechanism: the relative amount of marine shelf may influence global
heat budgets (large expanses of water tend to ameliorate the climate of adjacent land) and thus

1170 - FORDYCE

polar ice volume. All climate changes affect biotas, although only the effects of temperature
can be determined very easily. Climate changes may be direct or indirect, and density dependent
or independent. For example, temperature may directly affect metabolism, and hence
distribution, in many species (less so for endothermic marine mammals and birds than for
ectothermic reptiles and fish). Temperature and light may affect the distribution of food, and
thus, indirectly influence vertebrate distribution. Climatic changes which act independent of
population size or density are those such as temperature when it directly affects metabolism,
and poisons. Alternatively, climate change may have density dependent effects, e.g. any change
that affects the availability of food or habitat, the impact of which may vary depending on
species population density. Examples of the possible relationships between climate change and
the evolution of marine vertebrates will be given below.

GEOGRAPHY AND ITS INFLUENCE ON MARINE VERTEBRATES

Geography is the form of the earth's surface. Here, only the largest (i.e. continental-scale)
features need concern us. The subtleties of development of global geography are covered in
many general geology texts (e.g. Holmes 1978, Cocks 1981) and are not considered here. As
was the case for climate, above, a uniformitarian approach is used in interpreting the effects on
marine vertebrates of changing geography.

Geography changes through time. The study of plate tectonics over the last fifteen years or
so has given us a radically new insight into changing geography, including the formation of
new oceans and new barriers important to this discussion. Plate tectonic and climatic changes,
and perhaps changes in earth radius, all affect eustatic (global) sea level changes which, in turn,
affect the amount of physical habitat (e.g. shallow marine shelf) available to marine biotas.
Local tectonics may result in local transgressions and regressions, which locally dictate habitat
availability.

Geographic change affects both climate and biotas. Changing land to sea distribution,
mentioned earlier, can result in large-scale fluctuations in heat budgets (e.g. the physical, then
oceanic, then thermal, isolation of Antarctica). Changing geography can affect marine biotas
indirectly, via climate changes, or directly, by way of changes in barriers and dispersal routes
(Hallam 1981). The general effects of climate changes on biotas were outlined above, and
biogeography is considered below.

BIOGEOGRAPHY AND ZOOGEOGRAPHY

Biogeography is the study of the distribution of all living organisms, and zoogeography is
the study of the distribution of animals. Distribution patterns vary enormously in scale, from
local distribution (i.e. species ecology) through regional distribution (patterns from tens to
hundreds of kilometers) to geographical, or global patterns. Whatever is studied, and whether
fossil or living, it is critical to sample over a time-span that is short enough to be meaningful.
It is pointless to use so long a time-span that data are lumped and trends cannot be seen.

As with climate and geography above, a uniformitarian approach is used in
palacozoogeography. The distribution of animals changes spatially over the short times during
which we can observe al present, and it is assumed that this also occurred over geologic time.
At any given time (at present), the distribution of any species will reflect the vagility (or
ability to move) of individuals within the limits of the possible maximum range as defined by
limiting factors (e.g. food and climate). Temporal changes in distribution will reflect many
different factors, such as climate and geographic change (e.g. Hallam 1981), and evolutionary-
ecological change (adaptive radiation or extinction, which may be linked to climate and
geography).

THE AUSTRALASIAN MARINE VERTEBRATE RECORD - 1171
REPTILES
CHELONIA whe
CROCODILIA a -
PLESIOSAURIA - Plesiosauroidea
Pliosauroidea _

ICHTHYOSAURIA fling
MOSASAURIDAE SR,

BIRDS

SPHENISCIFORMES

MAMMALS

SIRENIA a,

CARNIVORA Otariidae
Phocidae

CETACEA Archaeoceti “ies.

Odontoceti <h.,

Mysticeti

Figure 2. Some marine vertebrates known as fossils from the Cretaceous and Tertiary of Australia and New
Zealand.

I do not wish to dwell on models of zoogeography at this stage, but it is worth briefly
summarising the two main modern theories to account for zoogeographic patterns. Dispersal
zoogeography stresses that animals are vagile and, until only a few years ago, also emphasized
the assumption that continents are stable. Dispersal events allow the multiple, separate, long-
distance dispersal of individuals which can colonise new habitat (e.g. Simpson 1940, Fleming
1979). In contrast, vicariance zoogeography stresses that animals are relatively sedentary, and
disperse little or not at all. Instead, ancestral biotas are split into subunits by relatively few
abiotic changes, which result in the formation of a barrier in the range of the ancestral taxon
(e.g. Cracraft 1980, Nelson & Platnick 1980, Patterson 1981). Some strict vicariance
zoogeographers restrict the concept of abiotic change only to continental fragmentation, while
others concede that climate change may only effect a vicariant event. Generally vicariance
events are stressed by those who study terrestrial biotas, while dispersal tends to be favoured by

1172 - FORDYCE

marine biogeographers. However the concepts are applied, it is noteworthy that both vicariance
and dispersal events result in allopatric speciation, and the formation of sister taxa on either
side of the barrier. For further reading on biogeography, see Chaps 19-21 in Forey (1981), and
Nelson & Rosen (1981, reviewed by Pielou 1981).

THE AUSTRALIAN MARINE VERTEBRATE RECORD AND ITS
IMPLICATIONS
MARINE REPTILES

Many types of marine reptiles occur in both the Cretaceous and Tertiary of Australia and
New Zealand (Tables 1, 2). Cretaceous species are known fairly well, but, because most
Australian species are of Early Cretaceous age, and most New Zealand species are of Late (in
fact, latest) Cretaceous age, few comparisons can be made between these areas.

The Australian Cretaceous Record
Most Australian marine reptiles (see Molnar, this volume) are from the Lower Cretaceous
of New South Wales and Queensland, although some Early Cretaceous and a few incomplete

Late Cretaceous specimens have been reported in the literature from South Australia and
Western Australia. Plesiosaurs constitute by far the most diverse group. According to Persson

TS te, ace

Figure 3. A., Kronosaurus queenslandicus (Pliosauroidea) from Cretaceous Toolebuc Fm. of Queensland;
About 13 metres in length; B, temporal bone of a phocid seal (NMV P160399); about 40 mm wide.

THE AUSTRALASIAN MARINE VERTEBRATE RECORD - 1173

(1960, 1963; but see Molnar 1982), as many as nine species are known of the long-necked
plesiosaurs and short-necked pliosaurs, of which the enormous Kronosaurus (Fig. 3A) is
undoubtedly the most spectacular (see Molnar, this volume). Incomplete plesiosaur remains
have been reported from the Lower Cretaceous of South Australia (Freytag 1964) and
apparently, the Upper Cretaceous of West Australia (Teichert & Matherson 1944).
Leptocleidus-like plesiosaurs of supposed Cretaceous age from fresh-water sediments in
Queensland (Bartholomai 1966) now are thought to be of Jurassic age (Molnar, this volume).

Table 1. Australian Cretaceous marine reptiles and their stratigraphic distribution. (Based on references
given below and in text).

Santonian (Late Cretaceous)
Mosasauridae genus and species indet., Lundelius and Warne 1960) (Squamata)
Plesiosauria genus and species indet., Teichert and Matheson 1944

Albian (-Santonian ?) (late Early to possibly Late Cretaceous)
Platypterygius australis (McCoy, 1867), McGowan 1972a, 1972b (Ichthyosauria;
Stenoptery giidae).

Albian (late Early Cretaceous)
Cratochelone berneyi Longman, 1915 (Chelonia, Cheloniidae)
Notochelone costata (Owen, 1882) (Chelonia, Cheloniidae)
Cimoliasauridae genus and species indet. (2), Persson 1960 (Plesiosauroidea)
Woolungasaurus sp. Persson 1960 (Plesiosauroidea, Elasmosauridae)
Kronosaurus queenslandicus Longman, 1924 (Pliosauroidea, Pliosauridae)

Albian-Aptian (early Cretaceous-late Early Cretaceous)
Elasmosauridae genus and species indet. (1-4), Persson 1960 (Plesiosauroidea)

Aptain (Early Cretaceous)
Cimoliasaurus maccoyi Etheridge, 1904 (Plesiosauroidea, Aptian)
Woolungasaurus glendowerensis Persson, 1960 (Plesiosauroidea, Elasmosauridae)
Dolichorhynchops ? sp. Persson 1960 (Pliosauroidea, Polycotylidae)
Polycotylid genus and species indet. (1), Persson 1960 (Pliosauroidea)

The many supposedly distinct species of Australian ichthyosaur apparently belong to only
one species, Platypterygius australis (McCoy 1867) (McGowan 1972a, 1972b). The reported
stratigraphic range is Albian (Early Cretaceous) and apparently, Santonian (Late Cretaceous).
Specimens are known from the Northern Territory, Queensland and West Australia. The only
described Australian mosasaur appears to be an indeterminate specimen from the Upper
Cretaceous of Western Australia (Lundelius & Warne 1960). Two species of marine turtle,
Cratochelone berneyi Longman 1915, and Notochelone costata (Owen 1881) are known from
the Albian (Lower Cretaceous) of Queensland (Molnar, this volume).

The New Zealand Cretaceous Record

Nearly all of the known New Zealand Cretaceous marine reptiles (Table 2) are Late
Cretaceous species. The only earlier records appear to be those of indeterminate ichthyosaurs
from the Upper Albian (Fleming et al. 1971). These specimens may belong to Platypterygius
(Stenopterygiidae) which, according to McGowan (1972a, 1972b), is the only known genus of

Cretaceous ichthyosaur.

1174 - FORDYCE
——=>>>>>>>S——_——EL——EEEEESESEESESESSSSS

Table 2. New Zealand Cretaceous and Tertiary marine reptiles and their stratigraphic distribution. Based on
Fordyce 1982a.

Otaian (Early Miocene)
Cheloniidae n. sp. Buckeridge 1982

Whaingaroan (Early Oligocene)
Chelonia genus and species indet., Fordyce, 1979.

Teurian (Early Palaeocene)
? Chelonia genus and species indet. Fordyce, 1979.

Haumurian (latest Cretaceous)
Mauisaurus haasti Hector, 1894, Welles & Gregg 1971 (Plesiosauroidea,
Elasmosauridae)
Elasmosauridae genus and species indet., Welles & Gregg 1971 (Plesiosauroidea)
Polycotylidae genus and species indet., Welles & Gregg 1971 (Pliosauroidea)
Tylosaurus haumuriensis (Hector, 1874), Welles & Gregg 1971 (Squamata,
Mosasauridae)

Taniwhasaurus oweni Hector, 1874, Welles & Gregg 1971 (Squamata, Mosasauridae)
Prognathodon waiparaensis Welles & Gregg, 1971 (Squamata, Mosasauridae)
Mosasaurus mokoroa Welles & Gregg, 1971 (Squamata, Mosasauridae)

Piripauan-Haumurian (Late Cretaceous)
Moanasaurus mangahouangae Wiffen, 1980 (Squamata, Mosasauridae)
Protostegidae genus and species indet., Wiffen 1981 (Chelonia)

Motuan (late Early Cretaceous)
Ichthyosauria genus and species indet., Fleming et al. 1971 (Stenopterygiidae)

ES65VGaP=a==NweaN—aoaoaoazqelqwqyvveeanananaeaea_e_e_l_emememlmlee—e_eeee

At least three species of plesiosaur and five species of mosasaur are known from the
Haumurian Stage (Maastrichtian, latest Cretaceous) and, in one case, the Piripauan Stage
(Campanian, Late Cretaceous) of New Zealand. Welles & Gregg (1971) reviewed in detail the
many nominal species that had been described up to 1971. Wiffen (1980) established a new
genus and species, Moanasaurus mangahouangae, for one specimen from a recently-discovered
rich vertebrate site in the North Island, and recently also described an indeterminate species of
protostegid turtle (Wiffen 1981). Latest Cretaceous elasmosaurid and mosasaurid teeth were
mentioned by Keyes (1981).

The New Zealand fauna is interesting for two reasons. Firstly, it is one of the most
diverse, and promises to become one of the best known, latest Cretaceous large marine reptile
faunas. Secondly boundary facies changes and unconformities are as marked as those in many
other localities. Thus, the New Zealand record eventually may help refine interpretation of the
Cretaceous Tertiary boundary event discussed below.

Interpretation of the Australian Cretaceous Record

None of the Australian Cretaceous marine reptile records has been shown to be of great
palaeozoogeographic importance. However, they do support the hypothesis that a catastrophic
event caused massive and worldwide terminal Cretaceous extinctions of large reptiles and many
other marine and terrestrial animals. Many hypotheses have been proposed to account for the

THE AUSTRALASIAN MARINE VERTEBRATE RECORD - 1175

extinctions, for example, a single cause or a random or cyclical coincidence of causes such as
warming, regression, lower continental relief and aridity, decreased oceanic nutrients, salinity
changes, magnetic reversals, increased vulcanism, a supernova, and impact of an extraterrestrial
body. The most recently proposed, that extinctions were caused by the impact of an extra
terrestrial body, also seems to be the most plausible. Such an impact could account for the
recently recognized anomalous enrichment of rare earth elements right at the Cretaceous -
Tertiary boundary. Alvarez et al. (1980) proposed that a large asteroid collided with the earth,
and the impact generated a vast amount of dust which, once it reached the upper atmosphere,
drastically reduced the amount of solar energy reaching the earth's surface. A consequent radical
decrease in photosynthesis eventually would have resulted in massive extinctions. Hsu (1980)
proposed, in an alternative model, that the impact of a comet generated cyanides and heat.
Cyanide would have caused poisoning of marine biotas which, in turn, would have upset CO?
equilibrium. Increased atmospheric CO? resulted in "greenhouse" heating, which affected both
plankton and terrestrial plants, and led to massive extinction. For references on the Cretaceous
- Tertiary extinctions in addition to those mentioned above, see Haq (1981) and Herman (1981).

The Australasian Tertiary Record

The published record of Australasian Tertiary marine reptiles is inconsequential. It is worth
noting that old New Zealand records of large marine reptiles supposedly from the basal Tertiary
are incorrect (Welles & Gregg 1971; Fordyce 1979), and this lends weight to the extinction
hypotheses mentioned above. Two marine turtles are known from fragments, as yet not
formally described, from the Palaeocene and Lower Oligocene of New Zealand (Fordyce 1979).
The description of a new species of cheloniid is in press (Buckeridge 1982). A supposed
crocodile from the New Zealand Tertiary was mentioned in print over a hundred years ago, but
has been shown to be a whale (Fordyce 1979).

BIRDS

Penguins (Order Sphenisciformes) constitute by far the most important group of Australian
marine birds, and they are the only ones discussed here. Other Australasian marine birds are
discussed elsewhere in this volume (Rich & van Tets 1982, Fordyce 1982). Important recent
references on Australasian fossil penguins are those of Jenkins (1974) and, particularly,
Simpson (1975). Simpson reviewed all major papers on Australasian fossils (e.g. those of
Marples 1952, and Simpson 1957, 1970, 1971a). The Australasian stratigraphic record shown
in Tables 3 and 4 is derived from these articles.

Summary of the stratigraphic Record of Penguins

The stratigraphic record of penguins extends from the Late Eocene to Recent, and is
exclusively Southern Hemisphere. Fossils are known from South Africa, South America and
the Antarctic Peninsula, as well as Australia and New Zealand. The earliest known specimens
are structurally similar to later penguins, and thus provide little clue as to the nature of their
ancestors. Australasian Late Eocene species are very large, and apparently are similar to those
from Antarctic Peninsula (see below). Only two indeterminate species have been reported from
the Oligocene of Australia (Table 3), whereas the New Zealand fauna (Table 4) is diverse and
well preserved. At least 13 species are known, of which four are Early Oligocene and nine Late

1176 - FORDYCE
—>>Il"™"l"liII—SI—S i S____—=a=——__

Table 3. Australian fossil penguins (Sphenisciformes) and their stratigraphic distribution. Based mainly on
Simpson (1975) : 22) and Jenkins (1974). Zone correlations follow Abele et al. (1976: Table 8.1).

Cheltenham (latest Miocene) or older
? Pseudaptenodytes minor Simpson, 1970
Pseudaptenodytes macraei Simpson, 1970
Spheniscidae genus and species indet., Simpson 1970

Longfordian (Globigerinoides trilobus trilobus zone, Jenkins 1974 : 292; Early Miocene)
Anthropodytes gilli Simpson, 1959

Janjukian (Globigerina labiacrassata zone?, Jenkins, 1974 : 292; early Late Oligocene)
Spheniscidae genus and species indet. (2), Glaessner, 1955; Simpson 1957

Aldingan (Late Eocene)
Anthropornis nordenskjoeldi Wiman, 1905 (= Pachydyptes simpsoni Jenkins, 1974;
R.J.F. Jenkins, pers. comm.)
Palaeeudyptes sp., Finlayson 1938, Glaessner 1955, Simpson, 1971a: 344, Jenkins
1974.

aaaaananananjap2jowsxwx#w00——=0—0—0——

Oligocene in age. As with Cetacea (see below), the Duntroonian (early Late Oligocene) was a
time of high diversity. The Australasian Miocene record is poor, particularly during the Middle
Miocene. No species of unequivocal Miocene age have been described from New Zealand, and,
from Australia, only one Middle Miocene and two latest Miocene species have been reported.
A few specimens of modern appearance are known from the Plio-Pleistocene of New Zealand.

Interpretation of the Penguin Record

Because penguins and dolphins share some ecological attributes (they are endothermic
marine carnivores and overlap in size), it would be expected that penguins would have been
influenced by climatic and geographic changes similar to those that influenced cetacean
evolution. These factors are discussed below, under Cetacea, and thus will not be elaborated on
here (see also comments by Simpson 1975: 37). There is certainly scope for further
investigation of parallels between penguin and cetacean evolution, for example, the high early
Late Oligocene diversities in both groups and their possible palaeoclimatic implications.
Temperature adaptations of penguins have been discussed widely by other authors, particularly
from the point of view of correlation of temperature with body size. It is possible that
penguins always have been even more cold-adapted than Cetacea, for the restriction of penguins
to the Southern Hemisphere suggests that tropical warm waters have always reinforced an anti-
tropical distribution. Simpson (1975: 38; cf. Jenkins 1974) considered it probable that known
Eocene-Miocene penguins were adapted to markedly higher environmental temperatures than
most Recent penguins, but it is difficult to reconcile this notion with the evidence of
apparently marked antitropical distributions or of low temperatures during Early Oligocene (see
section on Cetacea for discussion; also Fleming 1979; Fig. 13). More fossils and more
palaeotemperature data are needed in order to further elucidate the relationships between penguin
evolution and palaeoclimates.

Penguin palaeozoogeography has been discussed only in fairly general terms. Simpson
(e.g. 1971b: 384, 1975: 39) stressed that marked similarities exist at the generic level between
Late Eocene penguins from New Zealand and those of supposed Late Eocene age from Seymour

THE AUSTRALASIAN MARINE VERTEBRATE RECORD - 1177

Table 4. New Zealand fossil penguins (Sphenisciformes) and their stratigraphic distribution. (Based mainly
on Simpson 1975 ; 23).

Okehauan (Pleistocene)
Megadyptes antipodes (Hombron & Jaquinot), Fleming 1979 : 75.

Pleistocene (stage unspecified)
Eudyptula minor (Forster), Grant-Mackie & Simpson 1973 : 441.

Otaian (Early Miocene) - Waipipian (Late Pliocene) (see text)
Aptenodytes ridgeni - Simpson, 1972
Pygoscelis tyreei Simpson, 1972
Marplesornis novazealandiae (Marples, 1960)

Waitakian (late Late Oligocene)
Platydyptes amiesi Marples, 1952
Korora oliveri Marples, 1952

Duntroonian (early Late Oligocene)
Palaeeudyptes sp. (2), Simpson 1971a, 1975
Duntroonornis parvus Marples, 1952
Archaeospheniscus lopdelli Marples, 1952
Archaeospheniscus lowei Marples, 1952
? Platydyptes marplesi Simpson, 1971a

Duntroonian or Whaingaroan (Late or late Early Oligocene)
Genus and species indet. (3), Grant-Mackie & Simpson 1973

Whaingaroan (Early Oligocene) to Waitakian?
Palaeeudyptes antarcticus Huxley, 1859

Runangan (late Eocene)
Pachydyptes ponderosus Oliver, 1930
Palaeeudyptes marplesi Brodkorb, 1963
Palaeeudyptes sp. (2), Simpson 1971a, 1975 (includes one species reworked into
Waitakian).

ee ___ ee

Island, Antarctic Peninsula, but noted that no conspecifics are known (unfortunately,
synapomorphies were not identified.) He suggested that specific differences could reflect one or
all of the factors of different geological age, wide geographic separation, and ecological
differences (e.g. temperature adaptations). However, Zinsmeister (1979) concluded that
molluscan faunas from New Zealand and Seymour Island show evidence of gradual isolation
from the Palaeocene onwards, and indicate the complete isolation of Seymour Island from
Australasian influence before the Late Eocene. The discovery of accurately dated and more-
complete fossil penguins may help resolve the nature of Australasian-Seymour Island

relationships.

1178 - FORDYCE
MARINE MAMMALS

Fossil Cetacea (whales, dolphins and porpoises) constitute the most conspicuous
Australasian fossil marine mammals, and thus are dealt with in some detail. First, however, I
will consider the two other groups represented as fossils: the sea-cows and seals.

Sirenia

The only Australian fossil sea-cow (Order Sirenia - manatees, dugongs) reported in the
literature appears to be a scrap of skull on which was based the name Chronozoon australe de
Vis, 1883. The specimen, which is from the fresh-water Pliocene Chinchilla Drift, Darling
Downs, New South Wales, appears to be indeterminate (Reinhart 1976). Fossil and extant
sirenians are known to be both fresh-water and marine, but which habitat the Australian fossil
normally occupied is unknown.

Seals: the Stratigraphic Record and its Interpretation

Seals (Order Carnivora) encompass fossil and living true (or earless) seals, walruses, fur-
seals and sea lions. The true seals constitute the Family Phocoidea (sometimes placed in a
Superfamily Phocoidea), while the walruses (Family Obedenidae; not considered here) and fur-
seals and sea-lions (Family Otariidae) and other extinct groups are placed in the Superfamily
Otarioidea. Formerly, phocids and otariids were united in the Order Pinnipedia, but it is known
now that many similarities between phocids and otariids are convergent, and these two groups
are best placed within the Carnivora (e.g. Tedford 1977). There is not yet consensus on the
taxonomic ranks of phocids and otariids.

Fur-seals and sea-lions, which were reviewed recently by Repenning and Tedford (1977), are
represented in Australasia only as relatively young fossils. Partly on these grounds, it has been
proposed that otariids entered Southern Hemisphere waters as recently as the Early Pleistocene
(Repenning and Tedford 1977, Repenning et al. 1979). Gill (1968) discussed the occurrence
and palaeoclimatic significance of the skull of a fossil sea-lion, Neophoca cinerea, from the
Late Pleistocene of Queenscliff, Victoria. The specimen originally was described by McCoy
(1877) as a new species, Arctocephalus williamsi. Gill noted that the extant Australian sea-
lion, N. cinerea, presently lives in South and Western Australia waters to the north of and
warmer than those of Queenscliff, where the fossil was found. For this reason, he suggested
that the fossil occurrence could be interpreted as evidence of a southern migration during a
phase of interglacial warmth. Equally as likely, however, is that this single occurrence
represents a chance extralimital record, and is of no palaeoclimatic importance. A Late
Pleistocene species of Neophoca has been found in New Zealand (Fleming 1968). Another
New Zealand otariid, named _Arctocephalus caninus by Berry (1928), was thought for some
time to be of Pliocene age. In fact, it is a subrecent specimen (under 1,000 years old; Weston
et al. 1973) of the extant sea-lion, Phocarctos hookeri. To conclude this section on otariids, it
is noteworthy that the future discovery of only accurately dated pre-Pleistocene otariid from
Australasia could allow radical revision of the models of otarioid palaeozoogeography discussed
by Repenning & Tedford (1977) and Repenning er al. (1979),

Discoveries made over the last few years in Peru and South Africa (de Muizon & Hendey
1980), and within the last year in Australia, indicate that the history of true seals (Phocidae) in
the Southern Hemisphere extends back further and is more complex than recognized formerly.
The Australian specimens, which have not been described yet, encompass a few diagnostic
bones, mostly discovered by Timothy F. Flannery, that are held by the National Museum of
Victoria (NMV). These include P41759, two fused, eroded sacral vertebrae (from Beaumaris);

THE AUSTRALASIAN MARINE VERTEBRATE RECORD - 1179

P160399, a relatively complete right temporal (Beaumaris; see Fig. 3B); P160433, eight
vertebrae and four ribs (Beaumaris); and P160441, incomplete right temporal (from Hamilton).
Another specimen, P16198 (an isolated tooth; Beaumaris) was listed by Gill (1957: 181) as the
tooth of a squalodontid cetacean, but it may be a phocid incisor. The provenance of none of
these specimens is in doubt. All the Beaumaris specimens (see Gill 1957: text-fig. 11, for
map) came from within or just above a nodule bed at the base of the Black Rock Formation
(sensu Abele et al. 1976: 241). The age is probably Cheltenhamian, possibly Kalimnan, latest
Miocene-earliest Pliocene. The Hamilton specimen, P160441, was from a coquina (Gill
1957): Fig. 6) within the Grange Burn Formation (sensu Abele et al. 1976: 215), near locality
8 of Gill 1957: Fig. 3. The age is probably Kalimnan. The Grange Burn Formation is
succeeded by basalt that has been dated at about 4.45 myBP (revision of Turnbull er al. 1965,
by T. H. Rich, pers. comm). Of the above specimens, only the two temporals are definitely
phocid; the other material is probably phocid, but this has yet to be demonstrated. The two
temporals (Fig. 4) appear to be very similar to those of the only described Southern
Hemisphere temperate fossil, the South African Early Pliocene monachine, Homiphoca
capensis (Hendey & Repenning, 1972) (see also de Muizon & Hendey 1980). The exact
relationships of the Australian specimens have yet to be determined. If they do belong in or
close to Homiphoca, this indicates an important range extension, and, whatever the affinities,
the specimens are the first significant fossils reported from the Indo-Pacific region.
Presumably, the main dispersal mechanism of such forms in the Southern Hemisphere was by
means of the Circum-Antarctic Current.

The most significant described fossil phocid from New Zealand is the mandible of a Ross
seal, Ommatophoca rossi, of Early Pleistocene age (King 1973). The extant Ross seal lives in
pack ice around Antarctica, and this far northern Early Pleistocene record supports evidence
provided by other fossils that the New Zealand climate at that time was much colder than at
present. Other fossil phocids were mentioned by Fleming (1968; see also Fordyce, this
volume).

For further reading on the evolution and palaeozoogeography of seals, see Ray 1977,
Repenning 1977, Repenning &Tedford 1977, Repenning et al. 1979, and Tedford 1977.

Cetacea: the Stratigraphic Record and its Interpretation

Cetacea (whales, dolphins, porpoises) have a relatively good record in Australasia, although
much of this is still undocumented. Tables 5 and 6 summarise the stratigraphic record, based
both on published records and unpublished observations (those in Table 6 based on Fordyce
1982a). The tables give all necessary references to systematics and stratigraphic distribution,
and only the more important references will be given in the text below. The discussion here
will consider general trends in the global record of Cetacea from the Eocene to Recent, with
Australasian examples, then will consider palaeogeographic and palaeoclimatic implications of
the record. Some basic features of cetacean biology, which it is necessary to understand in
order to interpret fossils, are not outlined here, but are summarised, for example, in Fordyce
(1980a) and references therein.

The Stratigraphic Record

The oldest and most primitive whales, Suborder Archaeoceti - archaic toothed whales,
probably arose from an early ungulate group, the Mesonychidae. The earliest archaeocetes are
the protocetids, which are represented by teeth anda few skull and postcranial fragments from

1180 - FORDYCE

Table 5. Some Australian fossil Cetacea and their stratigraphic distribution. (Based on Fordyce 1982b and
personal observations, or on references cited).

Holocene ;
Delphinus delphis; Gill 1965 (Odontoceti, Delphinidae)

Age uncertain, but possibly Kalimnan (Early Pliocene)
Scaptodon lodderi Chapman 1918 (Odontoceti, Physeteridae)

Cheltenhamian to Kalimnan (latest Miocene to Early Pliocene)
Physetodon baileyi McCoy 1879 (Odontoceti, Physeteridae)
Scaldicetus macgeei Chapman 1912 (Odontoceti, Physeteridae)
Scaldicetus lodgei Chapman 1917 (Odontoceti, Physeteridae)
cf. Physeter sp., Fordyce 1982b (Odontoceti, Physeteridae)
"Steno" cudmorei Chapman 1917 (Odontoceti, Delphinidae)
Mesoplodon longirostris (Cuvier 1823) (= Mesoplodon compressus =
? Belemnoziphius compressus Auct.), Chapman 1917, Glaessner 1947 (Odontoceti,
Ziphiidae)
Mesoplodon spp. (2 or more), (Odontoceti, Ziphiidae)
“Ziphius sp., Sutherland and Kershaw 1971 : 159 (Odontoceti, Ziphiidae)
Odontoceti genus and species indet. (1 or more)
cf. Megaptera sp. (Mysticeti, Balaenopteridae)
cf. Balaenoptera spp. (2 or more) Mysticeti, Balaenopteridae)
cf. Balaena sp., Gill 1957 : 181 (Mysticeti, Balaenidae)

Batesfordian-Balcombian (latest Early to early Middle Miocene)
Rhabdosteidae genus and species indet. (Odontoceti)

Longfordian (Early Miocene)
Squalodontidae genus and species indet.
Prosqualodon davidis Flynn 1923, Flynn 1948 (Odontoceti, Squalodontidae)
Physeteridae genus and species indet.
Cetotheriidae genus and species indet. (= "Aglaocetus ? sp. nov." of Glaessner
1955) (Mysticeti)

Janjukian (Late Oligocene to earliest Miocene)

Mammalodon colliveri Pritchard 1939 (provisionally
Mysticeti Family incertae sedis)

Metasqualodon harwoodi (Sanger 1881); Pledge and Rothausen 1977 (Odontoceti,
? Squalodontidae)

Parasqualodon wilkinsoni (McCoy 1866) (possibly conspecific with
Prosqualodon davidis); (Odontoceti, Squalodontidae)

"Squalodon" gambierensis Glaessner 1955 (Cetacea incertae sedis).

—uquqoqKFR>>5S& ccc

Middle Eocene sediments of the Tethys Sea. No protocetids, nor any Middle Eocene whales,
have been reported from the Southern Hemisphere.

A more advanced family of archaeocetes, the Basilosauridae (or zeuglodons), was the dominant
Late Eocene group. Basilosaurids are well known as a result of Kellogg's (1936) work on
predominantly Northern Hemisphere material. The only reasonably well identified and
reasonably well dated Southern Hemisphere archaeocete is as-yet undescribed specimen of aff.
Dorudon of Late Eocene age, from New Zealand (Fordyce 1979: 739). Other specimens from
New Zealand may represent Late Eocene archaeocetes (Fordyce 1980b: 325). Neither
unequivocal archaeocetes nor Eocene Cetacea have yet been reported from Australia.

THE AUSTRALASIAN MARINE VERTEBRATE RECORD - 1181

Late Oligocene Cetacea are very rare, and all are problematic forms. The only taxon of as-
yet undisputed Early Oligocene reported outside Australasia appears to be the Ukrainian
archaeocete Platyosphys Kellogg, 1936. At least three Early Oligocene taxa are known from
New Zealand: one mysticete and two toothed Cetacea of uncertain age by Keyes (1973), but
descriptions of the other taxa have not been published formally.

Two Australasian specimens, both single isolated teeth, constitute the holotypes or named
species or uncertain late Early to early Late Oligocene age (for ages, see Pledge & Rothausen
1977: 287, and Fordyce 1980a: 33). "Squalodon” serratus Davis, 1888, is from New Zealand
while "S.” gambierensis Glaessner 1955, is from South Australian. Glaessner (1955, 1972)
indicated little doubt about the validity of assignment to Squalodon (which, in the strict sense
is known only from advanced squalodontids from the North Atlantic and European Miocene),
and this potentially has important biogeographic implications (mentioned below).

The start of the Late Oligocene saw the worldwide, seemingly sudden appearance of diverse
faunas of odontocetes (modern toothed whales) and mysticestes (baleen whales) (for reviews see
Whitmore & Sanders 1977, Fordyce 1980b). Species diversity, based on the minimum known
number of species, jumped from about four in the Early Oligocene to perhaps 30 in the early
Late Oligocene and about 45 in the later Late Oligocene (pers. obs.). Many of these Late
Oligocene species are Australasian).

The New Zealand earlier Late Oligocene fauna (Duntroonian Stage; see Stevens 1980 for
correlation) is diverse, It includes one supposed archaeocete (Kekenodon onamata), at least two
odontocetes, and at least four mysticetes (Table 6). The later Oligocene fauna (Waitakian
stage, which possibly may be partly earliest Miocene) encompasses at least eight species of
odontocete (including Squalodontidae, possible Kentriodontidae and possible Rhabdosteidae) and
at least two mysticetes, including Mauicetus parki. The abundance of fragmentary and mostly
unstudied specimens of Duntroonian - Waitakian age indicates that actual species diversity in
New Zealand at this time eventually will be found to be much higher than the present
published record suggests. Valid, described Australian species of Late Oligocene age (equals
local Janjukian Stage) encompass only three toothed Cetacea (Table 5.). Of these,
Mammalodon colliveri (a proto-mysticete?; Fordyce 1982b) is of very latest Oligocene or
earliest Miocene age, and Parasqualodon wilkinsoni may be conspecific with the earliest
Miocene Prosqualodon davidis. A higher species diversity than this for the Australian Late
Oligocene is suggested by the presence of other undescribed taxa in museum coilections (e.g.
Cetotolites spp. of McCoy 1879), the names of which, incidentally, are almost certainly
nomina dubia, and described material refereed to established species (e.g. some specimens
mentioned by Hall 1911).

On a global scale, Neogene and Quaternary Cetacea generally are known much better than
their Palaeogene antecedents. One could generalise that the Oligocene, as well as being the
time of first appearance of odontocetes and mysticetes, was a time of radical ecological
experimentation that resulted in many bizarre forms. In contrast, the Neogene saw a gradual
modemisation and a reduction in family diversity, as families which have persisted to the
present established themselves rapidly, presumably at the expense of other forms. For
example, the fossil record of many extant families (¢.g. Physeteridae, Ziphiidae, Platanistidae
and Balaenidae in the Early Miocene; Delphinidae, Phocoenidae, Stenodelphidae, and
Balaenopteridae in the Middle Miocene; Barnes 1977, Fordyce 1980b and personal observations)
indicates the rapid establishment of modern forms by the Middle Miocene, with radiations
occurring in the Early Miocene, if not the latest Oligocene, Some taxa of "archaic" appearance,
e.g. Squalodontidae and Rhabdosteidae, were widespread during the Miocene but disappeared by
the Pliocene, and no unequivocal archaeocete is known from the Neogene. Miocene Cetacea
probably constitute the greatest number of named taxa (see Orr & Faulhaber 1975: Fig. 4), and
this could give the impression that they are very well known. However, many named species
are based on fragmentary types, and are probably nomina dubia. For example, the recent

1182 - FORDYCE

a EE EEE ee
Table 6. Some New Zealand fossil Cetacea and their stratigraphic distribution. (Based on references cited in
Fordyce 1982a and this volume).

Holocene and Pleistocene, states uncertain.
Berardius sp. (B.. arnuxii) Odontoceti, Ziphiidae)

Nukumaruan (Early Pleistocene)
cf. Orcinus sp. (Odontoceti, Delphinidae)
Balaenidae? genus and species indet. (Mysticeti)

Opoitian (Early Pliocene) to Nukumaruan (Early Pleistocene)
Delphinus aff. delphis (Odontoceti, Delphinidae)
cf. Pseudorca sp. (Odontoceti, Delphinidae)
Physeteridae genus and species indet. (Odontoceti)

Opoitian (Early Pliocene)
Balaenopteridae genus and species indet. (Mysticeti)

Miocene and/or Pliocene, stages uncertain
Delphinidae genus and species indet. (Odontoceti)
Balaenidae genus and species indet. (Mysticeti)
cf. "Plesiocetus" dyticus Cabrera (=mysticete “allied to Balaenoptera", Hector 1881)
(Mysticeti).

Late Miocene or late Middle Miocene, stage uncertain
Ziphiidae genus and species indet., Fordyce and Cullen 1979 (Odontoceti)

Altonian (late Early Miocene)
Phocaenopsis mantelli Huxley 1859 (Odontoceti, ?Rhabdosteidae)

Otaian-Altonian (Early Miocene)
Tangaroasaurus kakanuiensis Benham 1935a (Odontoceti, ?Squalodontidae)

Waitakian-Otaian (late Late Oligocene-Early Miocene)
Rhabdosteidae? genus and species indet. (Odontoceti)

Waitakian (late Late Oligocene)

“Prosqualodon" hamiltoni Benham 1937b (possibly includes “Sgualodon" andrewi
Benham, 1942 (Odontoceti, Squalodontidae).

"Prosqualodon" marplesi Dickson 1964 (Odontoceti, ?Squalodontidae)

"Microcetus" hectori Benham, 1935b (Odontoceti, ?S qualodontidae)

Prosqualodon aff. davidis (Odontoceti, Squalodontidae).

cf. Phoberodon sp. (Odontoceti, Squalodontidae)

Kentriodontidae genus and species indet. (Odontoceti)

Odontoceti genus and species indet. (= "Squalodon" andrewi Benham 1942:
"Clarendon teeth")

Mauicetus parki (Benham 1937a) (Mysticeti, Cetotheriidae)

"Mauicetus" brevicollis Marples, 1956 (Mysticeti, Family incertae sedis)

Duntroonian (early Late Oligocene)
Kekenodon onamata Hector 1881 (Cetacea Incertae sedis)
Austrosqualodon trirhizodonta Climo & Baker 1972 (Odontoceti, ?Squalodontidae)
"Microcetus" aff. hectori (Odontoceti, ?Squalodontidae)
“Mauicetus" lophocephalus Marples 1956 (Mysticeti, Family incertae sedis)
"Mauicetus" waitakiensis Marples 1956 (Mysticeti, Family incertae sedis)

THE AUSTRALASIAN MARINE VERTEBRATE RECORD - 1183

Mysticeti genus and species indet., Fordyce 1982a.

Whaingaroan-Duntroonian (Early to Late Oligocene)
"Squalodon" serratus Davis 1888 (Cetacea incertae sedis)

Whaingaroan (Early Oligocene)
Cetacea genus and species indet., at least 2 species, including that described by Keyes
1973
Mysticeti genus and species indet.

Bortonian-Runangan (Middle to Late Eocene; probably Runangan, Late Eocene)
aff. Dorudon sp. (Archaeoceti, ?Basilosauridae)

thorough and useful appraisal by Barnes (1977) of Californian Neogene Cetacea indicates both
the need for revision of previously-named taxa and the presence of many new species. This
cautions against undue generalising about the Neogene record.

Cetacea from New Zealand Neogene (Table 6) have been studied relatively less than either
those from the New Zealand Oligocene or Australian Neogene, and thus warrant little
discussion. Described specimens encompass a squalodontid, a possible rhabdosteid (both Early
Miocene) and an indeterminate Middle or Late Miocene ziphiid. Many undescribed specimens
have been collected (Table 6, see also discussion of New Zealand fossil vertebrates by Fordyce
1982a and this volume), and there is great potential for future research.

Australian Neogene Cetacea encompass described species that range from important
complete and informative specimens to indeterminate fragments (Fordyce 1982a).
Prosqualodon davidis (Early Miocene, Tasmania; Flynn 1948) is one of the best-known species
of Squalodontidae. The only supposed cetothere reported from Australia, "Ag/aocetus? sp.
nov.” of Glaessner (1955) also may be of Early Miocene age. Two undescribed specimens, a
large squalodont and a sperm-whale (Physeteridae) have been collected from Batesford Quarry
(Longfordian, Early Miocene). Fragmentary specimens of small dolphins from the Middle
Miocene fluviatile-lucustrine Namba Formation of Lake Frome area represent an indeterminate
species of extinct "river-dolphin", Rhabdosteidae.

Latest Miocene assemblages from Beaumaris and Grange Burn (near Hamilton), Victoria,
are more or less contemporaneous and identical and can be considered together. The age of this
fauna was discussed earlier, under the section on phocid seals, The cetacean fauna is probably
the most diverse yet known from Australia, although study is difficult because nearly all
specimens are broken and eroded, and thus often not comparable. A list of known taxa is as
follows: (see also Table 5) beaked whales, Ziphiidae, known from rostra (Chapman 1917; see
also Glaessner 1947); large and small sperm whales, Physeteridae, known from teeth, earbones,
and skull fragments (three described nominal species based on isolated teeth, and a species close
to the extant Physeter macrocephalus); small odontocetes represented by teeth, mandibles and
earbones, including "Steno" cudmorei Chapman, 1917 (Delphinidae; probably not congeneric
with extant species of Steno); at least one species of rorqual or fin-whale, Balaenopteridae,
known from skull fragments and eroded earbones; and at least one species of right whale,
Balaenidae, known from eroded earbones. A similar, but younger fauna, from the Middle
Pliocene of Flinders Island, is also under study at present. It includes ziphiids (at least one
species of Mesoplodon, physeterids (cf. Physeter macrocephalus), and balaenopterids.

Interpretation of the Record

Outlined below are some of the palaeogeographic and palaeoclimatic implications of the
Australasian cetacean record. More detailed discussion was presented by Fordyce (1980b) and

1184 - FORDYCE

implications of the global record were considered by Barnes & Mitchell (1978) and Lipps &
Mitchell (1976).

Archaeocetes apparently reached New Zealand by the Late Eocene. Why, therefore, have
they not yet been reported from Australia? The paucity of Southern Hemisphere Eocene
Cetacea is difficult to interpret. It is unlikely that tropical warm waters formed an equatorial
barrier to dispersal, such as may effect living Cetacea, because Cetacea appear to have evolved
in the relatively warm waters of the Tethys. Furthermore, the absence of any significant polar
ice cap suggests that latitudinal climatic gradients were less pronounced than later in the
Tertiary.

The Early Oligocene cetacean record is truly problematic. The sudden appearance of diverse
faunas of odontocetes and mysticetes about the start of the Late Oligocene indicates that the
Early Oligocene was probably the time of origin and initial rapid diversification of the earliest
representatives of the modern whales. Mysticetes and odontocetes are unlikely to have appeared
before the Oligocene, as their structural, and probably phyletic, antecedents were the
dorundontines - a Late Eocene group. Why, then, is the Early Oligocene record so poor?
Simpson (1945) suggested that the absence of fossils representing some important stages of
cetacean evolution might reflect the fact the evolution occurred (for example, during the Early
Oligocene) in open ocean basins. However, the frequency with which extant open-ocean
species strand suggests that nearly all species should have a significant fossil record in
proximal marine sediments. This suggests that the lack of Early Oligocene Cetacea is
probably a preservation bias. It is known that a massive worldwide regression occurred very
early in Late Oligocene (at about 29 million years ago). This probably resulted in the
worldwide erosion of much of the proximal lower Oligocene, in which cetacean fossils were
preserved. Thus, the absence of many fossils is still compatible with the idea that odontocetes
and mysticetes arose during the Early Oligocene. Incidentally, a large sea-level drop in the Late
Oligocene has important ecological implications, which are considered briefly in the
conclusion of this article.

A climatic-change model can be used to explain the origins of the ancestors of modern
whales (Fordyce 1980b). In order to understand this, it is necessary to briefly consider the
adaptations of modern whales and the record of palaeoclimate change. Modern toothed whales,
the odontocetes, are active predators that catch single prey. They navigate and hunt by means
of echolocation, using high-frequency sound. The ability to echolocate is reflected in unique
skull-structures in living odontocetes, and the presence of similar structures in the skulls of
Oligocene odontocetes indicates that they, too, echolocated. Mysticetes, conversely, are
filterfeeding predators that strain their food in bulk from the water, and are not known to
echolocate. The filterfeeding mechanism is formed by baleen, the presence of which is
concomitant with a skull structure unique to mysticetes. Such a structure is present also in
Oligocene mysticetes, which indicates that they, like extant species, filterfed. These divergent
feeding strategies appear to have evolved in the Early Oligocene. Archaeocetes, from which
odontocetes and mysticetes evolved, lack any structures which could be interpreted as
adaptations for filter-feeding or echolocation. It is evident from this that a model for the
evolution of odontocetes and mysticetes must consider changes in food resources as a primary
factor, and such a model is outlined below.

Global oceans appear to have been relatively warm during the Late Eocene. In the very
latest Eocene, however, there was a dramatic drop in Southern Ocean bottom water temperature
of 4-5°C, and global temperatures generally fell (Savin 1977). This gives a clue to the chain of
events that influenced the evolution of odontocetes and mysticetes. Progressive physical
isolation of Antarctica during the Early Tertiary resulted in thermal isolation, which allowed
significant ice buildup on Antarctica by about the latest Eocene, Antarctica cooling resulted in
the development of the psychrosphere - cold, nutrient-rich deep water which today originates at
Antarctica to flow north into all major oceans of the world. The development of the

THE AUSTRALASIAN MARINE VERTEBRATE RECORD - 1185

psychrosphere had many effects (e.g. Kennett & Shackleton 1976, Keigwin 1980). The
Southern Oceans cooled dramatically. There was also general global cooling, probably as a
consequence of northward flow of the psychrosphere. The presumably novel nutrient-carrying
capacities of the psychrosphere probably caused major regional changes in nutrient turnover.
Polar cooling resulted in enhanced latitudinal thermal gradients which, in tum, may have led to
increased oceanic current activity. Most importantly, it seems that regional (southern?)
productivity changes, which resulted from changing nutrient cycles, resulted in the formation of
many new niches, and thus triggered the evolution of odontocetes and mysticetes (characterised
by new and divergent feeding adaptations) early in the Oligocene. For a more detailed climate
change, see Fordyce (1980b). Similar models for marine mammal evolution were also
discussed by Gaskin (1976) and Lipps & Mitchell (1976).

Southern oceanic climates modemised rapidly late in the Oligocene and in the Neogene (e.g.
Kennett 1980). The record of some fossil Cetacea is consistent with the notion of the early
development of a distinctive Southern (Austral) fauna. For example, closely related species of
Prosqualodon (sensu stricto) have been recorded from the earliest Miocene of New Zealand.
These are compatible with other evidence that the Circum-Antarctic Current was well
established before the Early Miocene.

In view of our presently limited knowledge of other Australasian Neogene Cetacea, little
else can be said about palaeoclimatic and palaeogeographic implications. Perhaps the one
exception is the case of the Middle Miocene rhabdosteids from the Lake Frome area, which are
interesting because they indicate external drainage of this region to the sea in or before the
Middle Miocene. They also indicate that the occupation of fluviatile-lacusirine environments is
not unique to the four extant species of "river dolphins". The relationships, and hence
palaeozoogeographic implications, of these dolphins are uncertain. Rhabdosteids are known
primarily from the Northern Hemisphere Miocene, but also from South America and possibly
New Zealand. The discovery of more-complete Australian rhabdosteids may allow us to tell if
they are part of a distinctive southern fauna, or if they are more closely related to northern
rather than to known southern species.

The nature of the relationships between Northern and Southern Hemisphere Cetacea
(primarily extant species) was discussed in detail by Davies (1963). He observed that many
Cetacea have antitropical distributions; that is, they are discontinuously distributed on either
side of the warm waters of the equator. Sometimes equatorial waters separate different
populations of one species while in other cases, closely related (sister) species are allopatric.
Davies proposed that changes in the extent of equatorial water during the Pleistocene would
have allowed variable interchange between north and south, with maximum exchange occurring
during the maximum (glacial) reduction in extent of tropical waters. Postglacial warming
would have resulted in the reestablishment of tropical barriers and renewed separation of
northern and southern faunas. This model accounts plausibly for the distribution of many
extant species and although Davies did not discuss this, it can probably be applied to the fossil
record. No Oligocene to Pleistocene southern cetacean appears to have reported to be
conspecific with any northern species. In fact, most Southern Hemisphere records of species
that have been placed in Northern Hemisphere genera are probably misidentifications (e.g.
"Squalodon" spp. from Australasia), and this strengthens the possibility that transequatorial
exchange has been limited since perhaps the Oligocene. Only once we have a much better idea
of the southern record, however, will it be possible to make any informed comments on this.

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1190 - FORDYCE

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Oregon

CHAPTER 26

A NEW LOOK AT THE

FOSSIL

VERTEBRATE

RECORD OF NEW ZEALAND

R. Ewan Fordyce!
TV GOPUCHON c-... 2: .sstsnnndnrasregeseeswenge sede 1192
Approach and Geographic Scope........... 1192
COMVENTIONS <<..cccesevertenenscrnentnserioesense 1195
AbDDreviatiOns.............cceeeeeeeeeeeeeeeeeeees 1195
Tiiter atures 2 cacditecadesdshovacuigentedee paSeinde 1195
Collection, Preparation and Curation .....1197
New Zealand Stratigraphy................006+ 1199
The Stratigraphic Record of
VETLCDIaleS..... eee eeeeeeceeeeneeeeeeenees 1203
Systematic Summary of New Zealand
Fossil Vertebrates..............:.666 1215
Fish: OVervieW...........:s::seeeeeeeeeees 1215
PIACOdEIMS 2.02.2... cee eeeeeeeeeeeeeseeees 1215
Chondrichthyes..............::s::sseeeeeeee 1215
FOLEOSIS. .d.ccscccsvssyteandedactacses cabeast’s 1218
Amphibia: Overview ................06++ 1220
Subfossil Frogs: Leiopelmatidae.....1221
Reptilia: Overview..............ceseeeee 1221
TUPtleS: sis ssccssertoushecaiectacscesabetee 1222
IchthyOSaufls............sceeeeseeeeeeeeeeeee 1223
Plesiosaurs (Sauropterygia)............. 1225
Tuatara (Sphenodonta
=Rhynchocephalia)................... 1227
Mosasaurs and Other Lizards
(Squamata) ...........eecceseeseeeesees 1228
Crocodiles (Crocodylia)..............06+5 1229
Dinosaurs (Saurischia and
Ornithischia)................:csseeeeeee 1230
Pterosaurs (Pterosauria)...........2.0006 1230
Miscellaneous and Problematic
Records of Reptiles................5. 1230

AveS: OVECLVICW ........0..ceece eee eeeeees 1231
A Miscellany of Tertiary Birds........ 1231
Bony-Toothed Pelicans
(Pelagornithidae).................006+ 1234
Moas - Dinornithiformes ............... 1236
Other Late Quaternary Bird............. 1241
Penguins (Sphenisciformes) ........... 1243
Mammalia ..........:eceeeeseeeeeeeeeeeeeees 1250
Seals - Pinnipedia ...............066+ 1250
Otartidae ...........ceeeeeee seco ees 1250
Phocidac............seseeeeeeeeees 1252
Other Records..............0066 1252
Whales, Dolphins, Porpoises -
ct: |<; «tO a 1253
Archaeoceti .........0.eeceeee ees 1253
Problematic Cetacea........... 1254
MYSIICCi...... eee eee eee ee eeees 1254
OdONtOcetl ..........cceseeeeeeees 1258
Other Work..........2.:0seeeeees 1262
Broader Issues in New Zealand
Vertebrate Palaeontology.......... 1263
Interpreting the Record ..............26 1263
Approaches to Taxonomy ............. 1263
Completeness of Records............++ 1263
New Zealand Type Specimens......... 1264
EXtiMCtiOn,..........:sssseeeseceeeeseeeseees 1265
Palaeozoogeography.........cceeceee 1267
Acknowledgements ...........s0ssessseeeeeeers 1269
REfCPENCES: oe ccveenveveiedvestentesved dates eaeene 1269
PP PORTIS Ts cicecteveseedaayyecertrniecabestcees 1296
PIA ao tateeceedaicnaaaaetenesatsereeees 1314

1 Department of Geology, University of Otago, Dunedin, New Zealand.

1192 - FORDYCE

INTRODUCTION

New Zealand has long been known to have an unusual modern biota which, like that of
Australia, reflects evolution in relative isolation. Indigenous vertebrates, although not diverse,
are a conspicuous clement of the biota, and there has been a great deal of interest in their
origins. This article reviews New Zealand's fossil and recently extinct vertebrates.

The fossil and recently extinct vertebrates have been the subject of study for just 150 years
by scientists in New Zealand and overseas (Figs 1, 2). Most effort has concentrated on moas,
species of large, flightless, recently-extinct ratite birds which have aroused considerable public
and scientific interest. Ironically, most of these birds disappeared so recently that in many
ways they can be regarded as members of the extant fauna. Other groups which might be more
revealing about ancient New Zealand have received less attention, perhaps because pre-Holocene
higher vertebrates are less common. There was a flurry of early work last century (1860s-
1880s) on marine vertebrates, but this was followed by a long hiatus. The work of Marples in
the 1940s perhaps properly signalled a renewed professional treatment of vertebrate fossils by
resident paleontologists. The prospects are better now for vertebrate palaeontology in New
Zealand than for many years for two reasons. Firstly, there is an increasing public awareness
in larger "educational" fossils such as dinosaurs and in scientific issues such as extinction,
long-term environmental change, and plate tectonics. Secondly, there seems to be increasing
interest from palacontologists overseas in what the New Zealand fossil vertebrate fauna might
tell about the evolution of life in the southwest Pacific.

The general approach here follows my previous review (Fordyce 1982c), which is expanded
and revised. My account is perhaps biased because of an avowed interest in marine mammals
and in South Island geology. Nonetheless, I have kept the text as wide-ranging as possible.
Most of the specimens discussed represent the diverse and sometimes abundant marine
vertebrates, which have been recovered from the relatively complete sequence of Late
Cretaceous to earlier Pleistocene marine sediments and a few older rocks. The enormous topic
of the extinct moa and other Holocene birds is summarised briefly. Millener (this volume)
gives a more detailed review.

APPROACH AND GEOGRAPHIC SCOPE

The aim of this article is to give a broad review of fossil vertebrates from New Zealand ina
manner useful to both undergraduate students and those with more specialised interests in
vertebrates, Thus, it considers the history of study, stratigraphy, preparation and curation,
aspects of morphology or systematics of major groups (under the traditional subdivisions of
fish, amphibians, reptiles, birds and mammals), evolution, extinction and zoogeography.
Published discussion of the latter three topics deals mainly with extant terrestrial birds, frogs,
and the tuatara, and is generally presented in broader context. Examples of more general
accounts that cover these latter topics include those of Cochrane (1973), Stevens (1985),
Stevens et al. (1988) and comments in McKenzie (1987) and Wards (1976). Understandably,
many articles on broader aspects of New Zealand's vertebrates are necessarily based on little
fossil evidence and much speculation. There is not space to give for all major groups details of
morphology, approaches to describing and diagnosing species, philosophy of classification, and
inferred ecology; rather, these are covered by some of the references cited.

This review presents some unpublished information which might better appear in formal
taxomomic works than in a general review. I have included this material to indicate more
realistically the potential breadth of New Zealand vertebrate palaeontology, and to caution about
possible errors or misinterpretations in past published articles, This saves the need to cover
unproductive ground.

FOSSIL VERTEBRATES OF NEW ZEALAND - 1193

F W Hutton

J von Haast

J Hector

Figure 1. Some contributors to the study of New Zealand fossil vertebrates. (Sketch of Hector by J.D.
Campbell; others by Jane Kerr).

1194 - FORDYCE

|W Keyes

Figure 2. Some contributors to the study of New Zealand fossil vertebrates. (Sketches by Jane Kerr).

FOSSIL VERTEBRATES OF NEW ZEALAND - 1195

The references cited are more comprehensive than those presented earlier (Fordyce 1982c).
Some minor references are omitted. It is possible that there are some major omissions, and I
will be grateful to receive any comments on these.

This article deals with fossils from New Zealand proper (Fig. 3): the North Island, South
Island, Stewart Island, and the Chatham Islands (not illustrated), The use and capitalisation of
geographical names follows those given by Wards (1976) or names cited in articles on New
Zealand fossil vertebrates. McKenzie (1987) gave a useful atlas and gazetteer.

CONVENTIONS

This review uses some widely accepted scientific conventions which provide a useful guide
to the status and origins of many names. Scientific names of many New Zealand species are
given as a formal binomen. The binomen (two-part name) has a generic name first and a
species name second, and these are always underlined. An example is Prognathodon
waiparaensis Welles & Gregg, 1971. In this case, Welles and Gregg proposed the new name
Prognathodon waiparaensis in 1971. In other cases, a species may have been redescribed and
placed in a different genus than used by the original author(s). The name of the author is then
placed in parentheses. For example, Mauicetus parki (Benham, 1937a) is a species described
originally by Benham in 1937 as Lophocephalus parki and later placed in the genus Mauicetus.
These taxonomic procedures, together with a useful glossary of taxonomic terms, were covered
in more detail by Ride et al. (1985).

Formal names of rock units are capitalised, e.g. Otekaike Limestone, while informal units
(those not yet properly defined and diagnosed in scientific literature) are not, €.g.
Maungataniwha sandstone. Formal stratigraphic subdivisions are capitalised (e.g. Early
Oligocene), while informal subdivisions are not (e.g. earliest Miocene, Middle Pliocene).

It appears to be ubiquitous in New Zealand to use the spellings Cenozoic and paleo- (as in
paleontology), but these conventions have had to be abandoned temporarily in this contribution
because of editorial requirements.

ABBREVIATIONS

The following abbreviations are used here for New Zealand institutions which hold
significant collections of fossil vertebrates: AUGD - Department of Geology, University of
Auckland, Auckland; CM - Canterbury Museum, Christchurch; NMNZ - National Museum of
New Zealand, Wellington; NZGS - New Zealand Geological Survey, Lower Hutt; OM -
Otago Museum, Dunedin; and OUGD - Geology Museum and Department of Geology,
University of Otago, Dunedin.

Webby et al. (1984) gave a guide to New Zealand institutions that hold fossils, but this
gave little insight into the fossil vertebrate collections of New Zealand.

LITERATURE

The most useful sources of literature on New Zealand fossil vertebrates (apart from that in
the earlier version of this article), including relevant aspects of geology and zoology (Fordyce
1982c), are the bibliographies of Adkin & Collins (1967) and Warren et al. (1977), together
with the indexes by Jenkins (1976, 1982). Noteworthy older works are those of Hamilton
(1903a, 1910), and Thomson (1913). The comprehensive, but un-indexed, bibliography of
fossil vertebrates by Romer et al. (1962) is also valuable.

Other bibliographies of vertebrate palaeontology which list papers published from 1929
(but are incomplete for New Zealand articles) include those of Camp & Allison 1961, Camp &

1196 - FORDYCE

Vanderhoof 1940, Camp et al. 1942, 1949, 1953, 1964, 1968 and 1972, Green et al. 1979, and
Gregory et al. 1973 and 1981. Not all modern bibliographies are available in New Zealand.
Such bibliographies are usefully supplemented by the Zoological Record, Science Citation
Index, and the Geological Society of America Bibliography and Index of Geology Exclusive
of North America, which was succeeded by the American Geological Institute Bibliography and

Index of Geology.

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Pahi O > Mangahouanga
Stream
2 Ohope
Lew4
Waik: . Tokomaru Bay
Kaawa Creek Ror Seay ‘ Raukumara
Ponganui Creek Glen Masse b, *
Meurant Island olulara
: Te Kauri 0
Waitomo Marakopa
Kiritehere
Waikawau Creek
Taihape Hanaroa R.
Tangarakau Waipunga
Atene Te Pohue
Waihi Beach Napier
Nukumaru Matapiro
Otokoa Stream Wanganul Cape Kidnappers
Stephens Island Waipukurau
Puponga
Paturau Coonoor
Aorere River 2 Hukanui
‘ ¥ Tinui district
Nelson Takaka OM
Wairoa " Martinborough
Murchison = Manaroa
Charleston _— Ward
Punakaiki
Reefton
Greymouth —— Kaikoura
Waimea Haumuri Bluff
Pascal Cheviot
arama Gore Ba’
Cc a oe Mot : Waipara River middle gorge
astle Hill basin otunau Weka Pass
Mt Potts Glenafric__
Curiosity Shop Waipara River lower gorge
St Bathans Wharekuri, Aviemore Karetu Quarry
Bannockburn ail Hakataramea
Nevis Valley Ss et
Wakatipu Duntroon, Kokoamu_ juganara
Oamaru gapa
a Kakanui Awamoa, Rifle Butts
SK BKanut Moeraki
# hag Point :
faa Waikoualti Middlemarch
ts 5 Caversham
Kg Burnside
Ceendsn, Milburn
S okomairiro
aaa Nugget Point
F- im Otamita Stream
Balfour
\4 Clifden
0 250 km

Figure 3. Some older fossil vertebrate sites in New Zealand. Only a few Late Quaternary sites are shown,

and the Chatham Islands are not shown.

Parengarenga Harbour

Oruwharo River

FOSSIL VERTEBRATES OF NEW ZEALAND - 1197

Local periodicals which were searched for articles include the Transactions and Proceedings
of the New Zealand Institute and its successors, the Transactions..., Proceedings..., and Journal
of the Royal Society of New Zealand; New Zealand Journal of Science and Technology; New
Zealand Journal of Geology and Geophysics; New Zealand Journal of Science; and various
publications of the New Zealand Geological Survey (Reports of Geological Explorations,
Bulletins, Paleontology Bulletins, Memoirs, Reports, and Records). I did not attempt to
thoroughly cover the periodicals issued by other local organisations (e.g. museums, natural
history societies). More has been published on New Zealand fossil vertebrates than is cited
here in the text, although most references not cited are mostly those on Quaternary birds
(innumerable titles). Page numbers are cited for the many articles that contain only brief or
incidental comment which might otherwise be overlooked because the title does not indicate
that they deal with palaeontology.

Some theses for higher degrees contain relevant material. I have cited some of these even
though they are not strictly publications, but have not referred to (or, in some cases, even read)
the body of the text to get information for this review. Information in such theses is best left
for the authors to publish. Besides, such items have no standing in nomenclature, and are
difficult to consult. Similarly, I have cited as few unpublished reports as possible, and mostly
have mentioned only those that have already been cited in the literature.

There is no thorough published bibliography of New Zealand fossil fish. Bibliographies
compiled for other groups include those of Welles & Gregg 1971 (marine reptiles), Archey
1941 and Hamilton 1894 (moas), Simpson 1971a (penguins), Oliver 1955 (other birds) and
Fordyce 1980b (cetaceans). The bibliography of this review contains those references cited in
the text and a few incidental titles. Full citations of the authors of extant species are not
always given in the text, and these articles are not cited in the bibliography.

Useful overviews of fossil vertebrates are available from a number of sources. Other
chapters in this volume give an Australasian perspective, and a great deal of local information
was presented by contributors to Archer & Clayton (1984). Carroll (1988) provided a key
reference for this review, and the older work by Romer (1966) still has more than historic
value.

COLLECTION, PREPARATION AND CURATION

Many New Zealand fossil vertebrates have been damaged (or are still being damaged!) by
improper collection, preparation and curation. This probably reflects a general lack of local
experience in collecting large fossils. Fortunately, recent books by Rixon (1976) and Croucher
& Woolley (1982) outline modern palacontological techniques applicable to work on New
Zealand fossil vertebrates. The handbook of Kummel & Raup (1965) deals little with
vertebrates specifically, but discusses many basic prob!ems. The Society of Vertebrate
Paleontology Newsletter often mentions new techniques, and the journals Curator and The
Geological Curator sometimes contain useful articles. A brief summary of important
techniques follows, and the chapter by Whitelaw & Kool in this book gives more details.
Further information can be supplied upon request. The Department of Geology at the
University of Otago, which currently has the best equipped laboratories in New Zealand for
preparation of fossil vertebrates, is open to visits from anyone interested in techniques.

New Zealand fossil vertebrates are covered under an Antiquities Act, and legal controls affect
the export of specimens. At present it is possible for private individuals to collect and
maintain private collections in New Zealand. For posterity, rare or unusual specimens are best
curated in an institution that can provide maximum public and scientific access.

It is essential that field data are recorded for each specimen collected. Collectors are urged
strongly to record vertebrate finds in the Fossil Record File maintained by the Geological

1198 - FORDYCE

Society of New Zealand, and it is mandatory to provide a fossil record number for some
purposes (e.g. publication in some journals), Curators in University geology departments, the
Geological Survey and museums can provide details. A detailed account of the file was given
by Sudlow & Edwards (1982).

Most pre-Quaternary vertebrates should be plaster jacketed fully (e.g. Rixon 1976: 41-53)
during excavation. This is particularly so for large or fragile specimens, those from relatively
uncemented sediments, or those from shattered or jointed rocks. It is tempting to consolidate
relatively uncemented matrix (e.g. greensand) in the field, but most New Zealand specimens are
too damp for consolidants to work well. Small concretions generally do not need to be
jacketed. Bones can be excavated in the field with needle, probe, toothbrush, paint-scraper,
knife, awl, hammer, cold chisel, and crowbar, depending on the nature of the matrix and
specimen. For efficient use of time and resources, a combination of chainsaw with masonry
blade, rotary percussion rock drill, and compressed air tools (chipping hammers, air scribes) are
attractive for large scale work.

Consolidants and glues must be used on most bones during laboratory extraction and
preparation. It is important to use agents whose action can be reversed e.g. water-based glues
(PVA - polyvinyl acetate - emulsion) or acetone-based glues and consolidants (polyvinyl
butyral - PVB, e.g. "Mowital" tradename). Polyesters and epoxies, which are strong but more
or less irreversible, should not be used unless it is certain that preparation is complete. A
summary on the application of these chemicals follows (based on Fordyce 1989c), and further
comment is given in the chapter by Whitelaw & Kool in this volume.

PVA emulsion - glue and consolidant: water-based; can be put on damp fossils; thinned
solutions do not penetrate well; flexible; not resistant to acetic acid; reversible (use acetone).

PVB in acetone - consolidant, but thick solution works as a glue; should not be put on
damp fossils; thinned solutions penetrate well, especially if dissolved in an acetone: ethanol
mix; limited flexibility; reasonable resistance to acetic acid; reversible (use acetone).

Polyethylene glycol (carbowax, e.g. molecular weight 4000) - consolidant; water-
compatible, so molten wax or thick aqueous solution can be put on damp fossils; molten wax
penetrates a limited extent unless the fossil is heated; inflexible; not resistant to acetic acid;
reversible (use water or heat).

Polyester resin (fibreglass resin) - glue and consolidant: organic base, with catalyst; works
as a glue for larger surfaces; should not be put on damp fossils; solutions penetrate well
especially if thinned slightly; limited flexibility; good resistance to acetic acid; caution
irreversible,

Epoxy resin (¢.g. "Araldite") - glue: organic base, with catalyst; works as an excellent glue
for small or larger surfaces; should not be put on damp fossils; limited flexibility; poor
resistance to acetic acid; more or less irreversible, but can be broken down slowly by acetic
acid.

Cyanoacrylate (e.g. “Superglue") - glue: organic, ready to use without mixing (catalysed by
residual water on surfaces to be glued); sets in a few seconds; good for field use, since it is
compatible with moisture in fossils; partly reversible, using acetone; limited flexibility and
resistance to acid; comes in several grades: 1, fast-setting, 2, slower, thicker, gap-filling form
(can be accelerated with suitable accelerant). Expensive, but can be bought in bulk.

Mechanical preparation may be carried out by hand with hammer, chisels and dental tools,
Pneumatic dental drills, airscribes (see Fordyce 1985c for outline of extended shanks for
airscribes), chipping hammers, and grinders allow faster and more efficient preparation. Fossils
can be extracted with acetic acid (Rixon 1976) which can be very effective, but expensive for
large blocks. Concentrated acid is dangerous to human tissues; these and other organic reagents
(such as organic solvents) can induce allergic reactions (or worse) in some people. Residual
dilute acid can damage fossil bones, if the specimen is not neutralised in dilute ammonia

FOSSIL VERTEBRATES OF NEW ZEALAND - 1199

solution and if it is not washed at least as long as the specimen was in acid (with multiple
changes of water). (See also the chapter by Whitelaw & Kool in this volume).

Specimens can be stored most safely in a plaster or fibreglass cradle, which allows even
distribution of weight and prevents breakage. Cotton wool and its fibres catch projections.
Polystyrene foam sheet and foam rubber or plastic are more useful for storing medium-sized
fossils. Cardboard boxes or glass vials are suitable for many smaller specimens.

NEW ZEALAND STRATIGRAPHY

New Zealand's sedimentary rocks, predominantly marine, range in age from Precambrian to
Recent (Fig. 4). The traditional view of New Zealand geology, as outlined in Suggate et al.
(1978) and commonly encountered in literature before the 1980s, long recognised three major
phases of sedimentation during the Phanerozoic. These phases supposedly spanned from the
Late Precambrian to the Early Devonian, from the Carboniferous to the Late Jurassic or Early
Cretaceous and from the Late Cretaceous to Quaternary. Apparent breaks between these three
sequences were thought to represent orogenies, the Tuhua Orogeny (Devonian), the Rangitata
Orogeny (earlier Cretaceous), and the Kaikoura Orogeny (Quaternary).

Research over the last decade, using a paradigm of a mobile earth, has overturned the notion
of a simple broadly three-fold sequence. There is strong evidence that rocks older than about
mid Cretaceous represent many independent terranes, "microplates" of tectonically-bounded but
otherwise unified lithostratigraphic sequences (Bishop et al. 1984, Howell 1980). What were
thought to be units deposited more or less in sequence (Suggate et al. 1978) cannot be shown
to have been originally juxtaposed, even though they are now. Some of the terranes indeed
could be far-travelled. It is understandable that because of modern physical proximity these
adjacent terranes were originally interpreted as contiguous. The recognition of terranes throws
doubt on the idea of a widespread earlier Cretaceous "Rangitata Orogeny", and the older Tuhua
Orogeny is best abandoned. The Kaikoura Orogeny might now be seen as encompassing the
events that stemmed from the migration of the Australian-Pacific plate boundaries into the New
Zealand area in the Neogene.

In summary, there are few published reports of early Palaeozoic vertebrates from New
Zealand (Fordyce 1982c wrongly stated none). Triassic to Late J urassic rocks have yielded (and
continue to yield) rather more, and there is great potential fer more discoveries. Vertebrates are
most common, although still little documented, in marine rocks of Late Cretaceous to
Quaternary age. This review concentrates on the Late Cretaceous-Cainozoic record.

Stratigraphy of New Zealand sedimentary rocks relies largely on biostratigraphy based on
marine invertebrates. Spores and pollen are used to a lesser extent. Local stage names are not
applied to the earlier Palaeozoic, but are used from the Permian onwards. The few relevant pre-
Late Cretaceous stages mentioned here are given with their international correlations. Late
Cretaceous and Cainozoic stages and their correlations are shown in Fig. 5. The sequence of
these younger stages is unequivocal (Finlay & Marwick 1940, Hornibrook et al. 1989,
Hoskins 1982). Although there is still debate about the definition of some stages and the
international correlations of their boundaries, this is one of the most precisely subdivided
marine Cainozoic sequences in the world.

Some problems exist with local stages, however. The stages were based originally on
locally abundant fossils with little regard to detailed overseas correlations. New Zealand has
been rather isolated in mid temperate latitudes since the early Cainozoic, and the fossil biota
reflects this. It has been difficult to establish correlations because of firstly, the endemic
element in the biota and, secondly, diachronous ranges in those taxa that also occur elsewhere.
Thus, it is difficult to match precise stratigraphic ranges for most New Zealand taxa with
ranges in nearby Australia, with localities nearer the equater. The problem is worse for more-

1200 - FORDYCE

distant northern temperate latitudes where type localities for stages of the Cainozoic are to be
found.

Ceno.
Agnatha
Teleostomei
Amphibia
Reptilia
Aves
Mammalia

Jurassic |Cretaceou
Elasmobranchiomorpha

Carb. Permian Trias.

Devonian

terrane boundaries
marked by vertical lines

e vertebrate horizon

WY ;

GY terrane

Cambrian

Buller Brook Street Caples Rakaia Pahau
Murihiku
Takaka DunMtMaita | [orlesse

Figure 4. Summary of spatio-temporal distribution of New Zealand tectonostratigraphic terranes, and
distribution of main vertebrate groups. (Terranes are simplified from those shown by Bishop et al. 1985).

FOSSIL VERTEBRATES OF NEW ZEALAND - 1201

INTERNATIONAL N Z Stage
Mp SUBDIVISIONS 7a —

Mangpanian
Waipipian
D
SS

Selected vertebrate-bearing units
loess, swamp and dune deposits, gravel
Kai-iwi Gp Waipaoa "Series"
Tewkesbury Fm
Te Aute Lst
Tangahoe Mst
Waiourr'Em Kaawa Shellbed

Greta Siltstone

Tongaporutuan Longford Fm Kaikorai Valley leaf beds

Waiauan
Lillburnian

Duntroonian
Whaingaroan

Runangan

| Porangan |

Teurian

Haumurian

Chatham Rise phosphorites
Double Corner Shellbeds
Nga Pari Fm
Southburn Sand
Manuherikia Gp
Rifle Butts Fm
Bluecliffs Silt

Gee Greensand

Milburn Lst Te Akatea Siltstone
Otekaike Lst Te Kuiti Gp

Weka Pass Stone Waitomo Sst
Takaka Lst Abel Head Fm

Kokoamu and Wharekuri Gsds and equivalents

Parengarenga Gp

20

Waikawau Fm
Tarakohe Mst

30

Glen Massey Fm

Whaingaroa Siltstone
McDonald Lst

OLIGOCEN

40 Kaiata Mst

Burnside Mst

Waihao Gsd Pahi Gsd
Tapui Glauconitic Sst

50 Abbotsford Mst

EoOoceENE

[mea [i

Waipara Gsd

AmuriLst (time transgressive)

Kauru Fm
Moeraki Fm

60

PALEOCENE

"Lingula beds”

Katiki Fm
Laidmore Fm

70

Conway Siltstone "Maungataniwha sst"

Figure 5. New Zealand latest Cretaceous and Cainozoic stages, approximate correlations, and examples of
some vertebrate-bearing units. This figure does not use all the revised nomenclature suggested by Browne &
Field (1985) and Field & Browne (1986).

Examples of specific correlation problems include those associated with the Eocene-
Oligocene and Oligocene-Miocene boundaries. These boundaries are difficult to correlate to
New Zealand because the main biostratigraphic markers (planktonic foraminifera) seem to have

1202 - FORDYCE

diachronous ranges. The problem is more broadly illustrated by comparing correlations shown
for the Landon Series (Fig. 5) and the Oligocene by Stevens (1980b), Hardenbol & Berggren
(1978), Loutit & Kennett (1981), and Vail & Hardenbol (1979). Kellogg (1956) summarised
one view of the stratigraphic implications of fossil Cetacea largely from the Landon Series. In
another example, Warren & Speden (1977) discussed correlation problems of the Piripauan and
Haumurian Stages. Irrespective of these issues, New Zealand Cainozoic stages are cited
throughout the text in order to indicate precise ages. International correlations are more
equivocal.

There are several general guides to New Zealand geology that might interest vertebrate
palaeontologists. Suggate et al. (1978) provided an account, from the viewpoint of the New
Zealand Geological Survey, which contains a broad summary of lithostratigraphy. Some
interpretations in this work are now widely acknowledged to be outdated. The series of
1:250,000 geological maps produced by the New Zealand Geological Survey for all of New
Zealand is valuable, although dated in places (the rocks have not changed, but more detail has
been added). Brown et al. (1968) presented a general review which is reasonably available
overseas but which is also now outdated, alarmingly sc in places. Fleming (1979), in a
revision of an earlier classic paper (1962a, and see Fleming 1949 and 1975), provided a useful
summary of the geology of New Zealand from a palaeontologist's point of view. Stevens
(1980c, 1983) interpreted New Zealand history in the light of plate tectonics and continental
drift, but provided little information on lithostratigraphy or biostratigraphy. Stevens (1985)
also reviewed New Zealand's changing geography from the Cambrian to present. Lillie (1980)
and Gage (1980) discussed the stratigraphy and geological evolution of New Zealand; Lillie's
account gives a particularly good guide to lithostratigraphy. Carter (1988) gave a singular
account of Cretaceous-Cainozoic stratigraphy noteworthy for its treatment of nomenclature.
Thornton (1985) provided a very readable field guide, and Burrows (1978), Beu et al. (1987) and
Gage (1979) summarised Quaternary geology.

Lithostratigraphic names are regarded by palaeontologists as important tools for information
retrieval. Most local stratigraphic names were discussed in the lexicon edited by Fleming
(1959b) or in the more recent and, regretfully, abbreviated, lexicon compiled by McGregor
(1987). As parts of the Cretaceous-Cainozoic project of the New Zealand Geological Survey
have been completed, it has been suggested that some old or long-used names should be
synonymised or abandoned. Articles that discuss lithostratigraphy for some of the Cretaceous-
Cainozoic project areas include those of Browne & Field (1985), Field & Browne (1986) and
H.J. Campbell et al. 1988. Some of the names used in these latter articles are used in this
review, but it was not possible to change all. The application of some names could be debated,
and it is likely that there will be disputes about the correct name(s) to apply for some years yet.
Few names can be gleaned from the New Zealand Geological Survey 1: 250,000 geological
maps which cover the country and otherwise provide a standard reference. Cainozoic and many
Mesozoic rocks are mapped therein as stages, while lithostratigraphic units are cited for the
Lower Palaeozoic and sporadically for younger rocks, Indeed, formation names are still not
widely used, or in some cases even established for some areas. For example, the enormous
body of Torlesse rocks which forms much of the axis of the South Island is largely too
complex structurally to yield sequences complete enough on which to clearly define and
diagnose formations, The Southland Syncline, for which many Permian-Jurassic stage
boundaries are known well, is lithologically too monotonous in places to allow different
formations to be recognised clearly (e.g. in Triassic volcanogenic sediments).

FOSSIL VERTEBRATES OF NEW ZEALAND - 1203

THE STRATIGRAPHIC RECORD OF VERTEBRATES

EARLY PALAEOZOIC

Palaeozoic rocks older than Permian crop out only in the South Island. Cambrian,
Ordovician, and Silurian marine rocks appear not to have yielded vertebrates. No non-marine
units of this age have been identified positively. Cambrian and Ordovician rocks have been
studied for acid-insoluble microfossils (e.g. Simes 1977), but no bone has been reported. Most
Lower Palaeozoic strata are deformed and metamorphosed. This, together with the lack of
serious attention from vertebrate palaeontologists, makes it seem unlikely that they will yield
vertebrates. Nevertheless, recent work on microvertebrates recovered from acid-insoluble
residues Overseas suggests that any finds might have profound biogeographic implications, and
careful prospecting seems warranted. For reviews of fossiliferous Lower Palaeozoic rocks, see
Cooper (1968, 1979, 1989) Cooper & Bradshaw (1985) and Suggate et al. (1978).

DEVONIAN

Marine Devonian rocks in the Baton and Reefton districts of the South Island have yielded
good invertebrate faunas but no significant bone. Jenkins (1967) reported "what appear to be
vertebrate fragments" from Waitahu River, Reefton. Later, Macadie (1985) identified
fragmentary bones of a possible arctolepid arthrodire ("resemble[s] Actinolepis"), acanthodians
and palaeoniscids from the Reefton sequence. An indeterminate bone fragment is also held in
the Canterbury Museum, Christchurch (M.A. Bradshaw, pers. comm.). Limestone in Lankey
Creek and mudstones in the Waitahu River near Reefton are lithologically similar to the units
of eastern Victoria from which fish have been collected, and could most profitably be
prospected. Articles on Devonian rocks were presented recently by Bradshaw & Hegan (1984)
and Cooper & Bradshaw (1985); see also Suggate (1957).

CARBONIFEROUS

The Carboniferous is known in New Zealand only by. conodont-bearing marine marble
associated with Torlesse-like rocks at Kakahu, South Canterbury (mentioned widely, most
recently by Hitching 1979). It is not clear whether the blocks of marble are autochthonous.
D.G. Jenkins & T.B.H. Jenkins (1971) stated that "rare fish scales" were present in conodont-
bearing acid-insoluble residues from the marbles. No vertebrates have been identified formally,
but further study seems justified.

PERMIAN

Widespread fossiliferous marine Permian rocks in Southland and Nelson have been studied
quite thoroughly by geologists familiar with invertebrate fossils, yet no significant bone has
been reported. The lack of identified bone may well represent observer bias, since Permian
sequences (as with most other Palaeozoic rocks) have yet to be prospected systematically by
vertebrate palaeontologists.

Wood (1956: 45, fig. 24E) noted the presence of an apparent lepidosteid (gar) fish scale
(New Zealand Geological Survey collections) in rocks of Arthurton Group, Gore district. H.J.
Campbell (pers comm., 1988) recently found fish scales in rocks of the Permian Takitimu
Group, western Southland.

1204 - FORDYCE

TRIASSIC AND JURASSIC

Triassic rocks in the South Island are included traditionally in two major sequences, the
relatively fossiliferous Murihiku Supergroup and the sparsely fossiliferous Torlesse
Supergroup. Torlesse rocks also occur in the North Island. Both sequences are predominantly
marine, and both also encompass Jurassic strata. The Torlesse Supergroup, for which
nomenclature has long been in a state of flux (e.g. Andrews et al. 1977), is probably formed by
a sequence of terranes (Bishop et al. 1984, Howell 1980, Retallack 1987), and it has been
suggested that parts of the Torlesse may have been deposited at low latitudes in the Northern
Hemisphere. Few vertebrates are known from either the Murihiku or Torlesse sequences.
Reviews of geology and palaeontology include those of Andrews et al. (1979), Begg et al.
(1985), H.J. Campbell & Johnston (1984), J.D.Campbell (1985b), Campbell & Warren (1965)
and Speden (1975, 1976).

Occasional incidental reference has been made to fish, but no reliable identifications to
species level have been published. T.B.H. Jenkins & D.G. Jenkins (1971) found teeth in acid-
insoluble residues of Warepan (Late Triassic) age from Okuku, North Canterbury, and Bradshaw
(1977) reported fish vertebrae from Arthurs Pass (New Zealand Geological Survey collections),
also probably of Warepan age. Strong (1984) mentioned microscopic teeth from Triassic
sediments at Roaring Bay, Nugget Point. HJ. Campbell & Cave (1987) noted fish remains
from the Otamitan of the Rolleston Range, Rakaia Valley. Marden et al. (1988: 392) reported
elasmobranch remains from the Oretian of Ruahine Ranges.

Teeth of "labyrinthodont" appearance, of uncertain affinities, have been found at Nugget
Point, South Otago (Hector 1878, 1886) and Wairoa, Nelson (Worley 1894). The Nugget
Point specimens could represent ichthyosaurs, which are known from recent finds in pebbly
sandstone of Otamitan Stage at Roaring Bay (J.D. Campbell 1987; fossils in Geology
Museum, University of Otago). McKay (1877b) reported “saurians" from Roaring Bay.
Positively identified ichthyosaurs are known from Mt Potts, Canterbury, and the Hokonui
Hills, Southland (J.D. Campbell 1965, Fleming et al. 1971; see also e.g. Anon. 1878b; Haast
1887; Hector 1874, 1877c, 1878; McKay 1878; Trechmann 1918; Wilckens 1927). Vertebrate
remains, including ichthyosaurs and other unspecified reptiles, occur in Torlesse rocks in
southern North Island (H.J. Campbell 1982, Stevens 1974).

H.J. Campbell et al. (1984: 283, 285; see also 1980) mentioned an undetermined small
limb bone, possibly that of a reptile, of Permian (or Triassic) age from Stephens Formation of
Stephens Island, Marlborough. The specimen appears to have been lost when sent to the
U.S.A. for identification (E.H. Colbert, pers. comm.). Of incertain age and relationships is a
large vertebrate, presumably a reptile, zecently collected from the lower Triassic (or uppermost
Permian) marine tuffaceous sandstone at Mossburn, Southland (Aitcheson et al. 1988). The
specimen, which is in the Geology Museum collections (University of Otago), is under
preparation at present.

Jurassic rocks are widespread, often conformable over the Triassic, but there are no
significant published reports of vertebrates. The only specimen reported is the poorly preserved
incomplete shaft of a long bone (New Zealand Geological Survey collections) of Jurassic age,
from Marybank Formation, Nelson (Johnston et al. 1980; 1987: 286, "resembles...reptilian
rib"). The specimen, which lacks condyles, has not been identified formally. Some outcrops
of terrestrial Jurassic sediment have well-preserved Icaf floras. Localities include Curio Bay on
the Catlins Coast in Southland, and Kawhia in Southwest Auckland. Curio Bay exposures
include possible channelled floodplains and palacosols in which terrestrial vertebrates might be
preserved, although field work by M.S. Pole and others (Department of Geology, University of
Otago) have not revealed bone. Further prospecting, especially in the nonmarine Jurassic (e.g.
axis of the Southland Syncline), would seem warranted, but the lack of good shelly fossils

FOSSIL VERTEBRATES OF NEW ZEALAND - 1205

Suggests that conditions in fresh water environments may have been too acidic for bone to
preserve.

INTERNATIONAL Approximate timing of oceanic,
Ma SUBDIVISIONS NZ Stage climatic and geographic events

-X—tastlectiian

EXT Mangpanian——]
Waipipian
Opoitian

Kapitean

Tongaporutuan

Waiauan
| —Lilburnian _

Duntroonian
Whaingaroan

Runangan

| _Porangan |

Teurian

Haumurian

increasing Milankovitch cycle amplitude

terrestrial glaciation in S temperate regions
Panama seaway closed - changed Pacific
zoogeography

increase in Antarctic glaciation
Messinian crisis

major southen cooling; regression

10

first? significant movement on Alpine Fault

cooling; E Antarctic ice sheet buildup

subtropical-tropical climate; global thermal
maximum; high sea level

evidence of local tectonism in NZ

warming

Oo C ECE

LY |

20

reduction in Antarctic ice?

Circum-Antarctic flow and zoogeographic
distributions established

extensive shallow shelf seas; minimal terrigenous
sedimentation

major sea level drop and/ or change in NZ tectonics

peak marine transgression; minimal NZ land area;
possible reduced terrestrial diversity?; ?archipelago

broad marine regression and increasing land area

with gradual then accelerating orogeny

30

major cooling and/or accumulation of Antarctic ice;
subantarctic cooling

cooling of southern surface and bottom waters;

psychrosphere developed; Antarctic
Convergence developed, affecting NZ waters?

? time of split, NZ-Australian ratites

warm equable climates; southern convergences
not developed; Antarctica relatively warm.

OLIGOCENE

| eary | tte |

40

50

EoOoceENE
oO

60

PALEOCENE

Cretaceous-Tertiary boundary extinctions

steady marine transgression reduces land area

70 peneplained NZ, with complement of reptiles and

birds, podocarps and beeches, separated from
Antarctica and Australia about mid Cretaceous

Figure 6. Summary of inferred major environmental changes in New Zealand in the latest Cretaceous and
Cainozoic.

1206 - FORDYCE

[ ] Quaternary

BB Late Cretaceous - Tertiary

VA) older than Late Cretaceous

Figure 7. Distribution of later Cretaceous and Cainozoic ("Notocene") rocks in New Zealand. (After
Suggate ef al. 1978).

THE CRETACEOUS-CAINOZOIC SEQUENCE

Upper Cretaceous rocks rest unconformably on basement over much of New Zealand. Such
younger Cretaceous sediments, as typified by those of Canterbury and Otago, are
compositionally more mature (quartz-rich) than the Torlesse-like older Cretaceous. They often
include quartzose nonmarine sands and gravels succeeded by marginal marine coal measures.

FOSSIL VERTEBRATES OF NEW ZEALAND - 1207

These sediments, and overlying more distal and terrigenous-poor sediments, indicate the
Start ofa broad transgression which continued without significant break into the middle of the
Cainozoic, about the Whaingaroan Stage. This broad transgression was followed by a more
rapid Miocene to Recent regression (Fig. 6), which marked uplift associated with the start of
the Kaikoura Orogeny and the movement of the boundary of the Australian and Pacific plates
into the New Zealand region. Relative fluctuations in sea-level, marked in the field by
alternations between more distal and more proximal sediments, were probably superimposed on
this broad transgressive-regressive sequence (e.g. Vella 1967; see also Haq et al. 1987, Loutit
& Kennett 1981, Vail & Hardenbol 1979) .

The thick, widespread Cretaceous-Cainozoic sequence (Figs 5, 7), the Notocene of Thomson
(1917), includes the main source horizons for vertebrates. Because there are many records of
vertebrates, the account below is only a guide.

Earlier Cretaceous

Earlier Cretaceous rocks have been regarded traditionally (e.g. Suggate et al. 1978) as part of
a different depositional sequence from later Cretaceous rocks, which are usually conformable
with the overlying Cainozoic. A variably developed stratigraphic gap somewhere in the middle
of the Cretaceous was long interpreted as evidence of the Rangitata Orogeny, but it is now
regarded as marking the base of a regionally widespread sedimentary sequence that caps an
amalgam of terranes and extends up to the Quaternary (fig. 4). In some areas, the upper part of
the Torlesse sequence is lithologically similar to quartzo-feldspathic sediments of the Jurassic
and Triassic, and is of Early Cretaceous age. Few vertebrates have been reported from the
Torlesse-like earlier Cretaceous. One, however, is Chapman's (1918) Diplomystus
coverhamensis from Cover Stream, in the Kaikoura ranges, and others might be expected.

Localised fluvio-lacustrine sediments of roughly mid Cretaceous age, such as the Hawks
Crag Breccia of Buller (Nathan 1978) and Kyeburn Formation of Otago (Bishop & Laird 1976,
Bishop 1979) have yet to be prospected extensively for bone. Large bones might be expected
in coarser infills of channels and scours in these lithologies. Finer-grained units such as the
Ohika Formation (lateral equivalent of Hawks Crag Breccia), parts of the Kyeburn Formation,
and parts of the Gridiron Formation (Suggate 1958, Crampton 1988) of the Clarence Valley
and nearby areas could also yield bones and, if suitable shallow water or floodplain facies are
present, footprints. Further study is needed. Productive nonmarine environments might be
expected from the broader setting of these units, which may have developed in broad fault-
bounded depressions associated with the initial rifting of New Zealand from other segments of
Gondwana. Traditionally the sediments have been identified as synorogenic (Laird 1980).

Late Cretaceous

Upper Cretaceous strata (Mata Series; Maastrichtian) have yielded locally abundant marine
vertebrates (Fig. 8). The best-known traditional localities are in the "saurian beds", better
recognized as Conway Formation of Browne & Field (1985; = Laidmore Formation of some
authors) of north-eastern South Island. This horizon occurs at Waipara River and the Cheviot
area in North Canterbury, and extends northwards to Haumuri Bluff in Marlborough.
Plesiosaurs and mosasaurs are known from these sites (Fleming 1963a, Fleming et al. 1959,
Gage 1969, Haast 1862 and 1871, Hector 1877, Hutton 1877, McKay 1877a, 1877e, 1883,
1892, Park 1888, 1913, Warren & Speden 1977, Welles & Gregg 1971, Wellman 1959,
Wilson 1963, Woods 1917). Most of the specimens are in the collections of the Canterbury
Museum (Christchurch), National Museum (Wellington), and New Zealand Geological Survey
(Lower Hutt).

1208 - FORDYCE

CRERICA 7

(SOUTH)
tenes 1 ZAMERICA) al Mangahouanga Stream,
possible reconstruction of south west Pacific,
after Stevens

Haumuri Bluff

some New Zealand localities for
Cheviot.

Late Cretaceous reptiles

tail vertebra of possible
theropod dinosaur

mosasaur
body outlines after Colbert

Figure 8. Some New Zealand localities for Late Cretaceous reptiles. (After Fordyce 1986).

Recently, Joan Wiffen collected and described a vertebrate fauna from informally-named
"Maungataniwha sandstone", at Mangahouanga Stream, Hawke's Bay. The age is uncertain
Piripauan or Haumurian stage (Moore 1987, Crampton 1989). Wiffen (1980,1981a, 1981b,

FOSSIL VERTEBRATES OF NEW ZEALAND - 1209

1983, 1984, 1986, Wiffen & Moisley 1986, Molnar & Wiffen 1988) described some of the
plesiosaurs, mosasaurs, turtles, a pterosaur and fish from Mangahouanga Stream. Molnar
(1981; see also 1980) described the caudal vertebra of a probable theropod dinosaur from this
locality, and Wiffen & Molnar (in press) are to report the discovery of ornithopod remains.
Molnar & Scarlett (1984) documented the phalanx of a large bird or dinosaur. The fossils are
in the New Zealand Geological Survey collections (Keyes 1984, 1989).

A recently-studied Cretaceous vertebrate site is at Shag Point, North Otago (Fordyce 1983c,
1986, 1987b), where plesiosaurs and mosasaurs occur. The fossils, including those in the
Geology Museum collections (University of Otago; Fig. 9; Pl. 1) have been found in
mudstones of the Katiki Formation, Haumurian Stage (latest Cretaceous). This is the most
southern site known at present for Late Cretaceous marine reptiles in New Zealand.

Other localities for Late Cretaceous vertebrates include many where isolated bones have
been collected. At the Waimakariri Gorge, "Lingula beds" (Keyes 1981a), of Teurian age in
part as well as Haumurian, have yielded chondrichthyans, elasmosaurs, and an undescribed avian
tarsometatarsus (collections include those at New Zealand Geological Survey, Lower Hutt;
Geology Museum collections, University of Otago, and the Department of Geology,
University of Auckland). Vertebrates from a phosphatic horizon in unconsolidated quartz sands
(Conway Formation?) near Cheviot, North Canterbury, inclade fish, reptiles, and a fragment of
avian femur (New Zealand Geological Survey, Lower Hutt and Geology Museum collections,
University of Otago) (Keyes in Feldmann 1984: 283). North Island localities include those in
southern Hawke's Bay (Adams 1983a and 1983b; fish), and Kaipara Harbour in Northland
(Evans 1983 and 1986: table 1; fish).

Cainozoic: Palaeocene

Few bones are known from the Palacocene (Teurian Stage, Dannevirke Series). The only
formally described specimen seems to be a possible turtle bone from Ward, Marlborough
(Fordyce 1980a; New Zealand Geological Survey collections). Undescribed bird and possible
reptile bones are known from Moeraki Formation, Moeraki (Hamilton 1902, Mantell 1850;
Geology Museum collections, University of Otago). From the Waipara Greensand, North
Canterbury comes a probable primitive penguin (New Zealand Geological Survey collections;
Fordyce et al. 1986). An undescribed turtle and an undescribed fragment of bird are known from
Chatham Island (New Zealand Geological Survey collections). There is an unsubstantiated
record of a plesiosaur also from Waipara Greensand (McKay 1877e: 37; see Fordyce 1980a,
Welles & Gregg 1971: 103).

Bone was discovered recently in shallow marine sediments of the Wangaloa Formation, by
J.D. Stilwell and others (Department of Geology, University of Otago), at Wangaloa in South
Otago. Material includes shark and chimaerid teeth, and scraps of bone apparently not those of
fish. The age is uncertain. The dominant fossil assemblage, of shallow water invertebrates, is
used to define the Wangaloan stage. The lack of age-diagnostic microfossils means that the age
can be defined only as latest Cretaceous or earliest Palaeocene. The depositional setting of this
unit (see Lindqvist 1986) is similar to units on Seymour Island, Antarctic Peninsula, which
have yielded Eocene terrestrial vertebrates, and more field work is needed. Gage (1957: 28)
mentioned fish teeth and vertebrae of Wangaloan (probable Palaeocene) age from marginal or
shallow marine Kauru Formation, North Otago, and other specimens from this site and a
similar lithology at Waihao River (South Canterbury) are also in the Geology Museum,

1210 - FORDYCE

Figure 9. Reconstruction (by Craig Jones) of part of skeleton of plesiosaur from North Otago; specimen in
the Geology Museum collections, University of Otago, scale bar, approx. 1 m.

FOSSIL VERTEBRATES OF NEW ZEALAND - 1211

University of Otago. Field & Browne (1986) also mentioned fish teeth from Kauru Formation,
as they defined it (a unit of broader geographic distribution than envisaged by Gage).

Other Suitable rocks and areas for future prospecting could include Katiki and Moeraki
formations in North Otago, and other shallow-water transgressive sediments further north in
eastern South Island. No vertebrates appear to have been reported from the thick distal
mudstones of the Whangai and Wanstead formations of Dannevirke area (Lillie 1953, Suggate
etal. 1978), although marine fish might be expected. None has been described from any of the
extensive marginal marine coals (Cretaceous and Eocene, as well as Palaeocene) of both the
North and South Islands.

Cainozoic: Eocene

Bones are rare in the Lower Eocene (upper Dannevirke Series), as for the Palaeocene. This
may reflect a collecting bias, since calcified body fossils of invertebrates are uncommon in
these rocks, and macropalaeontologists historically have concentrated on Bortonian and younger
Cainozoic rocks. It is possible that vertebrates have been recovered from rocks of this age but
that the age has not been recognized. Shallow water marine sediments in which bones might
be expected have an invertebrate fauna which, like that of Wangaloa Formation, is not
particularly age-diagnostic, and bones from the upper Dannevirke Series might not be
recognized as of this age.

A fish skull, the type-specimen of "Portheus" dunedinensis Chapman, 1934, comes from
the Abbotsford Mudstone, Heretaungan Stage (Dannevirke Series) near Abbotsford Railway
Station, Dunedin district (Geology Museum, University of Otago collections). This appears to
be the only formally described earlier Eocene macrovertebrate. Sediments of Bortonian Stage
(Arnold Series) and younger, however, have yielded fish otoliths (Schwarzhans 1980, 1984: 9;
types in New Zealand Geological Survey collections, Lower Hutt). Fish teeth, vertebrae and
scales are known from South Canterbury-North Otago (Geology Museum collections,
University of Otago), and a large fish skull was found at Hampden (New Zealand Geological
Survey collections, Lower Hutt). Penguins include types of Palaeeudyptes marplesi and
Pachydyptes ponderosus (see Simpson 1971a and Table 6) and cetaceans (aff. Dorudon and other
specimens; Fordyce 1985b and Table 7). Marshall (1917: 439) mentioned an apparent reptile,
still undescribed, from Opahi Group of Northland (see also Fordyce 1980a: 740). Turtles are
represented by specimens from the Pahi Greensand at Pahi, Kaipara Harbour (Department of
Geology, University of Auckland collections), the Kaiata Mudstone at Woodpecker Bay in
Westland, and the upper ?Abbotsford Mudstone at Boulder Hill near Dunedin. The latter two
specimens, of Kaiatan or possibly Bortonian age, are in the Geology Museum collections,
University of Otago; they are discussed below.

Future prospecting is certain to reveal other Eocene vertebrates; likely horizons and
localities could include productive ones mentioned above as well as others: the Pahi Greensand
(Northland), the time-transgressive Amuri Limestone (North Canterbury and Marlborough),
Kaiata Formation (North Westland-Buller), Iron Creek Greensand (North Canterbury), Waihao
Greensand and Tapui Glauconitic Sandstone (South Canterbury-North Otago), and Hampden
Formation (North Otago). In general, shallow marine Eocene rocks seem exposed poorly in the
North Island, but deep water sediments might yield undisturbed articulated skeletons.

The prospects for Paleogene nonmarine vertebrates seem poor. The most likely horizons in
which specimens might have been buried originally are the marginal-marine to non-marine
coal-measures of Eocene age (and older) which occur widely throughout New Zealand (e.g.
Westland, Southland, Waikato). They have been worked extensively for coal, but have yet to
provide any significant vertebrate remains. Indeed, not one bird has been reported from these
coals. Recent authors have noted, following Fleming (1962a: 93), that depositional
environments may have been too acidic to allow bone to preserve, although this suggestion is

1212 - FORDYCE

countered by the sporadic preservation of calcareous shells (e.g. Hyridella). Most nonmarine
sediments, even those of the Paleogene, when presumably there was no suggestion of axial
uplift on what is now New Zealand, are very much proximal. Most occur today within a short
distance of the axial mountains and thus have been subject to recent tectonism. At the outcrop
level, this has resulted in sheared, faulted sediments which have generally been leached
thoroughly by percolating ground water,

The above problems aside, fossiliferous shallow marine rocks, such as Tapui Glauconitic
Sandstone, and non-marine rocks, such as Papakaio [= Taratu] Formation in North Otago and
South Canterbury may also be good prospects for terrestrial vertebrates. Tapui sediments
appear locally similar to shallow marine mammal-bearing strata on Seymour Island, Antarctica
(Woodbume & Zinsmeister 1984), and Papakaio silts have yielded an insect fossil (Aitchison et
al, 1983), thus attesting to an environment favourable for the preservation of noncalcareous
fossils other than plants.

Cainozoic: Oligocene

Vertebrates are relatively common in Oligocene and Neogene rocks, and this review will
consider only a few localities. Oligocene sequences (Landon Series) are generally thin and
calcareous, with biogenic and authigenic sediments common but little terrigenous material.
They were deposited about the peak of a broad Cainozoic marine transgression, when land that
might have supplied terrigenous sediment was probably of low relief and/or distant.

Few earlier Oligocene vertebrates (Whaingaroan Stage) are known, These include:
penguins, from the Whaingaroa Siltstone (south-west Auckland); mysticete cetaceans and fish,
from the calcareous mudstone of Nile Group (near Karamea); and a cetacean, from the time-
transgressive foraminiferal ooze of Amuri Limestone (Waikari; Fordyce 1989a). These fossils
come from the rather characteristic fine-grained, biogenic and terrigenous-poor, distal facies of
the Early Oligocene, Sharks, turtles, penguins, and cetaceans (Keyes 1973) occur in bryozoan-
rich McDonald Limestone (Oamaru district), a presumably shallow water unit which formed
from reef debris that accumulated on one or more local highs. It is possible that coarser, more
proximal marine sediments, which should have preserved more-common vertebrates, were
eroded away by the event(s) that formed the widespread mid-Oligocene "Marshall
Unconformity".

Later Oligocene strata of the Duntroonian and Waitakian Stages (the latter Early Miocene in
part if not in whole) are rather well-known, as they are an important New Zealand-wide
commercial source of lime. Much of the Upper Oligocene limestone is flaggy limestone,
which has probably undergone significant diagenesis upon burial, and which has yielded rather
few vertebrates. Fish, penguins and cetaceans are known from these rather well-cemented rocks
near Te Kuiti (e.g. Grant-Mackie & Simpson 1973, Nelson 1978), Westhaven Inlet in North-
west Nelson, and Punakaiki in Westland (Geology Museum collections, University of Otago).

Rather soft and less-cemented bioclastic limestone and greensands on the eastern side of the
South Island are often rich in macrofossils. Such units have probably undergone shallow
burial at most, in contrast to the more flaggy horizons. Faunas include sharks, bony fish,
penguins (and rarely other birds) and cetaceans. Bones are locally conspicuous at the base of
the Duntroonian Stage at the "Marshall Unconformity" (Carter & Landis 1972 and 1982); they
are sometimes associated with phosphate nodules and may represent lag deposits. Indeed, some
concentrations within the greensands may reflect fluctuating sea levels (Fordyce 1987a). The
Kokoamu Greensand and Otekaike Limestone and their lateral equivalents have been the source
of most of the fossil penguins and cetaceans described from New Zealand (Fordyce 1980b,
1980c, 1983d, 1985d, 1987a, 1987b, Fordyce & Jones 1989). The flaggy Milburn Limestone
of South Otago was a noteworthy early source of vertebrates, probably because of large lime

FOSSIL VERTEBRATES OF NEW ZEALAND - 1213

and phosphate works in the area (Andrew 1906, Hamilton 1903, Park 1903, and other
references mentioned by Fordyce 1980b).

Cainozoic: Miocene

The Miocene, which is represented by the Waitakian Stage in part, and the Pareora,
Southland and Taranaki Series, saw the acceleration of a broad regression that ended late in the
Neogene with the uplift at the peak of the Kaikoura Orogeny. Sediments are generally thicker
than those of the Oligocene, with an increasing terrigenous component that effectively dilutes
the vertebrate fossils. Distal and proximal marine rocks and terrestrial rocks are exposed on
land. Often the earlier Miocene sediments are fine-grained and calcareous, such as the Tokama
Siltstone (as defined by Field & Browne 1986). This unit encompasses the Grey Marls of
North Canterbury, and the Bluecliffs Silt-Riflebutts Formation of South Canterbury-North
Otago. Coarser and presumably more proximal sediments occur higher in the column (e.g.
Brechin Formation and Double Corner Shellbed unit of Tokama Siltstone, of Canterbury).
There are important richly fossiliferous biogenic sequences in some areas, for example,
Southland (Wood 1969). However, increasing tectonism during the Miocene allowed localised
patterns of deposition to develop in progressively more-isolated basins, so that it is difficult to
generalise much more about patterns.

Localities in Northland, Wairarapa, East Coast, North Canterbury, South Canterbury, North
Otago, and Southland have yielded teleost otolith faunas of Otaian Altonian, Lillburnian,
Waiauan and Tongaporutuan ages (Grenfell 1981, 1983, 1984, Schwarzhans 1980, 1984).
Sharks, birds (penguins, pelagornithids) and cetaceans are known from the Greta Siltstone, a
rather ill-defined unit in North Canterbury which has been mapped as Miocene-Pliocene (Gregg
in Suggate et al. 1978: Fig. 7.77, Wilson 1963) and may range from Clifdenian to
Mangapanian (Browne & Field 1985). The vertebrates, which appear to have been recognised
as early as 1866 (Hector 1867a: 8), commonly occur in concretions, which are known to have
been reworked in some cases from older units. The concretions range in age from Otaian-
Altonian (Early Miocene) to Waiauan-Tongaporutuan (Late Miocene) and Waipipian (Late
Miocene) (Lewis 1976). The concretions are often dolomitised, thus making it difficult to
extract age-diagnostic microfossils. This area is favoured by amateur collectors (Anon. 1979).

Sharks’ teeth, teleost otoliths and rare cetacean bones occur in the Southburn Sand
(Altonian), Tokama Siltstone/Riflebutts Formation (Otaian-Altonian), Gee Greensand
(Waitakian-Otaian), Caversham Sandstone (Otaian) and Clarendon Sand (Otaian) of South
Canterbury to South Otago districts. Collections include those in the Geology Museum,
University of Otago (Fordyce 1980b, Fordyce et al. 1985, Gage 1957). Faunas in the
extensive bryozoan limestones and associated sediments of Southland include teleost otoliths
and sharks’ teeth (Fleming et al. 1969).

The oldest significant Cainozoic terrestrial bone assemblage is that of waterfowl from the
Manuherikia Group near St Bathans, Central Otago (Douglas et al. 1981; see also McKay
1894, 1897). The Manuherikia Group is of Altonian age, Early Miocene (Douglas 1977,
1986). Specimens are housed in the Geology Museum, University of Otago. Fragmentary
unidentified terrestrial bird bones also occur in a limestone, a possible lateral equivalent of
Manuherikia Group to the northeast in the Waitaki Valley (specimens in Geology Museum,
University of Otago). The sparse fossil fauna of the paralic to non-marine Longford Formation
of Murchison district includes a bird footprint ("kiwi") of latest Miocene age (Mildenhall

1974).

1214 - FORDYCE

Cainozoic: Pliocene

Pliocene rocks are generally not as widespread as those of the middle Cainozoic, as the seas
had moved off much of what is now New Zealand. There are few significant vertebrate sites in
the South Island other than the coastal North Canterbury Greta Siltstone sequences, mentioned
above, and blue-grey siltstones of the Blue Bottom Group, Westland (birds, cetaceans). Of note
is a possible bony-toothed pelican femur from the Nukumaruan (latest Pliocene-Early
Pleistocene), Motunau, North Canterbury (Canterbury Museum collections, Christchurch), A
moa, the type of Anomalopteryx antiquus Hutton, 1892, came from below Pliocene lavas at
Timaru (Forbes 1891a), and other earlier Pliocene or possibly latest Miocene moas are known
(see below). Pliocene sandstones, siltstones and, locally, limestones are widespread in the
North Island. Schwarzhans (1980, 1984) described teleost otoliths from Opoitian and
Waitotaran sediments in the North Island. Grant-Mackie (1983) mentioned shark teeth from
Kaawa. McKee (1984, 1985, 1986, 1987a, 1987b, 1988) noted the presence of sharks, a
"bony-toothed" pelican, a penguin and a dolphin in Waipipian sediments at Hawera, while
McKee & Fordyce (1987) documented a delphinid mandible from the Waipipian of Waihi
Beach. The "Te Aute Limestone" (Opoitian-Nukumaruan, Pliocene-Early Pleistocene; Beu et
al. 1980) of Hawkes Bay region has yielded a few undescribed and well-preserved vertebrates
including a skull of a delphinoid, cf. Pseudorca sp. (National Museum of New Zealand
collections). Bearlin (1985; see also Gaskin 1972: Fig. 3) figured a skull of the mysticete
whale, cf. Balaenoptera sp. (National Museum of New Zealand collections), from Opoitian
mudstones in Taihape. The North Island localities have been prospected little specifically for
vertebrates, and further study seems timely.

Galaxiid fish from the Dunedin district were stated to be of Pliocene age (McDowall 1976),
but they are probably from the Miocene (see J.D. Campbell 1985a for age).

Cainozoic: Quaternary

There are few Quaternary vertebrates clearly older than Holocene, that is, from Wanganui
Series sediments other than the Haweran Stage (sensu Beu et al. 1987). Most of the
Quaternary vertebrates are Holocene (Haweran) birds, extracted from widespread post-glacial
sediments (e.g. loess, peat, mud, dunes; Brewster 1987, Forrest 1987, McCulloch 1985,
Marwick 1937) that are within the roughly 36,000 year range of radiocarbon dating. Millener
(this volume) gives more details. There are a few records of moa-bearing sequences dated at
about 10 000 yBP, for example, Pyramid Valley (Scarlett 1955a) and Waipara (M. G. Harris
material, Geology Museum collections, University of Otago). An older sequence, quoted as
about 36,000 yBP, is the Cape Wanbrow (Oamaru) deposit (Grant-Mackie & Scarlett 1973).
This fauna includes birds, tuatara, and seals.

Older Pleistocene vertebrates in New Zealand are rare. Outcrops are restricted, and are
difficult to date because there are fewer biostratigraphic markers than earlier in the Cainozoic,
Fleming (1953) described the geology of the Wanganui district, whence came bones of fish,
moas including material referred to Dinornis robustus, the yellow-eyed penguin (Megadyptes
antipodes), and an undescribed otariid seal, all from the Nukumaruan (latest Pliocene-Early
Pleistocene) or Okehuan (Fleming 1953: 156, 175, 209, 1979: 75, Marshall 1919, Park
1887a). King (1983a) recently described an extinct new species of sea lion, Neophoca palatina,
based on a skull from Castlecliffian sediments at Ohope. Dolphin vertebrae (Geology
Museum, University of Otago) were found at Mowhanau Beach, Wanganui district, in the base
of the Kai-Iwi Siltstone; the specimens are probably of later Castlecliffian age (Late
Pleistocene). Fleming (1978) gave an overview of Quaternary faunal history.

FOSSIL VERTEBRATES OF NEW ZEALAND - 1215

SYSTEMATIC SUMMARY OF NEW ZEALAND FOSSIL
VERTEBRATES

FISH: OVERVIEW

The term "fish" is widely thought to describe a natural group of vertebrates, but it is an
artificial term of convenience which describes a nonmonophyletic (strictly paraphyletic) group.
Fish include all vertebrates except the tetrapods. Typically, they encompass jawless fish
(Agnatha; no reported New Zealand fossil records), placoderms, sharks and rays
(chondrichthyes), spiny sharks (acanthodians), bony fish (actinopterygians, including teleosts or
true bony fish), coelocanths (crossopterygians; no reported New Zealand records) and lungfish
(dipnoans; no reported New Zealand records). The characteristics of many of these groups are
discussed elsewhere in this volume, and are summarised in a variety of texts (e.g. Carroll 1988,
Colbert 1980, Romer 1966) and more advanced articles.

Only Chondrichthyes and teleosts are reasonably common in New Zealand, and almost all
are Cretaceous and Cainozoic (Table 1). None of the reported pre-Cretaceous fish appears to
have been described formally, and careful research would probably expand the record
significantly. Although fossil fish are relatively abundant in New Zealand, they have received
rather less attention than have the reptiles, birds and mammals.

There are many incidental references in literature to field occurrences of scales, teeth, and
isolated bones, but there are rather few formal descriptions published. Noteworthy older articles
include the major works by Davis (1888a; the first major review) and Chapman (1918), and
smaller articles by Newton (1876; on chimaeroids), Hector (1881b, 1894), Davis (1886,
1888b, 1894), Chapman (1934), Chapman & Pritchard (1904), Frost (1924, 1928, 1933; on
otoliths), Marples (1949a) and Stinton (1957). These, and more-recent works, are discussed
below.

PLACODERMS

It has been known since 1967 that scraps of fish occur in the Devonian at Reefton.
Recently Macadie (1985) observed that the fauna includes an arctolepid arthrodire (a primitive
placoderm) that "resemble[s] Actinolepis". Macadie reported scales and bone fragments of
arthrodires (another placoderm group), and of two teleostome groups, palaeoniscids and
acanthodians. Further study, especially field work, could be rewarding.

CHONDRICHTHYES

Chondrichthyes - sharks, rays, elephant fish and relatives (Carroll 1988: Fig. 5.2) - are
fairly well represented in Late Cretaceous and younger New Zealand sediments. There are no
significant formal descriptions of older material. Sharks, in particular, have received
considerable attention from amateurs because of the spectacular appearance and often good
preservation of their teeth. Early work, both in New Zealand and elsewhere, resulted in a
proliferation of names as each new tooth form was given a new species name. At present
many old names are being synonymised.

Mr I.W. Keyes (New Zealand Geological Survey) has done a great deal to review New
Zealand fossil sharks and rays. Articles published include a revision of New Zealand records of
species of Carcharodon, sensu lato (Fig. 10) (Keyes 1971, 1972) related to the living great
white shark. The very large robust triangular teeth of the extinct C. megalodon (=

1216 - FORDYCE

Procarcharodon megalodon of some authors) are known from the Whaingaroan to Opoitian in
both the North and South Islands. C. auriculatus (extinct) ranges from Porangan to Altonian,
and possibly Mangapanian. The extant C. carcharias ranges in New Zealand from Opoitian to
Recent.

Keyes (1977) reported the first Southern Hemisphere record of the sawfish Onchopristis
dunklei (Batoidea - skates and rays). This ganopristine sawfish is known from elongate rostral
teeth (Fig. 10) extracted, using acid, from richly fossiliferous Piripauan (to Haumurian?;
Maastrichtian, Late Cretaceous) shallow marine sediments at Mangahouanga Stream, Te Hoe
River, Hawke's Bay. The local record significantly post-dates the youngest (Cenomanian)
northern hemisphere records.

Keyes (1979) proposed a new genus of sawshark, /kamauius (Fig. 10) (Pristiophoridae) for
Pristiophorus ensifer (Davis). The gracile elongate barbed rostral teeth are reasonably common
in local Late Eocene to Early Pleistocene marine sediments. Keyes suggested that previous
reports of the related Pliotrema (also Pristiophoridae) from New Zealand are incorrect. Keyes
(1982) also reviewed the related Cainozoic sawshark Pristiophorus lanceolatus (Davis). This
smooth-toothed species also has a long range: Keyes reported it from the Middle Eocene to
Pleistocene of New Zealand, and the Early Miocene to Early Pliocene of Australia.

The first New Zealand records of the genera and species Megascyliorhinus cooperi,
Centrophorus squamosus, and Scymnorhinus [= Dalatias] licha were reported by Keyes (1984a),
who based the identifications on isolated teeth. All species are rather common, but are small
and only collected easily by screen-washing, a technique little-used in New Zealand before. The
extinct Megascyliorhinus cooperi (Scyliorhinidae - "catsharks"), which has "long, reflexed,
conical, flat-rooted teeth", ranges from Whaingaroan to Haweran. The extant Centrophorus
squamosus (Squalidae - "spiny dogfish") has a Haumurian to Mangapanian record in New
Zealand. Scymnorhinus [also known as Dalatias] licha (extant; deep water) includes Bortonian
to Mangapanian records.

Keyes also reviewed records of Eocene elasmobranchs from Chatham Island (Keyes 1987),
and commented on the stratigraphy of Late Cretaceous and Palaeocene species (Keyes 1981a;
identifications in Feldmann 1984; appendix in Wiffen 1980: 527),

Few recent contributions have been made by other authors. Grant-Mackie (1982) reported
the discovery of an articulated vertebral column, apparently from a shark, from the Whangarei
Limestone, Bream Bay, The specimen has not been described formally. Similar material is
known from the Landon Series of South Canterbury and Otago (fossils in Geology Museum,
University of Otago). Pfeil (1983) described a new species of Pseudoechinorhinus from the
Waimakariri "Lingula beds" (Teurian in part) and also (1984) listed 12 species of
elasmobranchs represented by teeth of possible Waitakian or younger age, from Chatham Rise.

Figure 10. Chondrichthyan fossils from New Zealand. A, B, [kamauius ensifer (Davis); specimen in the
Geology Museum collections, University of Otago, scale bar, 10 mm.; C, D, Lamna sp., specimen in the
Geology Museum collections, University of Otago;, scale bar, 10 mm; E, Myliobatis sp., specimen in the
Geology Museum collections, University of Otago, scale bar, 10 mm; F, Onchopristis dunklei, after Keyes
1977, scale bar, 10 mm; G, H, Dasyatis sp., specimen in the Geology Museum collections, University of
Otago, scale bar, 1 mm; I, Heptranchias sp., specimen in the Geology Museum collections, University of
Otago;, scale bar, 10 mm; J, Lamna sp., specimen in the Geology Museum collections, University of Otago,
scale bar, 5 mm.; K, Carcharodon megalodon; specimen in the Geology Museum collections, University of
Otago, scale bar,10 mm.

FOSSIL VERTEBRATES OF NEW ZEALAND - 1217

~>CECe Gecee- ==

—

1218 - FORDYCE

Gregory et al, (1979; 1983) described fossil traces that may have been made by eagle rays,
probably Myliobatis sp. Toothplates of Myliobatis sp., usually worn, occur in sandy facies of
Cainozoic age (e.g. Fig. 10), but they have not received serious attention in New Zealand.

TELEOSTS

Otoliths

Teleost otoliths were first studied over 50 years ago (Frost 1924, 1928, 1933) but, apart
from the work of Stinton (1957) it is only recently that they have been restudied seriously.
Schwarzhans (1980) completed a Ph.D. thesis on local Cainozoic otoliths which was recently
translated into English (Schwarzhans 1984). The roughly \60 nominal species described were
from mostly shallow-water sequences of Eocene or younger age, and some material was from
slage stratotypes. The taxonomic status of some species is not clear, as they are described in
open nomenclature under form-genus with the suffix -arum (see editorial note by Simes in
Schwarzhans 1984: 2, and Patterson 1987). In this case, such form genera are merely taxa of
nomenclatural convenience, and are used to hold specics that are thought to be new but for
which generic affinities are uncertain. Schwarzhans (1981a) gave further information on the
taxonomy of local species, and (1981b) published a summary of otolith palacozoogeography of
the New Zealand - South Australian region. The narrative historical zoogeography is
interpreted in light of changing continents and oceanic currents.

Grenfell (1984; abstracts published 1981, 1983) described an otolith fauna from Otaian-
Altonian (Early Miocene) Parengarenga Group of Northland, which included 55 species in 22
families. The fauna is predominantly deeper-water, and is dominated by the families Congridae,
Sternoptychidae, Myctophidae, Moridae, Bregmacerotidae, Macrouridae, Hoplichthyidae and
Gobiidac. The many genera identified on the basis of otoliths are listed in Table 1.

There is probably great potential for more work on otoliths, especially if detailed
stratigraphic research can be done, for example, through a succession of stages at one locality.
The existence of effectively a dual classification scheme, for otoliths on one hand, and taxa
known from complete skeletons on the other, is an impedi nent to phylogenetic studies using
fossils.

Marine Teleost Skeletons

Marine teleost fossils, other than those known from otoliths, have been studied little.
Chapman (1918) described a new species, Diplomystus coverhamensis, from Cover Stream,
Marlborough (?Ngaterian, late Early Cretaccous). He discussed ?Thrissopater sp. (Sawpit
Gully, Marlborough; Cretaccous or Early Tertiary) and Scombroclupea sp. (Weka Pass,
Canterbury; probably Duntroonian, Late Oligocene). Chapman (1934) later described two new
species: "Portheus" dunedinensis, trom Abbotsford, Dunedin, Heretaungan Stage (Early Eocene;
Geology Muscum collections, University of Otago) and Eothyrsites holosquamatus, from
Burnside, Dunedin, Bortonian-Runangan Stages (Late Eocene). The generic identification as
Portheus |strictly, Portheus = Xiphactinis), is unlikely, since the order Ichthyodectiformes is
only known from the Mesozoic. Chapman's work on fossil taxa other than teleosts (e.g.
foraminifera, chondrichthyans and cetaceans) is not regarded as reliable, and a review of his
research on teleosts seems long overdue.

Wiffen (1983) described a relatively complete specimen of the clupeiform, Pachyrhizodus
caninus (Clupeitormes: Pachyrhizodontidae). This was from informally-named Maungataniwha
sandstone (Piripauan or Haumurian, Late Cretaceous), at Mangahouanga Stream, Hawke's Bay.

FOSSIL VERTEBRATES OF NEW ZEALAND - 1219

Material of this large pachyrhizodontid includes elements of the skull, jaws i
Carroll (1988) placed Pachyrhizodus in the Elopiformes. ie aac) ie

Some groups that might be expected to be identified from distinctive elements have been
listed rarely. For example, the beaks or rostra of billfish (swordfish, marlin) seem not to have
been mentioned since the record of Parker (1892). An incomplete beak, not yet identified
formally, was recovered recently from the Otekaike Limestone of Waitakian Stage at Otiake
(Late Oligocene-earliest Miocene; Geology Museum collections, University of Otago).

Most institutions that house local fossils hold potentially important skeletal fish material.
For example, there is an undescribed flatfish in the New Zealand Geological Survey, collections
at Lower Hutt, and there are Cretaceous and Cainozoic teleosts in the Geology Museum at
University of Otago. In the course of field work mostly. on the middle Cainozoic, I have
occasionally noted articulated or little-disturbed fossils (e g. Landon Series, Oligocene: near
Karamea, Kaikoura, Rangitata River, Waihao River, and various localities in North Otago).
There seems to be scope for further studies on marine teleost skeletal fossils from the
Cretaceous-Cainozoic sequences, but whether such studies would be appropriate for higher
degree theses is uncertain. Studies of otoliths could be expanded beyond the valuable
contributions of Schwarzhans and Grenfell.

Freshwater Fish

The few fresh-water fish reported in print are all from the Cainozoic and all those mentioned
in the literature appear to be species of galaxiids. These are typically small, slender, primarily
fresh-water fish (Fig. 11) with cool-temperate Southern Hemisphere distributions. McDowall
(1970, 1978) reviewed extant New Zealand galaxiids.

McDowall (1976) briefly reviewed the best-known previous fossil discoveries, from Otago,
which had been discussed by Oliver (1936), Stokell (1945, 1972) and Whitley (1956a, 1956b:
34, as Galaxias kaikorai n. sp.). McDowall concluded that fish from Kaikorai Valley, near
Dunedin (Geology Museum, University of Otago) probably represent the extant species
Galaxias brevipinnis (Fig. 11). Autapomorphies (evolutionary novelties diagnostic at species
level), however, were not listed, and some doubt must remain about the identifications.
McDowall cited the age as Pliocene, and this has been followed by others (e.g. Carroll 1988:
605, Patterson 1967), although J.D. Campbell (1985a) indicated that plant fossils from
Kaikorai Valley are probably Late Miocene in age. McDowall also observed that galaxiids of
probably Taranaki-Waitotaran age (Late Miocene-Late Pliocene) are known from Foulden Hills,
near Middlemarch, Otago (Geology Museum, University of Otago). According to McDowall,
the Foulden Hills fossils probably represent the extant species Galaxias vulgaris.

McDowall noted that a supposed record of Oligocene Galaxtas from New Zealand (Romer
1966: 356) is wrong. Other fresh-water fish from lacustrine-fluvial Tertiary sediments in
Central Otago have been mentioned in print (e.g. Park 1906, 1908, Ferrar 1927, 1929,
Douglas 1986, Douglas et al. 1981 - Geology Museum collections, University of Otago), and
other material from St Bathans and Bannockburn is in the Geology Museum collections. None
of these fossils has been described formally yet.

Oliver (1928: 287) mentioned fossil fish similar in size to extant species of Gobiomorphus
(Galaxiidae). The fossils were obtained from the "Waipaoa Series" near Ormond, Gisborne
district, Hawke's Bay, and are probably Castlecliffian in age (Late Pleistocene; Suggate in
Suggate et al. 1978: 566). These specimens appear not to have been described.

1220 - FORDYCE

Figure 11. Galaxiid fossil from Dunedin, Galaxias sp.; specimen in the Geology Museum collections,
University of Otago, scale bar, 10 mm. (Body outline of Galaxias brevipinnis (length 185 mm) after
McDowall).

Galaxiid historical zoogeography has been discussed widely, without any significant
reference to fossils. References include those of Allan (1956) and McDowall (1978, 1980a).

Incidental Records of Fish

Many records of the occurrences of fish have been made incidental to work on Cretaceous-
Cainozoic marine rocks. The records, which attest to the abundance of fossils and the potential
for future work, include those of Adams (1983a and 1983b), Bell & Clark (1909: pl. 12),
Boreham (in Gage 1970: 554-555), Buchanan (1870: 165), Buckeridge (1984), Evans (1983 and
1986), Ferrar (1925; 38 and 1934: 33, 36), Fleming (1953: 192), Fordyce et al. (1985), Fraser
& Adams (1907: 55), Gage (1970: 515), Gair (1959: 274), Gudex (1918: 245-253), Haast
(1879: 306-307, 311), Hector (1881b), Henderson (1917: 94), Henderson & Grange (1926: 54),
Hutton (1887a: 399, and 1888a: 260), Mantell (1850: 329), McKay (1877d: 161, 1881a: 63,
68 and 70, 1881b: 82, 1888: 47 and 1890: 155-160), McKee (1984), Marwick (in Ongley
1939: 59, 61), Ongley & MacPherson (1928: 41, 48), Park (1886: 167, 1887a: 57, 1887b:
172-173, 1887c: 226), Parker (1897), Purnell (1875: 453), Purser (1961: 12-13), Rodgers &
Grant-Mackie (1978), Smith (1877: 576), Speight & Jobberns (1928: 223), Suter (1921: 35),
Thomson (1919: 314, 316, 1926a: 349), Uttley (1916: 21), Warren & Speden (1977: 22) and
Wells (1987: 107). This list is not necessarily comprehensive, since many other minor
references are likely to be in the literature.

AMPHIBIA: OVERVIEW

Amphibians were dominant and diverse terrestrial vertebrates for a time in the Paleozoic
(Carroll 1988), yet there is a negligible fossil record in New Zealand. Indeed, no pre-
Quaternary amphibians have been positively identified from New Zealand. "Labyrinthodont"
teeth from the Triassic (Hector 1886, Worley 1894) could represent amphibians, but are more
likely to represent ichthyosaurs (see below), and the whereabouts of the specimens is unknown,
so their identity cannot be checked. It is possible that pre-Quaternary amphibian remains
eventually will be found in New Zealand (e.g. in freshwater Miocene sediments of the
Manuherikia Group, Central Otago) but generally the lack of nonmarine sediment, and the
induration and deformation of Tertiary and older sediments counts against significant
discoveries.

FOSSIL VERTEBRATES OF NEW ZEALAND - 1221

SUBFOSSIL FROGS: LEIOPELMATIDAE

Three extant species of the endemic anuran genus Leiopelma (Leiopelmatidae, ?Lemnanura)
comprise some of the most unusual elements of the New Zealand terrestrial vertebrate fauna.
Leiopelma species are more semi-terrestrial than amphibious, and are limited in distribution.
They have been interpreted widely as a relict Gondwanan or archaic element of the New Zealand
biota, and have stimulated a great deal of discussion (e.g. Craw 1985, Fleming 1979: Fig. 32,
Robb 1986, Stevens 1985: 56) even in the absence of fossils older than Holocene.

Leiopelma is now much better understood than formerly as a result of the detailed modern
work by Worthy (1987a, 1987b). Worthy recognized the following species: 1, L. archeyi,
extant, Coromandel. 2, L. auroraensis, extinct; subfossil in South Island (Fiordland) only. 3,
L. hamiltoni, extant, Marlborough Sounds; subfossil elsewhere in South Island. 4, L.
hochstetteri, extant, northern North Island; also subfossil in South Island. 5, L. markhami,
recently extinct, North and South Islands. 6, L. waitomoensis, extinct; subfossil in North
Island only.

Worthy's work includes synoptic descriptions of the species based on osteology,
osteological comparisons of species, useful illustrations of bones, comparisons between
Ascaphus and Leiopelma, a cladistic analysis of Leiopelma, and a discussion of zoogeography.
The article also deals with the subfossil bones reported by Bull & Whitaker (1975) from
Paturau (Nelson), Martinborough and Coonoor (Wellington), Patoka and Hukanui (Hawke's
Bay) and Waitomo (Auckland) and by Bell (1978). The literature reviewed by Worthy obviates
the need for detailed references herein. According to Worthy, Leiopelma is a monophyletic
genus of probable Gondwanan origin, which encompasses two clades, a L. archeyi - L.
hamiltoni - L. waitomoensis group and a L. markhami - L. auroraensis - L. hochstetteri group.
The two groups may have diverged some 15 million years ago (Worthy 1987b: 409). Worthy's
work considered little about constructional morphology, and there may be scope for study here.

Worthy (1987b) discussed the palaeoecology of all species. Modern distributions are more
restricted than those of late Holocene fossils, and ranges may have contracted significantly
within the last 1000 years. The formerly widespread species were larger in body size in the
south.

There is potential for further work on Leiopelmatidae. No significant subfossils have been
found along the east of the South Island, and karst districts might well be searched, Older
(Tertiary) fossils might be expected in fluvio-lacustrine sediments, thus, it could be productive
to prospect in sediments such as those from the Miocene Manuherikia Group of Central Otago
(e.g. Douglas 1986). So far, concentrate from this unit (Geology Museum, University of
Otago) has yielded teleost and avian fossils down to microscopic sizes, but no frog bones.

REPTILIA: OVERVIEW

The fossil reptile record in New Zealand has long appealed to those interested in marine
reptiles, and within the last few years nonmarine fossils have increased the diversity.
Subclasses or orders of reptiles (classification after Carroll 1988) are represented locally as
follows:

Anapsida (extinct): no records.

Chelonia - turtles (extant): marine, Cretaceous and Tertiary; see below.

Ichthyosauria (extinct): Triassic, Cretaceous; see below.

Sauropterygyia (extinct): Cretaceous; see below.

Placodontia (extinct): no records.

Diapsida (Araeoscelida, Choristodera, Thalattosauria, Eosuchia; extinct): no records.

Sphenodonta (=Rhynchocephalia) (extant): Holocene; see below.

Squamata (extant): marine, Cretaceous; nonmarine, holocene; see below.

1222 - FORDYCE

Protosauria (extinct): no records.

Trilophosauria (extinct): no records.
Rhynchosauria (extinct): no records.

Thecodontia (extinct): no records.

Crocodylia (extant): no certain records; see below.
Pterosauria (extinct): Cretaceous; see below.
Omnithischia (extinct); Cretaceous; see below.
Saurischia (extinct): Cretaceous; see below.
Synapsida (extinct): no records.

By far the best known of the New Zealand fossils are two extinct fully marine groups from
the Late Cretaceous, the plesiosaurs and mosasaurs, Mesozoic ichthyosaurs and Cretaceous and
Tertiary turtles are known from scattered remains in marine rocks. Two species of dinosaur and
a probable pterosaur are known, but otherwise the only terrestrial reptile fossils are Holocene.
These groups are discussed below, and their records are summarised in Table 2.

TURTLES (CHELONIA)

The few fossil turtles found in New Zealand are from marine rocks, and there is little to
suggest that terrestrial or freshwater turtles might be expected in the fossil record. Since the
taxonomy of our local chelonians is understood poorly, the specimens are listed below in order
of decreasing age. All are from the the Cretaceous-Cainozoic sequence. Starting points for
further reading are articles by Gaffney (1981, 1984)

Wiffen (1981a) described chelonian plastron and carapace fragments from the informally-
named Maungataniwha sandstone, of Piripauan (possibly Haumurian) age (Late Cretaceous),
Mangahouanga Stream, Hawke's Bay. The bones appear to represent an indeterminate genus
and species of Protostegidae. This family includes mainly large taxa from the marine
Cretaceous (e.g. Protostega) of the Northern Hemisphere. Gaffney (1984) listed derived
characters (synapomorphies, or evolutionary novelties) of the family as including large stellate
plates in the hyoplastra and hypoplastra. With Dermochelyidae, protostegids have very reduced
dermal bone. Protostegids seem not to have been reported previously from the Southern
Hemisphere.

Fordyce (1980a) described the shaft of an incomplete limb bone, possibly a turtle humerus,
from the Amuri Limestone Formation (Teurian, Palaeocene), Ward, Marlborough (New Zealand
Geological Survey collections), Derived characters were not identified explicitly; the
provisional identification was reached through comparisons with figures provided by Romer
(1956). Strangely, the bone has large, possibly pathological, openings in what appear to be
the anterior and posterior faces. Another Teurian (Palaeocene) turtle, also undescribed, is
known from an opalised tuff, possibly the Red Bluff Tuff of H.J. Campbell et al. (1988), on
Chatham Island. The specimen includes long bones and carapace plates (New Zealand
Geological Survey collections, Lower Hutt).

At least three turtles are known from rocks of the Arnold Series. The partial carapace and
limb bones of a large turtle were collected recently by A. Kadar and B. Spdrli from the Pahi
Greensand, Opahi Group (Kaiatan, Late Eocene), on the shore platform at Pahi Peninsula,
Northland (Department of Geology collections, University of Auckland). The specimen is
currently (1989) under study at the Department of Geology, University of Otago. Marshall
(1917: 439) previously mentioned a reptile, still undescribed, from Opahi Group of Northland
(see also Fordyce 1980a: 740), but the whereabouts and identity of the fossil are unknown.
Another Eocene turtle was collected as a concretion by J. Goedert, from the Kaiata Mudstone
(probably Kaiatan, Late Eocene), near Woodpecker Bay, Westland. The small specimen
(Geology Museum collections, University of Otago) has only been prepared enough to reveal

FOSSIL VERTEBRATES OF NEW ZEALAND - 1223

that it includes carapace or plastron and limb elements. Of similar age is a recently discovered
large humerus, provisionally identified as that of a turtle, from a Bortonian or Kaiatan
mudstone at Boulder Hill, near Dunedin (Geology Museum collections, University of Otago).

_ Fordyce (1980a) mentioned undescribed bones of one individual, including fragments of
limbs and skull, and carapace or plastron (Canterbury Museum, Christchurch), apparently
quarried from a bryozoan limestone in the Oamaru district. Collection details are unknown, but
the style of the display case for the shell elements suggests late 1800s. Indeed, Hutton (1900b:
227) mentioned that New Zealand Tertiary reptiles on display at Canterbury Museum,
Christchurch "are represented by the bones of a turtle". The fossil was probably from the
McDonald Limestone (Runangan-Whaingaroan, latest Eocene-Early Oligocene), since the
lithologically similar but older Totara Limestone was quarried little for building stone. This
specimen was mentioned briefly by Marples (1949a: 104).

Buckeridge (1981) described a small femur, presumably that of a marine turtle, collected
from the Waikawau Formation (Otaian, Early Miocene) at Port Waikato (Department of
Geology collections, University of Auckland). He assigned it to a new species, ?Lepidochelys
waikatoica (Cheloniidae), even though the femur could not be placed positively at the generic
level. Such an action emphasises the possible value of nomenclature in communication but
not in identifying relationships. Alternatively, such material might better be described as
Cheloniidae genus and, therefore, species indeterminate.

Other published records of bones identified as those of Cainozoic reptiles have included at
least one of turtles (Hector 1876: 53, McKay 1877c:111), but the identity of none of these is
certain (Fordyce 1980a). The above records suggest that more finds should be made.
Surprisingly, some vertebrate-bearing units which have been studied intensively (e.g. Kokoamu
Greensand, Otekaike Limestone) have not yielded chelonians. The turtles have not been the
subject of a unified study by one individual, and a review seems timely.

ICHTHYOSAURS (ICHTHYOSAURIA)

New Zealand records of these highly specialised, dolphin-like marine reptiles (Fig. 12) are
rare. No relatively complete specimens are known. Local records were reviewed briefly by
J.D. Campbell (1965) and Fleming et al. (1971). The supposed Tertiary ichthyosaur,
Tangaroasaurus kakanuiensis Benham 1935a, is probably an Early Miocene shark-toothed
dolphin (Camp 1942, Fordyce 1978).

Previous Work

Campbell (1965) figured the partial jaws and teeth of a medium-sized indeterminate genus and
species of ichthyosaur from Otamitan (Late Triassic) sediments at Otamita Stream,Hokonui
Hills, Southland. Charig (in J.D. Campbell 1965) indicated that the specimen exhibits some
of the characters of the Family Shastasauridae.

Figure 12. Some Mesozoic reptiles from New Zealand. A, reconstruction of the skull of Prognathodon
waiparaensis, after Welles & Gregg, lower jaw is about 1110 mm long; B, reconstruction of plesiosaur,
Mauisaurus haasti (top) and mosasaur, Prognathodon waiparaensis (bottom), after Welles & Gregg; C,
vertebrae of indet. small mosasaur, specimen in the Geology Museum collections, University of Otago, scale
bar, 50 mm; D, exterior surface of left hypoplastron plate of protostegid turtle, after Wiffen, scale bar, 20
mm; E, reconstruction of an ichthyosaur, Platypterygius sp., after McGowan.

1224 - FORDYCE

FOSSIL VERTEBRATES OF NEW ZEALAND - 1225

Campbell (1965) also considered the status of Jchthyosaurus australis Hector, 1874 (not of
McCoy 1867), which later was reviewed in more detail by Fleming et al. (1971). I. australis is
based on vertebrae from the Torlesse Supergroup, Oretian Stage (Late Triassic), near Mt Potts,
central Canterbury (see Campbell & Force 1972). Fleming et al. concluded that /. australis
actually represents an indeterminate genus and species of ichthyosaur. Fleming ef al. also
reported that several other supposed ichthyosaurs had been collected from the Mt Potts area. It
may have been specimens from Mt Potts to which Hector (1879b: 77) referred.

New records of presumed ichthyosaurs from three localities in the Makirikiri Formation
(Motuan, Early Cretaceous) of the Tinui district, east Wellington (Fleming et al. 1971) are the
first from the North Island. All three specimens, unfortunately, are indeterminate. Stevens
(1974: 17, 20) reported a single ichthyosaur vertebra of presumably Triassic age, from
Houghton Bay, Wellington. Stevens also stated (1985: 57) that ichthyosaurs occur in the
Jurassic, but did not give details.

Fleming et al. (1971) mentioned the "teeth having Labyrinthodont characters” reported by
Hector (1886, 1880c) and Worley (1894) from unspecified formations (probably Triassic) at
Nugget Point, South Otago, and Wairoa district, Nelson. These could represent ichthyosaurs,
but could also have been amphibian teeth. The specimens were never described formally, and
their whereabouts are unknown, so their identity is uncertain. Park (1904) reported fragmentary
possible ichthyosaur remains from near Nelson.

New Records

Professor J.A. Grant-Mackie reported (pers. comm.) that an undescribed fragment of
ichthyosaur rostrum of presumed Late Cretaceous age was collected near Dargaville, North
Auckland (Department of Geology collections, University of Auckland). The only Cretaceous
ichthyosaurs known, according to McGowan (1972a), are species in the genus Platypterygius
(Family Stenopterygiidae, according to Romer 1966; Leptopterygiidae according to Carroll
1988). Whether this New Zealand specimen belongs here is uncertain.

Ichthyosaur vertebrae were collected recently from a pebbly sandstone (Otamitan Stage) near
Nugget Point, South Otago (Fordyce cited in J.D. Campbell 1987; Geology Museum
collections, University of Otago). J.D. Campbell and J.G. Begg also collected a tooth and a
series of ichthyosaur vertebrae from a pebbly sandstone (Etalian Stage, Middle Triassic) at Etal
Stream, Southland (Geology Museum collections, University of Otago). None of these
specimens has been prepared or described.

PLESIOSAURS (SAUROPTERYGIA)

Plesiosaurs are wholly aquatic, extinct reptiles. Later Mesozoic taxa represent two main
groups: the plesiosauroids, with a small head and long neck, and the pliosauroids, with a large
head and short neck. Both have oar-like limbs, and appear to have been quite streamlined.
General reviews were given by Carroll (1988) and Romer (1956, 1966), and more detailed
works are those of, for example, Brown (1981) and Persson (1963).

Plesiosaurs are perhaps the most conspicuous of New Zealand's fossil reptiles (e.g. Figs 8,
9; Pl. 1). Together with mosasaurs, the Late Cretaceous plesiosaurs were the subject of a
comprehensive review by Welles & Gregg (1971), who covered taxonomy, stratigraphy and
history of study.

Previous Work

Welles & Gregg (1971) presented a detailed history of discovery and description, and a
chronological review of previous discoveries; this information is not repeated in full here. As

1226 - FORDYCE

might have been expected, Alexander McKay was an important early collector. Most of the
specimens were described by Hector (e.g. Hector 1874, 1877b, Knight 1875), Haast (e.g. 1870)
and Owen (e.g. 1861) between about 1860 and 1890. There was no important systematic work
carried out between the 1890's and the late 1960's, when Welles & Gregg initiated their studies,
although New Zealand specimens were mentioned in review articles (e.g. Persson 1963). More
recent work is mentioned below.

Localities and Age

Welles & Gregg (1971; see also McKay e.g. 1877a, 1877e) indicated that nearly all
specimens of Late Cretaceous marine reptiles known from New Zealand had been collected from
three localitics in North Canterbury (Fig. 12). The source horizon is mostly or wholly the
Conway Formation as used by Browne & Field (1985) and Warren & Speden (1977), which is
equivalent in part or whole to the Laidmore Formation and "Saurian Beds" of older authors.
This unit is of Haumurian (latest Cretaceous) age. Localities include the upper gorge of
Waipara River (see Clark 1861, Haast 1870, Hood 1870, Hutton 1894, McKay 1877a), the
Cheviot district (see Hector 1893, Henderson 1921, Keyes in Feldmann 1984, Knight 1874) ,
and Haumuri Bluff (see Hector 1870b: 198, 1873b: 5, Knight 1874, McKay 1877a, and Warren
& Speden 1977).

More recent work has presented new information on occurences of Late Cretaceous
plesiosaurs. Warren and Speden (1977: Table 2, p. 22) documented the stratigraphic
distribution of plesiosaurs in the region of Haumuri Bluff and also discussed the stratigraphy of
the Late Cretaceous Piripauan and Haumurian Stages. Wiffen (1981la: 527; Wiffen & Moisley
1986) noted that plesiosaur remains (now New Zealand Geological Survey collections, Lower
Hutt) had been found at Mangahouanga Stream, The published records (e.g. Fordyce 1980a,
1987b, Hornibrook 1962, Keyes 1981a, Welles & Gregg 1971) emphasise that, despite early
reports to the contrary (McKay 1877e: 37, Thomson 1920), no New Zealand plesiosaur is
known positively to be younger than Haumurian (latest Cretaceous). Reports of plesiosaur-
like teeth from the Tertiary on the northern shores of Lake Wakatipu (Hector 1880a: 10,
McKay 1881d: 145, 1894: 13) probably refer to cetacean tecth (Benham in C.O. Hutton 1939).

The oldest local record of a supposed plesiosaur is that of a small problematic bone from
Oretian (Late Triassic) rocks at Marakopa, southwest Auckland (Campbell 1965). The
identification of the specimen, by W.E. Swinton, according to the specimen label (in the
Geology Museum collections, University of Otago), has not been confirmed.

Undescribed Specimens

Keyes (1981la) mentioned undescribed clasmosaur teeth from sediments of probable
Haumurian age in part, the "Lingula bed", at Otarama, on the Waimakariri River, Canterbury.
Neil Fowke collected much of the postcranial skeleton of a juvenile plesiosaur from Waipara in
about 1974, The prepared but undescribed specimen (now in the Geology Museum, University
of Otago) appears to have scale or skin impressions preserved. The most recently recognised
plesiosaur locality in New Zealand is Shag Point, North Otago, whence a relatively complete
¢. 7 m long specimen was recovered. The fossil is in the Geology Museum collections,
University of Otago (Fig. 9; Pl. 1; Fordyce 1983c, 1986, 1987b: 73).

Taxonomy

Many names have been proposed for the New Zealand specimens (Table 3). Welles &

FOSSIL VERTEBRATES OF NEW ZEALAND - 1227

Gregg (1971) and Wiffen & Moisley (1986) recognised five species. Of these, the elasmosaur
Mauisaurus haasti Hector, 1874 (see also Hector 1873b) (Plesiosauroidea) derives its identity
ultimately from the lectotype pelvis and paddle, although other elements, from both the classic
oe Island localities and Mangahouanga Stream, have been referred to it. The skull is
unknown.

Tuarangisaurus keyesi Wiffen & Moisley, 1986 (Plesiosauroidea: Elasmosauridae) is known
only from the well-preserved holotype skull and apparently associated cervical vertebrae from
Mangahouanga Stream. The species lacks the range of associated postcranial elements that
would allow easy comparison with many other nominal species. Wiffen & Moisley referred
some postcranial elements to it.

Welles & Gregg (1971) also recognized two indeterminate species, an elasmosaur
(Plesiosauroidea), which encompasses 6 nominal species of older authors, and a polycotylid
(Pliosauroidea), which encompasses 5 nominal species of older authors. Wiffen & Moisley
(1986), however, recognized only the family Pliosauridae, to which they referred
Mangahouanga Stream specimens and the indeterminate polycotylid of Welles & Gregg. In the
use of the family Pliosauridae they presumably followed Brown's (1981) abandonment of
Polycotylidae.

Future Work

It is difficult for those other than specialists on plesiosaur taxonomy to assess the
relationships of the New Zealand species, since the "evolutionary taxonomy" of the plesiosaur
workers has rarely provided explicit reasons for clustering specimens and species. Indeed, all
the published "diagnoses" are really synoptic descriptions which say little about similarities and
differences. One looks in vain for details of derived characters. Regrettably, these criticisms
can and should be aimed at much of the work on local fossil vertebrates. Future work on New
Zealand plesiosaurs might, therefore, usefully reassess relationships of described specimens
cladistically. The eventual description of specimens such as the large articulated plesiosauroid
from Shag Point may resolve the identity of nominal species based on non-comparable
elements. Further field work in promising localities (e.g. Northland, Raukumara Peninsula,
and eastern South Island) must eventually turn up new material.

TUATARA (SPHENODONTA = RHYNCHOCEPHALIA)

There are no fossil records of the tuatara (Sphenodon punctatus; Fig. 13), New Zealand's
largest and best-known extant reptile. Reviews of tuatara biology were given recently by
contributors in Newman (1982), in particular Dawbin (in Newman 1982: 152-3, on
palaeontology, and p. 164-5 on the skeleton). For figures of tuatara bones, see especially
Romer (1956, e.g. figs 60, 62, 63, 108); figures of odd bones were given, e.g. by T.H. Rich et
al. (1979) and Scarlett (1972b).

Crook (1975) cited a personal communication from T.H. Rich which indicated that Miocene
fossils of tuatara had been discovered, but these specimens are known now to be Recent,
probably younger than 1,000 years, and definitely no older than 10,000 years (T.H. Rich et al.
1979). Grant-Mackie & Scarlett (1973: 92) stated that the oldest tuatara bones known are those
from the Hillgrove Formation near Oamaru, North Otago. The age was given as Oturian, or
Last Interglacial, with a radiocarbon date of approx. 36,000 years old (at about the effective
limit of radiocarbon dating) . M.S. Pole (pers. comm.) reports that tuatara bones occur
sporadically in Late Quaternary horizons in Central Otago.

1228 - FORDYCE

Figure 13. Skull and body outline of extant Tuatara, Sphenodon punctatus. Skull after Romer, body outline
after Crook. Skull about 55 mm long.

Future work could involve prospecting in nonmarine Tertiary rocks, as suggested herein for
other terrestrial vertebrates. The youngest fossil records of Sphenodonta (rhynchocephalians)
other than the New Zealand Quaternary records are from the Cretaceous of North America
(Throckmorton ef al. 1981). However, there are enough differences between the Cretaceous
species and the living tuatara that the latter probably should not be called a living fossil
(Benton 1986).

MOSASAURS AND OTHER LIZARDS (SQUAMATA)

Lizards

There is no published record of significant fossils of terrestrial lizards, although bones are
known from cave deposits (Hutton 1899a, Worthy 1984) and occur in other Holocene
sediments (Gill 1985, Rich et al. 1979, Worthy 1987c). Any search for Cainozoic terrestrial
reptile fossils might profitably screen-wash finer-grained fluvial and lacustrine sediments.

As with tuatara, reviews of the biology of extant indigenous lizards were given by
contributors in Newman (1982) and by Robb (1986). These and other works include discussion
about the historical origins and zoogeography of New Zealand's terrestrial reptiles. Whitaker &
Thomas (1989) gave a guide to literature about New Zeeland lizards which lists articles on
subfossil lizards.

Snakes

There are no reports of fossil snakes. The occasional modern occurrences of sea-snakes
(Robb 1986) suggest that fossil sea-snakes might be expected in more northern marine
sediments. New Zealand has extant land vertebrates, as well as fossils (e.g. Theropoda), whose
ancestors must have been present here since New Zealand separated from adjacent landmasses
about the middle Cretaceous. This was some time after the earliest snakes appeared, so New
Zealand may well have had fossil land snakes (as well as fossil mammals - see below) which
did not persist to the present. The absence of snakes from New Zealand's modern vertebrate

FOSSIL VERTEBRATES OF NEW ZEALAND - 1229

fauna and/or the fossil record has been mentioned elsewhere (e.g. Fleming 1962a: 65, Caughley
1964: 51, Stevens 1980a: 174).

Mosasaurs

Mosasaurs, large aquatic Late Cretaceous reptiles, are the only members of the Squamata
known as fossils in New Zealand. Mosasaurs were reviewed by Welles & Gregg (1971) who
summarised the history of collection and study.

Mosasaur Taxonomy

Welles & Gregg recorded five species, all from Haumuri Bluff, Cheviot, and Waipara. Two
tylosaurines are known. The large Tylosaurus haumuriensis (Hector, 1874) has a skull
reportedly longer than 1 m. Taniwhasaurus oweni Hector, 1874, is known only from the
lectotype skull of an old individual. Prognathodon waipuraensis Welles & Gregg, 1971, a
plioplatycarpine, is represented only by the holotype (Fig. 12). This also apparently has a
skull longer than 1 m. A species of mosasaurine, Mosasaurus mokoroa Welles & Gregg,
1971, also represented only by the holotype, has a skull some 700 mm long. A fifth,
indeterminate, species includes specimens referred to 3 nominal species. Many of the old
names proposed by Hector, Hutton and Owen for both mosasaurs and plesiosaurs are synonyms
of the above or are nomina vana (Table 2), according to Welles & Gregg.

More recently, Wiffen (1981) described a new genus and species of mosasaurine,
Moanasaurus mangahouangae, based on a single specimen which includes a skull, vertebrae,
paddle and ribs. The fossil came from the informally-named Maungataniwha sandstone, of
Piripauan (or possibly Haumurian) age, at Mangahouanga Stream on Te Hoe River, Hawke's
Bay. It is the first mosasaur to be described from the North Island. Recently Wright (1989)
convincingly argued that Moanasaurus is a junior synonym of the problematic genus
Mosasaurus. In doing so, Wright reiterated the need for clear diagnoses of new taxa.

New Records of Mosasaurs

Keyes (1981a) mentioned that mosasaur teeth occur in a probable Haumurian unit, the
"Lingula bed", at Otarama, on the Waimakariri River in Canterbury. Vertebrae from both
small (probably juvenile) and large mosasaurs are known from sandstones and siltstones of the
Katiki Formation near Shag Point, North Otago (Fordyce 1987b; Geology Museum University
of Otago, Fig. 12). These fossils, not identified to spesies level, appear to be the most
southerly records of mosasaurs in New Zealand.

Future Work on Mosasaurs

The comments above, under plesiosaurs, apply to mosasaur work as well. In particular,
cladistic analyses might reveal more about the relationships of the New Zealand species with
overseas taxa. All the New Zealand species are described as endemic, and one or two are placed
in apparently monotypic genera. However, as with other vagile nektic vertebrates, there could
be close affinities with taxa overseas.

CROCODILES (CROCODYLIA)

Reports of fossil crocodiles from New Zealand appear to be erroneous. Welles & Gregg
(1971) noted the uncertain identity of Crocodilus novaezealandiae Hector, 1886 (1874: 334),
which was based on vertebrae of Haumurian (latest Cretaceous) age from Haumuri Bluff,
southern Marlborough. Procoelous vertebrae from Waipara, identified as crocodilian by Hood
(1870) and Haast (1870), may well have been those of mosasaurs.

Haast (1879: 311; see also Hutton 1887b, McKay 1887b) listed "Teeth of Crocodilus sp.
Waihao" from the “Oamaru Formation", South Canterbury. These teeth are not reptilian but

1230 - FORDYCE

are similar to those of species of the cetacean genus Dorudon (see Fordyce 1985b, and section
on Cetacea).

DINOSAURS (SAURISCHIA AND ORNITHISCHIA)

The apparent absence of dinosaur fossils in New Zealand was long regarded as sufficient
reason not to look for them or at least to assume that the absence was real. Indeed, discussions
of the New Zealand vertebrate fauna often stressed the lack of dinosaurs (e.g. Fleming 1962a:
93, with a caution that the lack of evidence does not always provide evidence of absence;
Stevens 1983: 53). Thus, it was of great interest when a single caudal vertebra (Fig. 8) of an
indeterminate species of apparent theropod dinosaur was discovered recently by Joan Wiffen at
Mangahouanga Stream. The vertebra was described and discussed by Molnar (1981; see also
1980), who noted that it resembles those of theropods, although ornithopod affinities cannot be
dismissed. Like the other vertebrates from the same site described by Wiffen (e.g. 1980,
1981a, 1981b, 1983) the bone is of Piripauan or Haumurian age (Late Cretaceous). Molnar's
(1981) description includes a provocative discussion of home range, land area and population
size.

Wiffen (1981a: 527) indicated that a phalanx of a bird or theropod was found at
Mangahouanga Stream. This is presumably the specimen described by Molnar & Scarlett
(1984) as that of a Late Cretaceous terrestrial bird or dinosaur. Scarlett recently indicated that
he now considers the phalanx to be that of a dinosaur (cited by B. McCulloch in an informal
newsletter, the Letter of information of the Society of Avian Paleontology and Evolution 1:
10, 1987). In the same article Scarlett indicated that an ornithopod dinosaur is now known
from Mangahouanga Stream, This is probably the specimen described by Wiffen & Molnar (in
press) as the partial ilium of a Dryosaurus-like ornithopod.

PTEROSAURS (PTEROSAURIA)

Wiffen & Molnar (1988; Wiffen 1986) reported that the distal end of a left ulna of a
pterosaur was recovered from Mangahouanga Stream. The bone is presumably of Piripauan or
Haumurian age (Late Cretaceous). The individual was estimated to have a wing span of 3.75 m
(Wiffen 1986). The inferred palaeolatitude is further south than any previously reported find of
pterosaurs.

It is reasonable to expect other specimens to be found in New Zealand, probably in
nonmarine or proximal marine sediments. Pterosaurs are one of the few groups that, perhaps
Surprisingly, have attracted no previous comment in the literature about local
palaeozoogeography. Most attention has focussed on terrestrial taxa, rather than volant groups.

MISCELLANEOUS AND PROBLEMATIC RECORDS OF REPTILES

There are other records of large bones, presumably reptilian, from Mesozoic rocks, but the
identity of these is uncertain (e.g. specimens in collections of the Geology Museum at
University of Otago, New Zealand Geological Survey, and Victoria University, Wellington).
Most rocks older than Late Cretaceous are so well cemented with silica and/or zeolites that
much effort is needed to prepare what are usually isolated bones, with little guarantee of
extracting diagnostic elements. Comments on some occurrences follow.

One specimen which may ultimately prove identifiable is that of a large vertebrate,
probably a reptile, from the Lower Triassic (or uppermost Permian) at Mossburn, Southland.
The specimen, discovered by paleontologists from the New Zealand Geological Survey, was
recovered for the Geology Museum, University of Otago. It covers perhaps 1.5 m2 of bedding

FOSSIL VERTEBRATES OF NEW ZEALAND - 1231

plane, and includes ribs and fragments of vertebrae. It is being prepared at present (1989). The
setting of the vertebrate was mentioned briefly by Aitcheson et al. (1988).

Johnston et al. (1980; 1987: 286) reported the incomplete shaft of a long-bone of apparent
Jurassic age, from Marybank Formation at Nelson. The bone is very weathered and lacks
condyles. Its identity is uncertain.

HJ f Campbell et al. (1984: 283) mentioned a small bone of a tetrapod from the Permian
(or Triassic) of Stephens Island, Cook Strait. Dr. E.H. Colbert (pers. comm., and cited in
H.J.Campbell et al. 1984: 283) reported that it may be a leg bone with "very much the
appearance of a Triassic reptile", but no diagnostic features were listed. The specimen was lost
about the time it was sent to the American Museum of Natural History, New York for
identification (E.H. Colbert, pers. comm.), thus continuing the history of New Zealand
specimens lost to or in overseas institutions (see also Haast 1870: 189, last para.; Welles &
Gregg 1971). Since it was incomplete, and was not identified formally, it is debatable that it
should be cited as evidence of a possible Triassic age (cf. H.J.Campbell et al. 1984: 288).

An anonymous author, almost certainly Hector (in McKay 1877g: 41) referred to the tooth
of a "megalosauroid . . . land saurian", apparently from the upper gorge of the Waipara River
(probably from the Laidmore Formation, Late Cretaceous). The specimen, which has not been
described, could have been the tooth of a marine reptile. Haast (1870: 189) also referred to the
"distal or lower part of the femur", possibly from a terrestrial reptile, from the Waipara region.

It is an understatement to say that further discoveries of terrestrial reptiles will be awaited
with interest.

AVES: OVERVIEW

Extinct birds are the best known of New Zealand's subfossil and fossil vertebrates, as the
literature and museum collections attest. Almost all of the many described species are from the
Quaternary. Indeed, few appear to be older than about 10,000 years. In contrast to the recently
extinct birds, there are a few species, mostly known from single specimens, older than Late
Pleistocene (Fig. 14). The fossil and/or recently extinct avifauna can be considered under these
subject headings, used below: a miscellany of Tertiary birds, bony-toothed pelicans -
Pelagornithidae, penguins - Order Sphenisciformes, moas, and Late Quaternary birds other than
moas.

General references useful for this section include those by Feduccia (1980), and Olson
(1985), and, on Australasian birds, Millener (this volume), Rich (1982), Rich &Van Tets
(1982, 1984), Williams (1973) and Williams & Millener (1981).

A MISCELLANY OF TERTIARY BIRDS

New Zealand's pre-Quaternary birds other than penguins, bony-toothed pelicans and moas
include few specimens, mostly incomplete and formally undescribed. Since most are, therefore,
of uncertain affinities, they are considered here according to geological age, rather than

taxonomic placement (Fig. 14).

1232 - FORDYCE

ms INTERNATIONAL N Z Stage
0 SUBDIVISIONS _/Haweran_| abundant bird bones ~10 000 yr bp - present
moa, Wanganui
odontopterygian?, North Canterbury; moa, Wanganui
Mangpanian , ; 4
penguin, Tereingaorinis, Hawkes Bay
D odontopterygian, Hawera
0 penguin, Kapitea Creek; moa, Taranaki
10 Tongaporutuan "kiwi" footprint, Murchison

20

30

40

50

60

70

1 O C E N

M

z
Ww
oO
@)
Oo
i
1@)
uw
z
et)
Oo
(e)
WW

PALEOCENE

Lillburnian

Clifdenian

Duntroonian

Runangan

Kaiatan

Bortonian

Porangan
Heretaungan
Mangaorapan

Waipawan

Teurian

Haumurian

odontopterygian, North Canterbury

Greta Siltstone fauna (age?), North Canterbury,

includes Aptenodytes, Pygoscelis, Marplesornis,
and Pseudodontornis

waterfowl, St Bathans

Otekaike Limestone fauna: penguins- Korora,
Platydyptes ; also undetermined non-sphenisciform
seabirds

Kokoamu Greensand fauna: penguins -

"Palaeeudyptes", Archaeospheniscus, Duntroonornis
and incertae sedis - Manu

indet. penguin, Glen Massey

large penguins, including Pachydyptes,
Oamaru district

early penguins (e.g."Palaeeudyptes" ), Burnside

undescribed "proto-penguin", Waipara
undescribed wing-propelled diver, Moeraki

undescribed volant bird, Chatham Island

undescribed bird, Waimakariri
undescribed diver, Cheviot

possible bird (probably dinosaur), Mangahouanga

no certain bird records in New Zealand
older than late Haumurian

Figure 14. Guide to stratigraphic occurrence of older bird fossils from New Zealand.

Olson (1985: 81) commented, presumably about avian paleontology in other lands, that
"The idea that every scrap of fossil bird bone is a priceless gift to be treasured with veneration
and treated as if diagnostic has infected avian palaeontology down to the present." Some
evidence of this is apparent in the plethora of names applied to moas and other Quaternary birds
from New Zealand. With this in mind, it is possibly fortunate that Tertiary and older birds are

FOSSIL VERTEBRATES OF NEW ZEALAND - 1233

rare enough in New Zealand to have discouraged much study. Nonetheless, some of the
undescribed specimens listed below should be described formally in future. They may form
important geographic and/or stratigraphic records even if they are too incomplete to warrant the
formal naming of new species.

Cretaceous-Palaeocene Records

The geologically oldest bird bones are undescribed. Keyes (1981a) mentioned that an avian
tarsometatarsus (New Zealand Geological Survey collections) was found associated with sharks
and invertebrates in lower Teurian (basal Palaeocene) or possibly Haumurian (latest Cretaceous)
marine sandstones of the "Lingula bed" at Otarama, Waimakariri River, Canterbury. Fleming
(1979: 40) had noted earlier that it might be of Cretaceous age. The bone has been under study
by J. Cracraft for some time. Earlier, an anonymous author, probably Hector (in McKay
1877g: 41), had referred to a possible bird bone from this or a nearby locality.

A broken distal end of a left femur came from Haumurian (latest Cretaceous) shallow
marine sands near Cheviot, North Canterbury (New Zealand Geological Survey collections,
Lower Hutt). The specimen has a marked angle in the shaft as seen in external view,
reminiscent of that seen in the North American Late Cretaceous diver Baptornis. It also has a
solidly-built shaft with only a small pneumatic cavity, and may well represent a flightless
form. It is currently under study at the Department of Geology, University of Otago.

Molnar & Scarlett (1984) described the isolated large phalanx of a what may be a bird
collected by Joan Wiffen from the Piripauan or Haumurian (Late Cretaceous) of Mangahouanga
Stream in Hawke's Bay. The element could not be assigned to any known taxon of bird, and
may represent a dinosaur. (See above, under reptiles.)

Two fragmentary specimens from the Moeraki Formation at Moeraki, North Otago are
probably Teurian, Palaeocene. Both are in the Geology Museum collections, University of
Otago. One is a crushed distal portion of a right tibiotarsus (Fig. 16) from a large individual
(condyle width exceeds 30 mm). The specimen has not yet been identified formally, and it it is
not known whether it represents a marine or non-marine species. or whether volant or
flightless. McKenzie & Hussainy (1968) referred to this as "a possible Cretaceous bird fossil,
now accepted as Palaeocene in age, from Moeraki". The second is a newly prepared specimen
(coracoid, fragment of head of humerus, scapula) also from Moeraki and also possibly from a
wing-propelled diver (Fordyce & Jones 1987, 1990) Previously, Mantell (1850: 326, Fig.7)
reported the discovery of an apparent bird bone at Moeraki.

Another Palacocene bird is represented by an undescribed and unprepared fragment of the
proximal end of a radius, from the Tahatika Grit (Teurian), Chatham Island (New Zealand
Geological Survey collections, Lower Hutt). The bone is pneumatic, and probably came from
a larger volant species. Its relationships are still uncertain.

There are no noteworthy Eocene records of birds other than penguins, which are discussed
below.

Oligocene

Apart from penguins, few younger Tertiary birds are known from New Zealand. Marples
(1946) described a new genus and species, Manu antiquus, based on a furcula from the
"Maerewhenua Greensand" [= Kokoamu Greensand Formation of Gage 1957; Duntroonian, Late
Oligocene] near Duntroon, North Otago. Marples regarded the holotype as similar to
albatrosses (Procellariiformes: Diomedeidae), although he did not clearly refer it to this family.
He also described and figured two incomplete isolated femora, which he speculated could be
related to M. antiquus. One femur was from a limestone of apparent Duntroonian age, while
the other was of uncertain provenance. No one seems to have formally revised Marples'

1234 - FORDYCE

species, nor reported other non-penguin bird remains from Duntroonian greensands. Some
authors (e.g. Kinsky 1970, and, by implication, Williams 1973: 304) have listed M. antiquus
as an undoubted albatross. Olson (1985: 208) stated that the species differs considerably from
albatrosses.

I have carried out extensive field work on the Kokoamu Greensand and Otekaike Limestone
and their lateral equivalents (e.g. Fordyce 1987a), with the recent assistance of Andrew Grebneff
and Craig Jones, but have found few birds other than abundant penguins. A battered small but
rather robust tibiotarsus, apparently not that of a penguin, was found associated with a cetacean
skull from the Kokoamu Greensand (Duntroonian; Late Oligocene) at the Waihao River,
Waimate district (fossil in Geology Museum, University of Otago). A small, well preserved
pneumatic and non-sphenisciform femur was found in the Otekaike Limestone (Duntroonian-
Waitakian; Late Oligocene-earliest Miocene), Hakataramea Valley. This specimen, in the
Geology Museum at University of Otago, has not yet been identified.

Miocene

Potentially one of the most important fossil bird sites ever found in New Zealand was
discovered in 1980 in Central Otago by Jon Lindqvist and Barry Douglas (Douglas et al. 1981;
see also Cotton 1919, Douglas 1986, McKay 1894). I made extensive further collections from
the source horizon, lacustrine sandy mudstones of the Manuherikia Group (Altonian, Early
Miocene) near Saint Bathans. Isolated avian elements include tarsometatarsals, humeri, ulnae
and metacarpals, probably anatid (Fig. 16), none of which has been identified positively to
order yet. Broken eggshell is also present. Material is held in the Geology Museum,
University of Otago. Further fieldwork and the formal description of the vertebrates is planned.

Fragmentary bird bones were recovered recently from fresh-water limestone in the Waitaki
Valley some 50 km northeast of Saint Bathans. The bones are associated with coal measures
of uncertain Southland-Taranaki (Middle to Late Miocene) age. The fossils are held in the
Geology Museum, University of Otago.

Mildenhall (1974: 47) figured a fossil "kiwi" footprint from Miocene mudstones near
Murchison. The specimen (New Zealand Geological Survey collections) was from the
predominantly non-marine Longford Formation (Fleming 1979: 66) and is of Tongaporutuan-
Kapitean (Late Miocene) age (1.W. Keyes, pers.comm.). The real identity of the specimen is
uncertain. Hutton (1899b) also described a footprint of a "kiwi-like bird” from a sandstone slab
at Manaroa, Pelorus Sound. As the rocks around Manaroa are metamorphic, it is likely that
the specimen either is Holocene or that it came originally from elsewhere,

BONY-TOOTHED PELICANS (PELAGORNITHIDAE)

"Bony-toothed" pelicans (Pelicaniformes: Family Pelagornithidae) have attracted some
attention in the world literature in the last few years. Because of their superficially unusual
appearance, these birds are sometimes placed in a suborder Odontopterygia or in their own
order, Odontopterygiformes. Harrison & Walker (1976) reviewed the group, as did Olson
(1985: 194-201; see also Steadman 1981), who provided comments on the former work.
Cracraft (1985) gave a cladistic overview of the Pelicaniformes without commenting on the
Pelagornithidae. Three pelagomithid specimens have been described from New Zealand,

A new species, Pseudodontornis stirtoni Howard & Warter, 1969, was established for a
partial skull found in a loose concretion of "Greta Siltstons" (sensu Lewis 1976) at Motunau
Beach, North Canterbury (see also Gregg 1974), A badly crushed femur associated with the
skull presumably belongs to the same individual. Howard & Warter placed the species in the
genus Pseudodontornis, previously reported from the Miocene of eastern North America, and in
the family Pseudodontornithidae, Harrison & Walker (1976) transferred P. stirtoni to a new

FOSSIL VERTEBRATES OF NEW ZEALAND - 1235

genus, Neodontornis, retained in the Pseudodontornithidae. Unlike Howard & Warter (1969),
but following Howard (1957), they recognized the bony-toothed pelicans as a distinct order,
Odontopterygiformes. In contrast, Olson (1985) suggested that all bony-toothed pelicans
belong in a single family of Pelicaniformes, the Pelagornithidae.

Figure 15. Tertiary birds from New Zealand. A, reconstruction of Pseudodontornis stirtoni; redrawn from
photo in Rich and Berra (1980, Pl. 1); B, ?anatid metacarpal from the Early Miocene, St Bathans, specimen in
the Geology Museum collections, University of Otago, scale bar, 10 mm; C, ?anatid tarsometatarsus from the
Early Miocene, St Bathans, specimen in the Geology Museum collections, University of Otago, scale as for
Fig. 16 B; D, ?anatid coracoid from the Early Miocene, St Bathans, specimen in the Geology Museum
collections, University of Otago, scale as for Fig. 16 B; E, crushed tibiotarsus, with tendinal bridge lost, of
indet. large bird, Palaeocene, Moeraki, specimen in the Geology Museum collections, University of Otago,
scale bar = 50 mm. (Figs B-E by Jane Kerr).

Howard & Warter indicated that the holotype of P. stirtoni could range in age from Early
Miocene (stage unspecified) to probably no later than Waitotaran (Late Pliocene). Scarlett
(1972a) cited a probable Waitotaran age, and Fleming (1979: 69) indicated no doubt about a
Late Pliocene age. It is possible that the concretion which encased the holotype was remanié,
and came from the Nukumaruan (latest Pliocene-Early Pleistocene) debris flows at Motunau
described by Lewis (1976). Lewis noted that the debris flows contain fossiliferous clasts of
"Greta Siltstone" (sometimes containing vertebrates, pers. obs.) of Waipipian (early Late
Pliocene), Waiauan to Tongaporutuan (Late Miocene) and Otaian to Altonian (Early Miocene)
age (see also Suggate et al. 1978: figs 7.77 and 8.15.) This cautions against citing a definite
age for P. stirtoni. The reconstruction of P. stirtoni given here (Fig. 15) is based on a figure
published by Rich & Berra (1980).

Scarlett (1972a) described the proximal (not distal) part of a right humerus as probably that
of a pelagornithid. The bone was from sediments of probably Waiauan (early Late Miocene)

1236 - FORDYCE

age, possibly the Double Corner Shellbed unit of the Tokama Siltstone, near the mouth of
Waipara River, North Canterbury. Harrison & Walker (1976) referred Scarlett's specimen
tentatively to Pelagornis miocaenus Lartet, 1857 (placed by them as Odontopterygiformes:
Pelagornithidae), which was described from the Middle Miocene of France. More material
would be needed to confirm this identification, and thereby its biogeographic implications.

McKee (1985) described a humerus and radius assigned to the family Pelagornithidae, to
which Pseudodontornis belongs. The specimens are from Tangahoe Formation (Waipipian,
"middle" Pliocene) near Hawera. The bones cannot be identified positively to species at
present. A possible younger record is provided by an isolated femur (Canterbury Museum,
Christchurch collections) which I collected from the Nukumaruan (latest Pliocene-Early
Pleistocene) at Motunau, North Canterbury. The femur compares closely with the femur
associated with the holotype of P. stirtoni; it may represent an indeterminate pelican or one of
the youngest global records of pelagornithids.

MOAS - DINORNITHIFORMES

Introduction

Moas are the extinct large flightless birds (Fig. 16) which are probably the best-known of
New Zealand's extinct vertebrates. They were first introduced to science by Richard Owen
(1840, 1843, 1844b, 1879, and other publications), who successfully employed Cuvier's
“principle of correlation" to predict that a scrap of femur represented a hitherto unknown large
"struthious" bird. The story of the late Quaternary moas and other birds is a complex one,
equal in magniiude or larger than that of all the other New Zealand fossil vertebrates, and is
better detailed elsewhere. I cover only a few aspects here for the sake of completeness in an
account of New Zealand fossil vertebrates. In particular, I have not attempted to incorporate
information from the archaeological literature. The reader is referred to the literature
summarised below for details. Recent popular accounts of moas include those by Brewster
(1987), Falla (1974) and McCulloch (1982). A starting point amongst more technical articles
was provided by Anderson (1984), Cassels (1984), Trotter & McCulloch (1984) and Worthy
(1987d, 1988a, 1988c, 1989a, 1989b, 1989c). For a guide to the earlier literature on moas, see
references in Anderson (1984), Cassels (1984), Trotter & McCulloch (1984) and especially in
Archey (1941) and Oliver (1949). The work of Owen provides a monumental historic series;
publications on moas include those of Owen 1840, 1843a, 1843b, 1844a, 1844b, 1846, 1848a,
1848b, 1856, 1865, 1866, 1870a, 1871, 1873, 1879, 1883a, and 1883b. The unindexed
bibliography of Romer et al. (1962) contains more references for varied authors, few of which
are included here.

Taxonomy

Moas (Order Dinornithiformes) and the extant kiwis are allied conventionally with the
ratites, traditionally accepted as a monophyletic group of Austral flightless palaeognathous
birds (e.g. Cracraft 1974b, Rich & Balouet 1984, Sibley & Ahlquist 1981). Two subdivisions
of Dinornithiformes are commonly recognized. Some authors use two subfamilies within one
family, Dinornithidae: Dinornithinae (greater moas), and the Anomalopteryginae (lesser moas)
(Cracraft 1976a, Worthy 1988e). Conversely, these taxa have sometimes been given separate
family status as Dinornithidae and Emeidae [= Anomalopterygidae] (Brodkorb 1963, Worthy
1989e).

FOSSIL VERTEBRATES OF NEW ZEALAND - 1237

reconstruction of Aptornis otidiformis, after
C, skull of Euryanas

Late Quatemary birds from New Zealand. A,
f Dinornis giganteus, after Wilson in Swinton (1975);

Figure 16. Some
Oliver; B, reconstruction o
finschi, after Van Beneden (1876).

The species of moa listed in Table 4 are those that were regarded as distinct by Cracraft
(1976a), with later emendations. Millener (1982) suggested that Anomalopteryx owent be
synonymised with A. didiformis (sce also Worthy 1987d, 1988a). Worthy (1989e) reinstated
Pachyornis australis. This classification reduces markedly the number of species recognised by

1238 - FORDYCE

earlier authors, such as Archey (1941) and Oliver (1949). It has been criticized in passing by
some other workers (e.g. Trotter & McCulloch 1984) but a detailed critique dealing with all
species has not appeared.

The most readily available summaries of the primary literature on moa taxonomy include
articles by Archey (1941), Brodkorb (1963), and Oliver (1949), Other articles which touch on
one or more species include those by, for example, Caughley (1977), Cracraft (1976a, 1976b,
1976c), Kinsky (1970), Oliver (1955), Scarlett (1972b, 1975), Worthy (1987d, 1988a, 1988c,
1989a, 1989b, 1989c, 1989e) and Yaldwyn (1959, 1979).

Recent studies have concentrated on diagnosing the species of moa more accurately than
before (e.g. Cracraft 1976b, 1976c; Worthy 1987d), but there is scope for more work. The
fragmentary nature of early type specimens (such as Dinornis novaezealandiae Owen, 1843)
makes it difficult to tell whether the types really are conspecific with the more complete
specimens on which many working definitions of species are based. This is a common
problem for nineteenth century types that are inadequate by modern standards (see comments
herein on the fossil penguin Palaeeudyptes antarcticus), and can be resolved several ways.
Some workers tacitly accept that the types are conspecific with better material, and use the
names attached to the types. In such cases, the matter of whether the type really is conspecific
with referred material is barely addressed. Alternatively, sophisticated biometrical analyses,
which can perhaps quantify subtle differences in degree, may help decide whether a type and
referred specimen are likely to be conspecific. In my opinion, however, it seems better to
determine differences in kind (presence-absence differences), using derived characters to justify
clustering. If such characters are absent on the types attached to long-used names, should the
names be regarded as nomina dubia? (names of dubious or doubtful application). Many workers
might disagree with such an approach, and here immunological techniques (Lowenstein 1986)
or comparisons of DNA (Wilson et al. 1987) might well be applied to moas to resolve
questions of the identity of important specimens.

Morphology

Moa anatomy is discussed in many works, such as the taxonomic articles listed above, and
illustrations of moa bones abound. Owen (1879, vol. 2; a synopsis of earlier works) provided
many figures. The papers of Archey (1941) and Oliver (1949) give a range of figures. Scarlett
(1972b; see also 1975) provided a general atlas of bones which includes many figures of moa,
as well as other bird bones that might be encountered in Holocene sediments. Worthy (1988c)
produced a valuable illustrated guide to leg bones. Worthy (1987d) and Yaldwyn (1979) also
gave figures and comments on describing morphology.

More recent articles on functional morphology include those of Alexander (1983a, 1983b,
1985), Reif & Silyn-Roberts (1987) and Worthy (1987d). All these deal with the legs. There
seems to be scope for further studies, particularly those that approach the broader topic of
constructional morphology.

Early Fossil Records of Moas

It is likely that moas have inhabited New Zealand since the Cretaceous (see a host of
references including, e.g. Fleming 1979, Hutton 1873, Stevens 1980a), or Eocene at the latest
(Sibley & Ahlquist 1981), but no fossils clearly older than Pliocene are known. Moa bones
are rare in Pliocene and younger marine rocks, so it seems unlikely that older Tertiary marine
rocks will be a significant source of bones. The most likely environments for fossilisation are
those represented by rare, completely non-marine sequences (e.g. Manuherikia Group, Saint
Bathans). The transgressive and regressive coal-measures that commonly bound local
Cainozoic marine sequences might yield bones, although they may have been deposited in

FOSSIL VERTEBRATES OF NEW ZEALAND - 1239

conditions too acidic for bones to preserve normally. Ironically, peat swamps have provided
some of the best selections of stratigraphically young moas (see e.g. accounts by Haast1869a,
1874b, 1874c; McCulloch 1985; Scarlett 1969b). Other comments on possible Tertiary
localities were given by T.H. Rich (1975).

Tertiary fossils of moas are known, despite Olson's (1985) suggestion that there are no
confirmed records. The oldest fossil record published appears to be that mentioned by Oliver
(1949: 65, 1955: 574), who stated that bones of Pachyornis mappini Archey, 1941, had been
found in Pliocene "papa rock" (mudstone) at Maungapurua, Taranaki. Fleming (1962a: 81)
gave the age as Kapitean. Oliver also mentioned that bones of P. mappini had been found in
the Pliocene at Nuhaka, Te Aute, Hawke's Bay. Fleming (1979: 66) commented that if the
bones really do represent the Recent species P. mappini, they suggest either that speciation
amongst moas was well advanced before the Pliocene, or that the bones are of Quaternary age.
However, as little is known of rates of evolution amongst moas, the similarity of the fossil
bones with those of a Recent species does not necessarily support a younger age.

Forbes (1891a) reported bones from clay which underlies basalt at Gleniti Valley, Timaru,
South Canterbury. The bones included the holotype of Anomalopteryx antiquus Hutton, 1892
[= A. didiformis (Owen, 1844a); fide Cracraft 1976a]. Matthews & Curtis (1966) cited a
radiometric date of 2.47+.0.37 myBP for the Timaru basalt, which suggests that the moa bones
are no younger than Late Pliocene. Drs P.R. Millener & J.A. Grant-Mackie cautioned (pers.
comm.), however, that it is not clear that these bones were truly in situ. No other bones have
been reported from this locality, and it is possible that the clay under the basalt might be a
fissure-fill (although Forbes’ sketch of the outcrop argues against the latter). To counter these
suggestions, pre-Pleistocene records of moas should be expected, and we should not have
preconceived ideas about the likely specific identity of such specimens. Amino acid
racemization (Weston et al. 1973) may resolve such problems about age, and immunological
tests might elucidate relationships.

Park (1887a: 63) mentioned moa bones from the Butlers Shell Conglomerate (Okehauan) of
Wanganui area (see Fleming 1953: 175, and pp. 156, 209 for other occurrences). Marshall
(1919) described a partial femur, apparently of Dinornis robustus Owen, 1846 [= D.
novaezealandiae Owen, 1843; fide Cracraft 1976a], from Nukumaru Beach, near Wanganui.
The specimen was from the Tewkesbury Formation (Fleming 1953: 156), and thus is of
Nukumaruan (latest Pliocene-Early Pleistocene) age.

Fleming (1953: 140) stated that the moa egg identified by Oliver as that of Anomalopteryx
didiformis from Tokomaru (= Kaiiwi?; Oliver 1949: 43, 45) is probably reworked. Fleming
discounted a Nukumaruan (latest Pliocene-Early Pleistocene) age. Eggs and eggshell are
common in younger sediments (e.g. Archey 1941, Field 1885, Hector 1867, Simpson 1955).

Hill (1889) noted the occurrence of supposed moa feathers from near Ormond, Gisborne
district, Hawke's Bay. They may have been from the "Waipaoa Series" (Castlecliffian, Late
Pleistocene; Suggate in Suggate et al. 1978: 566).

Moa bones are known from the sea floor around New Zealand (Fleming 1963c, Keyes &
Froggatt 1978). These are of unknown age, and could represent specimens washed in from
rivers or animals that lived on a then enlarged coastal plain during a phase of low (glacial) sea
level.

Footprints

Moa footprints have been mentioned or figured, for example, by Benham (1913), Collen &
Vella (1984), Hill (1895), McKay (1877d: 116), and Oliver (1949: 24-25). There seems to
have been no serious analysis of tracks in the manner seen for some extinct tetrapods elsewhere
(e.g. dinosaurs), where aspects of gait have been inferred from footprints.

1240 - FORDYCE

Late Pleistocene and Holocene Moas - Geography and
Stratigraphy

Much early literature (see Archey 1941 and Oliver 1949 for summaries) contains only
incidental or anecdotal comment on geographic distribution and stratigraphy. This probably
reflects the early inconsistent application of place names, lack of detailed maps, and absence of
an accurate Quaternary stratigraphy. Recent articles, some of which present important
radioisotopic dates, include those by Bell & Bell (1971), Brewster (1987), Burrows (1980b),
Cody (1979), Falla (1974; general review), Grant-Mackie & Scarlett (1973), Fleming (1963c),
Grant-Mackie (1965, 1979), Gregg (1972), Keyes & Froggatt (1978), McCulloch (1982;
general review), McCulloch & Trotter (1979), Medway (1971), Millener (1986), Millener &
Templer (1982), Nelson & Grant-Mackie (1980), Scarlett (1969b, 1972b; see other papers
listed by Anderson 1979 and Millener 1980c) and Worthy (1983, 1984, 1987d, 1988a, 1989a,
1989c, 1989e, Worthy & Mildenhall 1989).

Ecology and Behaviour

Attention has focused recently on ecology and behaviour. Burrows (1980a, 1980b; Burrows
et al. 1981, Burrows & Drake 1982, all on diet) concluded that moas were browsers that lived
along forest margins where they ate vegetation, including twigs of woody plants. Hamel
(1979) commented on inferred breeding behaviour. Hayward (1978; see also Smalley 1979)
discussed gizzard stones, and commented that moas may have been New Zealand's oldest
rockhounds. In fact, Cretaceous marine reptiles also carried gizzard stones. Older references on
gizzard stones include those of Hamilton (1892) and Hill (1890). Greenwood & Atkinson
(1977; see also Atkinson & Greenwood 1982) suggested that moa browsing behaviour might
account for the diversity in New Zealand of divaricating plants, plants in which the stems are
typically slender, leaves small and twigs at a high angle to the stem. This suggestion was
countered by McGlone & Webb (1981), and the matter of interaction between divaricating
plants and moas is still not clear.

Evolution and Extinction

Specifics of moa evolution have been discussed rarely, but the broader relationships have
been reviewed widely and inconclusively. For example, Cracraft (1974b) identified
synapomorphies that cluster moas and kiwis together with other ratites. Rich & Balouet
(1984) also united moas and kiwis, but cautioned that ratite interrelationships are not
understood well. They suggested that a longer fossil record might help resolve the
relationships of moas and kiwis. Patterson (1981b) cautioned, however, that the fossil record
may never help resolve relationships for many groups. Fleming (1980a) drew attention to the
now discredited record of a supposed ratite from New Caledonia; this megapode Sylviornis (see
Poplin et al. 1983), intially was a candidate for the possible sister taxon to moas. Sibley &
Ahilquist (1981) assumed that ratites, and moas and kiwis, are monophyletic. Olson (1985: 96-
97) cautioned that the palaeognathous palate, used for so long to diagnose ratites, may be a
shared primitive feature (see also Olson in James & Olson 1983: 40). Houde (1986) suggested
that ostriches and kiwis are not necessarily related to other ratites including moas; kiwis may
have evolved from volant palaeognaths that flew to New Zealand. These explicit or implied
problems of higher taxonomy are similar to those encountered in other relatively derived groups
(e.g. penguins, cetaceans). Ecological radiations often take place with the occupation of new
habitats and with the evolution of structures radically different from older taxa, so that all

FOSSIL VERTEBRATES OF NEW ZEALAND - 1241

ae are highly derived in comparison to likely sister groups, and homologies are difficult to
ine.

With the admitted benefit of hindsight, it seems odd that early workers did not question the
likelihood of some 26 species of moa evolving in New Zealand. Enviromental heterogeneity
has long been regarded as important in evolutionary processes, yet New Zealand would seem to
lack the ecological opportunities of larger landmasses. Indeed, Olson (1985: 102) commented
that it is difficult to explain currently accepted species diversities of moas, since speciation has
not produced comparable diversity amongst other terrestrial avian groups. This lack of
diversity amongst other groups may be apparent rather than real; note, for example, the new
and perhaps unexpected species of Leiopelma described recently by Worthy (1987a). The role
of altitudinal segregation of moa niches has been addressed little, but Worthy (1989c) gave
some comments on montane distributions.

The question of moa extinction has long been discussed. The two main hypotheses are that
extinctions were caused by changing climate or vegetation patterns, or by humans. The former
explanation was widely accepted until the advent of accurate radiocarbon dating, which showed
that most species overlapped temporarily with humans. Discussion includes that in articles by
Anderson (1982, 1984), Cassels (1982, 1984), Cracraft (1980b), Duff (1964), Falla (1974),
Fleming (1962c, 1969, 1973, 1979), Holdaway (1989), McCulloch (1982), Scarlett (1969b,
1974), Simmons (1968), and Trotter & McCulloch (1984), Some mummified specimens (e.g.
Brewster 1987, Forrest 1987; but cf. Worthy 1989a) and young radiocarbon dates reinforce the
idea of recent extinction. Millener (1981b) suggested that many North Island reports of moas
and other birds from middens may be of bones eroded from underlying ossiferous dunesands. If
this is so, such bones presumably were mixed subsequently with midden waste by downslope
sate movement before collection, and may not have been derived originally from the
middens.

OTHER LATE QUATERNARY BIRDS

Many Late Pleistocene-Recent birds (Fig. 16) are known apart from moas. Some of these
are listed in Table 5, which is based on published literature (other than archaeological works),
initially that of Brodkorb (1963, 1964, 1967, 1971, 1978). The table lists extant New Zealand
species which have a non-archaeological subfossil or fossil occurrence, and incorporates
subfossil and fossil birds from the Chatham Islands listed by R.J.Scarlett (in Fordyce 1982c).
The literature on these species is much more extensive than indicated by the citations in the
table.

Subfossil, according to Kinsky (1970), encompasses species that became extinct before
about 1800 A.D. Thus, some species are included that are extant now only outside New
Zealand.

Faunal Composition

At present, sea-birds and waterfowl predominate over forest birds, but it is likely that more
species of the latter will be found as work progresses. Cave faunas, in particular, give great
promise. Fleming (1962a: 93) commented that acid soils may well have destroyed evidence of
older terrestrial fossils, and this could explain the lack of fossil bush birds from other than cave
deposits. Recent work on cave faunas includes that of Millener (1983a, Millener & Templer
1982), and Worthy (1984, 1987a, especially figs 1 and 2, 1987b; Worthy & Mildenhall 1989).

1242 - FORDYCE

History of Work

As was the case for the moas, I do not intend to review the early history of work on ..ew
Zealand's non-ratite Quaternary birds. Some idea of this can be obtained from Brodkorb's
various catalogues and the New Zealand entries in Romer et al. (1962). Among the most
noteworthy recent work is that of R.J. Scarlett, former osteologist at the Canterbury Museum,
Christchurch, whose contributions span at least 30 years. Bibliographies of Scarlett's works
were published by Anderson (1979) and Millener (1980c).

Other non-archaecological work on faunas (cf. taxonomic articles - see Table 5) includes both
general and detailed comments. Examples include the articles of Bourne (1967), Dawson
(1949, 1958a, 1958b, 1959, 1961), Falla (1941), Grant-Mackie & Scarlett (1973), Horn
(1983), Medway (1967, 1971, 1974), P. Millener (National Museum of New Zealand),who
started studies on the North Island Quaternary avifauna in the late 1970's (Millener 1979,
1980b, 1981b - unpublished Ph.D.thesis, 1983a, 1983b, 1984, 1986, 1988b; Millener &
Templer 1982; see also Cody 1979), Olson (1975, 1977a, 1977b, 1984, 1985), Paulin (1973),
Reid & Williams 1975, T.H. Rich ef al. (1979), Worthy (1982, 1984, 1988b, 1989d, 1989e),
Worthy & Mildenhall (1989) and Yaldwyn (1956, 1958). Amongst older works, that of Forbes
(1891a, 1891b, 1882a-1892i, 1893a-1893c; see Dawson 1958b) stands out as a series that was
published quickly over a short time and thus has caused problems in nomenclature. R.
Holdaway (University of Canterbury) is currently working on the avifauna associated with
Harpagornis (Holdaway 1989).

Taxonomy

A limited selection of references to systematics of Late Pleistocene-Recent New Zealand
species is given in Table 5. Brodkorb (1963, 1964, 1967, 1971, 1978) listed many references
to early systematic articles, and also gave an idea of stratigraphic and geographic distribution.

Stratigraphy and Geographic Distribution

There is little good information about stratigraphy other than in a few recent articles. This
reflects several problems. Late Quaternary biostratigraphy has had a slow development,
although palynology now makes important contributions. Other techniques, such as amino
acid racemization, have promise. Many collectors have not been trained in the need for
stratigraphic rigour, and old collections in particular lack stratigraphic data. Cave deposits
contain material of mixed origin and age. Radiocarbon dating techniques are expensive and
destructive, although tandem accelerator mass spectrometry promises to be less destructive.
The lack of good stratigraphic information largely precludes temporal analyses, such as studies
of changing geographic patterns amongst the local birds. Nonetheless, there have been some
laudable attempts to make sense of the record (Beauchamp & Worthy 1988, Mills et al. 1984,
1988).

Most of the references in Table 5 also mention stratigraphy and geography. For further
information, see particularly Grant-Mackie & Scarlett (1973), Fieming (1979), Gregg (1972),
Millener (1981a; bibliography of cave occurrences), Rich et al. (1979), Scarlett (e.g. 1955,
1969, 1972; see also Anderson 1979 and Millener 1980c), and Worthy & Mildenhall (1989).

Evolution, Zoogeography and Extinction
The comments on moa evolution, above, apply very much to the other birds. Most

previous studies are descriptive and deal with morphology or faunas. There have been few
critical accounts of evolution at the species level or of issues in macroevolution based on

FOSSIL VERTEBRATES OF NEW ZEALAND - 1243

subfossil or fossil material. Worthy (1988b) discussed the Holocene origin of flightlessness in
the extinct Euryanas finschi, and other flightless species surely warrant attention. In general,
the most significant contributions to understanding of evolution of neospecies with a subfossil
record are based on studies of extant birds; examples are Turnagra capensis, discussd by Olson
et al. (1983), anatids including New Zealand taxa by Livezey (1986), Acanthisittidae by Sibley
et al. (1982), and kiwis by e.g. de Boer (1980) and Calder (1979).

Regional (as opposed to local) zoogeography has been discussed widely, recently in terms of
vicariance biogeography and earlier in terms of dispersalist biogeography. Probably the best-
known broader contributions on the evolution of New Zealand's avifauna are those of Fleming
(1962a - see 1949 for early version, 1962b, 1973, 1974, 1976, 1977, 1979), Cracraft's general
reviews (1973a, 1976, 1980a) have stressed the role of vicariance in the evolution of the
avifauna, but a detailed species-by-species consideration of the zoogeography and evolution of
New Zealand birds has yet to be compiled.

Richard Holdaway (Holdaway 1989) is presently looking at the recent history of the New
Zealand avifauna in terms of island-area biogeographic theory. This work involves a long-
overdue study of the chronology of recent extinctions

PENGUINS (SPHENISCIFORMES)

These medium-sized to large amphibious birds are skilled underwater fliers restricted today
and apparently in the past to the Southern Hemisphere. There they inhabit mostly cool-
temperate waters, and indeed they may have originated in the south. New Zealand has long
been an important area for the study of both extant and extinct penguins; the first fossil
penguin known to science was discovered near latitude 45°S in North Otago, and middle
Cainozoic penguins are diverse and abundant. For general references on penguins, see Simpson
(1976) and Stonehouse (1975). Fordyce & Jones (1990) gave a brief overview of New Zealand
and other fossil penguins.

Taxonomy and Morphology

The fossil species reported from New Zealand are listed in Table 6, and a brief history of their
description is given below. Noteworthy reviews of the taxonomy and morphology of local
fossil penguins are those by Marples (1952; 1974 - short account) and Simpson (1971a, 1972a,
1975).

Illustrations of some New Zealand fossil penguins are given in Figs 17, 18; Pl. 2. As with
other groups in this review, the morphology of the New Zealand fossil species is not detailed
here. As might be expected with a morphologically conservative group, taxonomically
important characters are more subtle differences in degree than in kind (e.g. Olson 1985: 216).

Approaches to Taxonomy

Some problems of taxonomy were discussed by Fordyce & Jones (1990). In summary, few
reasonably complete skeleton or skulls of fossil penguins are known (e.g. Fig. 17). Limb
bones, which are robust and non-pneumatised, preserve well, are easy to recognise and have
been used widely as type-specimens, but they occur commonly as isolated specimens. Indeed,
the first species of fossil penguin described, Palaeeudyptes antarcticus Huxley, 1859a (see also
Huxley 1859b-1859d), is known only from an isolated tarsometatarsus (Fig. 18) of uncertain
age, possibly Late Oligocene, from Kakanui. The skeletal structure of P. antarcticus is
unknown, however, as no other tarsometatarsus identical to that of the type specimen has been
found associated with a skeleton. The types of many nominal species described since 1859

1244 - FORDYCE

Cc»

ch

Figure 17. Relatively complete skeleton of "Palaeeudyptes"-like penguin from the Late Oligocene, South
Canterbury, specimen in the Geology Museum collections, University of Otago, scale bar, 1 m. (Drawn by
Craig Jones).

FOSSIL VERTEBRATES OF NEW ZEALAND - 1245

\, | Palaeeudyptes
antarcticus

Altonian

Duntroonian
Whaingaroan

Runangan

Bortonian

ih

|
)
}

L|
tyr
i
Korora oliveri

SNA00IN

LAO
Lr
new species? 3

new species? 2

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“Palaeeudyptes" marplesi | sf \(\ |
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Figure 18. Penguin tarsometatarsi as an index of species diversity; all the new specimens are in the
Geology Museum collections, University of Otago, scaie bar, 50 mm. (Tarsometatarsi drawn by Craig Jones.
Ages shown are approximate only)

have included the tarsometatarsus, which is not surprising since this is a robust leg element
that preserves well. Because the tarsometatarsus has been used widely in this way, it is easy to
make comparisons with P. antarcticus. However, the results of such comparisons are often
ambiguous. To cluster nominal species into higher taxa on the basis of "overall similarity” of
tarsometatarsi (let alone inferred age, geographic distribution or other non-taxonomic criteria)
will not necessarily result in the recognition of natural groups. Fordyce & Jones (1990) noted
that preliminary computer-aided cladistic analysis (using a cladistic program, PAUP) of this
bone provided, rather predictably, many alternative equally parsimonious cladograms. These
reflect: 1) the basically conservative form of this element; 2) a lack of good information about
ontogenetic, intra- and interspecific variation in extant species; 3) problems in identifying
character polarities; 4) the likelihood of widespread homoplasies. Accordingly, nominal taxa
which derive their identity from P. antarcticus, that is, other supposed species of Palaeeudyptes,
are not well founded. In general, should names that are poorly based by modern standards be
used? Fordyce (1988a), in reference to fossil whales, followed others in suggesting that they
should not. Many evolutionary taxonomists would disagree with such a strict reliance on
morphology, and R.J.F. Jenkins (1980), for example, suggested that P. antarcticus poses
problems "because it is unlocated stratigraphically", not because it is morphologically rather
uninformative.

Despite the widespread use of the tarsometatarsus in fossil penguin systematics, the form of
the element may be determined more by functional and structural (fabricational) constraints than
by phylogenetic ones. Although it may indicate body form, perhaps the tarsometatarsus alone
is not the most appropriate indicator of phylogeny. It should be possible to better define and
diagnose the species involved where tarsometatarsi are associated clearly with other bones.

1246 - FORDYCE

History of Research

Literature on the systematics of New Zealand fossil penguins, outlined in the following
chronological guide, spans 130 years. Huxley published the first account of a fossil penguin,
Palaeeudyptes antarcticus (Fig. 18), in 1859 (Huxley 1859a-1859d,1899). Hector (1870a)
announced the discovery of bones referred to P. antarcticus from Seal Rock, near Punakaiki,
North Westland, and later described the fossils in some detail (Hector1872). Hector (1873a)
also reported the discovery of bones, supposedly those of P. antarcticus, from near Oamaru.
Hector's specimens were assigned subsequently to other species (Simpson 1971a).

Other occurrences of fossil penguins, all identified as P. antarcticus or Palaeeudyptes, were
mentioned sporadically in the literature late last century. Hector (1876: 53), for example, stated
that part of a skeleton was recovered from Seal Rock, near Punakaiki. McKay (1877a: 585)
reported bones from a greensand conglomerate at Amuri (Haumuri) Bluff (possibly the
occurrence mentioned by Hector 1877a). Haast (1879: 311) observed that P. antarcticus had
been discovered in rocks of the "Oamaru Formation" (probably Oligocene calcareous sediments)
at Curiosity Shop (Rakaia River) and Broken River (Castle Hill Basin), both in Canterbury.
McKay also mentioned specimens from Castle Hill Basin (McKay 1881a:70) and Curiosity
Shop (McKay 1881b:82). Hector (1884: 539) reported “the giant fossil penguin" bones from a
limestone at Kaipuhe, North-west Nelson. Hutton (1885b: 272, 1885c: 549) documented the
apparent presence of Palaeeudyptes in the Weka Pass Stone and at Curiosity Shop. McKay
(1887a: 90) also reported bones of Palaeeudyptes from the Weka Pass Stone, North Canterbury.
Hector (in McLeod 1904: 524) commented that bones were recovered from limestone in a cave
at Greymouth. These obscure references appear not to have been discussed by later authors,
such as Marples (1952) and Simpson (1971a). The identity of most of the specimens is
unknown,

Minor incidental references were made to fossil penguins early this century (e.g.Henderson
1917; 94), but it was not until 1930 that the first important systematic article appeared, Oliver
(1930) described two new species, Pachydyptes ponderosus and Pachydyptes novaezealandiae,
the latter now placed in Platydyptes (see Simpson 1971a). Ovey (1939) commented on the
ages of some New Zealand specimens. Simpson (1946) briefly reviewed New Zealand
occurrences, but provided no new interpretation of systematics.

A new phase in the study of New Zealand fossil penguins (and, incidentally, whales) was
initiated by B.J. Marples, former Professor of Zoology at University of Otago. Marples (1946)
menwioned that he had recently discovered fossil penguin bones, and later (Marples 1949a)
indicated that work was in progress on New Zealand specimens, In his brief review of New
Zealand vertebrate palacontology, Marples(1949a) employed the generic names
Archaeospheniscus, Duntroonornis, and Platydyptes for the first time, but these names were
nomina nuda (improperly established names; see Ride ef al. 1985) until validated in 1952.
Marples (1952) reviewed Palaeeudyptes antarcticus, Pachydyptes ponderosus, and Pachydyptes
novaezealandiae (which he transferred to a new genus, Platydyptes), and also described the new
species Platydyptes amiesi, Archaeospheniscus lowei, Archaeospheniscus lopdelli,
Dunitroonornis parvus and Korora oliveri. A brief summary of this paper was given by Oliver
(1955).

Marples (1960) described a new species, Palaeospheniscus novaezealandiae, based on a
relatively complete skeleton from Motunau. This species is now placed in Marplesornis. Ina
later contribution, Marples & Fleming (1963) reported the discovery of the first fossil penguin
(an indeterminate, incomplete femur) from the North Island.

Simpson (1971a) presented a thorough review of New Zealand's pre-Pliocene penguins, in
which he described the new species ?Platydyptes marplesi and discussed penguin localities and
Stratigraphy. He also discounted the Heretaungan (Early Eocene) age reported for the earliest
New Zealand penguins (see, e.g. Fisher 1967; 734). Simpson (1972a) later considered the

FOSSIL VERTEBRATES OF NEW ZEALAND - 1247

North Canterbury specimens of supposed Pliocene age. He established a new genus,
Marplesornis, for Palaeospheniscus novaezealandiae, and described two new species referrable to
living genera, Pygoscelis tyreei and Aptenodytes ridgeni.

Grant-Mackie & Simpson (1973) redescribed the penguin originally described by Marples &
Fleming (1963), and also described new finds from Oligocene rocks south of Auckland. All of
the North Island specimens represent indeterminate genera and species. Simpson's (1975)
review of all fossil penguins provides a useful general summary of the New Zealand fossils,
even though it presents no new information on their systematics. This work is supplemented
by general comments in Simpson (1976, 1978: 231).

Scarlett (1983) described a new species, Tereingaornis moisleyi, based on wing elements
from the Waipipian (mid Pliocene) of Te Reinga Falls, Wairoa River, Hawkes Bay, North
Island. Scarlett originally intended to refer the material to Spheniscus, but ultimately
employed a new genus because of a referee's suggestion that Spheniscus should not be expected
in New Zealand. McKee (1986, 1987a) described other material which he referred to the
species.

Minor comment on the systematics of New Zealand species has appeared in articles on the
systematics of fossil penguins from other countries. These articles include those by R.J.F.
Jenkins (1974, 1980), Lowe (1933, 1939), Marples (1953), Olson (1985) and Simpson (1957,
1971b, 1972b, 1979).

New Specimens

Fossils collected since Simpson's major review of New Zealand taxa include specimens
potentially useful in the study of penguin phylogeny (Fordyce & Jones 1987, 1988 - both
abstracts, and 1990 review; Fig. 19). A few are mentioned below.

A penguin-like bird (New Zealand Geological Survey collections) discovered recently in the
Waipara Greensand, North Canterbury, is probably of Teurian or Waipawan age (Palaeocene or
Early Eocene). Elements include the interorbital region, mandibles, coracoids, scapula, furcula,
radius, ulna, synsacrum, vertebrae, and ribs (Pl. 2), which were mentioned briefly in a short
note (Fordyce et al. 1986). The fossil is probably a penguin, although it could represent some
other wing-propelled diver. Relationships are still uncertain, because of unresolved character
distributions and polarities.

Isolated elements, including partial humeri of a Pachydyptes-sized bird, are known from
Parkside Quarry at Weston, near Oamaru (Geology Museum collections, University of Otago).
The horizon is the McDonald Limestone, of Runangan - Whaingaroan age (Late Eocene - Early
Oligocene).

A suite of specimens (Geology Museum collections, University of Otago) has been found
in the Kokoamu Greensand and its lateral equivalents in the Waitaki Valley and nearby areas.
They are of possible late Whaingaroan and certain Duntroonian age (late Early - Late
Oligocene). One includes much of a skeleton (Fig. 17): rostrum, quadrates, vertebrae,
humerus, radius, ulna, more-distal wing elements, a partial synsacrum, femora, tibiotarsus,
tarsometatarsus, phalanges and unguals from near Waimate (Geology Museum collections,
University of Otago). Another is a newly discovered and still unprepared second skeleton from
this site, and articulated elements (generally two or more limb bones) are known from varied
localities. Assessments of comparable elements suggest that at least 3 new species (Fig. 18)
are represented. Also from the Waitaki Valley area are a few specimens (Geology Museum
collections, University of Otago) from the Otekaike Limestone, upper Duntroonian - Waitakian
Stages (Late Oligocene - earliest Miocene). These include a small humerus and ulna similar in

1248 - FORDYCE

INTERNATIONAL NZ No. PENGUIN SPECIES

Ma SUBDIVISIONS Stage 10

QUATERNARY
puios (bate

CENE Early

Tongaporutuan

Walauan

Lillburnian
Clifdenian

Altonlan

Waltaklan

Duntroonian
Whalingaroan

Runangan

Bortonlan

Porangan
Heretaungan

Teurlan

Figure 19. Summary of species-level diversity over time of penguins and possible penguins from New
Zealand.

10 <— ages uncertain

20 Early

30

40

50

Ww
z
Ww
1)
Oo
=
ud
z
Ww
oO
ce)
oS
_
fe)
Ww
z
Ww
oO
fe)
Ww

<@— ages uncertain
60

PALEOCENE

FOSSIL VERTEBRATES OF NEW ZEALAND - 1249

size and profiles to Eudyptula minor (extant Little Blue penguin), a humerus, and articulated
hindlimb elements from a larger bird.

Stratigraphic and Geographic Distribution

_ Middle Cainozoic marine rocks of Otago and South Canterbury, in the South Island, have
yielded a remarkable number of fossil penguins (mostly in Otago Museum collections and in
Geology Museum collections, University of Otago; Fig. 19). Details appear in Table 5. In
summary, the most productive units and localities include the Burnside Marl (Kaiatan, Late
Eocene) and the Concord Greensand (Duntroonian-Waitakian, Late Oligocene-earliest Miocene)
at Burnside Quarry, Dunedin; the Totara Limestone (Runangan, Late Eocene) and the McDonald
Limestone (Runangan-Whaingaroan; Late Eocene-Early Oligocene) near Oamaru; the condensed
sequence of the Kokoamu Greensand (late Whaingaroan-Duntroonian; late Early-Late
Oligocene) and the Otekaike Limestone (late Duntroonian-Waitakian, Late Oligocene-earliest
Miocene) in the Duntroon, Wharekuri and Waimate districts. Surprisingly, no penguin fossils
seem to have been found in the southern South Island in rocks younger than Waitakian Stage
(latest Oligocene-Early Miocene), for example, otherwise richly-fossiliferous units such as the
Gee Greensand, the Tokama Siltstone (encompasses Riflebutts Formation and Bluecliffs Silt)
the Caversham Sandstone, and the Southburn Sand.

Other information on the stratigraphy of New Zealand penguins appears in the major
systematic articles listed previously, especially Simpson (1971a, 1972a - but see comments
above on stratigraphy around Motunau - and 1975; also cf. Fisher 1967 and Simpson 1970).

Zoogeography

The palaeozoogeography of New Zealand fossil penguins has been discussed widely, often
in association with comments on likely habitat preferences and palaeoecology. References
include those of Baubier (1919), Cracraft (1973), Fleming (1979, 1980b), Fordyce (1982b),
R.LF. Jenkins (1974, 1980), Marples (1962), Rich (1975a, 1975b, 1979), Simpson (1975),
Stevens (1980a) and Stonehouse (1969).

Similarities between Late Eocene penguins of New Zealand, Australia, and Seymour Island
(Antarctic Peninsula), and Oligocene penguins of New Zealand and South America were noted
or implied, for example, by R.J.F. Jenkins (1974), Marples (1963), Millener (1988a), Rich
(1979) and Simpson (1971b, 1972). Discussion of zoogeography rests entirely on taxonomy,
and, because penguin taxonomy has been based on traditional "evolutionary taxonomy" rather
than "phylogenetic systematics" (cladistics), it is difficult to evaluate palaeozoogeographic
hypotheses at present. Nonetheless, it is startling to handle roughly contemporaneous
specimens from New Zealand and Seymour Island and find few if any significant differences
between them.

Penguins, Palaeotemperatures and Gigantism

The topic of penguin palaeoecology has dealt mostly with the possible relationships
between body size and the physical climate. There has been much discussion, some rather
fanciful, about the size of New Zealand and other fossil penguins (see Simpson 1975 for a
summary), and this is important in systematics as well as palaeoecology. Because fossils
(particularly complete skeletons) are relatively rare, there is little information available on
ontogenetic and intraspecific variation in size. Yet, size is widely regarded as taxonomically
important. Sizes of fossil penguins are extrapolated from isolated bones on the basis of ratios
derived from extant species. Are such extrapolations reliable? There is little evidence of radical

1250 - FORDYCE

variation in gross morphology amongst early penguins, but the lack of complete specimens
means that we have no idea if body proportions did differ from those of extant species, for
example, because of different ecology. Further study is required. Simpson suggested, possibly
because of the similar body sizes of the organisms involved, that large penguins disappeared
through ecological displacement by cetaceans and seals.

MAMMALIA

New Zealand is noted for its lack of terrestrial mammals. The only extant indigenous
mammals, other than marine mammals, are three species of bat. Mystacina tuberculata and M.
robusta are the sole members of the endemic family Mystacinidae and Chalinolobus
tuberculatus belongs in the Vespertilionidae. All are known from a few subfossil occurrences
(e.g. Holdaway 1989, Millener 1980b) but have no significant fossil record.

Other terrestrial mammals known to have been introduced by man (the dog, Canis
familiaris, and the rat, Rattus exulans), which sometimes occur in middens, are not discussed.
The only mammals considered in detail here are marine mammals, the cetaceans and seals.

It is possible that terrestrial mammals once lived in New Zealand, and it would seem
appropriate to explore likely fossiliferous horizons. It was long thought that the absence of
older mammal fossils from Australia indicated a short history of mammals there, but the recent
discovery of a mid Cretaceous monotreme (Archer et al. 1985) indicates otherwise. The
presence of theropod and omithopod dinosaurs in the New Zealand fossil fauna supports the idea
of physical links with Australia until or about the middle of the Cretaceous, and the ancestors
of other faunal elements are regarded widely as having reached New Zealand before the split
with Australia-Antarctica: Sphenodon, Leiopelma, Dinornithiformes, and many plants.
Placental mammals may have reached New Zealand then. The absence of Recent mammals or
younger fossils could reflect such events as reduced land area during the middle Cainozoic,
fragmentation of land into an archipelago, or climatic extremes during the late Cainozoic.

Seals: Pinnipedia

Seals are amphibious carnivores that encompass two main groups: true seals (Phocoidea;
traditionally Phocidae and Odobenidae), and fur-seals and sea-lions (Otarioidea: Otariidae).
Nomenclature, especially the number of higher taxa and their ranks, is in a state of flux,
particularly as Wyss (1987, 1988) reassessed relationships and suggested that seals are
monophyletic. For other recent reviews of these groups, see Berta & Deméré (1986), King
(1983b), de Muizon (1981), Repenning & Tedford (1977; includes many figures of bones), and
Repenning et al. (1979). Illustrations of seal bones, which may help in identifying fossils,
have been provided in many articles. For examples, see de Muizon (1981), Repenning &
Tedford (1977), and Scarlett (1972b).

New Zealand fossils represent both phocids and otariids. None of the published specimens
is certainly older than Pleistocene. The few specimens known were reviewed by Fleming
(1968), and further commen was provided later by King (1973) and Weston et al. (1973). The
outline below is based mainly on these articles.

Otariidae

A supposed new species of Otariidae, Arctocephalus caninus Berry, 1928, was established
from a mandible of supposed Opoitian (Early Pliocene) age from near Cape Kidnappers,
Hawke's Bay. Weston et al. (1973) demonstrated that the mandible is probably less than 1000
years old. It probably represents the extant Phocarctos hookeri - Hooker's sea-lion (Berry &

FOSSIL VERTEBRATES OF NEW ZEALAND - 1251

King 1970, Beu & Grant-Taylor 1975, Weston et al. 1973). An "almost complete skull" from

Figure 20. Fossil pinnipeds and cetaceans from New Zealand. A, Lateral view of fossil mandible of Ross

seal, Ommatophoca rossi, latest Pliocene, after King (1973); scale bar, 20 mm, body outline, after Walker
1975, shows living Ross seal; B, dorsal view of reconstruction of skull of "Prosqualodon” hamiltoni

Benham, after Fordyce (1978), scale bar, 200 mm; C, dorsal view of reconstruction of skull of
"Prosqualodon" marplesi, after Dickson, scale bar, 200 mm. Rostrum may have been longer than shown.

1252 - FORDYCE

Castlecliffian (Upper Pleistocene) sediments, Ohope, near Whakatane, represents another
otariid, which King (1983a) recently described as the new species Neophoca palatina, placed in
a formerly monotypic genus. King gave detailed comparisons, but did not explicitly identify
synapomorphies which might be of use in otariid systematics.

A third otariid is represented by four as-yet undescribed thoracic vertebrae collected before
1887 from the Butlers Shell Conglomerate (Okehuan, Early Pleistocene), near Kai-iwi,
Wanganui (Fleming 1953:175, 1968). Park (1887a: 63) suggested that the vertebrae may have
been derived from older sediments, but the reason for this idea is not clear. I have not examined
the specimen.

Phocidae

King (1973) described a mandible of the extant phocid, Ommatophoca rossi (Fig. 20), from
limestone (Hautawan, Late Pliocene) near Napier. The Ross Seal now occurs mainly in pack-
ice around Antarctica. This fossil record represents a marked range extension which may have
been a response to expanded cool water belts in the Southern Ocean. Another phocid is
represented by a few skull fragments of probable Nukumaruan (latest Pliocene or Early
Pleistocene) age from near Waipunga, Hawke's Bay. The skull originally was complete, but
was broken by its discoverers (Fleming 1968). Fleming noted that J.A.Berry had concluded
that the fragments are those of a species in a new genus close to the extant elephant seals,
Mirounga .

Other records

There are only a few other records apart from those listed by Fleming (1968). Of older
possible records, Park (1910: 128) stated that "bones and vertebrae" of a seal were collected
from blue estuarine mudstone (Burnside Marl, mainly Kaiatan, but Bortonian and Runangan in
part, Middle and Late Eocene) near Dunedin. This record was mentioned without further
discussion by Service (1934) and Paterson (1941). Marples (1949a) stated that the bones,
which were never described, appear to have been lost. If mammalian, this supposed seal was
probably cetacean (see discussion below), but it is possible that it could have been a penguin or
teleost. Mutch (in Suggate et al. 1978: 520) stated that the remains of seals had been found in
the basal Clarendon Sand (Otaian, Early Miocene) near Milburn, South Otago. The identity
and whereabouts of these specimens are unknown, and this seems a dubious record. I have
examined the "vertebra from the upper miocene beds at Castle Hill, which appears to belong to
the tail of a seal" (Hutton 1900b: 227). It is probably Oligocene in age, and may be from a
small odontocete.

Hutton (1885a: 212) stated that a "skull of a large Sea Elephant" (Mirounga sp.) had been
found in Pleistocene gravels near Oamaru, Hector (1880b) also mentioned Mirounga sp.,
presumably Quaternary. Grant-Mackie & Scarlett (1973:98) listed bones of the extant species,
Arctocephalus forsteri, from the Hillgrove Formation (Oturian, Last Interglacial, ¢. 36,000
years old) near Oamaru, and J.A. Grant-Mackie (pers. comm.) reported recently that bones of
elephant seals, Mirounga sp., have been identified provisionally from these strata.

Whitten (1973, unpublished thesis, per J.A,Grant-Mackie, pers. comm.) collected a small
bone, apparently of a phocid seal, from the Tangahoe Formation (Waipipian, Late Pliocene)
on the coast west of Hawera. What appears to be part of a seal flipper is known from apparent
Late Miocene rocks near Mangaweka (Department of Geology collections, University of
Auckland). These specimens have not been described formally yet.

Other authors have indicated the presence of supposedly fossil seals in New Zealand (e.g.
Mantell 1850: 337, Kellogg 1922: 107-108, table at page 46). These almost certainly are
specimens from archaeological sites. Inasmuch as Miocene phocids now are known from
Victoria, Australia (Fordyce & Flannery 1983), other finds of pre-Pleistocene seals can be
expected in New Zealand.

FOSSIL VERTEBRATES OF NEW ZEALAND - 1253

Whales, Dolphins, Porpoises: Cetacea

New Zealand fossil Cetacea are diverse, and locally abundant (Figs 20-23). For many years
they were little studied, were regarded as difficult to work with, and were considered an
impediment to better things. For example, J. Marwick (in letter to H.D. Skinner, 26 July
1938; letter in Otago Museum) commented ". . . the bones are too fragmentary to be identified
other than marine mammal. . . I don't think the specimen is worth keeping, even if the locality
were known for certain. Best build up that low place at the back of the section". Cetacean
fossils are perhaps most conspicuous in thin calcareous sediments of the Landon Series
(Oligocene), and are known to range back to the Late Eocene in New Zealand. Most of the
described species are of Late Oligocene age, but there are noteworthy Early Oligocene and
Neogene records. The fossil cetacean fauna has not been the subject of monographic treatment,
although an unpublished thesis (Fordyce 1978, 1979c) reviewed the morphology and
systematics of many specimens. A guide to the history of study up to 1978 was given by
Fordyce (1980a). More recent work has been mostly that of Bearlin (1985, 1987a, 1987b,
1988) and Fordyce (e.g. 1977a, 1979a, 1982a, 1983a, 1984a, 1985a, 1985b, 1985d, 1987a,
1987b, 1988b, 1989a, 1989b). Some of these recent summaries cover new finds that have not
been described formally yet. Otherwise, most of the published information comprises either
incidental records, or older descriptions which require revision in the light of modern concepts
of systematics. Many identifications remain suspect (including a few of Fordyce 1978), largely
because of the difficulty involved in making comparisons with taxa described overseas.

All three cetacean suborders are represented in New Zealand: Archaeoceti (extinct;
paraphyletic group), archaic toothed whales; Odontoceti (extant), modern toothed whales,
dolphins and porpoises; and Mysticeti (extant), filter-feeding baleen whales. The systematic
a of the more important specimens are summarised in Table 6 and are discussed in the
text below.

Archaeoceti (Archaic Toothed Whales)

There are provisional records of Archaeoceti from New Zealand, but clearly diagnostic skull
remains have not been found yet. One individual is represented by two teeth reminiscent of
those of the Northern Hemisphere genus Dorudon (Fig. 22). The teeth, originally identified by
Haast (1879: 311) as those of a crocodile, were probably from the Waihao Greensand,
Bortonian-Kaiatan Stages (Middle-Late Eocene) at Waihao River, South Canterbury (Fordyce
1985b). Park (1910: 128) noted that undescribed "bones and vertebrae" of a "seal" had been
found in the Burnside Mudstone (Bortonian-Runangan, Middle-Late Eocene) at an unspecified
site (probably Burnside Quarry) near Dunedin. As odontocetes and mysticetes probably did not
evolve until the Early Oligocene, and seals not until late in the Oligocene, it is possible that
the "seal" was an archacocete (Fordyce 1980c: 325).

The problematic cetacean Kekenodon onamata Hector, 1881a, is known positively only
from the holotype skull fragments (Fig. 21) from the Wharekuri Greensand, Whaingaroan or
Duntroonian Stage (Early or Late Oligocene; Marwick 1959), at Wharekuri, North Otago. The
most detailed description of the holotype is that of Kellogg (1936). Other specimens have been
referred to the species, but either these are known to represent other species of Cetacea (e.g. the
specimens discussed by Hector 1894: 119 and McKay 1882a: 67-68, 1882b: 103-104) or their
identity is unknown (e.g. specimens mentioned by Gudex 1918: 253, 258, Hutton 1888a: 259,
and Park 1905, 1910: 129, and 1911: 545-548). Kekenodon onamata has been identified as an
archacocete (e.g. Hector 1881; Kellogg 1936; Fordyce 1979b, 1980c), although recently this
status was questioned (Barnes & Mitchell 1978, Fordyce 1982c). The holotype does not
exhibit characters that presently warrant its placement in the Odontoceti or Mysticeti, but is

too incomplete to place confidently. It is more specialised than other species normally placed

1254 - FORDYCE

in Archacoceti, but is retained there provisionally in the Basilosaurinae as a late-persisting
relict species (cf. Fordyce 1982c: 669).

Problematic Cetacea

Marples (1949b) described a large natural cranial endocast probably from the Milburn
Limestone (Waitakian, Late Oligocene-earliest Miocene) from the Milburn area, South Otago.
Marples believed that the endocast represented an archacocete-like cetacean, but until more
comparative material is available, odontocete or mysticete affinities cannot be discounted.

Mysticeti (Baleen Whales) - Problematic Mysticetes

Mysticetes are usually thought of as toothless, and such representative baleen whales are
discussed below. However, some toothed fossil taxa are known. For some time these were
allied with Archacoceti or Odontoceti because of the plesiomorphic presence of teeth. The only
family currently available for toothed mysticetes is the Aetiocetidae, which is not yet reported
formally from New Zealand and is strictly known so far from only two north-east Pacific
species. The toothed taxa which occur in New Zealand probably do not belong here.

B AA

\ \ f
F a aan Mn (\

' *)y Pay
‘yee

WG

Figure 21, Harbones of fossil Cetacea from New Zealand. A, periotic of Kekenodon onamata Hector,
ventral view, from Fordyce (1978) scale bar, 10 mm; B, periotic of Kekenodon onamata Hector, dorsal view,
from Fordyce (1978), scale bar, 10 mm; C, tympanic bulla of "Mauicetus" lophocephalus Marples, internal
view, from Fordyce (1978), scale bar,1O mm; D, periotic of Mauicetus parki (Benham), ventral view, scale
bar, 10 mm; E, periotic of new species of Mysticeti (= Kekenodon onamata of Benham), ventral view, from
Fordyce (1978), scale bar,10 mm. F, Periotic of new species of Mysticeti (= Kekenodon onamata of
Benham), dorsal view, from Fordyce (1978), scale bar, 10 mm; G, periotic of "Prosqualodon" hamiltoni
Benham, ventral view, from Fordyce (1978), scale bar, 10 mm; H, periotic of "Prosqualodon" hamiltoni
Benham, ventral view, from Fordyce (1978), scale bar, 10 mm.

The primitive toothed mysticete Mammalodon colliveri is known from the holotype skull
and other material from the Jan Juc Formation, latest Oligocene, Victoria, Australia (Fordyce
1984a, and manuscript A). It is not an aetiocetid, but represents a group of bizarre small
mysticetes in which the rostrum is short. Recently, two New Zealand localities in the

FOSSIL VERTEBRATES OF NEW ZEALAND - 1255

eee Greensand (Duntroonian, Late Oligocene) have yielded remains of a Mammalodon-
like cetacean (Geology Museum collections, University of Otago). One specimen, an
incomplete periotic from Waihao River (South Canterbury), closely matches Australian )
material, and could be conspecific with M. colliveri . Another, a fragment of skull, teeth and a
bulla from Hakataramea from (South Canterbury), is less certainly Mammalodon.

A third toothed specimen, collected recently from Waihao River (South Canterbury) and
still not fully prepared, has denticulate heterodont teeth reminiscent of those of some
dorudontine archaeocetes. The periotic has features that are difficult to interpret cladistically but
which are seen both in Mammalodon and Kekenodon. This specimen probably represents

Figure 22. Teeth of fossil Cetacea from New Zealand. A, cf. Dorudon sp., of Fordyce (1985b), lingual
view, from Fordyce (1978), scale bar, 10 mm; B, "Squalodon" serratus Davis, buccal view, from Fordyce
(1978), scale bar, 10 mm. C, "Squalodon" serratus Davis, anterior view, from Fordyce (1978), scale bar, 10
mm; D, ?Mysticeti indet. (= protosqualodont of Keyes 1973), anterior cheek-tooth, lingual view, from
Fordyce (1978), scale bar, 10 mm; E, ?Mysticeti indet. (= protosqualodont of Keyes 1973), posterior cheek-
tooth, buccal view, from Fordyce (1978); scale bar,10 mm.; F, Tangaroasaurus kakanuiensis Benham, cheek-
tooth, anterior view, from Fordyce (1978), scale bar, 10 mm; G, Tangaroasaurus kakanuiensis Benham,
cheek-tooth, lingual view, from Fordyce (1978), scale bar, 10 mm.

1256 - FORDYCE

another taxon of toothed mysticete. The teeth are larger and smoother than, but are otherwise
similar to those of another probable toothed mysticete (Fordyce 1985d, 1989a), known only
from a partial mandible which retains apparent homologues of archaeocete premolars P1 to P4
and molar 1. The cylindrical jaw and lack of symphysis at P1 level suggest mysticete
affinities, although odontocete affinities were mentioned previously (Fordyce 1980c). The jaw
is from the lower Whaingaroan (earlier Oligocene) Amuri Limestone, which may make it one
of the oldest known mysticetes.

Other specimens are even more problematic. One, "Squalodon" serratus Davis, 1888a, is
known only from the holotype, an isolated cheek-tooth (Fig. 22). This was probably from the
Omihi Formation (possibly Weka Pass Stone Member, later Whaingaroan-Duntroonian) at
Karetu River, North Canterbury. The holotype, which Glaessner (1972) redescribed, exhibits
no features that indicate previously suggested close relationships with either Squalodon spp. in
the strict sense (cf. Glaessner 1972) or Kekenodon onamata, and "Squalodon" serratus is
probably a mysticete. The isolated holotype tooth is similar to one from an incomplete
specimen from Oregon, north-east Pacific (Fordyce 1980d), and the Oregon specimen in turn
has skull features reminiscent of Mammalodon colliveri.

Keyes (1973) described two "protosqualodont" teeth from the McDonald Limestone, near
Oamaru. ‘The source horizon is probably Whaingaroan Stage (Early Oligocene), but could be
Runangan (latest Eocene). Fragments of skull and two more teeth of this specimen (Fig. 24;
Fordyce 1978) have not been properly described yet. The specimen is probably a primitive
mysticete. In turn, it is similar to the Oregon specimen mentioned above. Whether the
similarities between these fossils are derived characters that indicate close relationships, or
whether they are primitive or homoplasous characters, is uncertain.

Family Cetotheriidae

Cetotheres comprise a diverse range of toothless carly mysticetes which have been classified
together mainly because they lack characters typical of extant mysticetes but are more derived
than the toothed mysticetes. Cetotheres differ from Balaenopteridae (extant) in the primitive
lack of an abruptly depressed supraorbital process. The family, therefore is strictly
nonmonophyletic, and requires revision. Family Cetotheriidae is used here more broadly than
by Fordyce (1982c), which acknowledges both the nomenclatural convenience of the group and
the difficulty of revising it.

There are many Oligocene to earliest Miocene cetotheres from New Zealand, but there are
no significant younger Neogene records (Bearlin 1985, 1988). The first formally named New
Zealand species, Mauicetus parki (Benham, 1937a) is known only from the holotype, an
incomplete skull and earbone (Fig. 23) in the Geology Museum collections, University of
Otago. The holotype probably came from the Milburn Limestone (Waitakian, Late Oligocene-
earliest Miocene), at or near Milburn, South Otago. Benham (1937a, 1939, 1942) referred
many other specimens to M. parki but none is demonstrably conspecific (see Fordyce 1980b:
28-32). M. parki is primitive amongst Mysticeti in its relatively long intertemporal region
and is one of the older mysticetes described (Fordyce 1980c, Rothausen 1971). The species
does seem to represent a distinct genus, in contrast to the suggestion of Glaessner (1955).

Marples (1956) described three new species which he referred to Mauicetus. These species
are based on rather incomplete specimens, and their relationships are not clear. Recently,
however, more Oligocene mysticetes have been collected from the Kokoamu Greensand
(Fordyce 1987b, manuscript A), and these should help resolve some of the problems in
taxonomy. The Kokoamu Greensand is mostly of Duntroonian (Late Oligocene) age but is
locally late Whaingaroan (later Early Oligocene), Information about this important unit was
given by, for example, Fordyce et al. (1985), Gage (1957, 1959) and Hornibrook (1966).

"Mauicetus" lophocephalus Marples, 1956, is known only from the holotype skull (Fig.
25; now lost), mandible, carbones (Fig, 21), vertebrae, and forelimb elements, which came

FOSSIL VERTEBRATES OF NEW ZEALAND - 1257

from the Kokoamu Greensand near Duntroon, North Otago. "M." lophocephalus is not
congeneric with M. parki, the type-species of the genus (Benham 1939), is more primitive than
this species, and is more primitive than other species usually assigned to the Cetotheriidae.
Fordyce (1982c) placed it in family incertae sedis (of uncertain relationships at family level),
but, for reasons mentioned above, Cetotheriidae is used here. Another species, "Mauicetus"
waitakiensis Marples, 1956, is known positively only from the holotype skull fragments,
earbones, and a few vertebrae. These were also from the Kokoamu Greensand at Kokoamu,
near Duntroon, North Otago. It represents a species similar to and possibly congeneric with
"M." lophocephalus, and it appears not to be congeneric with M., parki.

Marples’ third species, "Mauicetus" brevicollis Marples, 1956, is known only from the
holotype vertebrae and limb fragments, which came from the Otekaike Limestone (Gage 1957;
Waitakian, latest Oligocene-earliest Miocene), west of Duntroon, North Otago. This species
lacks skull bones or earbones, which makes comparisons difficult. It is not clearly congeneric
with Mauicetus parki,"M." lophocephalus or "M." waitakiensis, but it will be difficult to
diagnose the species or determine its relationships until a skull is found. At present, the
relationship of "M." brevicollis to other specimens from the Duntroonian or Waitakian
(Fordyce 1980b, 1983d, 1987b) is uncertain.

Many new mysticetes have been collected from the Kokoamu Greensand and laterally
equivalent sediments, mainly in south-eastern South Island (Fordyce 1987a, 1989b, and
manuscript A) but only a few have been mentioned in print. These specimens indicate an
unsuspected early diversity amongst mysticetes, and should ultimately expand the record of
mysticetes significantly. Several skulls are known. Of note is one large skull with an original
length estimated at 2 m, which has well-developed vascular grooves for baleen, forward-placed
nares and long parietals (Geology Museum collections, University of Otago). This skull
(figured in Fordyce manuscript A) is reminiscent of Marples’ reconstruction of "Mauicetus"
lophocephalus, but is not conspecific. Other skulls are known. The as-yet unpublished study
of the periotics and/or bullae of these species (e.g. Fig. 23) indicates the presence in the
Kokoamu Greensand, laterally equivalent greensands, and basal Otekaike Limestone, of at least
thirteen cetothere species that differ from "M." lophocephalus (Fordyce 1978, 1987a, 1989b).
These are not clearly conspecific with "M." waitakiensis either. Two of these mysticetes were
described in part by Benham (1937c) who identified them wrongly as Kekenodon onamata.
Both specimens were collected by McKay (1882a, 1882b; see Fordyce 1980b). Benham's
specimen "2" was misidentified by Fordyce (1980c: 327) as a balaenid. Some of these thirteen
species might ultimately be placed in families other than Cetotheriidae sensu stricto.

Another undescribed mysticete is represented by mandibles and skull fragments from the
Abel Head Formation (Duntroonian, Late Oligocene), at Puponga. This was identified by
Whitmore & Sanders (1977) as a probable cetothere,

Fordyce (1980c: 326-7) briefly mentioned the incomplete rostrum of an as-yet undescribed
mysticete from Whaingaroan (Early Oligocene) limestones in southeast Nelson. This provides
a firm early record of baleen-bearing mysticetes. Other undescribed specimens, from the basal
upper Whaingaroan part of the Kokoamu Greensand, may be equally as old.

Family Balaenopteridae

Rorquals (Middle Miocene to Recent) are represented by at least two skulls from the
Neogene (Bearlin 1985, 1988; Fordyce 1980b). Hector (1881a) mentioned one in abstract,
where he identified it as similar to Balaenoptera and stated that it was from the Miocene of
Westland (Fig. 23). Neither the age nor the locality is known certainly. Bearlin (1985)
identified it as a species of Balaenoptera, and discounted relationships (¢f. Fordyce 1982a) with
the cetothere "Plesiocetus" dyticus Cabrera, 1926 from South America.

1258 - FORDYCE

Figure 23. Dorsal views of skulls of fossil Mysticeti from New Zealand. A, "Mauicetus" lophocephalus
Marples, after Marples (1956), scale bar, 500 mm; B, cf. Balaenoptera sp. (of Hector), reconstruction by
Bearlin (1987a), scale bar, 500 mm; C, Balaenoptera sp. (of Bearlin 1985), after Bearlin (1985), scale bar,
500 mm.

The second specimen is a relatively complete skull (National Museum of New Zealand
collections) from Opoitian (Early Pliocene) mudstone of the Waiouru Formation in Taihape.
Bearlin (1985) indicated that this may represent a new genus and species close to Balaenoptera,
but later (1988) identified it as a probable new species of Balaenoptera (Fig. 23). Previously,
Gaskin (1972: Fig. 3) identified it as a cetothere.

Balaenopterids are represented by at least one indeterminate earbone from Chatham Rise
phosphorites, which are of Middle or Late Miocene age (Fordyce 1984b).

Family Balaenidae

Right whales (Early Miocene to Recent) have not been described formally from New
Zealand. No positively identified specimens are known, in contrast to the rather common
occurrences in Australia (Bearlin 1985, Fordyce 1982b). Kingma (1971: Fig. 24) figured the
isolated mandible of a large mysticete, possibly a balaenid, in Nukumaruan (latest Pliocene-
Early Pleistocene) gravels at Matapiro, Hawke's Bay. An undescribed mysticete periotic and
associated bone scraps from Middle-Late Miocene mudstone (stage uncertain) at Gore Bay,
previously thought to represent a balaenid (Fordyce 1982c), appears to be a balaenopterid
(Bearlin 1987). The supposed balaenid of Duntroonian age (Late Oligocene), mentioned by
Fordyce (1980c: 327) was misidentified.

Odontoceti - Family Agorophiidae

Primitive odontocetes of the type traditionally classified in the Family Agorophiidae have
not been reported from the southwest Pacific, Conversely, many specimens occur in Oligocene
rocks in the north-east Pacific (Oregon and Washington States) and those bordering the west

FOSSIL VERTEBRATES OF NEW ZEALAND - 1259

Atlantic. Their absence from otherwise fossiliferous rocks in New Zealand could reflect biases
in collecting and preservation, or real differences between northem and southern faunas.
Family Squalodontidae

"Shark-toothed dolphins", usefully reviewed by Rothausen (1968; see also Pledge &
Rothausen 1977), are represented by at least three species in New Zealand (Table 6). Other
nominal species of Squalodontidae cannot be referred positively to family at present, but some
probably do represent Squalodontidae.

"Prosqualodon" hamiltoni Benham, 1937c, is known with certainty only from the lectotype
skull (Figs 20, 21) and associated elements of Waitakian (Late Oligocene-earliest Miocene) age,
reportedly from Caversham, in Dunedin (Fordyce 1980b). "Prosqualodon" hamiltoni had a
relatively long and broad rostrum, and probably does not belong to Prosqualodon in the strict
sense (Fordyce 1980b, 1980c, Rothausen 1970). "Prosqualodon" hamiltoni may be the
unspecified New Zealand squalodontid which Dal Piaz (1977) suggested was related to a new
species of Italian squalodontid.

"Squalodon" andrewi Benham, 1942 is a problematic species based on an inadequate type
from the Milburn Limestone (Waitakian, Late Oligocene-earliest Miocene), at Milburn
(Andrew 1906). The species, as represented by the holotype, could be conspecific with "P."
hamiltoni. However, since the type of "S." andrewi is poor, the name is probably a nomen
dubium - a name of uncertain application. Benham referred other material to the species, but
none of the referred material is clearly conspecific with the holotype. For example, the
strongly ornamented nominal paratype teeth referred to "Squalodon" andrewi represent another
cetacean, possibly a species of Prosqualodon.

Another squalodontid which is close to or conspecific with the Tasmanian Prosqualodon
davidis, is represented by isolated teeth apparently from the Milburn Limestone (Waitakian,
Late Oligocene-earliest Miocene), in South Otago (Fordyce 1984a: Fig. 20). Isolated cetacean
teeth are generally difficult to identify, but in this case the teeth are strikingly similar in size,
shape and ornament to those of Prosqualodon davidis Flynn, 1923 (Early Miocene, Tasmania;
Flynn 1948). Prosqualodon davidis in turn may be conspecific with Prosqualodon australis
from the Late Oligocene-Early Miocene of Patagonia (M.A. Cozzuol, pers. comm.).

Long-beaked Squalodontidae are represented by an undescribed large skull and jaws from the
Otekaike Limestone, Waitakian Stage, near Duntroon (Late Oligocene-earliest Miocene). The
specimen, which was figured by Fordyce (manuscript B), is in the Geology Museum
collections, University of Otago.

Two species known from incomplete jaws and robust, heterodont teeth are referred
provisionally to the Squalodontidae. However, it will not be possible to positively identify the
family until complete skulls are discovered. Tangaroasaurus kakanuiensis Benham, 1935a, is
known only from the holotype teeth (e.g. Fig. 22) and jaws. These came from the Rifle Butts
Formation (= Tokama Siltstone; Otaian or Altonian, Early Miocene), Kakanui, North Otago.
Benham originally described the specimen as an ichthyosaur (see Anon. 1935a, Camp 1942,
Heune 1936). A second possible squalodontid is represented by a fragment of mandible with
supernumerary teeth, described by Fordyce (1983a). This is from the Milburn Limestone
(Waitakian, Late Oligocene-earliest Miocene), South Otago, and is in the Geology Museum
collections, University of Otago. Another specimen, part of a large rostrum (Otekaike
Limestone, Ngapara, North Otago), was listed as a possible squalodontid ("P hoberodon-like
species" of Fordyce 1982c). However, this is more likely to represent a physeterid (sperm
whale).

ae described nominal species of Squalodontidae may belong in that family, or may
represent other heterodont, early odontocetes. They are placed in Squalodontidae in Table 6
merely for convenience. Of these, Microcetus hectori Benham, 1935b (Anon. 1935b), was
based on a right mandible, which Benham identified as the left maxilla. It was collected by

1260 - FORDYCE

McKay (1882b) from the Otekaike Limestone, Waitakian Stage (Late Oligocene-earliest
Miocene), probably from near the Otekaike River or near Wharekuri, North Otago. The
mandible was associated with a skull (Fordyce 1980b: 16), which was prepared in 1986 but has
not been described yet. Rothausen (1961) noted that "M." hectori is probably not congeneric
with Microcetus in the strict sense, and I support this conclusion.

Undescribed teeth, a periotic, skull fragments and vertebrae collected by Marples from the
Kokoamu Greensand (Duntroonian, Late Oligocene), Duntroon, North Otago, may represent a
species ancestral to "M." hectori. This specimen was briefly mentioned by Fordyce (1980c:
327). The structure of the holotype teeth of "M"." hectori, and of the teeth and periotic of the
older specimen, is unlike those of more-typical squalodontids (e.g. species of Squalodon), and
the specimens may not be Squalodontidae. Nomenclature is further complicated by the
observation that "M." hectori could be conspecific with the contemporaneous, possibly
topotypic, and similar sized species "Prosqualodon" marplesi Dickson, 1964. Comparisons are
difficult because there are not enough common elements preserved. "P." marplesi was based on
a rather complete skull (Fig. 20) and associated elements from the Otekaike Limestone
(Waitakian), near Otekaike River, North Otago. The dotted reconstruction on Figure 20 is
speculative, and there is no clear idea of the length of the rostrum. No referred specimens have
been described. The holotype appears not to represent a species of Prosqualodon (sensu Stricto)
(Fordyce 1980b, 1980c). It does not show any features that indicate clear affinities with
Squalodontidae.

Austrosqualodon trirhizodonta Climo & Baker, 1972, is known only from the holotype
mandibles from the Abel Head Formation (Duntroonian), Puponga, Northwest Nelson. Climo
& Baker assigned the species to the Squalodontidae because its empty tooth sockets indicate a
heterodont dentition (plesiomorphy), but the incomplete holotype mandibles exhibit no
squalodontid features. A relatively complete skull which lacks teeth and mandibles was
collected from the nearby Aorere River, but whether it is related to A. trirhizodonta is
uncertain. The skull is in the Geology Museum collections, University of Otago. Park (1890)
also mentioned fossil Cetacea from Aorere area.

Family Eurhinodelphidae

Eurhinodelphids (for a time called Rhabdosteidae), which are typically small to medium-
sized odontocetes with medium to long rostra, may be represented by at least two specimens.
Phocaenopsis mantelli Huxley, 1859a (see also 1859b-1859d), is known only from the
holotype, an isolated humerus from the Rifle Butts Formation (= Tokama Siltstone; Altonian,
Early Miocene) at Awamoa or Old Rifle Butts, near Oamaru. Previously the species was
identified wrongly as a porpoise (family Phocoenidae) of Pleistocene age (Walker 1975).
Eurhinodelphid affinities are not certain, and the species is best placed family incertae sedis
until new material is found (Fordyce 1982a; also see Abel 1905). Earbones and skull
fragments of a small odontocete from the Waima Formation (Waitakian or Otaian, latest
Oligocene or Early Miocene) at Kaikoura, Marlborough, probably represent a eurhinodelphid.
This specimen, which is in the Canterbury Museum collections, had been identified as a
kentriodontid (Fordyce 1980c: 328) because of its similarity to eurhinodelphid material
described overseas and there misidentified as kentriodontid. It is possible that some worn
periotics from Chatham Rise, which Fordyce (1984b) identified provisionally as ziphiid, are
from eurhinodelphids.

Family Ziphiidae

Beaked whales (Ziphiidae) are represented by several specimens. Fordyce & Cullen (1979)
described a worn ziphiid mandible recovered by dredging from the sea-floor on the Chatham
Rise, east of the South Island. The mandible is not identifiable to generic level. It is probably
of Middle or Late Miocene age. At least two ziphiid periotics (including cf. Hyperoodon sp.)
are also known from the Chatham Rise (Fordyce 1984b), and undescribed ziphiid rostra with

FOSSIL VERTEBRATES OF NEW ZEALAND - 1261

ossified mesethmoids were recently (mid-1988) dredged from the Rise. Hector (in Gray 1871)
mentioned that skulls of the extant genus Berardius (species not stated) were relatively common
in "the old alluvial deposit" from unspecified localities, and Dieseldorff (1901) mentioned a
skull of Ziphius, but none of these specimens nor their locality and age appear to have been
described.
Family Physeteridae

Sperm whales are known only from a few isolated elements. A worn tooth is reportedly
from the Te Aute Limestone at Waipukurau, Hawke's Bay (Fordyce 1978; Canterbury
Museum, Christchurch collections). It is probably of Mangapanian age (Late Pliocene; Beu et
al. 1980), Isolated teeth which match those of form-genus Scaldicetus are known from Middle
or Late Miocene phosphorites on the Chatham Rise (Fordyce 1984b).

: Part of a large rostrum from the Otekaike Limestone (Waitakian, latest Oligocene - earliest
Miocene) near Ngapara in North Otago, probably represents an early sperm whale. The
specimen was earlier listed as a possible squalodontid ("Phoberodon-like species" of Fordyce
1982c) because it has remnants of a heterodont dentition. Its large size and the thick layers of
cementum on the incomplete teeth strongly suggest physeterid affinities. This raises the
question as to whether the fragment of mandible with supernumerary teeth (Milburn
Limestone, South Otago) described by Fordyce (1983a) might also be a physeterid.

Family Kogtidae

Pygmy sperm whales, which are sometimes placed as a subfamily of Physeteridae, have
been reported only recently from New Zealand. Only one specimen is known, a periotic from
Middle or Late Miocene phosphorites on Chatham Rise (Fordyce 1984b, and figure in
manuscript A). It is possible that this or some other Chatham Rise specimens are younger
than Late Miocene, since bones may have accumulated on the rise over millions of years.

Family Kentridontidae

Family Kentridontidae was resurrected by Barnes (1978) to encompass dephinoids which
have essentially symmetrical crania and from which the true dolphins (Delphinidae; below)
apparently arose. There are three possible New Zealand records. One specimen, a partial skull
with procumbent teeth, mandibles and earbones represents an undescribed species probably in
the genus Kentriodon. It was found in the Te Akatea Siltstone (Waitakian, latest Oligocene -
earliest Miocene), just south of Port Waikato, Auckland (Grant-Mackie 1970; Department of
Geology collections, University of Auckland). This Kentriodon-like specimen includes part of
a natural endocranial cast similar to the "odontocete-like" cetacean endocranial cast described by
Marples (1949b). The latter cast was from the Milburn Limestone (also Waitakian), Milburn,
South Otago.

A third specimen is represented by a recently prepared skull (lacking part of the rostrum),
periotic, part mandible, and vertebrae, from the coast between Cape Farewell and Kahurangi
Point, northwest Nelson. The fossil, which is in the Geology Museum collections, University
of Otago, is from the Kaipuke Siltstone, and is probably of Otaian age (Early Miocene).

Isolated periotics and tympanic bullae from Chatham Rise, east of New Zealand, may
represent Middle or Late Miocene kentriodontids (Fordyce 1984b), but I have not yet been able
to distinguish these specimens positively from those of Delphinidae, since earbones of the two
families intergrade in morphology. One supposed kentriodontid from Kaikoura, in
Marlborough (Fordyce 1980c: 328) is probably a eurhinodelphid (see above).

Family Delphinidae

True dolphins are distinguished from kentriodontids by their asymmetrical crania and more
extensive basicranial air sinuses (Barnes 1978). They have an increasingly significant New
Zealand record. For example, at least four species, probably all extinct, are known from
earbones derived from Middle or Late Miocene phosphorites on Chatham Rise, east of New

1262 - FORDYCE

Zealand (Fordyce 1984b). The assemblage from the Rise also includes unphosphatised
specimens that are likely to be younger than Miocene. Weston et al. (1973) mentioned a
partial scapula of probable Waipipian (Late Pliocene) age from Atene, Wanganui, which was
identified as that of Tursiops sp. The specimen is probably too incomplete to allow its
accurate identification (Fordyce 1978). McKee & Fordyce (1987) described a jaw similar to that
of Delphinus or Stenella from the Pliocene of Waihi Beach, Taranaki.

Amongst undescribed fossils, a tooth similar to those of the extant killer whale, Orcinus
orca, was collected from the Greta Siltstone (Nukumaruan, latest Pliocene-Early Pleistocene) at
Motunau, North Canterbury. The worn cranium of a Globicephala-like broad-beaked delphinid
was found in a float concretion at Glenafric, near Motunau. The age is uncertain, but the likely
source sediments in the immediate area are no younger than Miocene. Both fossils are in
Canterbury Museum collections, Christchurch. The small, partial basicranium of another
possible delphinid is also known from this site (Geology Museum collections, University of
Otago). A skull of Delphinus aff. delphis from blue clays (Waitotaran) at Waihi Beach, near
Hawera (F.M. Climo, pers. comm.) has not been described formally yet. An undescribed skull,
possibly that of a species close to the extant false killer whale, Pseudorca crassidens, is known
from the Te Aute Limestone (Opoitian-Nukumaruan, Late Pliocene-Early Pleistocene; National
Museum of New Zealand collections) at Napier. Other delphinid fossils are in the collections
of the Geology Museum (University of Otago), Canterbury Museum, National Museum of
New Zealand, New Zealand Geological Survey, and Department of Geology collections,
University of Auckland.

Family Phocoenidae

Porpoises (Phocoenidae) are known provisionally from a single periotic of an unknown
species from Middle or Late Miocene phosphorites on Chatham Rise, east of New Zealand
(Fordyce 1989b). Fordyce (1984b) identified the specimen provisionally as Kentriodontidae or
Delphinidae indeterminate.

Other Families of Odontoceti

There are not yet any certain fossil records of families Plantanistidae, Lipotidae, Iniidae, or
Pontoporiidae from New Zealand. There is no reason to expect to find them, since fossil and
extant representatives of these groups are found far from the Southwest Pacific.

Other Work

There are many other incidental references to local fossil Cetacea, many of which were listed
by Fordyce (1980b) together with brief comment on the content of the articles involved. These
references give information about localities and taxonomy. Examples include those of Adams
(1910), Anon. (1873), Anon. (1875), Anon. (1878a), Benson (1968), Carlson et al. (1980),
Dal Piaz (1977), Fordyce (1977b, 1983b, 1988b, 1989b, manuscript A, manuscript B), Gage
(1952), Grange (1927), Gregg (1959), Hall (1911), Hamilton (1903b and 1904), Hector (1876a,
1882a, 1882b, 1887, 1889, 1892a, 1892b, 1900), Henderson & Grange (1926), Hill (1900),
Hochstetter (1864; see Fleming 1959a), Hornibrook (1966), Hutton (1888b, 1900a), Hutton &
Ulrich (1875), Kellogg (1923), McKay (1877b, 1877f, 1877g, 1881d), Marshall (1910, 1912a,
1912b), Mason (1941), Morgan (1911), Park (1890, 1918, 1921, 1923), Speden & Keyes
(1981), Speight (1912), Speight & Wild (1918), Stromer (1915), J.A. Thomson (1906, 1920,
1926b), J.T. Thomson (1874), and Ward & Lewis (1975).

FOSSIL VERTEBRATES OF NEW ZEALAND - 1263

BROADER ISSUES IN NEW ZEALAND VERTEBRATE
PALAEONTOLOGY

INTERPRETING THE RECORD

Most palaeontologists are more interested in broader patterns of the fossil record than in
minor details. Indeed, developments in macroevolution promise to increase both public and
scientific interest in palaeontology. However, the New Zealand fossil vertebrate record has so
far contributed few specifics to discussion of macroevolution. Potentially, the New Zealand
fossils might help interpret aspects of evolution, extinction, and their possible correlation with
environmental change such as changing sea-levels (Vail & Hardenbol 1979, Vella 1967),
temperatures, elevation (Fleming 1963b) and geography (Fig. 6). There is great scope for
speculative interpretation, in the absence of a detailed vertebrate record more typical of some
northern continents, although in fairness much of the past speculation has probably usefully
explained known patterns and directed future work. More modern examples that discuss
specifically New Zealand fossil vertebrates include articles by Schwarzhans (1980, 1981b,
1984, on fish), Simpson (e.g. 1975, on penguins), Fordyce (e.g. 1979a, 1979b, 1980c, on
Cetacea), Fordyce & Jones (1989, on penguins) and Fleming (many articles, listed by Keyes
1987, mostly based on neospecies). Quaternary birds, in particular, have got a lot of attention.
I do not intend in this review to cover specific case studies, but touch on a few general points
of interpretation.

APPROACHES TO TAXONOMY

Some general aspects of systematics are relevant here, since they govern approaches to the
local fossil, recently extinct and extant vertebrates. For example, few local palaeontologists
have considered principles of cladistics in their work (but cf. Fordyce & Jones 1989, Worthy
1987a: Fig. 40). Most taxa, therefore, have been defined by an amalgam of primitive and
derived characters (for a recent example see Wiffen & Moisley 1986: 213-4, 250). These
definitions of taxa have been supported at times by non-morphological attributes, such as
inferred ecology and inferred geographic distribution. For example, Fleming (in Newman
1982: 342) commented on the role of likely geographic distribution in making taxonomic
decisions, as did Scarlett (1983: 419) when establishing a new genus, and Worthy (1987a)
commented on the geography of Ascaphidae and Leiopelmatidae. Stratigraphic range seems not
to have been used as an integral part of definitions of New Zealand fossil vertebrate taxa.

Presumably, while some commonly discussed groups are monophyletic, others are
paraphyletic or polyphyletic. Caution may be required, then, in interpreting the record of many
commonly accepted taxa.

COMPLETENESS OF RECORDS

Many higher taxa have marked gaps in their stratigraphic records. Are the gaps real (taxa
have not been in the New Zealand region), or are they a result of biased study? For example,
Keyes (1984) noted new local records of chondrichthyan taxa that are really quite common but
were not recognized because they are small and best recovered by screening. Few teleostean
skeletons have been described. No local pre-Quaternary specimens of Sphenodon or Leiopelma
are known, and there are a few scraps of pre-Quaternary moa, yet ancestors of these taxa are
thought widely to have been represented in New Zealand since the Mesozoic (Cracraft 1975,

1264 - FORDYCE

1980a, Fleming 1979, Hutton 1873). Were terrestrial mammals once present? Amongst the
rather better represented marine vertebrates, there are many Palaeogene penguins but few that
are stratigraphically well-placed from the Miocene. Odontocete and mysticete cetaceans, which
have a good local Late Oligocene record, probably evolved during the Early Oligocene (Fordyce
1980c). It is ironic that Cetacea of this apparently important age are known poorly locally and
globally (e.g. Fordyce manuscript A). And, we are profoundly ignorant about the stratigraphic
distribution of New Zealand fossil vertebrates in units older than the relatively complete later
Cretaceous-Cainozoic sequences (Fig. 4).

NEW ZEALAND TYPE SPECIMENS

New Zealand workers, like those from some other former colonies, face the problem of type
specimens that are housed overseas. Some moas, Cretaceous reptiles and one cetacean, for
example, are currently in Britain where they may well be curated more securely than some New
Zealand collections are at present. Regretfully, material has not only gone overseas
permanently for study but some important specimens have been lost in transit or upon arrival
overseas. Examples include Cretaceous reptiles (see Welles & Gregg 1971), an apparently
rather complete skeleton of a Neogene penguin from North Canterbury (R.J. Scarlett, pers.
comm.), and the Permo-Triassic reptile long bone from Stephens Island which was sent to
Colbert (see above), New Zealand institutions, including those museums with a statutory
requirement to house types, at times have been no more responsible, because of a lack of funds
and staff and perhaps because the short history of study here has instilled little appreciation of
vertebrate fossils. Thus, parts of the holotypes of the cetaceans Kekenodon onamata,
Mauicetus lophocephalus and Tangaroasaurus kakanuiensis have been lost (Fordyce 1980b), and
some material has been poorly curated.

Many type-specimens of New Zealand fossil vertebrates are inadequate because they are
dubiously diagnostic or are incomplete. They are, therefore, easily misinterpreted. Some of
these have been alluded to above, and other examples abound. Stinton (1957), for example,
reported fossil ice-fish, Notothenia, from New Zealand. This identification was rightly regarded
as important (Holdgate 1970: 352), but it is a misidentification (Schwarzhans 1984). Many
holotypes of nominal species of moa are based on one or a few elements (e.g. Archey 1941,
Brodkorb 1963, Oliver 1949), The holotype of the penguin Palaeeudyptes antarcticus is an
isolated tarsometatarsus of uncertain (?Oligocene) age. No other specimens of this species are
known, so the relationships of P. antarcticus to all other fossil penguins are debatable.
Nonetheless, Hector referred many other specimens to this species, and other species have been
identified as congeneric with P. antarcticus. The Late Oligocene seabird, Manu antiquus, is
based on a furcula, and its affinities are uncertain. Cetaceans include dubiously diagnostic
elements such as teeth (e.g. "Squalodon" andrewi), forelimb fragments (e.g. Phocaenopsis
mantelli), and frustratingly incomplete skulls (Mauicetus parki). Such nominal species based
on incomplete specimens are sometimes identifiable at all taxonomic ranks down to species,
but there are many cases where affinities are vague even at ordinal level (see comments by
Fordyce 1988a), Such uncertainty reflects ignorance about homologies and thus about the
taxonomic significance of characters, and reflects the undiagnostic nature of many types.
Biochemical techniques (e.g, comparisons of amino acids, immunological comparisons;
Lowenstein 1986, Wilson ef al. 1987) show promise for helping determine the biological
affinities, and thus the nomenclatural problems, of many fragmentary types. These techniques
could be used to see whether incomplete types (of little value other than as name-bearers) are
likely to be conspecific with more complete specimens which are useful in interpreting the
biology of species. In the meantime, the palacontologist must decide whether such names are
based on such incomplete types that they are nomina dubia - names of uncertain biological
affinities. Cladistic approaches to palacontology will focus further attention on such types.

FOSSIL VERTEBRATES OF NEW ZEALAND - 1265

Uncertainties about systematics of New Zealand fossil vertebrates reflect a lack of good
specimens in most groups. For marine vertebrates, for example, many fish are known only
from otoliths, which may be classified differently than complete skeletons. Fossil penguins
are known by many isolated bones but few partial skeletons. Recent local discoveries of
articulated specimens may eventually solve this problem, but at present it is difficult to deal
with incomplete specimens such as fragmentary bird bones (Manu antiquus, above) and isolated
teeth of Cetacea which may have some stratigraphic, geographic or ecological value. Isolated
fragments are difficult to deal with cladistically since we rarely have much idea of the polarities
and thus the potential value of many characters. Thus, comparisons can be based on little more
than gross similarity. This approach may allow one to eliminate the taxa to which a scrappy
specimen does not belong. However, it will not necessarily allow specimens to be placed in
clades, especially at, say, genus or species level.

Some higher taxa (e.g. families, suborders) are not well defined, and this makes it hard to
place some specimens (see comments by Fordyce 1989a). For example, amongst Cetacea, the
Suborder Archaeoceti is paraphyletic, as it does not encompass all descendants of the most
recent common ancestor of the group. Archaeocetes are conventionally defined as Cetacea that
lack the derived characters of odontocetes and mysticetes (e.g. Barnes & Mitchell 1978). Thus,
they are defined by default. Many otherwise problematic Cetacea are referred to the Odontoceti
or Mysticeti because they seem more derived than archaeocetes, but the recent discovery of
remingtonocetids is a reminder that highly specialised archaeocetes can be encountered. Ratites
form another widely recognised group which, in contrast to traditional views, may be
polyphyletic (Houde 1986, James & Olson 1983).

EXTINCTION

The global fossil record indicates that extinction of species is the rule. Most species seem
to have disappeared without evolving into others. Despite its acknowledged importance in
paleobiology, there has been little discussion of extinction amongst New Zealand fossil
vertebrates other than moas.

Causes of Extinction

The last decade has seen renewed interest in general processes of extinction over geological
time and their role in evolution. Many sorts of environmental change have been identified as
important causes of extinction, For example, large scale habitat loss, perhaps through changes
in climate or sea-level, may be critical. Attention has focused on the role of the transgression
which affected New Zealand from the later Cretaceous to about the middle of the Cainozoic
(e.g. Rich & Rich 1982). But interpretations differ about the extent of land at peak
transgression, and this is important since the absolute area available possibly affected evolution
of faunas. Alternative Oligocene reconstructions are those of Fleming (1962a, 1979) and of
Kamp (1986). How much loss of area is too much for a particular taxon to cope with, and do
area models of biogeography adequately explain diversity changes? Presumably, profound
habitat loss would lead to reduced population size, so that a genctic bottle-neck was reached to
prevent effective interbreeding. What constitutes a minimum effective population size is
uncertain, and for how many generations must populations be at low levels before bottle-neck
effects act? Breeding programmes with extant endangered New Zealand taxa, admittedly mostly
distinct only as subspecies, suggest that small isolated populations can be maintained
adequately over some years, these examples, however, are short even in terms of ecological
time. One of the main problems with such broad explanations is that they are so general as to
explain little about the disappearance of individual species.

1266 - FORDYCE

It is axiomatic that extinctions will result if environments change more quickly than
species can change, but this explanation tells little. So-called "genetic senescence and the
sporadically revived idea of orthogenesis (e.g. Grehan 1984) have been suggested as possible
explanations of inability of organisms to keep pace with environmental change, although they
do not seem to have been applied seriously to New Zealand fossil vertebrates. Such ideas have
been used as well to explain the persistence of apparently ancient and morphologically
conservative taxa, but I am unaware of examples amongst the vertebrates where orthogenesis
has provided a better explanation of apparent tightly constrained evolution than has critical
analysis of clades and interpretation in light of constructional morphology.

Sorts of Extinction

Phyletic extinction (pseudoextinction), where one species becomes extinct as it evolves into
another, seem not to have been recorded for New Zealand fossil vertebrates. Marine teleosts,
well represented by otoliths, might reveal extinction associated with such ancestor-descendant
relationships. Schwarzhans (1980, 1984) alluded to such relationships, as shown by otoliths,
but did not provide details.

Examples of individual extinction (of a single species) abound amongst many recently
extinct local species, e.g. huia (Heteralocha acutirostris) and piopio (Turnagra capensis), and
individual extinctions have been discussed widely in the literature (e.g. Anderson 1984,
Diamond & Veitch 1981, Fleming 1962b, 1962c, 1969, Holdaway 1989, McDowell 1969,
Millener this volume, Williams 1960, 1962, 1964, 1973), There are no clearly documented
examples for fossil vertebrates in the New Zealand literature. No fossil taxa are known well
enough to say that a perceived (apparent) absence above a particular horizon is a real absence
which indicates the extinction of the species. Only those fossil taxa that are sampled over
close stratigraphic intervals through relatively invariant facies and over a wide geographic range
(e.g. marine teleosts, and some continental mammals elsewhere) might provide conclusive
examples. However, one might expect this type of extinction to have been the rule in New
Zealand. There is evidence of local extinction (change in range) for some species. For
example, the extant Takahe (Porphyrio mantelli) has experienced a major recent range
contraction (Beauchamp & Worthy 1988, Mills eg al. 1988), and the extant Ross Seal
(Ommatophoca rossi), which is an obligate ice-dweller in Antarctica, occurs as a Late Pliocene
fossil in New Zealand (King 1973).

Taxonomic extinction affects many or all members of a taxon, and is not necessarily
coincidental with other extinctions. A taxon could disappear, however, as part of a mass
extinction, €.g. moas. Taxonomic extinction could result from climate changes and changing
geography, and ecological displacement through competition could also be important. New
Zealand examples include moas (Dinornithiformes), and the tuatara (Sphenodon) is the last of
the Sphenodonta, a taxon extinct elsewhere,

The disappearance of moas was probably part of a mass (or ecological) extinction. Such
events see the coincident extinction of many unrelated groups in response to a single cause, or
perhaps coincident causes, and are often followed by ecological replacements. The role of
humans in the extinction of moas seems clear enough in New Zealand (Holdaway 1989) where
it might be called overkill, but climatic change has been invoked to account for the
disappearance of some large Quaternary land vertebrates elsewhere (see Murray, this volume).
There appears to have been continued invasion by Australian species during the Quaternary,
which might indicate unfilled ecological opportunities, and/or a lack of resistance to invasion
and perhaps indicates a depauperate earlier Quaternary avifauna. Fleming (1962b) touched on
the historical context of invasions, Moas were discussed by Anderson (1984), Cassels (1984)
and Trotter & McCulloch (1984) (see also references cited under moas, above)

FOSSIL VERTEBRATES OF NEW ZEALAND - 1267

The mass extinction at the Cretaceous-Tertiary boundary, which affected m
worldwide, is reflected amongst local vertebrates. tn New Zealand no aeons care
ichthyosaurs or representatives of some key invertebrate groups are clearly found above the
boundary. The reports by McKay (1877e: 37) and Thomson (1920: 346) of plesiosaurs from
above the boundary at Waipara have not been substantiated. There is scope for more study on
marine vertebrates, especially fish. Studies are likely to be complicated because of problems of
biostratigraphy in the common nearshore marine facies, in which good age-diagnostic species
may be rare. Extraterrestrial object impacts have recently been invoked as a possible cause of
the extinctions, see innumerable recent articles including those by Alvarez et al. (1980),
Jablonski (1984), Nitecki (1984), and Officer et al. 1987..

PALAEOZOOGEOGRAPHY

Studies of the geographic distribution of fossils tell much about evolutionary processes,
past geographies, and ancient environments, given that environment controls distribution.
Studies can be on an outcrop, local, regional or global scale. Results of such studies may
indicate change in continental arrangement, in climate or in the evolution of clades. Problems
arise because palaeobiogeography relies on information from diverse sources. For example,
species and higher taxa are the raw data of zoogeography, but different taxonomists may
classify the same organism in different ways depending on their preference for phenetics,
evolutionary systematics, cladistics or other approaches. Furthermore, the taxonomy of many
New Zealand vertebrates has not been reviewed recently. It can be difficult to assimilate the
results of studies based on “evolutionary taxonomy", in contrast to cladistics, where characters,
polarities and their significance must be stated explicitly. Thus, it is difficult at present to
assess previous hypotheses about the origins of many taxa. For example, moas and kiwis are
of uncertain relationships with other ratites, and the ratites are not known positively to be
monophyletic (Houde 1986, P.V. Rich 1982, cf. Cracraft 1974b). New Zealand ratite
zoogeography, formerly discussed by Cracraft (1980a) and others, requires further study.

Conflicts also arise in biostratigraphy and thus in the timing of events in
palacobiogeography. Examples were given above.

The fossil record for vertebrates is probably quite biased geographically. A map of
occurrences of a fossil species will generally show a map of outcrops or even a single outcrop.
Such a map may not represent the range of the species as a whole, especially for taxa that are
rare. For example, large vertebrates, such as whales, are conspicuous in outcrop but are rare,
and the distribution maps for such taxa will show a single occurrence. In contrast, many
invertebrates are inconspicuous yet common at one or a number of localities. Absence of
evidence is not the same as evidence of absence.

Factors Controlling Distribution

The little that is known about the New Zealand fossil record suggests that it has its share of
endemic taxa. Indeed, few fossil species known in New Zealand have been recorded elsewhere.
One could claim that the record is biased for some taxa, such as cetaceans and perhaps some
marine reptiles which were probably rather vagile and should have been recorded elsewhere. But
why are some taxa not more cosmopolitan? It is usually stated that interacting biological,
physical and geological factors govern distributions, but this general explanation actually

explains very little.
Amongst physica
(e.g. the Tasman Sea,
topography (e.g. walter depth, elevation on |
temperatures, Or seasonality), and currents. G

1 factors, barriers that conspicuously limit distribution include distance
the effects of which have been discussed widely for local vertebrates),
and), temperature (e.g. absolute limits, range of
eological factors, such as the timing and nature of

1268 - FORDYCE

tectonic movements, must also have played a role. New Zealand has been isolated since about
the mid-Cretaceous, by which stage dinosaurs, flightless birds and other terrestrial vertebrates
perhaps got here. Does the absence of snakes and land mammals reflect early isolation?

Biological processes interact in a complex fashion with physical and geological factors.
Examples include the relationship between biological diversity and habitat area, and the link
between the evolution of species and geographic change, Empirical observations on island
biotas (e.g. birds) suggest that the diversity of species bears some constant relationship to
habitat area (Case & Cody 1987, Hoffman 1985) Given this, changes in habitat area should
influence evolution and extinction (e.g. Rich & Rich 1982). Such changes in area should be
expected through sea-level change during the broad transgression from the Cretaceous to the
Oligocene, and through uplift associated with continental collision.

It has long been thought that the evolution of species relates to geographic change.
Intraspecific geographic variations can be influenced by incipient barriers. The range of a
species may be disrupted physically, so that originally contiguous populations are split and
genetically isolated from each other. Indeed, whole biotas may be disrupted in this way by
abiotic vicariant (continental fragmentation) events, and this probably happened in proto-New
Zealand about the mid Cretaceous. Populations may then diverge so that sister species are thus
formed through allopatric speciation. Possible New Zealand examples of allopatric speciation
include those of the extant kea and kaka, Nestor spp. (Fleming 1979; see also Lambert 1982),
but until recently there has been little discussion of fossil vertebrates other than passing
mention of possible north-south species pairs of moas (Worthy 19874).

Evolutionary divergence might also follow dispersal events, where members of a single
species disperse to previously unoccupied areas. The large number of indigenous birds that
have apparent close (congeneric or even conspecific) relatives in Australia, and historical
evidence of successful invasions, suggest ongoing trans-Tasman dispersal. Presumably, sister
species in New Zealand have diverged from Australian counterparts since dispersal (e.g. Falla
1953, Fleming 1962b). There are important questions to address here: has this process been
significant in the distant past, what has prevented gene flow across the Tasman after such
dispersal events and why has New Zealand not been more resistant to the invasion of new
species?

Amongst the more important, and certainly the most readable, articles on the influence of
changing geography on the evolution of New Zealand's biota are those by Fleming (e.g. 1962a,
1975, 1979, and others listed by Keyes 1981b and 1988). Fleming's approach to historical
biogeography was mostly, but certainly not wholly, that of the dispersalist, which emphasises
that animals are vagile and that multiple, separate, long-distance dispersal of individuals occurs
to aliow new habitats to be colonised. Dispersal events plausibly account for the origins of
many of New Zealand's fossil and extant marine invertebrates and vertebrates, and many extant
and extinct birds. Vicariance events, in contrast to dispersal, occur when abiotic changes (e.g.
continental fragmentation) result in a disjunction in the ancestral range of relatively sedentary
animals that might not disperse easily. There seems little doubt that vicariance events did
account for the origin of some species in New Zealand's extant fauna (e.g. Leiopelmatidae,
Sphenodon), but this has received little discussion until recently (e.g. Caughley 1964, for pre-
drift interpretations). Although dispersal and vicariance together might account for the origins
of New Zealand's biota, this has been ignored or underemphasised by some vociferous disciples
of pure dispersal (who are becoming increasingly rare), of pure vicariance (who are becoming
increasingly common), and of panbiogeography (who seem to disown all others). Some would
argue that panbiogeography holds the key to resolving problems in the historical biogeography
of New Zealand, but I am unaware of any serious discussion of fossil vertebrates, especially the
marine vertebrates, that contribute in such a large way to the local record. Local approaches to
panbiogeography were revealed by contributors in Craw & Gibbs (1984; for review see Stoddart
1985). For discussion of more theoretical aspects of New Zealand biogeography, see, e.2.

FOSSIL VERTEBRATES OF NEW ZEALAND - 1269

Caughley (1964), Cracraft (1973, 1974a, 1980a), Craw (1978, 1980, 1985), McDowall (1980a,
1980b), McGlone (1985), Worthy (1987a) and many references cited in these papers. It must
be stressed that hypotheses in historical biogeography stand or fall on taxonomy and, to a
lesser extent, stratigraphy. For example any track analyses in biogeography which consider
only the "similarity" of species involved, are more debatable than "cladistic vicariance"
hypotheses for reasons discussed by Patterson (1981a). Finally, it seems appropriate to heed a
comment by Pielou (1981): " an explanation for any given disjunction (whether it reflects
palaeogeography or palaeoclimate) is more likely to be obtained by common-sense than by
abstract theorising”,

References on evolution, extinction and biogeography of the New Zealand biota include
articles by Caughley (1964), Cracraft (1975), Diamond & Veitch (1981), Fleming (1949,
1962b, 1963d, 1975, 1976a, 1976b, 1980b), Forbes (1893c), Fordyce (1980c, manuscript A),
Gaskin (1975), Hutton (1872), Keast (1971), Molnar (1981), McDowall (1969, 1973), P.V.
Rich (1975a, 1975b, 1979), Scarlett (1957), Simpson (1975) and Stevens (1973, 1976,
1980a), other contributors in Ballance (1980) as well as those listed above.

ACKNOWLEDGEMENTS

A compilation such as this reflects inputs from diverse sources, and it is difficult to
acknowledge all of them adquately. Were it not for the efforts of a long list of sharp-eyed
prospectors and collectors over a span of 150 years, this review could not have been produced.

I thank many colleagues, especially Doug Campbell, Hamish Campbell, Mario Cozzuol,
the late Charles Fleming, Neil Fowke, Jack Grant-Mackie, Andrew Grebneff, Craig Jones, Ian
Keyes, Dave MacKinnon, Phil Millener, Pat Rich, Tom Rich, Ron Scarlett, and Frank
Whitmore, for their general interest in discussing and/or commenting on New Zealand fossil
vertebrates over the years, or for comments on the manuscript. Chris Paulin gave useful
comment on the table of fish. The following institutions provided longer-term research
facilities: the University of Canterbury, the National Museum of Natural History (Smithsonian
Institution), Monash University, the Museum of Victoria, and, most importantly, the
University of Otago. In addition, I acknowledge valuable access to the collections of, and the
loan of specimens from, the Department of Geology at Auckland University, the New Zealand
Geological Survey, the National Museum of New Zealand, Canterbury Museum, and Otago
Museum. Work reported here was supported variously by a New Zealand University Grants
Committee Postgraduate Scholarship, a grant in aid from New Zealand Geological Survey, a
Smithsonian Institution Postdoctoral Fellowship, a Monash University Postdoctoral
Fellowship (all 1975-1982), and more recently the Research Committee of the University of
Otago (1982-1991), the National Geographic Society (1987-1991), and the New Zealand
Lottery Board (1988-1990).

Important preparation was carried out by Andrew Grebneff and Craig Jones, with help from
others including David Bevin, Lew Kerr, Stuart Munro, Fiona Rayns and David Wood. I thank
Robert Bearlin, Craig Jones, Brian Jamieson, Jane Kerr and Fiona Rayns for help with
illustrations and/or literature. June Crawford and Adrien Dever helped with typing.

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1270 - FORDYCE

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FORBES, H.O., 1891a. On avian remains found under a lava-flow near Timaru, in Canterbury. Trans. N. Z.
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FORBES, H.O., 1892a. Evidence of a wing in Dinornis. Nature 45: 257.

FORBES, H.O., 1892b. New extinct rail. Nature 45: 416.

FORBES, H.O., 1892c. On a recent discovery of the remains of extinct birds in New Zealand. Nature 45:
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FORBES, H.O., 1892d. Aphanapteryx in the New Zealand region. Nature 45: 580-581.

FORBES, H.O., 1892e. Preliminary notice of additions to the extinct avifauna of New Zealand (Abstract).
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FORBES, H.O., 1892f. Aphanapteryx and other remains in the Chatham Islands. Nature 46: 252-253.

FORBES, H.O., 1892g. Remarks on a series of extinct birds of New Zealand, recently discovered. Rep.
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FORDYCE, RE, 1977a. The development of the Circum-Antarctic Current and the evolution of the Mysticeti
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FORDYCE, R.E., 1979a. Cetacean evolution and late Paleogene environments of the Southern Oceans.
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FORDYCE, R.E., 1979c. The morphology and systematics of New Zealand fossil Cetacea. (Abstract of
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FORDYCE, R.E., 1980a. Records of two Paleogene turtles and notes on other Tertiary reptilian remains from
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FORDYCE, R.E., 1980b. The fossil Cetacea of New Zealand (A catalogue of described papers and species with
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FORDYCE, R.E., 1980c. Whale evolution and Oligocene Souther Ocean environments. Palaeogeog.
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FORDYCE, R.E., 1982b. The Australasian marine vertebrate record and its climatic and geographic
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FORDYCE, R.E., 1985b. Late Eocene archaeocete whale (Archaeoceti: Dorudontinae) from Waihao, South
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FORDYCE, R.E., 1986. The Otago plesiosaur: what might one learn from fossils? N. Z. Sci. Teacher 51:
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FORDYCE, R.E., 1988a. Taxonomic status of Victorian fossil whales assigned to the genus Cefotolites
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1278 - FORDYCE

FORDYCE, R. E., 1989a. Problematic Early Oligocene toothed whale (Cetacea, ?7Mysticeti) from Waikari,
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FORDYCE, R.E., 1989b. Origins and evolution of Antarctic marine mammals. LGeol. Soc. Lond. sp. publ.
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FORDYCE, R.E., manuscript A. Evolution and zoogeography of cetaceans around Australia and New Zealand.
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FORDYCE, R.E., manuscript B. Fossil whales, dolphins and porpoises. Submitted to Earth Sct..

FORDYCE, R.E. & CULLEN, D.J., 1979. A Miocene ziphiid whale (Odontoceti: Cetacea) from central
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FORDYCE, R.E. & FLANNERY, T.F., 1983. Fossil phocid seals from the Late Tertiary of Victoria. Proc. R.
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FORDYCE, R.E., HORNIBROOK, N. DE B. & MAXWELL, P.A., 1985. Field trip guide to Cenozoic geology
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FORDYCE, R.E. & JONES, C.M., 1987. New fossil penguin material from New Zealand. [Abstract.]
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FORDYCE, R.E. & JONES, C.M., 1988. New fossil penguin material from New Zealand and the early history
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FORDYCE, R. E. & JONES, C. M., 1990. The history of penguins, and new fossil penguin material from
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FORDYCE, R.E., JONES, C.M. & FIELD, B.D., 1986. The world's oldest penguin? Geol. Soc. N. Z. newslett.
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FORREST, R.M., 1987. A partially mummified skeleton of Anomalopteryx didiformis from Southland. J. R.
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FROST, G.A., 1933. Otoliths of fishes from the Tertiary formations of New Zealand. Trans. N. A. Instit. 63:
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GAFFNEY, E.S., 1981. A review of the fossil turtles of Australia. Am, Mus. Novit. 2720: 1-38.

GAFFNEY, E.S., 1984. Historical analysis of theories of chelonian relationship. Syst. Zool. 33: 283-301.

GAGE, M., 1952. The Greymouth coalfield. N.Z. Geol. Surv. Bull. ns. 45: 1-240. [Cetacea, p. 49.]

GAGE, M., 1957. The geology of Waitaki subdivision. N. Z. Geol. Surv. Bull. ns. 55: 1-135.
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GAGE, M., 1959. Kokoamu Greensand, In Océanie. Fascicule 4. New Zealand. Lexique Stratigraphique
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GAGE, M., 1969. Rocks and landscape. In The Natural History of Canterbury. G.A. Knox, ed., A.H. & A.W.
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GAGE, M., 1970, Late Cretaceous and Tertiary rocks of Broken River, Canterbury. N. Z. Jl. Geol. Geophys.
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GAGE, M., 1979. Ice age earth: New Zealand was strictly for birds and bats. Geol. Mag. 51: 838-843.

GAGE, M., 1980. Legends in the Rocks. Whitcoulls, Christchurch: 1-426,

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WIFFEN, J. & MOISLEY, W.L., 1986. Late Cretaceous reptiles (families Elasmosauridae, Pliosauridae) from
the Mangahouanga Stream, North Island, New Zealand. N. Z. JI. Geol. Geophys. 29: 205-252.

WIFFEN, J. & MOLNAR, R.E., 1988. First pterosaur from New Zealand. Alcheringa 12: 53-59.

WIFFEN, J. & MOLNAR, R.E., in press. An Upper Cretaceous ornithopod from New Zealand. Geobios.

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WILLIAMS, G.R., 1964. Extinction and the Anatidae of New Zealand. In The Wildfowl Trust Research and
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FOSSIL VERTEBRATES OF NEW ZEALAND - 1295

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WORTHY, T.H., 1987c. Osteological observations of the larger species of the skink Cyclodina and the
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WORTHY, T.H., 1988c. An illustrated key to the main leg bones of moas (Aves: Dinomithiformes). natn.
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Mosasauridae). J. Palaeontology 63: 126-127.

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31.

WYSS, A., 1988. On "retrogression” in the evolution of the Phocinae and phylogenetic affinities of the
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YALDWYN, J.C., 1956. A preliminary account of the sub-fossil deposits of the Martinborough Caves. Rec.
Dom. Mus. 3: 1-7.

YALDWYN, J.C., 1958. Notes on the environment and age of the sub-fossil avifauna of the Martinborough
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YALDWYN, J.C., 1959. Recent name changes in the moa genus Dinornis. J. Polynesian Soc. 68: 80-82.

YALDWYN, J.C., 1979. The types of W.RB. Oliver's moas and notes on Oliver's methods of measuring moa
bones. In Birds of a Feather, A. Anderson, ed., N. Z. Archaeolog. Assoc. Mono. 2: 1-24

1296 - FORDYCE

APPENDIX I

Table 1. Genera of fish reported as fossil from New Zealand. Classification based on Ayling & Cox (1982), Grenfell
(1984), Romer (1966) and Schwarzhans (1980, 1984). Not all older names (e.g. of Frost) have been synonymised herein; see
Schwarzhans (1984: 217-223). Authors of genera are not cited unless the genus was described from New Zealand. Selected
references to descriptions and/or age follow the generic names. Only the more significant unpublished records are included.
Repository cited only for specimens not previously mentioned in print.

Class CHONDRICHTHYES

ORDER SELACHII
Family Heterodontidae - horn sharks

Heterodontus [= Cestracion, Romer 1966]; Chapman 1918, "Miocene"; Keyes 1987, Bortonian-Runangan (Middle-Late
Eocene).

Synechodus,; Chapman 1918, "Cretaceous" (Romer 1966 cites "Eocene"); Keyes in Wiffen 1981a, Piripauan or Haumurian
(Late Cretaceous), Keyes1981a, Teurian (Paleocene)

Family Odentaspididae - sand sharks

Eugomphodus [includes Odontaspis, Carcharias]; Davis 1888b; Chapman 1918, "Oligocene, Miocene, Lower Pliocene";
Keyes in Wiffen 1981a, Piripauan or Haumurian (Late Cretaceous); Keyes 1987, Mangaorapan-Runangan (Early-Late
Eocene); Pfeil 1984, ?Waitakian (latest Oligocene-earliest Miocene). (Romer 1966 cites "Upper Cretaceous")

Family Hexanchidae - cow sharks, six gill sharks
Hexanchus; Romer 1966, "Upper Cretaceous” (includes in part Notidanus of Chapman 19187).

Notorynchus; Chapman 1918 ("Notidanus"); Kemp 1978,"Miocene”; Keyes in Wiffen 1981a, Piripauan or Haumurian (Late
Cretaceous); Keyes 1981a, Teurian (Paleocene); Keyes 1987, Bortonian-Runangan (Middle-Late Eocene).

Family Carcharhinidae [= Carchariidae] - requiem sharks
Galeocerdo, Davis 1888b, Chapman 1918; Oligocene, Miocene, Pliocene (I.W. Keyes records, pers. comm.)

Scapanorhynchus; Davis 1888, Chapman 1918, "Upper Cretaceous".

Family Lamnidae [= Isuridae] - mackeral sharks

Carcharodon (sensu lato); Keyes 1971, 1972, 1975, 1987, various stages from Porangan to Castlecliffian (Middle Eocene-
Late Pleistocene); Pfeil 1984, ?7Waitakian (latest Oligocene-earliest Miocene).

Tsurus; Chapman 1918, "Cretaceous, Miocene" (Romer 1966 cites "Pliocene"); Pfeil 1984, ?Waitakian (latest Oligocene-
earliest Miocene).

Lamna; Chapman 1918, “Cretaceous, Eocene, Miocene"; Keyes 1987, Mangaorapan-Runangan (Early-Late Eocene).

Family Scyliorhinidae - catsharks

Megascyliorhinus, Keyes 1984, Whaingaroan (Early Oligocene)-Recent; Pfeil 1984, ?Waitakian (latest Oligocene-earliest
Miocene)

Family Squalidae - spiny dogfishes (includes Dalatlidae - spineless dogfishes of some authors)

Centrophorus; Keyes 1981a, 1984, ?Haumurian (Late Cretaceous), Teurian (Paleocene), Bortonian (Middle or Late Eocene)
sporadically to Recent, Pfeil 1984, ?Waitakian (latest Oligocene-earliest Miocene)

Scymnorhinus [= Dalatias]; Keyes 1984, Bortonian (?Middle or Late Eocene) sporadically to Recent; Pfeil 1984,
?Waitakian (latest Oligocene-earliest Miocene)

Squalus; Keyes 1981a, Teurian (Paleocene); Keyes in Wiffen 1981a and in Craw & Watt 1987, Piripauan or Haumurian
(Late Cretaceous).

Family Echinorhinidae - bramble sharks

Pseudoechinorhinus Pfeil 1983; Teurian (Paleocene)

FOSSIL VERTEBRATES OF NEW ZEALAND - 1297

Family Pristiophoridae (see Keyes 1979 for discussion)

aa apace ch 1979, Bortonian-Nukumaman (Late Eocene-Late Pliocene; full range of stages shown by Keyes 1982:
ig.

Pristiophorus; Chapman 1918, "Miocene, Pliocene"; Keyes 1987, Mangaorapan-Runangan (Early-Late Eocene); Pfeil
1984, ?Waitakian (latest Oligocene-earliest Miocene). (Report by Keyes 1981a of Pristiophorus from Paleocene is
erroneous; Keyes 1982)

Family Squatinidae
Squatina; Keyes 1981a, Teurian (Paleocene)
ORDER BATOIDEA
Family Sclerorhynchidae
Onchopristis; Keyes 1977, Piripauan or Haumurian (Late Cretaceous)

Family Dasyatidae

Dasyatis; Pfeil 1984, ?)Waitakian (latest Oligocene-earliest Miocene). Unpublished records are known, I.W. Keyes pers
comm., e.g. Altonian (Early Miocene), based on OUDG collections.

Family Myliobatidae - eagle rays
Myliobatis; Davis 1888a, Chapman 1918, “Tertiary, Miocene”; Hector 1881b. Unpublished records include Bortonian
(Middle? or Late Eocene), Otaian and Altonian (both Early Miocene), e.g. OUDG collections. On ray trace fossils,
see Gregory et al. 1979, 1983.
ORDER CHIMAERIFORMES
Family Chimaeridae - ghost sharks
Chimaera; Romer 1966, "Tertiary". Original source not found.
Ischyodus, Newton 1876, "Cretaceous"; Keyes in Wiffen 1981a, Piripauan or Haumurian (Late Cretaceous);

Family Callorhinchidae - elephant fishes

Callorhynchus; Newton 1876, "Cretaceous"; Keyes in Wiffen 1981a, Piripauan or Haumurian (Late Cretaceous); Keyes
1981a, Teurian (Paleocene); Wangaloan (latest Cretaceous or earliest Paleocene), based on OUDG collections.

Class OSTEICHTHYES
ORDER ELOPIFORMES

Family Elopidae
Elops [? = Chloropthalmus; Schwarzhans 1984: 47, 218]; Frost 1924, 1928, "Oligocene or Miocene”
Pterothrissus, Schwarzhans 1984, Mangaorapan - Heretaungan (Early-Middle Eocene)
Family Pachyrhizodontidae
Pachyrhizodus, Wiffen 1983, Piripauan or Haumurian (Late Cretaceous)
Thrissopater?; Chapman 1918, "Cretaceous"
ORDER ANGUILLIFORMES
Genus uncertain; includes otoliths in OUGD collections, Altonian (Early Miocene).
Family Heterenchelyidae

Heterenchelys; Stinton, 1957, Lillbumian-Waiauan (Middle-Late Miocene). (Schwarzhans 1984 placed material in
Gnathophis.)

Heterenchelyidarum;, Grenfell 1984, Otaian-Altonian (Early Miocene)

Family Congridae - conger eels
Conger; Romer 1966, "Miocene" (possibly specimens described by Frost 1933)
Congridarum; Grenfell 1984, Otaian-Altonian (Early Miocene); Schwarzhans 1984

Gnathophis; Grenfell 1981 (but not cited by Grenfell 1984); Schwarzhans1984, Manzaorapan - Heretauangan (Early
Eocene) sporadically to Opoitian (Pliocene)

1298 - FORDYCE

Maxwelliella Schwarzhans, 1980 (1984); Mangaorapan/Heretauangan (Early Eocene)
Pseudoxenomystax, Schwarzhans1984, Opoitian (Pliocene)
Scalanago; Schwarzhans1984, Mangaorapan - Heretauangan (Early Eocene)

Genus indet.; Schwarzhans 1984, Kaiatan (Late Eocene), Waitakian (latest Oligocene-earliest Miocene), Clifdenian and
Lillbumian (Middle Miocene)

Family Ophichthidae - snake eels

Mystriophis?; Stinton 1957, Clifdenian (Middle Miocene)

ORDER CLUPEIFORMES

Family Clupeidae - herrings
Anchoa; Schwarzhans1984, Altonian (Early Miocene)
Diplomystus, Chapman 1918, “Cretaceous”
Scombroclupea; Chapman 1918, "Eocene"

Genus indet.; Schwarzhans 1984, Altonian (Early Miocene)

ORDER SILURIFORMES
Family Ariidae
Tachysurus, Schwarzhans1984, Opoitian (Pliocene)

ORDER OSTEOGLOSSIFORMES
Family Ichthyodectidae

Xiphactinus [= Portheus; Romer 1966}; Chapman 1934, "Upper Cretaceous"

ORDER SALMONIFORMES
Family Argentinidae

Argentina; Schwarzhans1984, Waitakian (latest Oligocene), Altonian , (Early Miocene)
Family Galaxiidae - whitebaits

Galaxias; McDowall 1976, Miocene; Pliocene records unsubstantiated
Family Photichthyidae - lighthouse fishes

Polymetme; Schwarzhans1984, Altonian (Early Miocene)
Family Gonostomatidae

Gonostoma; Schwarzhans1984 [as Gonostomatidae], Altonian (Early Miocene)
Family Auloplidae

Aulopus, Schwarzhans1984, Mangaorapan - Heretauangan (Early Eocene)
Family Myctophidae - lantern fishes

Benthosema; Grenfell 1981, Otaian (Early Miocene) (but not cited by Grenfell 1984); Schwarzhans 1984, various stages
from Altonian (Early Miocene) to Tongaporutuan (latest Miocene)

Bolinichthys; Schwarzhans1984, Lillbumian (Middle Miocene)

Diaphus; Grenfell 1984, Otaian (Early Miocene); Schwarzhans 1984, various stages from Whaingaroan (Early Oligocene)
to Tongaporutuan (latest Miocene)

Diogenichthys; Schwarzhans1984, Altonian (Early Miocene)

Hygophum; Grenfell 1984, Otaian-Altonian (Early Miocene); Schwarzhans 1984, Altonian (Early Miocene),
Tongaporutuan (latest Miocene)

Lampanyctodes; Grenfell 1984, Otaian (Early Miocene); Schwarzhans 1984, Clifdenian (Middle Miocene), Tongaporutuan
(latest Miocene), Opoitian (Pliocene)

FOSSIL VERTEBRATES OF NEW ZEALAND - 1299

Lampanyctus; Stinton 1957, Lillbumian-Waiauan, Tongaporutuan (Middle-Late Miocene) (identification = Lampanyctodes,
fide Schwarzhans 1984: 62)

Lampichthys, Schwarzhans 1984, Tongaporutuan (latest Miocene)
Myctophidarum, Grenfell 1984, Otaian-Altonian (Early Miocene)

Myctophum [= Scolepus; Romer 1966]; Frost 1928, 1933, Stinton 1957, Clifdenian (Middle Miocene) (see comments by
Schwarzhans 1984: 59)

Notoscopelus; Schwarzhans 1984, Altonian (Early Miocene), Tongaporutuan (latest Miocene), Opoitian (Pliocene)

Symbolophorus; Grenfell 1984, Otaian-Altonian (Early Miocene); Schwarzhans 1984, Tongaporutuan (latest Miocene),
Opoitian (Pliocene)

Family Sternoptychidae - hatchet fishes
Maurolicus; Grenfell 1981, 1984, Schwarzhans1984, Otaian-Altonian (Early Miocene)
Polyipnus, Grenfell 1984, Otaian-Altonian (Early Miocene)

Family Chlorophthalmidae - greeneyes

Chlorophthalmus; Grenfell 1984, Otaian-Altonian (Early Miocene); Schwarzhans 1984, Kaiatan (Late Eocene),
Duntroonian (Late Oligocene), Waitakian (latest Oligocene-earliest Miocene), Altonian (Early Miocene)

Family Scopelarchidae - pearleyes [Schwarzhans 1984 used Scopelosauridae]

Scopelarchus; Grenfell 1984, Otaian (Early Miocene); Schwarzhans 1984, Kaiatan (Late Eocene)

ORDER LOPHIIFORMES
Family Lophiidae
?Lophius; Grenfell 1984, Altonian (Early Miocene)

ORDER GADIFORMES

Family Moridae - morid cods (sensu lato, as used by Romer 1966; see Karrer 1971. See Schwarzhans
1984 for alternative ordinal position.)

Actuariolum; Karrer 1971; Grenfell 1984, Otaian-Altonian (Early Miocene); Schwarzhans 1984, Waitakian (latest
Oligocene-earliest Miocene), Altonian (Early Miocene), Clifdenian (Middle Miocene)

Austrophycis; Schwarzhans 1984, Altonian (Early Miocene)
Laemonema; Schwarzhans 1984, Lillbumian (Middle Miocene)
Lotella; Grenfell 1984, Altonian (Early Miocene)
Physiculus, Grenfell 1984, ?Otaian, Altonian (Early Miocene)
Raniceps; Frost 1924, “Miocene” [? = Euclichcthys, fide Schwarzhans 1984: 217]
Tripterophycis; Schwarzhans 1984, Kaiatan (Late Eocene)
Family Melanonidae - pelagic cods

Karrerichthys Schwarzhans, 1984 (= Melanonidarum), Grenfell 1984, Otaian (Early Miocene); Schwarzhans 1984,
Tongaporutuan (latest Miocene)

Family Bregmacerotidae

Bregmaceros, Romer 1966, "?Eocene"; Grenfell 1984, Otaian-Altonian (Early Miocene); Schwarzhans 1984, Kaiatan (Late
Eocene), Clifdenian (Middle Miocene)

Family Bythididae - cuskeels
Oligopus; Grenfell 1984, Otaian-Altonian (Early Miocene)
Bythitidarum, Grenfell 1984, Otaian (Early Miocene)
Saccogaster, Schwarzhans 1984, Opoitian (Pliocene)
Genus indet.; Schwarzhans 1984, Clifdenian (Middle Miocene)

1300 - FORDYCE

Family Euclichthyidae

Euclichthys; Grenfell 1984, Otaian-Altonian (Early Miocene); Schwarzhans 1984, Kaiatan (Late Eocene), Altonian (Early
Miocene)

Family Gadidae (= Family incertae sedis of Grenfell 1984)
Gadiculus, Schwarzhans 1984, Altonian (Early Miocene)
Gadus; Frost 1924, "Miocene"

Macruronus; Schwarzhans 1984, Duntroonian (Late Oligocene), Altonian (Early Miocene), Tongaporutuan (latest
Miocene), Opoitian (Pliocene)

Merluccius; Frost 192A, "Miocene" [? = Macruronus, fide Schwarzhans 1984: 218]
Micromesistius; Schwarzhans 1984, Opoitian (Pliocene)
Family Carapidae - pearlfishes [= Ophidiiformes, fide Schwarzhans 1984]

Carapus [= Fierasfer, = Jordanicus; Romer 1966]; Frost 1924, Stinton 1957, Clifdenian (Middle Miocene); Grenfell 1984,
Otaian-Altonian (Early Miocene); Schwarzhans 1984, Waitakian (latest Oligocene-earliest Miocene), Altonian (Early
Miocene)

Family Macrouridae - rattails, grenadiers [= Coryphaenidae; Romer 1966]
Bathygadus, Grenfell 1984, Otaian (Early Miocene)

Coelorinchus, Grenfell 1984, Otaian-Altonian (Early Miocene); Stinton 1957, Clifdenian and Waitotaran (Middle Miocene
and Late Pliocene); Schwarzhans 1984, various stages from Duntroonian (Late Oligocene) to Waitotaran (Pliocene).
Romer 1966 included this genus in Scombridae but cited no New Zealand record)

?Hymenocephalus, Grenfell 1981 (but not cited by Grenfell 1984)

Lepidorhynchus, Grenfell 1984, Otaian-Altonian (Early Miocene); Schwarzhans 1984, Tongaporutuan (latest Miocene),
Opoitian (Pliocene)

Macrouridarum, Grenfell 1984, Otaian (Early Miocene)
Macrourus; Frost 1924, 1928, 1933, "Miocene"
Macrurulus Schwarzhans, 1980; Schwarzhans1984, Mangaorapan - Heretauangan (Early Eocene), Bortonian (Late Eocene)
Maorigadus Schwarzhans, 1980, Schwarzhans1984, Mangaorapan - Heretauangan (Early Eocene)
Nezumia,; Schwarzhans 1984, Tongaporutuan (latest Miocene)
Trachyrincus, Grenfell 1984, Otaian (Early Miocene); Stinton 1957 and Schwarzhans1984, Tongaporutuan (Late Miocene)
Ventrifossa; Schwarzhans1984, Altonian (Early Miocene), Tongaporutuan (Late Miocene)
Family Ophiidae - lings [Ophidiiformes, fide Schwarzhans1984]
Ampheristus; Schwarzhans1984, Mangaorapan - Heretauangan (Early Eocene)
Genypterus,; Schwarzhans1984, Opoitian (Pliocene)
Monomitopus, Schwarzhans1984, Kaiatan (Late Eocene), Tongaporutuan (Late Miocene)
Neobythites,; Schwarzhans1984, Waiauan (Middle Miocene)
Nolfophidion; Schwarzhans1984, Bortonian (Late Eocene)
Ophidium; Frost 1924, "Miocene-Pliocene"
Ophidiidarum; Frost 1924, "Miocene"
Sirembinorum, Grenfell 1984 (as Sirembo in Grenfell 1981), Altonian (Early Miocene)
Genus indet.; Schwarzhans 1984, Lillbumian (Middle Miocene)

ORDER LOPHITFORMES
Family Ogcocephalidae
2Dibranchus; Schwarzhans1984, Kaiatan (Late Eocene)

FOSSIL VERTEBRATES OF NEW ZEALAND - 1301

ORDER BERYCIFORMES

Family Melamphaeidae - bigscale fishes
Melamphaes; Grenfell 1984, Otaian (Early Miocene); Schwarzhans1984, Tongaporutuan (Late Miocene)
?Scolpeloberyx;, Schwarzhans1984, Tongaporutuan (Late Miocene)

Family Trachichthyidae - rougheyes
Egregioberyx Schwarzhans,1980; Schwarzhans1984, Bortonian (Late Eocene)

Hoplostethus; Grenfell 1984, Otaian-Altonian (Early Miocene); Schwarzhans1984, Tongaporutuan (latest Miocene),
Opoitian (Pliocene)

Paratrachichthys; Grenfell 1984, Altonian (Early Miocene); Schwarzhans1984, Waiauan (Middle Miocene)
Trachichthodes, Schwarzhans1984, Kaiatan (Late Eocene)
Family Berycidae - alfonsinos
Centroberyx; Grenfell 1984 (as Trachichthodes, Grenfell 1981), Otaian-Altonian (Early Miocene)
Family Holocentridae
Adioryx; Schwarzhans1984, Mangaorapan - Heretauangan (Early Eocene)
Genus indet.; Schwarzhans 1984, Bortonian (Late Eocene)
ORDER ZEIFORMES
Family Caproidae
Antigonia; Schwarzhans1984, Bortonian (Middle Eocene)
ORDER INCERTAE SEDIS [SEE COMMENTS BY SCHWARZHANS1984]
Family Platycephalidae

Platycephalus; Schwarzhans1984, Bortonian (Middle Eocene), Duntroonian and Waitakian (Late Oligocene-earliest
Miocene)

Family Mugiloididae

Parapercis; Schwarzhans 1984, Duntroonian, Waitakian (Late Oligocene-earliest Miocene), Altonian (Early Miocene),
Clifdenian (Middle Miocene) (includes Cottus of Frost)

Family Uranoscopidae
Uranoscopus, Schwarzhans1984, Bortonian (Middle Eocene)
Family Leptoscopidae

Leptoscopus; Schwarzhans 1984, Waitakian (latest Oligocene-earliest Miocene), Altonian (Early Miocene), Tongaporutuan
(latest Miocene)

Family Hemerocoetidae
Hemerocoetus, Schwarzhans 1984, Altonian (Early Miocene), Opoitian (Pliocene)
Krebsiella Schwarzhans, 1980; Schwarzhans1984, Bortonian (Late Eocene), Waitakian (latest Oligocene-earliest Miocene)
Waitakia Schwarzhans, 1980; Schwarzhans1984, Duntroonian (Late Oligocene), Altonian (Early Miocene)

ORDER SCORPAENIFORMES

Family Cottidae - sculpins
Cottus, Frost 1928, "Oligocene or Miocene"

2Cottoideorum; Grenfell 1981, ? = ?Cotteidei genus indet. of Grenfell 1984, Otaian and/or Genus indet.; Schwarzhans
1984,

Cottidarum; Grenfell 1981, ? = ?Cotteidei genus indet. of Grenfell 1984, Otaian and/or Altonian (Early Miocene)
Genus indet.; Schwarzhans 1984, Kaiatan (Late Eocene), Altonian (Early Miocene)

Family Cyclopteridae
Cyclopterus; Frost 1933, "Miocene"

1302 - FORDYCE

Family Hoplichthyidae - ghost flatheads
Hoplichthys; Grenfell 1984, Otaian-Altonian (Early Miocene)
Praehoplichthys Schwarzhans, 1980; Schwarzhans 1984, Bortonian (Middle Eocene), Kaiatan (Late Eocene)

Family Trichodontidae [as placed by Schwarzhans 1984; relationship to Trachinidae, below,
uncertain.]

Trichodon; Schwarzhans 1984, Altonian (Early Miocene)

ORDER PERCIFORMES
Family Percidae
Percaletes; Romer 1966, "Miocene". Original source not found.
Family Ambassidae
Dapalis,; Schwarzhans 1984, Mangaorapan - Heretaungan, Bortonian (Early-Middle Eocene)
Family Sparidae - breams
Dentex; Frost 1924, 1928, "Miocene"; Schwarzhans 1984, Mangaorapan- Heretaungan (Early-Middle Eocene)
Pagellus; Frost 1928, "Miocene"
Sargus; Chapman 1918, "Miocene"
Family Serranidae - groupers, sea perches
Serranus; Frost 1924, “Miocene”
Genus indet.; Schwarzhans 1984, Bortonian (Middle Eocene), Altonian (Early Miocene)
Family Labridae - wrasses
Labrodon; Chapman 1918, "Miocene" (Paleocene-Pliocene, according to Romer 1966)
Labridarum (as Pseudolabrus; Grenfell 1981); Grenfell 1984, Otaian (Early Miocene)
Family Nototheniidae - icefishes

Notothenia; Stinton 1957, Lillburnian-Waiauan (Middle-Late Miocene) (Misidentification, according to W. Schwarzhans,
pers. comm. and Schwarzhans 1984: 14)

Family Trachinidae
Trachinus; Frost 1924, "Miocene"
Family Gempylidae - snake mackerels
Eothyrsites; Chapman 1934, "Oligocene"

Family Epigonidae [ = Apogonidae, cardinalfishes, fide Ayling & Cox (1982); = Lactariidae in part,
fide Schwarzhans 1984: 192.]

Epigonus; Grenfell 1984, Altonian (Early Miocene); Schwarzhans 1984, Clifdenian (Middle Miocene)
Family Apogonidae

Apogonidarum, Grenfell 1984, Altonian (Early Miocene)
Family Acropomatidae [ = Lactariidae in part, fide Schwarzhans 1984: 192.]

Acropoma; Grenfell 1984, Otaian-Altonian (Early Miocene); Schwarzhans 1984, Duntroonian and Waitakian (Late
Oligocene-earliest Miocene), Altonian (Early Miocene), Clifdenian (Middle Miocene)

Genus indet.; Schwarzhans 1984, Kaiatan (Late Eocene)

Family Lactariidae [includes Acropomatidae, Epigonidae, Scombropidae in part; Schwarzhans 1984:
192.)

Lactarius, Grenfell 1984, Otaian-Altonian (Early Miocene); Schwarzhans 1984, Duntroonian and Waitakian (Late
Oligocene-earliest Miocene), Altonian (Early Miocene), Clifdenian (Middle Miocene), Opoitian (Pliocene)

Neoscombrops; Schwarzhans 1984, Mangaorapan - Heretauangan (Early Eocene)

FOSSIL VERTEBRATES OF NEW ZEALAND - 1303

Paralactarius Schwarzhans, 1980; Schwarzhans 1984, Mangaorapan - Heretauangan (Early Eocene), Kaiatan (Late Eocene),
Whaingaroan (Early Oligocene), Waitakian (latest Oligocene-earliest Miocene)

Family Sillaginidae

Sillago; Schwarzhans 1984, Bortonian (Middle Eocene), Altonian (Early Miocene)
Family Gerreidae

?Gerreidarum; Grenfell 1984, Altonian (Early Miocene)
Family Pomadasyidae

Pomadasyidarum; Grenfell 1984, Early Miocene

Genus indet.; Schwarzhans 1984, Mangaorapan - Heretaungan, (Early Eocene) Waitakian (latest Oligocene-earliest
Miocene), Altonian (Early Miocene)

Family Cepolidae - bandfishes
Cepola; Grenfell 1984, Altonian (Early Miocene)
Family Mugiloididae - weevers, grubfish
Parapercis; Frost 1924, "Oligocene"; Grenfell 1984, Otaian-Altonian (Early Miocene)
Genus indet.; Schwarzhans 1984, Bortonian (Middle Eocene)
Family Scombridae
Genus indet.; Schwarzhans 1984, Altonian (Early Miocene)
Family Gobiidae - gobies
Gobiidarum (?as Gobius; Grenfell 1981); Grenfell 1984, Altonian (Early Miocene)
Family Eleotrididae - gudgeons
Gobiomorphus, Oliver 1928, Castlecliffian (Late Pleistocene)
Paradiplospinus; Grenfell 1984, Otaian-Altonian (Early Miocene)
Family incertae sedis

Percidarum; Stinton 1957, Tongaporutuan (Late Miocene)

ORDER PLEURONECTIFORMES
Family incertae sedis

Genus uncertain; includes skeletal material in NZGS collections, age uncertain.
Family Bothidae - lefteye flounders

Arnoglossus;, Grenfell 1984, Otaian-Altonian (Early Miocene); Schwarzhans 1984, Kaiatan (Late Eocene), Waitakian
(latest Oligocene-earliest Miocene), Altonian (Early Miocene)

Genus indet.; Schwarzhans 1984, Altonian (Early Miocene)
Family Soleidae
Achirus, Schwarzhans 1984, Lillbumian/Waiauan (Middle Miocene)
Solea; Frost 1928, "Miocene"
Soleidarum; Grenfell 1981 (but not cited by Grenfell 1984)
Family Eucitharidae
Citharus; Frost 1928, "Miocene" [? = Trachiniodei indet. , fide Schwarzhans 1984: 218]

Pleuronectidarum; Frost 1924, "Miocene"

ORDER TETRAODONTIFORMES
Family Trigonodontidae

Trigonodon, Romer 1966, "Miocene". Original source not found.

1304 - FORDYCE

Incertae_ sedis

Sparidarum; Frost 1924, "Miocene"
ooo—————a—a—e—e——a OOOO
~~ ———[—EOSSEEEESEEEE——————E

Table 2. Fossil Reptilia recorded from New Zealand. Taxa listed here as incertae sedis will probably be placed firmly after
more detailed study, and it should not be assumed that currently uncertain relationships are irresolvable. Only the more
significant unpublished records are included. Repository cited only for specimens not previously mentioned in print.

Order CHELONIA

Chelonia Incertae sedis
Family, genus and species indet. (Fordyce 1980a), Teurian (Palaeocene), Ward.
Family, genus and species indet., 7Teurian (?Palaeocene). Chatham Island. See text. NZGS collections.
Family, genus and species indet., Bortonian or Kaiatan (Late Eocene), Boulder Hill. See text. OUGD collections.
Family, genus and species indet., ?7Kaiatan (Late Eocene), Woodpecker Bay. See text. OUGD collections.
Family, genus and species indet., Kaiatan (Late Eocene), Pahi. See text. AUGD collections.

Family, genus and species indet. (Fordyce 1980a, Marples 1949a); Runangan-Whaingaroan (latest Eocene - Early
Oligocene), Oamam.

Protostegidae

Genus and species indet. (Wiffen 1981); Piripauan or Haumurian (Late Cretaceous), Mangahouanga Stream.

Cheloniidae

"Lepidochelys" waikatoica Buckeridge 1981; Otaian (Early Miocene), Port Waikato.

Order SQUAMATA

Mosasauridae (all Late Cretaceous) (synonymies follow Welles & Gregg 1971)

Mosasaurus [= Moanasaurus?] mangahouangae (Wiffen, 1980). Holotype: skull, vertebrae, paddle bones; Piripauan or
Haumurian, Mangahouanga Stream. Keyes 1989, Wright 1989.

Mosasaurus mokoroa Welles & Gregg, 1971. Holotype: skull, vertebrae; Haumurian, Cheviot. Includes Taniwhasaurus
oweni Hector, 1874, in part, and Leiodon haumuriensis of Hutton 1895.

Prognathodon waiparaensis Welles & Gregg, 1971. Holotype: skull, vertebrae, ribs; Haumurian, Waipara.
Taniwhasaurus oweni Hector,1874. Lectotype: skull, jaws; Haumurian, Haumuri Bluff.

Tylosaurus haumuriensis Hector, 1874. Lectotype: skull and jaws; Haumurian, Haumouri Bluff.

Genus and species indet. of Welles & Gregg 1971; includes Mauisaurus haasti Hector, 1874 (in part), Leiodon

haumuriensis Hector, 1874 (in part), Taniwhasaurus oweni Hector, 1874 (in part); Haumurian, Waipara and Haumuri
Bluff. Undetermined material includes fossils from Shag Point, OUGD collections.

Order SPHENODONTA
Sphenodontidae
Sphenodon punctatus (Gray). Late Pleistocene to Recent, various localities; Rich et al, 1979.
Order SAURISCHIA

Theropoda incertae sedis

Theropoda genus and species indet. (Molnar 1981); Piripauan or Haumurian (Late Cretaceous), Mangahouanga Stream.

FOSSIL VERTEBRATES OF NEW ZEALAND - 1305
Order ORNITHISCHIA

Ornithopoda incertae sedis

Omithopoda genus and species indet. (Wiffen & Molnar in press); Piripauan or Haumurian (Late Cretaceous),
Mangahouanga Stream.

Order SAUROPTERYGIA

Plesiosauroidea (synonymies follow Welles & Gregg 1971; see Brown 1981 for altemative classification.)
Elasmosauridae

Mauisaurus haasti Hector, 1874. Lectotype: pelvis, paddle; Haumurian (Late Cretaceous), Jed River near Gore Bay.
Includes Plesiosaurus australis of Hector 1874 (in part), Mauisaurus brachiolatus Hector, 1874, Cimoliosaurus caudalis
Hutton, 1894 and Mangahouanga Stream specimens discussed by Wiffen & Moisley 1986.

Tuarangisaurus keyesi Wiffen & Moisley, 1986. Holotype: skull and apparently associated vertebrae; Piripauan or
Haumunan (Late Cretaceous), Mangahouanga Stream.

Genus and species indet. of Welles & Gregg 1971; Haumurian (Late Cretaceous), Waipara and Haumuri Bluff. Includes
Plesiosaurus australis Owen, 1861 (in part), Plesiosaurus crassicostatus Owen, 1870b (in part), Plesiosaurus hoodi
Owen, 1870b, Plesiosaurus holmesi Hector, 1874 (in part), Plesiosaurus mackayi Hector, 1874, and Mauisaurus haasti
Hector, 1874 (in part). Undetermined material includes fossils from Shag Point, OUGD collections; Figs. 11, 12.

Plesiosauroidea: Polycotylidae

Genus and species indet. of Welles & Gregg 1971; Haumurian (Late Cretaceous), Waipara and Haumuri Bluff. Includes
Plesiosaurus australis Owen, 1861, Plesiosaurus crassicostatus Owen, 1870b (in part), Plesiosaurus holmesi Hector,
1874 (in part), Plesiosaurus traversi Hector, 1874, and Polycotylus tenuis Hector, 1874 (in part).

Plesiosauria incertae sedis

Genus and species indet. of Welles & Gregg 1971; Haumurian (Late Cretaceous), Waipara and Haumuri Bluff. Includes
Polycotylus tenuis Hector, 1874 (in part), and Mauisaurus haasti Hector, 1874 (in part).

Genus and species indet. of Campbell 1965; Oretian (Late Triassic), Marakopa. Dubious identification; ordinal position
uncertain.

Order ICHTHYOSAURIA
Ichthyosauria incertae sedis
Genus and species indet.; Etalian (Middle Triassic), Etal Stream. See text. OUGD collections.
Genus and species indet. of Campbell 1965; Otamitan (Late Triassic), Otamita Stream. (Shastasauridae?).
Genus and species indet., Otamitan (Late Triassic), Nugget Point. See text. OUGD collections.

Genus and species indet. of Fleming et al. 1971; Oretian (Late Triassic), Mt Potts. Includes Ichthyosaurus australis
Hector, 1874, Ichthyosaurus pottsi Hector, 1886, and Mixosaurus (2) hectori Lydekker, 1889.

Genus and species indet. of Fleming ef al. 1971, Motuan (Early Cretaceous), Tinui (possibly Stenopterygiidae).
Genus and species indet.; ?Late Cretaceous, Dargaville. See text. AUGD collections.
———— SS _
—S ____ eee

Table 3. Summary of species of moas (Dinomithiformes) recognised by Cracraft (1976a), Millener (1982), and Worthy
(1987d, 1988a). See text for other recent references.

Emeldae [= Anomalopterygidae = Dinornithidae: Anomalopteryginae]

Anomalopteryx Reichenbach, 1852.
A. didiformis (Owen, 1844a). Synonyms: A. parvus (Owen, 1883a), A. antiquus (Hutton, 1892), A. oweni (Haast, 1886b).
See Millener 1982.
Megalapteryx Haast, 1886a.
M. didinus (Owen, 1883b). Synonym: M. hectori Haast, 1886a, M. benhami Archey, 1941; see Haast 1884, Worthy
1988a.

Pachyornis Lydekker, 1891.

1306 - FORDYCE

P. australis Oliver, 1949. See Worthy 1989e.
P. elephantopus (Owen, 1856). Synonym: P. murihiku Oliver, 1949; see Scarlett 1968b, Worthy 1989e.
P. mappini Archey, 1941. Synonym: P. septentrionalis Oliver, 1949; see Worthy 1987d.
Eurapteryx Haast, 1874b. Synonym: Zelornis Oliver, 1949.
E. curtus (Owen, 1846). Synonyms: E. exilis Hutton, 1897, E. tane Oliver, 1949.
E. geranoides (Owen, 1848). Synonyms: E. gravis (Owen, 1870a), Zelornis haasti (Rothschild, 1907).
Emeus Reichenbach, 1852.
E. crassus (Owen, 1846). Synonym: E. huttonii (Owen, 1879).
Dinornithidae [= Dinornithinae of some authors]
Dinornis Owen, 1843.
D. struthoides Owen, 1844a. Synonyms: D. gazella Oliver, 1949; D. torosus Hutton, 1891.

D. novaezealandiae Owen, 1843. Synonyms: D. ingens Owen, 1844b, D. robustus Owen, 1846, D. hercules Oliver,
1949.

D. giganteus Owen, 1844a. Synonym: D. maximus Owen, 1867 [= 1866].

Table 4. Guide to species of Late Quatemary birds recorded from New Zealand, other than moas (Dinomithiformes) and
penguins (Sphenisciformes). Full authorships are not cited for extant species. Based on Brodkorb (1963, 1964,
1967, 1971, 1978), Scarlett in Fordyce 1982c and the limited selection of references cited; this is not intended to be
a comprehensive guide to literature. Recent literature is mentioned in the text.

APTERYGIFORMES - kiwis

Apterygidae

Apteryx australis Shaw; Scarlett 195Sa.

Apteryx haastii Potts; Lydekker 1891.

Apteryx owenii Gould; Scarlett 1962.

Pseudapteryx gracilis Lydekker 1891.

PODICIPEDIFORMES - grebes

Podicipedidae

Podiceps rufopectus (Gray); Scarlett 1955a, 1969b.

Podiceps cristatus (Linnaeus); Hom 1983.

PROCELLARIIFORMES - albatrosses, petrels

Diomedeidae (see Boume 1974, as well as references below).
Diomedea exulans Linnaeus; Harrison & Walker 1978.
Diomedea chlororhynchus Gmelin; Lydekker 1891.

Procellariidae (see Bourne 1974, as well as references below).
Pterodroma hypoleuca (Salvin); Millener 1980b.

Pterodroma inexpectata (Forster); Worthy & Mildenhall 1989.
Pterodroma cookii (Gray); Worthy & Mildenhall 1989.
Macronectes giganteus (Gmelin); Boume 1974, Lydekker 1891.
Puffinus tenuirostris (Temminck); Scarlett 1976a.

Puffinus cf. P. assimilis Gould; Millener 1980b.

FOSSIL VERTEBRATES OF NEW ZEALAND - 1307

Puffinus gavia (Forster); Grant-Mackie & Scarlett 1973, Millener 1980b, Worthy & Mildenhall 1989.
Puffinus griseus (Gmelin); Grant-Mackie & Scarlett 1973.

Pachyptila turtur (Kuhl); Millener 1980b, Worthy & Mildenhall 1989.

Pachyptila sp., Grant-Mackie & Scarlett 1973.

Pelecanoides urinatrix (Gmelin); Worthy & Mildenhall 1989.

Pelecanoides sp.; Rich et al. 1979.

Hydrobatidae

Pelagodroma marina (Latham); Millener 1980b.

PELECANIFORMES .- pelicans

Pelecanidae
Pelicanus novaezealandiae Scarlett, 1966 (as P. conspicillatus novaezealandiae); Archey 1931, Rich & Van Tets 1981.

Phalacrocoracidae
Phalacrocorax carbo (Linnaeus); Worthy & Mildenhall 1989.

ANSERIFORMES - swans, ducks, geese

Anatidae

Anas aucklandica Gray; Worthy & Mildenhall 1989.

Anas chlorotis Gray; Scarlett 1969b.

Anas gibberifrons Muller; Scarlett 1969b.

Anas superciliosa Gmelin; Grant-Mackie & Scarlett 1973, Millener 1980b.

Anas sp., Grant-Mackie & Scarlett 1973.

Aythya novaeseelandiae (Gmelin); Lydekker 1891.

Biziura lobata (Shaw). Synonyms: Biziura delatouri Forbes, 1892a - see Howard 1964; Olson 1977b, Scarlett 1969a.

Cereopsis sp., Rich 197Sa: 101.

Chenonetta jubata (Latham); Lydekker 1891.

Cnemiornis calcitrans Owen, 1865. Synonyms: Cereopsis novaezealandiae Forbes, 1891b, and Cnemiornis minor
Forbes, 1891d; see Brodkorb 1964: 218; Worthy & Mildenhall 1989.

Cnemiornis septentrionalis Oliver, 1955.

Cygnus atratus (Latham); Dawson, 1958.

Cygnus sumnerensis (Forbes, 1892e). Synonym: Cygnus chathamicus Oliver 1955; see Brodkorb 1964: 210.

Euryanas finschi (Van Beneden, 1875); Van Beneden 1876, Scarlett 1969b, Worthy 1988b, Worthy & Mildenhall 1989.
Hymenolaimus malacorhynchus (Gmelin); Worthy & Mildenhall 1989.

Malacorhynchus scarletti Olson, 1977b; equals Malacorhynchus membranaceus (Latham) of Oliver 1955, Scarlett 1969a,
and others.

Mergus australis Hombron & Jaquinot,; Kear & Scarlett 1970, Olson 1977b.

Oxyura australis Gould; Horm 1983.

Oxyura cf. O. australis Gould; Millener 1979.

Pachyanas chathamica Oliver, 1955.

Tadorna variegata (Gmelin); Scarlett 1969b, Grant-Mackie & Scarlett 1973.
FALCONIFORMES - birds of prey

Accipitridae

1308 - FORDYCE

Circus approximans Peale; Scarlett 1955a.
Circus eylesi Scarlett, 1953; Scarlett 1969b, Worthy & Mildenhall 1989.
Circus teauteensis Forbes, 1892e. Synonym: Circus hamiltoni Forbes, 1892e; see Brodkorb 1964: 279.

Harpagornis moorei Haast, 1872. Synonym: Harpagornis assimilis Haast, 1874a; see Brodkorb 1964: 273; Hector 1875,
Oliver 1955, Worthy & Mildenhall 1989.

Haliaeetus australis (Harrison & Walker, 1973). Dawson 1961, Olson 1984.

Falconidae

Falco novaeseelandiae Gmelin; Scarlett 1955a, Worthy & Mildenhall 1989, Yaldwyn 1956.

GALLIFORMES - game birds, fowl-like birds

Phasianidae

Coturnix novaezealandiae Quoy & Gaimard; Scarlett 1969b, Grant-Mackie & Scarlett 1973, Worthy & Mildenhall 1989.

GRUIFORMES - cranes and rails.

Rallidae (based on Olson 1975a, 1975b, 1977a; but see Scarlett 1976b).

Gallinula (Tribonyx) hodgeni (Scarlett, 1955b). Synonym: Gallirallus hartreei Scarlett, 1970a; Scarlett 1976b. See also
Olson 1976, 1986, Worthy & Mildenhall 1989.

Capellirallus karamu Falla, 1954; Scarlett 1970b.

Fulica chathamensis Forbes, 1892f (includes F. prisca Hamilton, 1893, as a subspecies), see also Millener 1980a, 1981a,
Worthy & Mildenhall 1989.

Gallirallus australis (Sparrman); Brodkorb 1967: 133, Millener & Templer 1981, Worthy & Mildenhall 1989.
Gallirallus dieffenbachii (Gray); Forbes 1893a, Scarlett 1979.

Gallirallus modestus (Hutton); Forbes 1893a.

Gallirallus minor (Hamilton, 1893) (doubtfully distinct from the extant G. australis; Olson, 1977a); Rich et al. 1979.
Gallirallus insignis (Forbes, 1892).

Rallus philippensis Linnaeus; Worthy & Mildenhall 1989.

Diaphorapteryx hawkinsi (Forbes, 1892b); Andrews 1896, Forbes 1892f, 1893a.

Porphyrio mantelli (Owen); Beauchamp & Worthy 1988, Gurr 1952, Mills et al. 1984, 1988, Oliver 1955, Owen 1848a,
Parker 1882, Worthy & Mildenhall 1989.

Aptornithidae (see Oliver 1955, and Olson 1975, 1977a, 1985: 162 for status); Apteromithidae and Apertornis have
priority according to Olson 1985: 162. See Forbes 1890, Worthy 1989d.

Aptornis otidiformis (Owen, 1844b); Worthy & Mildenhall 1989.

A. defossor Owen, 1871; Brodkorb1967: 132; Olson 1985: 163; synonymous with Aptornis otidiformis (Owen, 1844b),
according to Kinsky 1970.

CHARADRIIFORMES - shorebirds

Haematopodidae
Haematopus unicolor Forster, Forbes 1893a.
Scolopacidae

Coenocorypha aucklandica (Gray); Hom 1983, Millener 1981b, Miskelly 1987, Worthy 1987e, Paulin 1973, Worthy &
Mildenhall 1989.

Coenocorypha chathamica (Forbes, 1893a).
Recurvirostridae

Himantopus himantopus (Linnaeus); Scarlett 1969b.

FOSSIL VERTEBRATES OF NEW ZEALAND - 1309

Laridae

Larus dominicanus Lichtenstein; Grant-Mackie & Scarlett 1973.
Larus sp., Grant-Mackie & Scarlett 1973.

Sternidae

Sterna striata (Gmelin); Grant-Mackie & Scarlett 1973.

COLUMBIFORMES - pigeons
Columbidae

Hemiphaga novaeseelandiae (Gmelin); Scarlett 1969b, Grant-Mackie & Scarlett 1973.
Hemiphaga sp., Grant-Mackie & Scarlett 1973.

PSITTACIFORMES - parrots and parakeets
Nestoridae

Nestor meridionalis (Gmelin); Oliver 1955, Grant-Mackie & Scarlett 1973, Scarlett 1969b.
Nestor notabilis Gould; Dawson 1952, Worthy & Mildenhall 1989
Nestor sp., Dawson 1952.
Strigops habroptilus Gray; Worthy & Mildenhall 1989.
Platycercidae
Cyanoramphus novaezelandiae (Sparrman); Scarlett 1955a, Yaldwyn 1956, Millener & Templer 1981.
Cyanoramphus sp., Grant-Mackie & Scarlett 1973, Worthy & Mildenhall 1989
Cacatuidae

Strigops habroptilus Gray; Scarlett 1955a, 1969b, Millener & Templer 1981.

CAPRIMULGIFORMES - nightjars and frogmouths
Aegothelidae

Megaegotheles novaezealandiae Scarlett,1968a; Rich & Scarlett 1977, Millener & Templer 1981.

STRIGIFORMES - owls
Strigidae
Ninox novaeseelandiae (Gmelin); Scarlett 1955a, 1969b.

Sceloglaux albifacies (Gray); Scarlett 1955a, 1969b, Grant-Mackie & Scarlett 1973, Millener 1983.

PASSERIFORMES - songbirds
Acanthisittidae

Acanthisitta chloris (Sparrman), Worthy & Mildenhall 1989.

Pachyplichas yaldwyni Millener, 1988.

Pachyplichas jagmi Millener, 1988.

Traversia lyalli (Rothschild); Worthy & Mildenhall 1989.

Xenicus longipes (Gmelin); Worthy & Mildenhall 1989.

Xenicus gilviventris Pelzeln, Worthy & Mildenhall 1989.
Motacillidae

Anthus novaeseelandiae (Gmelin); Worthy & Mildenhall 1989.
Corvidae

Palaeocorax moriorum Forbes, 1892f (Includes Corvus moriorum antipodum (Forbes, 1893a) as a subspecies; Brodkorb
1978: 160); Forbes 1892h, Grant-Mackie & Scarlett 1973, Scarlett 1972, Worthy & Mildenhall 1989.

1310 - FORDYCE

Paradisaeidae

Turnagra capensis (Sparrman); Scarlett 1955a, 1969b, Medway 1971, Worthy & Mildenhall 1989. See Brodkorb 1978:
171 and Olson et al. 1983 for systematic position.

Muscicapidae
Bowdleria punctata (Quoy & Gaimard); Millener 1980b.
Mohoua albicilla (Lesson); Medway 1971, Millener & Templer 1981, Worthy & Mildenhall 1989.
Mohoua novaeseelandiae (Gmelin); Worthy & Mildenhall 1989.
Mohoua ochrocephala (Gmelin); Worthy & Mildenhall 1989.
Gerygone igata (Quoy & Gaimard); Millener 1980b, Worthy & Mildenhall 1989.
Rhipidura fuliginosa (Sparrman); Millener & Temple 1981; Worthy & Mildenhall 1989.

Petroica australis (Sparrman); Scarlett 1955a, Yaldwyn 1956, Grant-Mackie & Scarlett 1973, Millener 1980b, Worthy &
Mildenhall 1989.

Petroica macrocephala (Sparrman); Worthy & Mildenhall 1989.
Callaeatidae

Creadion carunculatus (Gmelin); Medway 1971, Scarlett 1955a, Grant-Mackie & Scarlett 1973, Worthy & Mildenhall
1989.

Heteralocha acutirostris (Gould); Medway 1971, Williams 1973.

Callaeas cinerea (Gmelin); Scarlett 1955a, Medway 1967, 1971, Grant-Mackie & Scarlett 1973, Millener 1980b, Worthy
& Mildenhall 1989.

Meliphagidae
Anthornis melanura (Sparrman); Medway 1971, Millener & Templer 1981, Worthy & Mildenhall 1989.
Notiomystis cincta (du Bus); Millener & Templer 1981.

Prosthemadera novaeseelandiae (Gmelin); Scarlett 1955a, Yaldwyn 1956, Grant-Mackie & Scarlett 1973, Medway 1967,
1971, Worthy & Mildenhall 1989.

————————aaaeea

Table 5. Species of fossil penguin (Sphenisciformes: Spheniscidae) reported from New Zealand. Based partly on Simpson
(1971a, 1972b).

Palaeeudyptes Huxley, 1859a. Type-species: Palaeeudyptes antarcticus Huxley, 1859a. Nominal species also known from
Australia and Seymour Island.

Palaeeudyptes antarcticus Huxley, 1859a. Holotype: tarsometatarsus, stage uncertain, but possibly from remanié base of
Gee Greensand, Waitakian (latest Oligocene-earliest Miocene), Kakanui. Does not include specimens referred by
Hector (1872) and Marples (1952). See Fordyce & Jones 1987.

"Palaeeudyptes" marplesi Brodkorb, 1963, Holotype: wing and leg bones, vertebrae, Kaiatan or Runangan (Late Eocene),
Bumside, Dunedin,

"Palaeeudyptes" sp(p). indet. Includes specimens referred to P. antarcticus by Hector (1872) and Marples (1952).
Specimens: isolated bones of some individuals, associated limb bones of others, Runangan (Late Eocene) to
Duntroonian (Late Oligocene), possibly Kaiatan to Waitakian, from Seal Rock (North Westland), Bumside, Duntroon.

Pachydyptes Oliver, 1930. Type-species: Pachydyptes ponderosus Oliver, 1930.

Pachydyptes ponderosus Oliver, 1930. Holotype: wing and trunk bones, Runangan (Late Eocene), Oamaru. Includes
specimens identified as Palaeeudyptes antarcticus by Hector (1873) in part.

Platydyptes Marples, 1952. Type-species: Pachydyptes novaezealandiae Oliver, 1930. See Fordyce & Jones 1987.

Platydyptes novaezealandiae (Oliver, 1930). Holotype: wing bones and other fragments, Duntroonian or Waitakian (Late
Oligocene-earliest Miocene), Oamaru.

Platydyptes amiesi Marples, 1952. Holotype: wing bones, Duntroonian; possibly Waitakian (Late Oligocene-earliest
Miocene), Duntroon.

"Platydyptes" marplesi Simpson, 1971. Holotype: wing, leg and thorax bones, Duntroonian (Late Oligocene), probably
Wharekuri. Includes Platydyptes novaezealandiae of Marples (1952) in part.

FOSSIL VERTEBRATES OF NEW ZEALAND - 1311

Archaeospheniscus Marples, 1952. Type-species: Archaeospheniscus lowei Marples 1952. Species also are known from
Seymour Island.

amar a a lowei Marples, 1952. Holotype: wing, leg and a few trunk bones, Duntroonian (Late Oligocene),
uuntroon.

se aa lopdelli Marples, 1952. Holotype: mainly wing and leg bones, Duntroonian (Late Oligocene),
untroon.

Duntroonornis Marples, 1952. Type-species: Duntroonornis parvus Marples, 1952.
Duntroonornis parvus Marples, 1952. Holotype: tarsometatarsus, Duntroonian (Late Oligocene), Duntroon.
Korora Marples, 1952. Type-species: Korora oliveri Marples, 1952.

Oe te Marples, 1952. Holotype: tarsometatarsus, Waitakian (latest Oligocene-earliest Miocene), Hakataramea
ey.

Marplesornis Simpson, 1972a. Type-species: Palaeospheniscus novaezealandiae Marples, 1960.

Marplesornis novaezealandiae (Marples, 1960). Holotype: relatively complete skeleton, Miocene-Pliocene, possibly
Waipipian, Waiauan-Tongaporutuan, or Otaian-Altonian, Motunau.

Tereingaornis Scarlett. Type-species: Tereingaornis moisleyi Scarlett, 1983.

Tereingaornis moisleyi Scarlett, 1983. Holotype: wing elements, Waipipian, mid Pliocene, Te Reinga, Wairoa River.
See also McKee 1986, 1987a.

Pygoscelis Wagler. Type-species: Pygoscelis papua (Forster), extant Gentoo penguin.
Pygoscelis tyreei Simpson, 1972a. Holotype: partial skeleton; age as for Marplesornis novaezealandiae, Motunau.
Aptenodytes Miller. Type-species: Aptenodytes patagonica Miller, extant King penguin.

Aptenodytes ridgeni Simpson, 1972a. Holotype: leg bones and fragments of trunk, age as for Marplesornis
novaezealandiae, Motunau.

Megadyptes Milne-Edwards. Type-species: Megadyptes antipodes (Hombron & Jaquinot), extant Yellow-eyed penguin.
Megadyptes antipodes (Hombron & Jaquinot), Extant around New Zealand. Okehuan (Early Pleistocene) fossils reported
by Fleming (1979: 75) and J.C. Yaldwyn (pers. comm.). Holocene specimens noted by Grant-Mackie & Scarlett
(1973).
Eudyptula Bonaparte. Type-species: Eudyptula minor (Forster), extant Licle Blue or Fairy penguin.

Eudyptula minor (Forster). Extant around New Zealand. Pleistocene fossils (stage unspecified) reported by Grant-Mackie
& Simpson (1973: 441). Holocene specimens noted by Grant-Mackie & Scarlett (1973).

Eudyptes Vieillot. Type species: Eudyptes chrysocome (Forster), extant Rockhopper penguin.
Eudyptes pachyrhynchus Gray. Extant around New Zealand. Holocene, Grant-Mackie & Scarlett (1973).
Eudyptes sp. Holocene, Grant-Mackie & Scarlett, 1973.

Genus or genera and/or species indeterminate; not demonstrably congeneric with above.

Genus and species indet. Femur, Whaingaroan (Early Oligocene), Motutara. Marples & Fleming, 1963; Grant-Mackie &
Simpson, 1973.

Genus and species indet. Leg and wing bones, Duntroonian (Late Oligocene), Te Kauri. Grant-Mackie & Simpson, 1973.

Genus and species indet. Mainly leg bones, Whaingaroan (Early Oligocene), Glen Massey. Grant-Mackie & Simpson,
1973.

Genus and species (5 or more species) still undetermined, of Fordyce & Jones 1987, 1988. Mostly leg and wing bones, but

includes partial skeleton (Fig. 18), Duntroonian (Late Oligocene) and Waitakian (latest Oligocene-earliest Miocene).
OUGD collections.

a ae—emom—m—_——

1312 - FORDYCE

eee
Table 6. Fossil Cetacea recorded from New Zealand. Only the more significant unpublished records are included.
Repository cited only for specimens not previously mentioned in print.
Suborder ARCHAEOCETI
Basilosauridae
Kekenodon onamata Hector, 1881a. Holotype: skull bones, teeth; ?later Whaingaroan (late Early Oligocene) or

Duntroonian, Late Oligocene, Wharekuri. Provisional family assignment.

aff. Dorudon sp. of Fordyce 1985b; Bortonian or Kaiatan, Middle or Late Eocene, Waihao. Provisional identification.
Suborder INCERTAE SEDIS

Genus and species indet. ("archaeocete-like” cetacean, Marples 1949b). Specimen: cranial endocast; Waitakian?, latest
Oligocene-earliest Miocene?, Milburn?

Suborder MYSTICETI
Family incertae sedis
"Squalodon" serratus Davis, 1888b. Holotype: tooth; late Whaingaroan or Duntroonian, late Early or Late Oligocene,
Karetu River. Probably but not certainly Mysticeti.

Mammalodon sp. Periotic, skull fragments, Duntroonian, Late Oligocene, Waihao and Hakataramea.

Genus and species indet. of Fordyce 1989a. Specimen: mandible; Whaingaroan, Early Oligocene, Waikari. Probably but
not certainly Mysticeti.

Genus and species indet. ("third toothed specimen" of text). Periotic, skull fragments, Duntroonian, Late Oligocene,
Waihao.

Genus and species indet. ("protosqualodont" of Keyes 1973). Specimen: skull bones, teeth; Whaingaroan, Early
Oligocene (possibly Runangan, Late Eocene), Oamaru. Probably but not certainly Mysticeti.

Cetotherlidae
Mauicetus parki (Benham, 1937a) (Benham 1939, 1942). Holotype: skull; probably Waitakian, latest Oligocene-earliest
Miocene?, Milbum?.

"Mauicetus” lophocephalus Marples, 1956. Holotype: skull, mandible earbones, vertebrae; Duntroonian, Late Oligocene,
Duntroon.

"Mauicetus" waitakiensis Marples, 1956. Holotype: skull fragments, earbones, vertebrae; Duntroonian, Late Oligocene,
near Duntroon.

"Mauicetus” brevicollis Marples, 1956. Holotype: vertebrae, limb fragments; Waitakian, latest Oligocene-earliest
Miocene, near Duntroon.

Genus and species indet. ("Kekenodon onamata .. . specimen 2" of McKay 1882b and Benham 1937c). Specimen:
earbones, fragments of skull and other bones; ?later Whaingaroan (late Early Oligocene) or Duntroonian, Late
Oligocene, Wharekuri.

Genus and species indet. ("Kekenodon onamata . . . specimen 4" of McKay 1882b and Benham 1937c). Specimen:
earbones, fragments of skull and other bones; ?later Whaingaroan (late Early Oligocene) or Duntroonian, Late
Oligocene, Wharekuri.

Genus and species indet. ("non-arched rostrum” of Fordyce 1980a). Specimen: rostrum and fragments of skull bones; late
Whaingaroan, later Early Oligocene, Southeast Nelson. OUGD collections.

Genus and >11 species indet. in addition to above (see text), ?later Whaingaroan (late Early Oligocene) or Duntroonian
Kokoamu Greensand and lateral equivalents, Late Oligocene, Waitaki Valley area. Includes “large skull” of Fordyce
1987 and manuscript A. OUGD collections.

Balaenidae (provisional identification)

Genus and species indet. (Kingma, 1971), Nukumaruan, Late Pliocene, Matapiro.

FOSSIL VERTEBRATES OF NEW ZEALAND - 1313

Balaenopteridae (provisional identification)

Balaenoptera sp. of Bearlin 1985, 1988 ("cetothere" of Gaskin 1972: Fig. 3), Opoitian, Early Pliocene, Taihape.
cf. Balaenoptera sp. (of Hector 1881; Bearlin 1988), Neogene, Westland.

Genus and species indet., balaenopterid of Bearlin 1987; ?Middle or ?7Late Miocene, Dovedale. OUGD collections.
Genus and species indet., 7Middle or ?Late Miocene, Gore Bay. OUGD collections.

Suborder ODONTOCETI

Family incertae sedis
"Squalodon" andrewi Benham, 1942. Holotype: wom tooth, now lost; Waitakian?, latest Oligocene-earliest Miocene,
Milbum. Not Squalodon, but possibly Squalodontidae; nomen dubium?.
Squalodontidae

"Prosqualodon" hamiltoni Benham, 1937b. Lectotype: skull; Waitakian, latest Oligocene-earliest Miocene, ?Caversham,
near Dunedin.

Prosqualodoa species, cf. P. davidis Flynn, 1923. Waitakian, latest Oligocene-earliest Miocene, Milburn; Fordyce 1984a.
OM collections.

Genus and species undetermined ("longirostral skull” of text; Fordyce ms B), Waitakian, latest Oligocene-earliest
Miocene, Duntroon. OUGD collections.

Tangaroasaurus kakanuiensis Benham, 1935a. Holotype: teeth, incomplete jaws; Otaian-Altonian, Early Miocene,
Kakanui. Family position uncertain, but probably Squalodontidae.

Genus and species indet. ("Squalodon" andrewi Benham, 1942, in part, nominal paratype). Specimen: teeth; Waitakian?,
latest Oligocene-earliest Miocene?, Clarendon. Generic and family position uncertain, but possibly Squalodontidae.

Genus and species indet. (squalodont with supernumerary teeth, of Fordyce 1983a), Waitakian, latest Oligocene-earliest
Miocene, Milbum. Generic and family position uncertain, but probably Squalodontidae.

"Microcetus" hectori Benham, 1935b. Holotype: mandible, skull; Waitakian, latest Oligocene-earliest Miocene, Otiake
River or Wharekuri. Generic and family position uncertain.

"Microcetus” aff. hectori, Duntroonian, Late Oligocene, Duntroon. Generic and family position uncertain. OUGD
collections.

"Prosqualodon" marplesi Dickson, 1964. Holotype: skull, mandibles, vertebrae; Waitakian, latest Oligocene-earliest
Miocene, Otiake River. Generic and family position uncertain

Austrosqualodon trirhizodonta Climo & Baker, 1972. Holotype: mandibles; Duntroonian, Late Oligocene, Puponga.
Generic and family position uncertain.

Genus and species indet. (odontocete of Fordyce 1987), Whaingaroan or Duntroonian, late Early or Late Oligocene,
Aorere. Generic and family position uncertain. OUGD collections.

Eurhinodelphidae
Phocaenopsis mantelli Huxley, 1859. Holotype: humerus; Altonian, Early Miocene, Awamoa or Old Rifle Butts, near
Oamaru. Probably Eurhinodelphidae, but family position uncertain, Fordyce 1982a.

Genus and species indet. Waitakian-Otaian, latest Oligocene-Early Miocene, Kaikoura. Provisional identification. CM
collections.

Genus and species indet., Fordyce 1984b. Middle or Late Miocene, Chatham Rise. Provisional identification.

Ziphiidae

cf. Hyperoodon sp., Fordyce 1984b. Middle or Late Miocene, Chatham Rise.
"Berardius sp.", Hector in Gray 1871. Recent?, localities unknown.

Genus and species indet., Fordyce & Cullen, 1979. Middle or Late Miocene, Chatham Rise.

1314 - FORDYCE

Physeteridae
Genus and species indet. ("Phoberodon-like species” of Fordyce 1982b), Waitakian, latest Oligocene-earliest Miocene,
Ngapara. Family identification provisional. OM collections.
cf. Scaldicetus sp., Fordyce 1984b. Middle or Late Miocene, Chatham Rise.

Genus and species indet., Mangapanian, late Pliocene, Waipukurau. CM collections.

Kogiidae

cf. Kogia sp., Fordyce 1984b. Middle or Late Miocene, Chatham Rise.

Kentriodontidae
Genus and species undescribed ("porpoise", Grant-Mackie 1970), Waitakian, latest Oligocene-earliest Miocene, Port
Waikato.
Genus and/or species undescribed, Otaian, Early Miocene, northwest Nelson coast. OUGD collections.

Genus and species indet. ("odontocete-like" cetacean, Marples 1949b). Specimen: cranial endocast; Waitakian?, latest
Oligocene-earliest Miocene?, Milburn? Family position uncertain, but possibly Kentriodontidae.

Delphinidae

Delphinus aff. delphis, Waitotaran, Late Pliocene, Waihi Beach, Hawera. Wanganui Museum collections.
cf. Pseudorca sp., Waitotaran, Late Pliocene, Napier. NMNZ collections.
cf. Orcinus sp. , Nukumaruan, Early Pleistocene, Motunau. CM collections.

Genus and species indet. (close to Delphinus sp. or Stenella sp.; McKee & Fordyce 1987), Waitotaran, Late Pliocene,
Waihi Beach, Hawera.

Globicephala sp. , Neogene, Glenafric. CM collections.

Genus and species indet. (21 spp. based on skulls), Neogene, Motunau. CM collections.
Genus and species indet. Castlecliffian, Mowhanau Beach, Wanganui. OUGD collections.

Genus and species indet. (>4 spp. based on periotics, Fordyce 1984b). Middle or Late Miocene, Chatham Rise.

Phocoenidae

Genus and species indet. (kentriodontid or delphinid of Fordyce 1984b). Middle or Late Miocene, Chatham Rise.

PLATES

Plate 1. Partly-prepared skeleton of large Late Cretaceous plesiosaur from North Otago, with Ewan Fordyce
(left) and Craig Jones (right). The anterior is towards the bottom, the vertebrae run towards the bottom, and
a flipper is apparent mid left; specimen in the Geology Museum collections, University of Otago. (From
Fordyce 1986).

Plate 2. Probable early penguin from the Palaeocene or Early Eocene, North Canterbury; from left to right,
coracoid, clavicle, ulna, humems, specimen in New Zealand Geological Survey collections, smallest divisions
on scale bar, 1 mm. (Photo by D.V. Weston; from Fordyce et al. 1986).

PLATE 1 FOSSIL VERTEBRATES OF NEW ZEALAND - 1315

1316 - FORDYCE PLATE

CHAPTER 27

THE QUATERNARY
AVIFAUNA OF NEW
ZEALAND

Phil R. Millener!

FMPOGUCHEME: fechas cee tr cerssishinsengesese 1318 PLOCEHATTITONINES sess evens shaseresvectevs 1327
New Zealand's Geologic Background...... 1318 PelecaniFORMes.S vis....cceceverecsrevekss 1327
Possible Centres of Origin, Dispersal Routes CNEONTOLMOS. cosy acinepa sativa ce oelvages 1327
and Times of Colonisation ........ 1320 PUSCUIPOTINCS ES a3 | sos anf cbtsaes tebe evens 1329
THE Fossil RECORD ac cct cous Secu cesvivsesacenes 1321 Parcons borne Sess ced o bet sscckoctal agen 1329
RAVES a eicr ie ie purictssodalaseels 1322 Galliformes. 0 cndeacsseccatiaeewsbrede 1330
Sy PRIS 2.5 sh Cldtetastrlechboateteleeny tense 1322 GrULPOTMES sco. l oa. fosca looses vives 1330
PUI cs kako cacwsehaaesysheroauednes beens 1322 CharddrifOrMess.nivetpnaieAastehansnide 1331
1117374 (2) ee SOO BOC Pe 1323 Columbiformes..............c00000eeeeees 1331
Significance of Quaternary Avifaunal PSIMACTFOTMOSS: ssccw flag sieisde sees setts 1331
PRCA NIE, 4 test ret died avandtoabearactnake 1323 CuculifOrmes..........cseeceecceeeeeeeeeees 1332
The New Zealand Quaternary Avifauna: a Caprimulgifornes reds cnatscaaseanes 1332
Resumé of its Composition, SOMSUOTIMNEE ie eesaahaaseestisctawdondacas 1332
Distribution CoraciifOrmes ...........ceccseeeeeeeeeeeeee 1332
and Relationships ...............0066 1323 PasserifOrmes ...........ssccseceseceeeeees 1333
Dinornithiformes ...............c0csc0eeeee 1325 SUMMA Ye 2 la cvednseni ele sdetaredeenctlet 1334
ADLCEY BIT ORMI GS 0... snccaverevasven senesced 1326 REPELENCES. on. cicvnstesdiedueonee'vaveeed 1334
SPNEMISCHONIMNES .6-.cevsivnreesecssdrersrer 1326 Appendix Tek, crc .ske tery. cone daioded pases 1340
Podicipediformes..........cccccceeeeees 1326

es
1 National Museum, P.O. Box 467, Wellington, New Zealand.

1318 - MILLENER

INTRODUCTION

Some 380 living and fossil avian taxa have been recorded from New Zealand and its
outlying islands (Turbott 1990), Seventeen named and at least five as yet unnamed species are
known only as Tertiary fossils, some 34 species (the exact number depending upon the
classification adopted) found in Pleistocene to subrecent deposits, are presumed to have become
extinct during Polynesian settlement and a further eleven, at least, have been exterminated
within the last one hundred and fifty years. Of the living species, 125 (largely migrant
shorebirds, wide-ranging oceanic species or wind-blown strays) do not breed in the New Zealand
region. The breeding species include 91 sea and shore birds, 62 land and freshwater birds and 39
introduced birds.

In terms of evolutionary history the New Zealand avifauna, fossil and living, shows some
intriguing features. There are relatively few land bird species, and those are largely endemic.
With some notable exceptions few groups have shown a marked degree of adaptive radiation; an
unusually high proportion of the species is flightless or has weak powers of flight. A
remarkable number of species have become extinct within the last 1000 years. The
complexities of the origin and evolution of this largely unique avifauna must rely for their
explanation on many lines of evidence, not least an adequate fossil record (see T. Rich 1975).
Unfortunately, this record in New Zealand is so incomplete that it is of little use in clarifying
the ancestry of the New Zealand fauna, or its geographic origins. Thus, speculation must rely
largely upon indirect evidence and be guided by knowledge of fossil faunas from other areas of
more complete record.

NEW ZEALAND'S GEOLOGIC BACKGROUND

It has been argued (Cracraft 1973) that palaeogeographic and palaeoclimatic events in the
Mesozoic and Cainozoic have greatly influenced present day bird distribution and evolution, and
that these events need to be considered when discussing the centres of origin and pathways of
dispersal of avian groups.

A succession of episodes in the geologic history of New Zealand, the nature and timing of
which have profoundly influenced the development of other segments of the biota (Climo
1975, Godley 1975, Forster 1975), have been no less crucial to the evolution of its avian
fauna.

New Zealand, at the beginning of its geologic history, formed a segment of Gondwana,
roughly equidistant between the Pacific coast of West Antarctica and the east coast of Australia
(Fleming 1979). It was, thus, contiguous with other southern continents and shared to some
degree their faunal and floral assemblages. Palaeontological evidence, both faunal and floral,
indicates that an equable climate prevailed in both East and West Antarctica during the late
Mesozoic and early Tertiary, and thus this region could have served as a dispersal route for
organisms amongst the segments which later formed the southern continents (Rich 1975a).
Geophysical data (Kennett 1977, Weissel, Hayes & Herron 1977) indicate that sea-floor
spreading on the South-east Pacific Rise in the Late Cretaceous, and differential movement
between Australia, New Zealand and Antarctica produced first the separation of New Zealand
from West Antarctica (82 myBP) and then the opening of the Tasman Sea (82-60 myBP -
Weissel & Hayes 1977). There is some evidence, at best equivocal, that during the Cretaceous,
land may have extended north along the Lord Howe Rise to New Caledonia and south to the
Campbell Plateau (Fleming 1979). The increasing isolation of the northward and eastward
drifting New Zealand landmass would obviously have decreased the opportunities for dispersal
of terrestrial organisms from this time on. The timing of New Zealand's separation is

NEW ZEALAND QUATERNARY AVIFAUNA - 1319

particularly critical to the subsequent evolution of its avian fauna, in that separation apparently
pre-dated the arrival in the Australian region of both snakes and placental mammals.

Throughout the Tertiary New Zealand as part of a mobile Pacific margin became a changing
archipelago as tectonically induced, small scale troughs and folds resulted in frequent changes in
geography (Fleming 1979). Tertiary climates, although fluctuating considerably, were
generally warm, temperate to subtropical until the Late Miocene-Pliocene deterioration which
culminated in the Pleistocene ice-ages. Forests were, for much of the Tertiary, subtropical to
tropical and probably clothed most of the land area. In the absence of land connections,
dispersal of faunal elements to New Zealand during the Tertiary must have been dominantly
trans-Tasman, undoubtedly assisted since the Oligocene by the circum-Antarctic current and its
associated west wind drift (Barker & Burrell 1977).

The Pleistocene was a period of marked tectonic and climatic fluctuation. Continued
mountain building, which began in the Pliocene, altered topographic relief. Repeated glacial
and interglacial episodes produced marked geographic, climatic and biological changes. At each
interglacial, high sea-levels inundated low lying areas with the consequent separation of islands
leading to varying degrees of endemism in the biota isolated upon them. Particularly in the
Chatham Islands, changing Pleistocene sea-levels profoundly affected physiography. At the
peak of the Upper Castlecliffian transgression the sea, which left evidence of fossil reefs on the
summit of 296 m high Pitt Island (Hay et al. 1970: 52, 54) would have almost completely
inundated the group. Thus much of the terrestrial biota may have been eliminated. The diverse
subfossil and recent faunas of the Chathams must, necessarily, have resulted largely from
adaptive radiation among post-Castlecliffian colonists (see Meredith this volume; Millener in
press a).

During glacial periods, retreat of the sea allowed considerable land extension and many
islands previously separated during the Pliocene became again joined to the mainland.
Extensive climatic change, assisted by violent volcanic activity in the North Island, decimated
fauna and flora alike, although forest refugia undoubtedly persisted in many regions, especially
in the north. Almost certainly, many Tertiary elements, especially those adapted to warm-
temperate forest habitats, became extinct during cold phases of the Pleistocene. Some typical
Australian plants, such as Eucalyptus, Casuarina and Acacia formerly well established in New
Zealand, were victims of the Pleistocene decimation (Stevens 1979).

The post-glacial period saw a general amelioration of climate, although some fluctuations
may have been sufficiently severe to have significantly affected certain biotic elements
(Fleming 1963a, McGlone & Moar 1977). Areas previously deforested became revegetated and
a rapid rise in sea-level resulted in the inundation of former straits which once again became
barriers to dispersal of terrestrial organisms. Shorelines readjusted to a sea which stabilized at
its present level some 6,500 years ago (Thom & Chappell 1975; Gibb 1979). Except in
limited alpine, arid or wetland regions and in areas subjected to recent volcanism, forest and
scrub would have dominated the landscape.

The arrival of man about 1,000 years ago initiated a series of ecological changes of
unprecedented rapidity and magnitude. The introduction, for the first time in New Zealand's
history, of carnivorous mammals - firstly Polynesian rats and dogs, then European cats, rats
and mustelids - as well as the extensive clearance of indigenous forest has had a continuing,
profound ecological impact upon the more ancient elements of the avian fauna especially
(Fleming 1962c).

1320 - MILLENER

POSSIBLE CENTRES OF ORIGIN, DISPERSAL ROUTES AND
TIMES OF COLONIZATION

The recent revolution in biogeography, initiated by the general acceptance of continental
movement, has provided the impetus for the development of new hypotheses regarding the
origins of many groups of birds, especially those with present southern distributions. Mayr
(1944), Oliver (1945, 1955) and Darlington (1957) sought a Palaearctic origin, followed by
southward dispersal, for most groups of birds. The implicit assumption of stable continents
prevailed in these theories. It has become increasingly apparent, however, that the history and
distribution of many avian families can be better reconciled with "drift" rather than "stabilist"
interpretations of geological events.

The persistence of land connections in the Southern Hemisphere until the Late Cretaceous
and Tertiary and of climates which supported sizable forests until the late Tertiary in South
America, Australia, New Zealand and the Antarctic, facilitated avifaunal interchange along both
continental and archipelagic routes. Avian faunas on the major land masses became
increasingly distinctive as the continents became more isolated and climates more zoned
(Cracraft 1973).

Cracraft (1973, 1974) has strongly supported a Gondwanan origin and dispersal for the
ratites, penguins, some galliforms such as the megapodes, and the suboscines, and, with
somewhat less certainty pigeons, parrots, cuckoos and their allies (see also Rich 1975a,b).

Six main geographic elements have been recognized in the New Zealand avifauna (i.e.
archaic, Malayo-Pacific, austral, Holarctic, Australian and cosmopolitan - see Fleming 1963b).

Of those taxa inferred to have been in New Zealand long enough to be classed as archaic
elements (the ratites and the two endemic families of passerines) only the ratites need have
reached New Zealand directly via a southern route. Some groups, especially those of ancient
dispersal, may have originated in southern (Gondwana) regions but reached New Zealand, in
effect secondarily, via Australia. Equally, groups of northern (Malayo-Pacific) affinity may
have come either direct from the north and/or via Australia.

The southern or austral element in the avifauna consists exclusively of sea birds, especially
penguins, albatrosses, petrels and cormorants, many of which have circumpolar distributions.

A few New Zealand birds (the Holarctic element) are related to North Temperate forms but
are absent in the intervening tropics, e.g. the Scaup (Aythya novaeseelandiae), the Merganser
(Mergus australis), the Black-billed gull (Larus bulleri) and the South Island Pied Oystercatcher
(Haematopus ostralegus finschi).

The dominance of the Australian element in the New Zealand avifauna, especially amongst
terrestrial and freshwater birds, has long been recognized (Falla 1953, Fleming 1962a, b,
Williams 1962). As a result of trans-oceanic "sweepstakes" dispersal, Australia has been a
prolific source of plants and animals from the early Tertiary to the present.

Fleming (1962a) used the morphologic differences that distinguish New Zealand birds from
their overseas relatives as a rough indication of the time that has elapsed since their first
successful colonization. Despite the problems arising from changes in systematic ranking of,
and differing evolutionary rates in particular groups, Fleming's yardstick, in default of other
evidence, is used here. Thus, the New Zealand ratites, the moas (Dinornithidae and Emeidae)
and kiwis (Apterygidae), possibly, but not necessarily (see Houde 1986), were here in the Late
Cretaceous (see Sibley & Ahlquist 1972, Sibley & Frelin 1972). The two remaining endemic
families, the Wrens (Acanthisittidae) and the Wattlebirds (Callaeatidae) are presumed to be early
Tertiary, and the 25 endemic genera later Tertiary colonists. The New Zealand thrushes
(Turnagra), formerly considered to comprise an endemic family Tumagridae have recently been
placed in the Paradiseaidae (Olson et al. 1983). The endemic species of overseas genera are

NEW ZEALAND QUATERNARY AVIFAUNA - 1321

probably Pleistocene arrivals whilst endemic subspecies may be no older than post-glacial;
certainly well-differentiated subspecies have developed across Cook and Foveaux straits and on
the Auckland Islands in less than 10,000 years. Those taxa inseparable from their overseas
counterparts are probably all Holocene arrivals - many, such as the Pukeko (Porphyrio p.
melanotus), Pied Stilt (Himantopus h. leucocephalus), Banded Rail (Gallirallus philippensis
assimilis) and Kingfisher (Halcyon sancta vagans) to judge by their lack of subfossil records
may, in fact, have colonised only within the last few hundred years. If the known historic
colonists and the increasing variety of stragglers which may yet breed in New Zealand, are
added to this list of inferred trans-Tasman migrants, it is clear that the composition of New
Zealand's past and present avian fauna has been markedly influenced by "sweepstakes dispersal".

THE FOSSIL RECORD

; The major inadequacy of the New Zealand avifaunal fossil record is that only two disparate
time periods are represented: the Tertiary and the Late Pleistocene and Holocene (Fig. 1). A
very limited number of taxa, representative of only three orders, have solely Tertiary records
(see Fordyce 1982, and this volume); far more are known from Quaternary deposits, but the
majority of them are found only in post-glacial to sub-recent deposits (Fig. 2).

Fossil bones of most of the extant and recently extinct bird species known in New Zealand
occur in deposits of Plio-Pleistocene to sub-recent age. The extinct species (see Brodkorb
1963, 1964, 1967, 1971, 1978) represent eight orders: one (Dinornithiformes) includes the
extinct, endemic families, Dinornithidae and Emeidae; another (Caprimulgiformes) has no
living New Zealand representatives but is known both living and fossil in Australasia; the
remainder (Pelecaniformes, Anseriformes, Falconiformes, Gruiformes, Charadriiformes,
Passeriformes) are cosmopolitan, with some living New Zealand species (Appendix I).

Plio-Pleistocene deposits have so far yielded only a few isolated bones, none of them older
than about 2.5 million years (Worthy et al., in prep), and all attributable to recently extinct or
extant taxa; moas, of all the known genera except Megalapteryx, a kiwi (Apteryx sp.), an
anseriform (Cnemiornis gracilis) and an eagle (Harpagornis moorei) have been identified
(Millener 1981b).

One of the few extensive collections dating from the last (Otira) glacial period is from Cape
Wanbrow, Oamaru, where a transgressive shallow marine/beach/dune/loess deposit has yielded a
considerable array of faunal remains (Grant-Mackie & Scarlett 1973). Some of those from the
lower part of the sequence have given radiocarbon dates ranging from >26,100 (NZ3093) to
>17,300 yBP (NZ 3092). Representatives of twelve orders of marine and terrestrial birds,
reptiles and marine and land mollusca have so far been recovered.

Honeycomb Hill Cave, northwest Nelson, is another site with subfossil deposits which are,
at least in part, as old as the Otira Glacial. Here the extraordinarily diverse faunal assemblages,
from which more than fifty bird species have been identified, have yielded dates ranging back to
more than 20,000 yBP (Millener 1984a, Worthy 1987a, 1988a).

Other terrestrial deposits, which form the major sources of subfossil remains, are largely of
post-glacial age. The four main types of deposit (cave sediments, dune sands, swamp alluvium
and occupation middens) are widely distributed throughout New Zealand and on some outlying
islands. The total number of subfossil sites known in New Zealand exceeds 800, of which
almost 550 are in the North Island (see Millener 1981b), more than 220 in the South and
Stewart Islands, and some 50 in the Chathams (see Millener 1981a, Meredith this volume);
Throughout this paper the term "fossil" is used for faunal remains from consolidated deposits
(generally Early Pleistocene or older), while the convenient term "subfossil" is applied to all
remains from essentially unconsolidated sediments (dunesands, cave silts, buried soils) typically

of late-glacial or post-glacial age.

1322 - MILLENER

Caves

Sediments within caves, most formed within limestone (Whangarei, Waikaremoana,
Waitomo, Martinborough, Karamea, Charleston, Te Anau, Otago) or marble (Takaka) but
occasionally in lava (Auckland) have long been known as significant sources of subfossil
faunal remains (Hamilton 1892, Bartrum 1924, Archey 1941, Oliver 1949, Yaldwyn 1956,
1958, Medway 1967, 1971, Paulin 1973). The avian species found in cave deposits are, most
frequently, flightless forest-dwellers. Those found most consistently and in greatest abundance
include many species of moa (e.g. Anomalopteryx didiformis), kiwis (e.g. Apteryx australis
and A. owenii), rails (e.g. Gallirallus australis), ducks (e.g. Euryanas finschi), parrots (e.g.
Strigops habroptilus) and passeriformes (e.g. Callaeas cinerea). Frequently sediments within
caves also contain stream-washed accumulations of land snails (Dell 1955, Climo 1975), many
of them species confined to forest habitats.

The age of the majority of such remains is unknown but a iew radiocarbon determinations
from the Waitomo region (North Island) and North-west Nelson (South Island) have yielded
dates within the range 1,000-30,000 years BP (McCulloch & Trotter 1979, Cassels & Millener
1985, Worthy 1987b).

Swamps

Among the most prolific sources of subfossil avian remains have been peat swamps usually
in, or adjacent to, limestone country. Localities such as Lake Poukawa and Te Aute in the
North Island and Herbert, Enfield, Kapua and Pyramid Valley in the South Island have been
amongst the most important subfossil sites so far investigated, and many of them figure
prominently in the early literature (Hamilton 1889, Forbes 1892, Falla et al. 1941, Archey
1941, Oliver 1949). The vast majority of avian remains recovered from such swamps have
been those of moas, although at Te Aute (Hamilton Joc. cit.), Pyramid Valley (Scarlett 1955)
and Lake Poukawa (Price 1963, 1965; Horn 1980, 1983) significant numbers of carinate taxa
have also been found. In the northern North Island most of the extensive peat swamps are too
acidic to preserve bone material. Sedimentological and palynological data obtained from a
number of stratified swamps, notably Pyramid Valley (Moar 1970, Gregg 1972), Scaiffes
Lagoon (Trotter 1970) and Lake Poukawa (McGlone 1978, Pocknall & Millener 1984) have
allowed interpretations of their depositional vegetational and climatic histories to be made.
Bone material from only a very few swamps has so far been radiocarbon dated and as yet no
ages greater than approximately 9500 yBP have been obtained (Trotter 1970, McCulloch &
Trotter 1979 ),

Dunes

Extensive Late Pleistocene dunes exist in many parts of New Zealand, but none so far
investigated appears to have maintained conditions suitable for long term preservation of bone
material. Dunes developed during post-glacial time, however, in some localities have retained a
remarkable array of generally well preserved avian remains, as well as those of associated
reptilian and landsnail faunas. A significant feature of the faunas within such dune deposits is
that they consist for the most part of species which utilise or have utilised forested habitats
almost exclusively. The consistent occurrence of such species as Kaka (Nestor meridionalis),
parakeets (Cyanorhamphus spp.), Kokako (Callaeas cinerea), Saddleback (Philesturnus
carunculatus) and Tui (Prosthemadera novaeseelandiae), of extinct taxa with inferred forest-
dwelling habits such as moas (Dinornithiformes) and rails (Rallidae) and of landsnails such as
Rhytida spp. and Serpho kivi (an obligate arboreal species: Powell 1979) attest to the former
existence of extensive vegetative cover at, or in close proximity to, their sites of deposition.

NEW ZEALAND QUATERNARY AVIFAUNA - 1323

Radiocarbon determinations on both moa bone collagen and landsnail shell carbonate from
sites on the Aupouri Peninsula, North Island, have yielded ages within the range of about 600-
6,000yBP (Millener 1981b), while those for birds from Chatham and P. H. Island dine sands
range from 1500 to 8000 y.B.P. (Millener, unpubl. data).

Middens

Occupation middens frequently contain the discarded or worked bones of a wide variety of
birds hunted by the early Polynesians. Such middens are most common on coastal dunes but
are also present in inland rock shelters. Most dated sites rich in avian bones are between 400
and 800 years old (Scarlett 1974, 1979, Moore & Tiller 1975, 1976, McCulloch & Trotter
1975, Sutton 1979). In younger sites, bird remains are generally scarce (Simmons 1968). Of
the 33 avian species known to have become extinct during the Polynesian period, the remains
of at least 28 (including all 11 moa species) have been found in association with Archaic Maori
occupation sites. However, it seems that for none of them can extinction be attributed solely
to direct hunting. In addition to these extinct taxa, the remains of a considerable array of the
living species have also been found in middens (e.g. Hamel 1977, Davidson 1979, Leach 1979,
Scarlett 1979, Sutton 1979, Foley 1980, Cassels 1984). The species composition of midden
faunas varies considerably from one site to another. Petrels (Procellariidae) and cormorants
(Phalacrocoracidae) are usually the most common marine or quasi-marine birds, while of the
terrestrial species, Kaka (Nestor meridionalis), parakeets (Cyanorhamphus spp.), pigeon
(Hemiphaga novaeseelandiae), Weka (Gallirallus australis) and Tui (Prothemdera
novaeseelandiae) are generally the most abundant.

SIGNIFICANCE OF QUATERNARY AVIFAUNAL REMAINS

The great quantity and variety of remains from all these subfossil sources, are unfortunately
of little use in clarifying origins and evolutionary trends since the majority are geologically
very recent. Nonetheless they are of considerable use in taxonomic studies and for indicating
former distributions, probable times of colonization and extinction.

Distributional data should be interpreted with caution, however, because the location of
remains is surely more indicative of the distribution of sites with conditions suitable for the
preservation of bone material, than of the actual distributions of faunal populations. Similarly
due to the vagaries of chance preservation, indicated times of colonization can only be minima,
and those of extinction maxima, as it is highly unlikely that one would find the first or last
fossil specimens of a species.

THE NEW ZEALAND QUATERNARY FAUNA - A RESUME OF
ITS COMPOSITION, DISTRIBUTION AND RELATIONSHIPS

New Zealand's living avian fauna contains members of eighteen orders of which all but one
(Apodiformes: swifts) appear in the Quaternary fossil record. Two additional orders
(Dinornithiformes: moas and Caprimulgiformes: owlet-nightjars, etc.) are known only as
fossils. Key references to, and the subfossil distributions of all known taxa are given, for
mainland New Zealand by Millener (1981b), and for the Chatham Islands by Meredith (this
volume) and Millener (in press a); see also Table 1.

Figure 1: Biostratigraphic and geomagnetic subdivisions of the Quaternary.

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NEW ZEALAND QUATERNARY AVIFAUNA - 1325

DINORNITHIFORMES: MOAS

This unique and diverse group of ratites, known to science only since 1837 (Owen 1839),
has been the centre of focus for avian palaeontology in New Zealand (Archey 1941, Oliver
1949, Cracraft 1974, 1976).

On biogeographic and taxonomic grounds, in the absence of fossil evidence it has
traditionally been inferred that the ancestors of both moas and kiwis (Apterygiformes) were
present, and flightless, on the New Zealand segment of Gondwana prior to its Late Cretaceous
separation (Fleming 1975). However, more recent research (Houde & Olson 1981, Olson
1985, Houde 1986) suggests the possibility of post-separation colonisation by volant
ancestors. During the Tertiary the archipelagic nature of the New Zealand landmass would have
tended to encourage adaptive radiation in the group (Hutton 1892). It is possible that the
dichotomy of moas into two families may have occurred during the Oligocene when marine
transgression divided New Zealand into two or more disparate land masses. The proliferation
into the very large number of forms known from post-glacial deposits was probably due to a
later (Pleistocene) radiation (Cracraft 1976).

The taxonomy of moas, especially at the species level, has been the subject of considerable
research but has still to be satisfactorily resolved. Haast (1874), Archey (1941) and Oliver
(1949) all admitted over 20 species, Scarlett (1972) somewhat fewer. In a major revision,
Cracraft (1976) accepted only thirteen species but Millener (1982) then synonymised
Anomalopteryx oweni with A. didiformis. More recently Worthy (1987a, 1988a, 1989) has
reduced to synonymy Megalapteryx benhami (with M. didinus) and Dinornis torosus (with D.
novaezealandiae), while resurrecting Pachyornis australis, thus leaving for current acceptance a
total of eleven moa species. The degree of diversity implied by classical taxonomic schemes,
in which as many as 38 species of moa were accepted, has meant that the group has long been
upheld as an almost unparalleled example of adaptive radiation. Morphologically and,
therefore, probably ecologically, the moas appear to have been more conservative than have
been, for instance, the New Zealand rails, a group with roughly the same number of endemic
genera. In hardly more than a dozen species the rails show a comparable range of variation in
size and structure, and especially in bill shape, considerably greater diversity. It would seem
that the more conservative classification, proposed by Cracraft and modified by Millener (1982)
and Worthy (loc. cit.) reflects better the true degree of evolutionary adaptation achieved by the
moas.

Two of the six genera of moas appear to have been restricted, at least in Quaternary time, to
the South Island. None is known from the Chatham Islands. Stewart Island records are rare,
there being only one from natural dunes (Benham 1909) and few more from middens (Scarlett
1979). Most of the moas appear to have had fairly broad distributional ranges and are known
from cave, swamp, dune and midden sites. Megalapteryx and Anomalopteryx appear to have
been more common in steeper country, while Euryapteryx was more confined to coastal
lowlands. To judge by midden occurrences Dinornis was absent from the north-eastern South
Island in Polynesian times (Scarlett 1974).

The apparent abundance of moa remains in early occupation sites led to the first settlers in
New Zealand being called Moa-hunters. A closer assessment of the importance of moa in such
sites has led to the recognition that this is an inappropriate economic designation. It appears
that in no site where food resources have been analysed has moa been the mainstay of the
Archaic Maori diet (Green 1975, but see McCulloch & Trotter 1984).

The frequent occurrence of moa remains in swamps on what are currently open plains led
early investigators to conclude that most species were predominantly inhabitants of grassland
filling a niche similar to that of ungulate grazers. The similarity of moas, in size and form, to
other large ratites of open country habitat (e.g. the emu, Dromaius novaehollandiae and ostrich,
Struthio camelus) doubtless to some extent predicated this assumption. Many lines of evidence

1326 - MILLENER

such as gizzards containing berries, fruits and shoots of forest trees (Gregg 1972, Burrows
1980, Burrows et al. 1981); moa bones associated with remains of avian and molluscan species
of obligate forest habitat (Yaldwyn 1958); and palaeobotanical data indicating that most of the
areas in which moa remains have been found were, until relatively recently forested (Molloy et
al. 1963, Moar 1970, McGlone 1978), strongly indicate that moas were forest or forest fringe
dwellers (Hamel 1979) and were much more the ecological counterparts of cassowaries than of
the savannah dwelling ratites.

APTERYGIFORMES: KIWIS

The kiwis are amongst the most primitive and specialised of the living ratites. Some
workers suggest that they, and their closest allies the extinct moas, developed from a common
stock (Houde 1986). It is unknown whether the split into the two sister groups occurred before
or after New Zealand's separation from Gondwana. Despite the long phyletic history of the
group the living species of kiwi appear to have developed only recently, probably in the
Pleistocene. No fossil kiwi bones are known before the Quaternary, although footprints
attributed to this group have been found in mudstones of ?Late Miocene-?Pleistocene age
(Mildenhall 1974, Fleming 1979). The only palaeospecies of kiwi so far described
(Pseudapteryx gracilis Lydekker, 1891) was regarded by Reid & Williams (1975), following
Storer (1960), as "the earliest known kiwi". However, Millener (1987) reduced P. gracilis to
junior synonymy with Apteryx owenii and, further, indicated that far from being the “earliest
... kiwi" the specimen in question was "almost certainly no older than late Holocene." Bones of
the three extant species have been found in numerous subfossil deposits, an indication that all
formerly had much wider distributions than they do at present (Reid & Williams 1975). The
Little Spotted Kiwi, Apteryx owenii, from subfossil evidence formerly widespread in both
main islands (Scarlett 1962, 1967b), is known to have become extinct in the North Island
about 1875 and now apparently survives only on Kapiti Island, to which it was introduced in
1913,

SPHENISCIFORMES: PENGUINS

Many of the extant species of penguin which at present inhabit the New Zealand coasts are
known from Late Pleistocene to subrecent deposits. Two essentially Antarctic species (King
Penguin, Aptenodytes patagonicus and Royal Penguin Eudyptes chrysolophus schlegeli) have
been found in subfossil deposits of mid-Holocene age at Macquarie Island (McEvey & Vestjens
1974) while the rare, mainland-breeding Yellow-eyed penguin (Megadyptes antipodes) has been
recorded subfossil from the Chathams (Millener, in press a). A. patagonicus is also known
from subfossil/midden deposits on Chatham Island (Marshall et al, 1987, Millener, in press a).
Scarlett (pers. comm.) has suggested that a penguin from the Oamaru fauna (Grant-Mackie &
Scarlett 1973), known only from a single undescribed tibiotarsus, may be a species ancestral to
Eudyptes pachyrhynchus.

PODICIPEDIFORMES: GREBES

The Crested Grebe (Podiceps cristatus) and New Zealand Dabchick
(Poliocephalusrufopectus), the only two species of this order recorded subfossil in New
Zealand, are both closely allied to Australian forms. Neither is known from deposits older than
Late Holocene and, indeed, their rarity even in occupation middens indicates that both are,
perhaps, very recent colonists.

NEW ZEALAND QUATERNARY AVIFAUNA - 1327

PROCELLARIIFORMES: ALBATROSSES, PETRELS AND RELATED
FORMS

Although the Procellariiformes are an ancient group which probably originated along the
Gondwana coastline in the Late Cretaceous (Harper 1978), the only pre-Quaternary fossil
known from New Zealand is the very tentatively and perhaps incorrectly assigned Manu
antiquus from the Oligocene. Many of the modern procellariiform families are known to have
been in existence in the Miocene, and it is highly probable that the New Zealand region, close
to their centre of origin, has long been occupied by representatives of the group. The living
species, which form the bulk of the austral component in the New Zealand avifauna (Fleming,
1979) are predominantly of southern origin, although some such as Pterodroma hypoleuca may
be of tropical affinity. The genus Procellaria is of particular biogeographic interest, as it
shows close alliance to, and possibly shares a common ancestry with, the Mediterranean
shearwater Calonectris (Harper 1978). Many of the 41 procellariiform species now breeding in
the New Zealand region are known from Quaternary deposits. The subfossil evidence for
former distributions indicates that many species, which now breed only on offshore islands,
formerly did so on a number of inland mountain ranges (Millener 1980a, 1981b, 1984a).

PELECANIFORMES: PELICANS, GANNETS, CORMORANTS AND
RELATED FORMS

Apart from the solely Tertiary record of the extinct Pseudodontornis, the fossil record of this
order in New Zealand includes just one extinct taxon, Pelecanus novaezealandiae (see Rich &
van Tets 1981), which has been found subfossil in various localities in both main islands
(Scarlett 1966). The Phalacrocoracidae (shags, cormorants) with thirteen living species is the
dominant pelecaniform family in New Zealand, and van Tets (1976) has inferred an Australasian
centre of adaptive radiation and dispersal for this group. Many of the living New Zealand
species of this family are known from subfossil deposits.

CICONIIFORMES: HERONS AND RELATED FORMS

No extinct taxa of this order are known from New Zealand. Subfossil distributions indicate
that the White Heron (Egretta alba modesta) formerly had a much wider breeding range than it
does at present. Okarito in South Westland has been, since breeding was first discovered there
in 1865, the only known nesting area. The finding of disproportionately high numbers of
mature and immature bones in subfossil dune deposits near North Cape strongly indicates that
for at least a period during the late Holocene this species also bred there (Millener 1981b).
Many of the ciconiiform taxa in New Zealand are inseparable from their Australian
counterparts, implying that they are probably recent, certainly post-glacial colonists. The
rarity of some species (e.g. bittern, Botaurus poiciloptilus and Royal Spoonbill, Platalea regia)
in subfossil deposits tends to support this conclusion. Scarlett (1979) attributed a number of
subfossil bones from the North Cape area to Plataiea, but these have since been found to be
those of Egretta alba (Millener 1981b).

Identification of subfossil bones of the Black Bittern, Jxobrychus (Dupetor) flavicollis, from
Lake Poukawa (Horn 1980) is considered invalid. These bones are referrable to xobrychus
novaezealandiae, the extinct New Zealand Little Bittern (Millener & Bartle, in prep.).

Figure 2. Localities producing fossil birds in the late Cainozoic of New Zealand.

1328 - MILLENER
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NEW ZEALAND QUATERNARY AVIFAUNA - 1329

ANSERIFORMES: SWANS, GEESE, DUCKS

The anseriform fauna, past and present, shows strong Australian affinities (Williams 1964,
Livezey 1986). The various degrees of endemism shown by the New Zealand taxa indicate that
colonization has been occurring over a prolonged period. The only pre-Quaternary fossil
anseriform remains comprise a few, as yet undescribed, duck bones from the Miocene of
Central Otago (Fordyce, pers. comm.) but subfossil remains of many of the living, and of at
least seven extinct taxa are known from numerous Pleistocene to sub-recent deposits.

The extinct taxa include three endemic genera; Cnemiornis; Pachyanas; and Euryanas.
Cnemiornis gracilis and C. calcitrans were primitive anseriformes (see Livezey, 1989) known
from the North and South Islands respectively. Pachyanas chathamica, a stoutly built duck,
was restricted to the Chatham Islands while Euryanas finschi (Finsch's Duck) was widely
distributed throughout mainland New Zealand. Both species of Cnemiornis were completely
flightless, while neither Pachyanas nor Euryanas (contra Worthy 1988b) would seem to have
been capable of powered flight. An endemic species of swan (Cygnus sumnerensis), closely
allied to the Australian Black Swan (Cygnus atratus) was formerly distributed throughout
mainland New Zealand, Stewart Island and the Chathams. A new species of pink-eared duck
Malacorhynchus scarletti, congeneric with the Australian M. membranaceus has been described
from sub-fossil remains at Pyramid Valley, South Island (Olson 1977). A musk duck,
originally described by Forbes (1892) as Biziura delautouri, is known from a number of
subfossil localities. Although Scarlett (1969) and Harrison & Walker (1970) assign all New
Zealand specimens to the living Australian species (B. lobata), Olson (1977) suggests that the
New Zealand Biziura may be at least subspecifically, if not specifically, distinct. Millener
(1984b) favoured the retention of full specific status (B. delautouri) for New Zealand Biziura.

Subfossil remains from Lake Grassmere, Marlborough and from two sites in the North
Island (Millener 1981b) indicate that a Mergus, possibly, but not necessarily, conspecific with
the recently extinct, endemic Auckland Island merganser (M. australis) was formerly present on
the mainland (Kear & Scarlett 1970). Several subfossil bones of Mergus were reported (some
in error) from the Chathams by Marshall et al. (1987), and this record has recently been further
substantiated through the excavation, from a limestone cave near the Te Whanga lagoon, of
several associated skeletons (see Millener, in press a).

The Blue-billed Duck (Oxyura australis), a living Australian species, was reported as a
subfossil at Lake Poukawa by Horn (1983) but my examination suggests that the material is
more correctly referable to Aythya.

The extant anseriforms also show varied degrees of endemism. Hymenolaimus
malacorhynchus (Blue Duck) belongs to an endemic genus, Aythya and Tadorna each have one
species and Anas one species and two sub-species which are endemic. The remaining ducks are
inseparable from their overseas counterparts.

FALCONIFORMES: FALCONS, EAGLES, HAWKS AND RELATED

FORMS

The oldest known New Zealand fossil falconiforms are of Pleistocene age. Three extinct
taxa are known; Harpagornis moorei is an eagle which in my view is most closely related to,
and perhaps should be placed in, Aguila, rather than retaining its present status as a monotypic
endemic genus (see also Shufeldt 1895); Circus eylesi, an endemic species, was a harrier very
much larger than C. approximans (Scarlett 1953); Haliaeetus australis (see Olson 1984) is an

1330 - MILLENER

endemic species of sea-eagle from the Chatham Islands. In the absence of carnivorous
mammals, these large raptors, especially Harpagornis, may have been the only significant
predators of moas and other avian taxa, prior to human settlement.

The living Falco novaeseelandiae (New Zealand Falcon) is widely known from deposits of
Late Pleistocene to subrecent age, but the Australasian Harrier (Circus approximans) is so
rarely found subfossil that it may well be a very recent colonist.

GALLIFORMES: QUAIL AND RELATED FORMS

The sole native galliform is the recently extinct Coturnix n. novaezelandiae, closely allied
to the Australian Stubble Quail (C. n. pectoralis). The New Zealand Quail, to judge by
subfossil remains, was formerly widely distributed throughout the country but is presumed to
have become extinct in the 1870's (Oliver 1955).

GRUIFORMES: CRANES, RAILS

The Rallidae, a family with a fossil record stretching back to the Late Oligocene in the
Northern Hemisphere (Olson 1985) is represented in New Zealand by a diverse assemblage of
living and fossil taxa. The occurrence of at least nineteen taxa of living and recently extinct
rails, covering a wide range of morphologic and, thus, taxonomic distinctiveness from endemic
genera to forms inseparable from subspecies outside New Zealand, offers convincing evidence
of multiple invasion via trans-oceanic dispersal during much of the Tertiary. All the extinct
forms were flightless, as are most of the living ones.

Diaphorapteryx hawkinsi, a particularly aberrant rail is known only from subfossil remains
on the Chatham Islands. It appears to have been derived from a Gallirallus ancestor perhaps
resembling G. sylvestris of Lord Howe Island (Olson 1975).

Capellirallus karamu which had wings proportionately smaller than those of any known
rail, and an elongated, almost kiwi-like bill, was first described by Falla (1954) from Karamu
Cave, near Hamilton. It may also have been derived from the Gallirallus group, perhaps
through an ancestral stage somewhat resembling G. modestus of the Chatham Islands (Olson
1975). Capellirallus is known only from the North Island (Scarlett 1970b).

Gallinula (Tribonyx) hodgenorum (includes Gallirallus hartreei, Scarlett 1970a) is known
from both the North and South islands (Scarlett 1970b). This flightless form was probably
derived from the volant ancestor which also gave rise to G. ventralis of Australia and the
flightless G. mortierii surviving in Tasmania (Olson 1975).

Gallirallus dieffenbachii and G. modestus, both recently extinct, are known only from the
Chatham Islands. The larger species, G. dieffenbachii, is the less modified and is therefore
presumably a later colonist than G. modestus. There is clear evidence that the two species were
formerly sympatric on all three islands (Chatham, Pitt, Mangere) from which they are known
as subfossils (Millener, in press a).

Gallirallus minor is a species which has never been properly defined or illustrated and may
be merely a small variant of G. australis (New Zealand Weka). G. australis has four living
subspecies and is probably an early derivative of the same stock which later gave rise to G.
philippensis, a species widely distributed in the southwestern Pacific. The New Zealand form
G. philippensis assimilis is no more than subspecifically separable from its Australian
counterpart, and its rarity in subfossil deposits suggests that it is perhaps a recent colonist.

Fulica (= Nesophalaris) chathamensis and F. prisca, very large, flightless and now extinct
coots were presumably derivatives of F. atra stock (Olson 1985), The two species known from
subfossil remains, F. prisca in the North and South islands and E. chathamensis in the
Chatham Islands must, necessarily have become flightless independently (Millener 1980b,
198 1a).

NEW ZEALAND QUATERNARY AVIFAUNA - 1331

Subfossil remains of Porphyrio (= Notornis) mantelli (Takahe) attest to its former wide
distribution in both main islands of New Zealand (Williams 1960). The nominate subspecies
is known only from subfossil bones in the North Island, but the South Island form (P. m.
hochstetteri) still persists in very limited numbers in Fiordland.

The remaining living New Zealand rails, the Pukeko (Porphyrio p. melanotus) and the
crakes (Porzana spp.), to judge by their virtual absence from subfossil and midden deposits, are
probably very recent colonists indeed.

The aberrant Aptornis otidiformis and A. defosser which appear to have affinities with
Rhynochetos jubatus, the Kagu, of New Caledonia are justifiably placed in their own family
Aptornithidae (Olson 1975). Through priority a name change to Apterornis (Apterornithidae)
seems likely (Olson & Zusi in prep.)

CHARADRIIFORMES: WADERS, GULLS, TERNS AND RELATED
FORMS

Only 12 of the 59 species of Charadrii listed for New Zealand (Turbott 1990) are indigenous
breeders. Members of this suborder are extremely rare in subfossil deposits, and only a single
extinct species, Coenocorypha chathamica, a snipe from the Chatham Islands, is known. The
endemic genus Coenocorypha belongs to a primitive group of Charadriiformes of probable
Northern Hemisphere origin. Of the other species in the genus, C. pusilla, now persists only
on South-east Island (Chathams) and LC. aucklandica on several subantarctic islands (see
Miskelly 1987a,b, 1988, Worthy 1987b). Bones tentativelly assigned to this latter species are
common in some cave deposits, particularly those in the Waitomo-Mahoenui area, North Island
(Medway 1967, 1971). The two remaining endemic charadriiform genera, the Shore Plover
(Thinornis) and the Wrybill (Anarhynchus), the former now restricted to South-east Island, are
virtually unknown as subfossils.

The two species of stilt in New Zealand possibly provide a further example of double
invasion from a single stock. The Black Sult (Himantopus novaezealandiae) is an endemic
species and thus assumed to be a Pleistocene colonist, while the Pied Stilt (H. Ah.
leucocephalus) is conspecific with the Australian form and may well be a post-glacial
immigrant (Fleming 1979).

COLUMBIFORMES: PIGEONS

Pigeons reach their greatest diversity in Australia and the southwestern Pacific. Because of
this Cracraft (1973) and Rich (1976) have suggested a southern (Gondwanan) origin for the
group. New Zealand's only living species is the Woodpigeon (Hemiphaga novaeseelandiae).
Both the mainland and Chatham Island subspecies are commonly found as subfossils,
especially in midden deposits. No extinct New Zealand taxa have been described, although
Hemiphaga n.sp. is listed from the Oamaru fauna (Grant-Mackie & Scarlett 1973).

PSITTACIFORMES: PARROTS

The Psittaciformes are possibly closely allied to the Columbiformes (Cracraft 1973) and
like the latter exhibit their greatest diversity in the Australasian region. All three New Zealand
genera are endemic. Strigops habroptilus (Kakapo), a flightless ground parrot, is perhaps most
closely related to Geopsittacus of Australia (Forshaw 1978). Bones of Strigops are amongst
the most commonly found in cave deposits throughout New Zealand. The species has probably
been extinct in the North Island since about 1906 (Williams 1956) and now survives naturally
only in very small numbers on Stewart Island (introduced on Little Barrier and Maud Island).

1332 - MILLENER

Nestor has two extant New Zealand species, the South Island alpine N. notabilis (Kea) and the
much more widely distributed N. meridionalis (Kaka). Fleming (1962b) has suggested that the
two forms became ecologically isolated during the early Pleistocene. There can be little doubt
that N. productus of Norfolk Island is a derivative of New Zealand Nestor stock (Fleming
1979). Subfossil Nestor meridionalis bones (not subspecifically identified) are known from the
Chatham Islands (Dawson 1952, 1959), while an undescribed small Nestor, doubtfully a new
species, is known only from subfossil bones in the South Island (Dawson 1952, Grant-Mackie
& Scarlett 1973). Based on the abundance of its bones in midden deposits it would appear that
N. meridionalis was a favourite quarry of the early Maori.

Cyanorhamphus is a genus for which New Zealand appears to have been the centre of
dispersal (Fleming 1979), Three species are resident in New Zealand and a number of South
Pacific islands have been colonised by their derivatives. Most of these insular populations are
weak subspecies, but some, presumably the results of earlier colonizations, are specifically
distinct. Some islands (e.g. the Chatham Islands, the Auckland Islands and the Antipodes
Islands) support two species which have colonized at different times.

CUCULIFORMES: CUCKOOS

Two species of cuckoo are regular migrants to and breed in New Zealand, but both are rare
as subfossils. The Long-tailed Cuckoo (Eudynamys taitensis) is known from two sites in the
North Island and one in the South Island, while the Shining Cuckoo (Chrysococcyx lucidus)
has been identified from a single site in the Chatham Islands.

CAPRIMULGIFORMES: OWLET-NIGHTJARS AND RELATED FORMS

Megaegotheles novaezealandiae, the only species in an extinct endemic genus, is the sole
member of this order known in New Zealand. The species is most closely allied to Australian
members of the order, both living and fossil (from the Miocene of New South Wales - Rich &
McEvey 1977) and was first recognised and described only in 1968 (Scarlett 1968). Subfossil
remains of Megaegotheles have been recorded from widespread cave and dune deposits in both
main islands (Rich & Scarlett 1977). This genus has recently been included within Aegotheles
by Olson, Balouet & Fisher (1989).

STRIGIFORMES: OWLS

Two species of owl, both with their closest relatives in Australia, are native to New
Zealand. The Laughing Owl (Sceloglaux albifacies), based on its subfossil distribution, was
formerly common in both the North and South Islands but has been presumed extinct since
about 1915 (Williams & Harrison 1972). The Morepork (Ninox novaeseelandiae), very closely
allied to the Australian Boobook Owl, is extremely rare in subfossil sites and is possibly a
very recent colonist. Subfossil bones considered to be those of Tyto alba by Scarlett (1967a)
have been shown to be those of Scleoglaux (Millener 1983).

CORACIIFORMES: KINGFISHERS

The sole New Zealand species, Halcyon sancta, is virtually unknown in subfossil sites and
could well be another of the very recent colonists. It is closely allied to others of the genus in
the Southwest Pacific.

NEW ZEALAND QUATERNARY AVIFAUNA - 1333

PASSERIFORMES: PERCHING BIRDS

Twenty-five species of native passerines are known from New Zealand and, despite the
relative fragility of their bones, most are known as subfossils. Palaeocorax moriorum (New
Zealand Crow) is known from subfossil remains only, in dune, swamp and midden deposits, in
both mainland New Zealand and the Chathams, but has seldom been found in cave deposits.
As Virtually all New Zealand caves are in relatively rugged inland country, this strongly
indicates that the former distribution of the Extinct Crow was restricted to lowland, essentially
coastal areas. The Stephen's Island Wren (Traversia lyalli) first discovered, but also
exterminated, in 1894, the Chatham Island Fernbird (Bowdleria rufescens) not recorded since
about 1900 and the Huia (Heterolocha acutirostris) last reliably reported in 1907, have become
extinct within historic time. A number of other passerines, including the North Island Bush
Wren (Xenicus longipes stokesi), Stead's Bush Wren (X. 1. variabilis), and both New Zealand
thrushes (Turnagra capensis subspp.), are, likewise, probably now extinct (see Turbott 1990).

New Zealand's endemic passerine families are of particular taxonomic interest. Recent
classifications place the wrens (Acanthisittidae) with the suboscines, implying a relationship
with some South American group (Raikow 1987). They may, however, have been an early
arrival of Australian suboscine stock (which itself may have reached Australia via East
Antarctica - Cracraft 1973) - but see Sibley et al, (1982). There have been three genera of
wrens described from living specimens: Acanthisitta, Xenicus and Traversia (sometimes
synonymised with Xenicus). All four species in these genera are now known from subfossil
deposits in both the North and South Islands. Fleming (1979) suggested that the Rock Wren
(X. gilviventris) was an alpine derivative of the forest-dwelling Bush Wren (X. longipes), thus
displaying the same type of speciation pattern seen in the Nestor parrots. A new genus of
acanthisittid, Pachyplichas, has recently been described from subfossil remains, there being two
distinctive species one (P. jagmi) from the North Island the other (P. yaldwyni) from the South
(Millener 1988). A further new acanthisittid genus (not yet formally described - see Millener &
Worthy, in press) is known from two South Island subfossil locations.

Various relationships have been suggested for two other endemic passerine groups. The
Wattlebirds (Callaeatidae) have been placed with a group of Australian and Papuan corvid-like
families (Mayr & Amadon 1951, but see also Williams 1976), while the New Zealand
Thrushes are now placed in the Paradisaeidae (Olson et al. 1983). They, together with the
Wrens, probably represent at least three colonisations during the Tertiary (Fleming 1962a).

The majority of the remaining endemic New Zealand passerines are likely to be of
Australian origin. Some of the endemic genera still show clear relationship to Australian
forms (e.g. Anthornis to Meliphaga, Notiomystis to Meliornis - Fleming 1962a).

Many are endemic species assigned to Australian genera (e.g. Petroica, Gerygone and
Rhipidura), which themselves are the products of adaptive radiation within Australia from a
limited number of ancestral sources (Sibley 1976). Fleming (1950) suggested that the New
Zealand representatives of Petroica have resulted from two invasions by birds of the same
ancestral stock. The robins (subgenus Miro) seem to be derived from an early, perhaps
Pliocene, invasion of Petroica stock, while the tits (Petroica s. str.) represent a later, perhaps
Early Pleistocene, one.

A notable feature of the forest-dwelling passerines is the large number of species which
have developed distinct island forms on either side of Cook and Foveaux straits or on outlying
islands. Mohoua with M. albicilla (Whitehead) and M. ochrocephala (Yellowhead), Gerygone
with G. igata (Grey Warbler) and G. albofrontata (Chatham Island Warbler) and Bowdleria with
B. punctata (Fernbird) and B. rufescens (Chatham Island Fernbird) are three genera with
specifically distinct derivatives.

Some species (e.g. the Fernbird, Bowdleria punctata, and the Tit, Petroica macrocephala)
have as many as five subspecifically distinct island races.

1334 - MILLENER

The passerine taxa found most often in subfossil deposits are usually the larger forms,
whose more robust bones render them more readily preserved. To some extent collecting
techniques may also exaggerate this bias toward the larger taxa, but it would seem that in many
cases the more frequent occurrence of certain species does truly reflect a greater natural
abundance. The Crow (Palaeocorax moriorum), Kokako (Callaeas cinerea), Tui (Prosthemadera
novaeseelandiae) and Saddleback (Philesturnus carunculatus) are the predominant passerines in
natural assemblages, while the Tui, due no doubt to pronounced selectivity of hunting, is by
far the most abundant in midden deposits. In contrast, the Huia (Heterolocha acutirostris),
although second only to the Crow in size, is much less frequently found (see Williams 1976,
Millener 1981b).

SUMMARY

The evolutionary development of New Zealand's avifauna has been influenced by the
interplay of geologic, geographic, climatic and ecological factors. At the outset, New Zealand's
position as a segment of the temperate Gondwanan supercontinent during the Mesozoic enabled
it to take part in faunal and floral exchanges which occurred among the then contiguous
southern continents. New Zealand's separation in the Late Cretaceous came at a time that
allowed some of the archaic elements of its biota to establish themselves, but still early
enough to prevent colonization by snakes and predatory mammals. New Zealand's drift north
and its consequent isolation since the early Tertiary has meant that any subsequent colonization
had to be entirely transoceanic. “Sweepstakes dispersal", assisted since the Oligocene by the
circum-Antarctic current and its associated Westwind Drift, has long been and continues to be
important for colonists of Australasian origin. The archipelagic nature of the New Zealand
landmass throughout the Tertiary and Pleistocene doubtless fostered adaptive radiation and
speciation, but also caused numerous extinctions amongst the original inhabitants and
incoming colonists alike. The monotypic and relict nature of many of New Zealand's endemic
genera is more likely an indication of Pleistocene extinction of their congeners, than of
conservatism and lack of speciation in the Tertiary. The climatic extremes of the ice ages,
which must have decimated the warm-adapted Tertiary biota, also provided stimulus for
speciation among those elements which survived. The post-glacial fauna must have been a
remarkable melange, comprizing survivors of the Pleistocene decimations and new transoceanic
colonists, expanding to fill niches left vacant by those which had succumbed. This fauna with
its many aberrant, specialised and frequently flightless forms had evolved in splendid isolation
on arichly forested, predator-free landmass and would have been ill-adapted to survive what was
to be for many of its members the final episode in their evolutionary history - the arrival of
man,

Man, by his extensive clearance of forests, by his introduction of mammalian predators and
to some limited extent by his own predation, wrought ecological changes of such great
magnitude and rapidity that almost half New Zealand's complement of terrestrial birds was
exterminated in less than a thousand years.

Man's influence continues to have an almost universally detrimental effect on the remaining
bird fauna, although natural immigration continues to partially offset depletion.

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OLSON, S.L., 1984. The relationships of the extinct Chatham Island eagle. Notornis 31(4): 273-277.

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OLSON, S.L., PARKES, K.C., CLENCH, M.H. & BORECKY, S.R., 1983. The affinities of the New Zealand
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OLSON, S.L. & ZUSI, R.L., in prep. The anatomy and taxonomic status of Apterornis (Aves:

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OWEN, R., 1839. Notice of a fragment of a femur of a gigantic bird of New Zealand. Trans. Zool. Soc. Lond.
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PAULIN, C.D., 1973. Sub-fossii avian remains from two limestone caves in North , Taranaki. Mauri Oral:
95-98.

POCKNALL, D.T. & MILLENER, P.R., 1984. Vegetation near Lake Poukawa prior to the Taupo Eruption. J/.
R. Soc. N.Z. 14(2): 151-157.

POWELL, A.W.B., 1979. New Zealand Mollusca. Collins, Auckland.

PRICE, T.R., 1963. Moa remains at Poukawa, Hawkes Bay. NZAA Newsletter 6(4): 169-174.

PRICE, T.R., 1965. Excavations at Poukawa, Hawkes Bay, New Zealand. NZAA Newsletter 8(1): 8-11.

RAIKOW, R.J., 1987. Hindlimb myology and evolution of the Old World suboscine passerine birds
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1338 - MILLENER

REID, B. & WILLIAMS, G.R., 1975. VII. The Kiwi, In Biogeography and Ecology in New Zealand. G.
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RICH, P.V., 1975a. Antarctic dispersal routes, wandering continents and the origin of Australia's non-
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RICH, P.V., 1975b. Changing continental arrangements and the origin of Australia’s non-passeriform
continental avifauna, Emu 75(3): 97-112.

RICH, P.V., 1976. The history of birds on the island continent, Australia. Proc. 16th Int. Ornith. Congress.
Canberra: 53-65.

RICH, P.V. & MCEVEY, A.R., 1977. A new owlet-nightjar from the eariy to mid-Miocene of eastern New
South Wales. Mem. natn. Mus. Victoria 38: 247-254.

RICH, P.V. & SCARLETT, R.J., 1977. Another look at Megaegotheles, a large owlet-nightjar from New
Zealand. Emu 77(1): 1-8.

RICH, P.V. & VAN TETS, G.F., 1981. The fossil pelicans of Australasia. Rec. S. Aust. Mus. 18(12): 235-
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RICH, T.H.V., 1975. Potential pre-Pleistocene fossil tetrapod sites in New Zealand. Mauri Ora 3: 45-54.

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SCARLETT, R.J., 1962. Subfossil records of the Little Grey Kiwi in the North Island. Notornis 10(2): 84-
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SCARLETT, R.J., 1966. A pelican in New Zealand. Notornis 13(4): 204-217.

SCARLETT, R.J., 1967a. A subfossil record of a Barn Owl in New Zealand. Notornis 14(4): 218-219.

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WEISSEL, J.K., HAYES, D.E. & HERRON, E.M., 1977. Plate tectonics synthesis: the displacements between
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NEW ZEALAND QUATERNARY AVIFAUNA - 1339

WILLIAMS, G.R., 1962. Extinction of the land and freshwater inhabiting birds of New Zealand. Notornis
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New Zealand.

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Dinomithiformes) older than the Otira Glaciation. J. Roy. Soc. N. Z.

YALDWYN, J.C., 1956. A preliminary account of the subfossil avifauna of the Martinborough caves. Rec.
Dominion Mus. 3: 1-7.

YALDWYN, J.C., 1958. Notes on the environment and age of the subfossil deposits of the Martinborough
caves. Rec. Dominion Mus. 3: 129-135.

1340 - MILLENER

APPENDIX I
AVIAN FOSSIL REMAINS FROM PLIO-PLEISTOCENE
DEPOSITS OF NEW ZEALAND

[SOUTH | [CHATHAM ]
ISLAND || iSLANDS
DINORNITHIFORMES
Anomalopteryx didife-mis E
Megalapteryx didinus E i+ +/4)4
Pachyornis mappini e|[+{+]+ +[] [ 1
P. australis El! [[+[+[+
P. elephantopus E| [[+[+l+[+
Emeus crassus E +/+/+/ +1)
Euryapteryx curtus E[[+[+/+]+ | i| |
E. geranoides Ej [+{+(+] +] |+/+|+|+
Dinornis struthoides Ell+ +)4l4ll/+4)4)4l4+
D. novaezealandiae E}[+)+{[+}+])+]+]/4]+
D. giganteus E)|+/+}+/4+][+{+]) +] + |
APTERYGIFORMES
Apteryx owenii [[+[+|+ +[[+ +]+]4]]
A. australis +] +/+] +) [+/+] +] +] |
A. haastii A | [+L [+
SPHENISCIFORMES
Aptenodytes patagonicus [] +
Megaayptes antipodes {| +14 +
Eudyptula minor +(4+/4+ +44 +l4l4
Eudyptes cf. pachyrhynchus te: +/+ +) + +] +]
PODICIPEDIFORMES
Podiceps cristatus af] [+ [+[+
Poliocephalus rufopectus + +1]+ 4/4] +
PROCELLARIIFORMES
Diomedea exulans / epomophora +{ [+/+ za ou
D. epomophora +/+
2D. chrysostoma T [+
2D. chlororhynchos i] +
D. bulleri +/+] | Tol +
D. cauta C+ t [+]+0) +/+ +/+
Phoebetria palpebrata [ {I U +
Macronectes halli +/+ +
Fulmarus glacialoides +
Daption capense +/+ +/+]
Pterodroma macroptera +\+ ry +
P. lessonii + T T41 r+ [+
?P. externa | | | +/
P. inexpectata Trel+lalel) [4l4l4+ +/+
P. neglecta a [| Ty
+ 4 +— 2 | f— 4
P. magentae IT | [ [+[+]+]
2P. ultima | +
P. leucoptera oT Al + TT 1
P. cookii +l+[4{] [+ 1] |
P. nigripennis TT [|
P. axillaris th +/+
?Halobaena caerula [+ | [ lpet

NEW ZEALAND QUATERNARY AVIFAUNA - 1341

NORTH SOUTH CHATHAM |
ISLAND ISLAND ISLANDS

iS] S A
PROCELLARIIFORMES TE dfn br] fl

Pachyptila vittata +)+ +1414 ope
P. salvini [|
P. turtur i+]: |
P. crassirostris iM TY 1

+

i

i+{+[+
+

+
+
+
+
+
+
Bs

?Procellaria cinerea
P. parkinsoni

P. westlandica

P. aequinoctialis a
Puffinus carneipes [+[+
P. bulleri
P. griseus

fl
P. tenuirostris {|
++
44

+[+]+]+[+[+/+]

+|+[+

P. gavia/huttoni

P. assimilis
Oceanites oceanicus
Garrodia nereis
Pelagodroma marina
Fregetta tropica |
Pelecanoides urinatrix +/+ +l4]+

PELECANIFORMES
Pelecanus novaezealandiae El|+/+)/+}+ +/+
Sula bassana | ++ + +
S. dactylatra +
Phalacrocorax carbo +4) + +| i+

P. vari Tal tel+ aE
. varlus |

oe fe +]

+

+

+

|
cE

Bn
+
+
+
+

+/+|+]+
+

=
b

P. melanoleucos Lt | +] |
Leucocarbo carunculatus i {|
Stictocarbo punctatus +{+]

CICONIIFORMES
Egretta alba +| f+}+ +
E. sacra I +
Botaurus stellaris + | LO |
Ixobrychus novaezelandiae E||+ + ml

ANSERIFORMES
Cygnus summerensis
Cnemiornis calcitrans
C. gracilis

Pachyanas chathamica
Euryanas finschi
Malacorhynchus scarletti
Biziura delautouri
Tadorna variegata
Anas superciliosa

2A. gibberifrons

A. aucklandica

A. rhynchotis

1342 - MILLENER

—$<$< {= “= ——
NORTH SOUTH CHATHAM
ISLAND ISLAND ISLANDS

ANSERIFORMES
Hymenolaimus malacorhynchos
Aythya novaeseelandiae
?Oxyura australis

Mergus australis

FALCONIFORMES
Circus approximans

C. eylesi

Harpagornis moorei
Haliaeetus australis
Falco novaeseelandiae

GALLIFORMES

Coturnix novaezelandiae El] +[+ +[+ ; 2] +i + +] |
GRUIFORMES a rat! Pe
Gallirallus philippensis TL FRET ]
G. dieffenbachii e|| | | IT | ial [+[+]+
G. modestus E|| : [ Vela]
G. australis +++ +++ [T+]
Capellirallus karamu Ell+ (+i +) +] ] ja ipl
Diaphorapteryx hawkinsi E ater
Porzana pusilla [ (+t | Tele TT +|+
P. tabuensis “T+t | [+f [+ lecieald te
Porphyrio porphyrio mili +| + +{] oh {|
P. mantelli ++ leit

: 4} —_| jt} —+
Gallinula hodgenorum Eli t+{t+) [+l +) +{+] +] | |
Fulica prisca El] [++] [[+[+[+]+
F. chathamensis Ell | Tete sled TT +[+]+
Aptornis otidiformis El|+/+{+] +1] 7 [
A. defossor E [+ +fals al
CHARADRIIFORMES 7 et -
Haematopus ostralegus LE} [+] Y [+ [+ [+] | Lal
H. unicolor ee a
H. chathamensis mime in | id
Charadrius obscurus +t | Ei asa ta teak IT im

Charadnus bicinctus

+
. . . cond a a a pains S| A eal Pa jae
Thinornis novaeseelandiae mill {ff il dt aril

Anarhynchus frontalis oo | | |
4 ae = rset — aa
Numenius phaeopus | +) + +/+] |
; = = ’ + ++
Limosa lapponica + il lier: inet Pay
. . —TT a "i ~y ‘of
Arenaria interpres eee: IT el Eli (pak

C. chathamica

Calidris canutus
Himantopus himantopus /
novaezelandiae
Catharacta skua

?C. maccormacki
?Stercorarius longicaudus

Coenocorypha aucklandica Se Gad fea Vail cba laa bc
rpsiers et TEAC
na

NEW ZEALAND QUATERNARY AVIFAUNA - 1343

NORTH || SOUTH | |CHATHAM
ISLAND || ISLAND || ISLANDS
CHARADRIIFORMES ag ca ad LF dt en
Larus dominicanus LI +[+i{ [+]+/+ +/+
L. scopulinus TET [+]+tT [+]4 + T+]+
9L. bulleri el
Hydroprogne caspia [[+l+lt [ [+l+ [
Stema vittata / paradiseae TT +P
S. albostriata EES eT +[+
S. nereis Tier TT =
S. stnata LL | [+{+ +/4]4T] +] 4
COLUMBIFORMES
Hemiphaga novaeseelandiae T[+]+]+[+][+]+]+]+ +/+) 4+
PSITTACIFORMES
Strigops habroptilus +[+]+]+[[+[+]+]+
Nestor meridionalis +/+) 4) 4+!) +) 4+)4)+ +/+
N. notabilis [ +[40 [| |
Cyanorhamphus novaezealandiae [[+[+[+]4+] [+/+ +|+| +/+
C. auriceps +) +/+) + +/+] + +/+
CUCULIFORMES
Chrysococcyx lucidus [ TI [| +[ |
Eudynamys taitensis +/+ Lt
STRIGIFORMES
Ninox novaeseelandiae [FGeETAeE ETT
Sceloglaux albifacies E{/+/+l+l+i/+l+[+/+ +?
CAPRIMULGIFORMES
Megaegotheles novaezealandiae E|] +{+ +/+
CORACIIFORMES _
Halcyon sancta IL [+L [+I [+ il
PASSERIFORMES -
Acanthisitta chloris T +L TT eT T+] |_|
Xenicus longipes LI +/+]+ +] |
X. gilviventris | +| +/ | |
Traversia lyalli JE] | [+ [| [+ |
Pachyplichas jagmi Ell [++] | |
P. yaldwyni el | TT it f+] | UT
Anthus novaeseelandiae [ [[+f+f+i+]) [+l f+ip | i+i+
Bowdleria punctata | [+/+ [+] [+i] | |
B. rufescens El | |_| {| |+| +/+
Mohoua novaeseelandiae Ltt | LL t+ | | |
M. albicilla LI Le ia
M. ochrocephala | _|
Gerygone igata mut
G. albotrontata LI | 1
Rhipidura fuliginosa i |
Petroica macrocephala | {||
P. australis | | L 1 4
P. traversi L | |

1344 - MILLENER

NORTH SOUTH. | |CHATHAM
ISLAND ISLAND || ISLANDS
PASSERIFORMES
Notiomystis cincta [+|+
Anthornis melanura +/+) + + [+] [++] +
Prosthemadera novaeseelandiae tl +l+ +[ [+ +[+1+]/ +/+ +
Philesturnus carunculatus ti+}+i+] [+i titi +i
Heterolocha acutirostris E +{+{+7 | |
Callaeas cinerea +{+[4+[+] [+ +] 4) +1 |
Turnagra capensis ET] +/+) 4+] [4+) 4+] 4+]+
Palaeocorax moriorum E|i+}+ +[4[]+ +) +) + +/+

Legend: Nomenclature follows Turbott (1990) except where more
recent revisions apply

Extinct species

Australian species now extinct in New Zealand

Identification uncertain

Swamp / alluvium / colluvium

Cave

Dune sand / loess

Midden - these data should be interpreted with caution as, especially
in eroding dune-midden sites, unequivocal evidence for primary
association of avian remains with midden debris is frequently lacking.

mm
*

=Zo00a~

CHAPTER 28

VERTEBRATE FOSSIL
FAUNAS FROM ISLANDS IN
AUSTRALASIA AND THE
SOUTHWEST PACIFIC

Charles Meredith!
INtrOductiONn ...... 0. cece cee cee ec eececceseuccceuscescecs 1346
Coastal Islands (tases sca deasdee Beldeciote doses 1346
The Bass Strait Islands.............c.ccccecesceeee 1346
Kangaroo Island ............cceeecccseeeecceeeeeceees 1349
Other Southeastern Coastal Islands............ 1350
Western Australian Islands...........e.ececceeee 1350
OGeariic ISAS wre Soe Use heb dood weee veloc s 1350
Norfolkelsland: :Scvctes ar, testa: Woes db 1350
Lord Howerlslangd). 23. ieeies cescssseies ovtdase 1354
Kermadec Islamds.i.c. ccc iccssaccsscd ccctevsevectede 1355
New Caledonia .............ccccsscsesecesccececesace 1355
Fig t STAM Sb, ioige Suivetesecitastitites cone sed Mestwecb ec 1355
COOK ISIAMES rst 3. deltilendts nth death sect thse tae 1355
Chatham Islands ................sccecsecececceceeees 1356
NGWZGALAT | satis sins ane sincier odteinewc ne andl Pekebhde's 1359
Macquarie Island .............ccecceeceesceeeceeeeees 1360
DUS CUSSION 3 ardor ies rere Bie 6 dele bs soe eede ceil en edou> 1360
Taxonomy and Phylogenetic Relationships. 1362
Taphonomic Studics...........ceeceeeeeeeeee scene 1362
Seabird Biogeography .............cc cece eeeeeee ees 1362
Biogeography of Insular Landbirds............. 1363
ROLERENCES cist icsse aches Satenass deodeultettecbeies cate 1365
ADDON GI cach wislvncelierat aban able abet eGoant sh bd 1369
ATC Se terpt totes 52m is Sheep lia steal salto whe te isi eM ede 1374

1 Biosis Research Pty Ltd, 127 Queens Parade, Clifton Hill, Victoria 3068), Australia.

1346 - MEREDITH

INTRODUCTION

The study of extant island faunas has occupied a special place in biology since the time of
Wallace and Darwin, contributing significantly to modem theories of speciation, biogeography
and community structure. It is, therefore, surprising that the fossil and sub-fossil faunas of
islands have received little attention, even though the existence of some has been known for
over a century. This is now changing. For example, recent work on the rich fossil faunas of
the Hawaiian chain (Olson & James 1982a, b) and the West Indies (Olson 1978, Pregill &
Olson 1981) has shown how dramatically different Late Pleistocene and Holocene island faunas
can be compared to those of the present. Such studies have cast doubts on the interpretation of
neontological data from islands and have generated interest world-wide in island palacontology.

In this Chapter, I review the results of palacontological studies of vertebrates on
Australasian and Southwest Pacific islands, discuss their relevance to a variety of biological
problems, and make some suggestions for future investigations. The section on the most
outstanding island fauna of the region, that of the Chatham Islands (Appendix I), is contributed
by Phil Millener.

The Chapter is strongly biased towards avian fossils. For the purpose of this review I
classify islands into two types: (i) coastal - those that were joined to the Australian mainland
during Pleistocene sea level lows, and (ii) oceanic - those that lacked such a mainland
connection, The fossil faunas of New Guinea (Plane, this volume) and Tasmania (Baird, this
volume) are not dealt with as they are covered elsewhere in this book and are best considered as
continental faunas. The fossil faunas of New Zealand (Fordyce, this volume) and New
Caledonia (Balouet, this volume) are also dealt with in detail elsewhere in this volume, but, as
they are relevant to the analysis presented here, I have briefly summarised them.

COASTAL ISLANDS

Fig. 1 shows the localities of Australian coastal islands that have produced fossil faunas.
These faunas are summarised in Appendix I. It is immediately clear that all the major sites are
off the southeastern coast. This reflects the intensity of exploration in that region. The small
amount of material known from the western Australian islands suggests that their systematic
exploration would prove fruitful. Elsewhere, there are hundreds of islands along the Great
Barrier Reef and in Torres Strait, some of which are likely to produce fossil material. Further
to the north, the islands around New Guinea merit exploration.

THE BASS STRAIT ISLANDS

The Bass Strait islands are granitic remnants of a Late Pleistocene landbridge which
connected Australia and Tasmania. They occur along two submarine ridges, the Bassian Rise
in the east (the Furneaux Group and other small islands) and the King Island Rise in the west
(King Island and the Hunter Group). The Bassian Rise extends from Wilsons Promontory to
northeast Tasmania and is presently submerged to a depth of less than 60m along most of its
length, with its greatest depth close to Victoria. The King Island Rise, extending from
northwest Tasmania to King Island, is submerged to similar depths but is separated from
Victoria by the western extension of the Bassian Depression (Hope 1973; Fig. 2). With rising
sea levels after the Last Glacial Maximum, the Bassian Depression would have been rapidly
resubmerged after only brief exposure, cutting the connection between King Island and Victoria.
Around 12,000 to 13,500 years before present (yBP) the Bassian Rise would have been

VERTEBRATE FOSSIL FAUNAS FROM ISLANDS - 1347

inundated by the rising sea between Flinders Island and Wilsons Promonto
ry (Hope 1973). The
land between King Island and northwest Tasmania would have been flooded between 10,000 and

12,500 yBP, and that between the Furneaux Group and northwest Tasmania be
10,000 yBP (Hope 1973). P mania between 8,500 and

Figure 1. Map showing the location of Australian coastal islands that have produced vertebrate fossil
deposits: 1, Bowen Is.; 2, Deal Is.; 3, Erith Is.; 4, Flinders Is.; 5, East Kangaroo Is.; 6, Cape Barren Isig 7,
Preservation Is.; 8, Long Is.; 9, Maatsuyker Is.; 10, Hunter Is.; 11, Three Hummock Is.; 12, King Is.; 13,
Kangaroo Is.; 14, Brothers Is.; 15, Salisbury Is.; 16, Houtman-Abrolhos Ids.; 17, Dirk Hartog Is.; 18, Bernier
Is.

Nearly all the larger islands and some of the smaller have produced Late Pleistocene and
Holocene fossil material of reptiles, birds and mammals from sand-dunes, caves and swamps.
The mammal material has been reviewed by Hope (1973). She found that, of the 24 species of
mammals found as fossils on the islands, 14 were unknown there in historic times. None were
endemic species. She concluded that the composition and past distribution of the mammal
fauna was consistent with an initial colonisation of the developing landbridge in the Late
Pleistocene from Tasmania, and this Tasmanian-derived fauna was then resistant to
displacement by mainland elements once the landbridges finally reached Victoria.

The avian material from King Island and the Furneaux Group has received little attention.
Most deposits are dominated by bones of the Short-tailed Shearwater Puffinus tenuirostris,

1348 - MEREDITH

Figure 2. The bathymetry of Bass Strait.

which is a summer-breeding seabird, extremely abundant throughout Bass Strait. The
abundance of material of this species has tended to mask the variety of other bones that are
present (C. Meredith, pers. obs.). Hope's (1969) references to unidentified bird bones from
most of her sites indicate that further excavations and analyses of already collected material
would be profitable. Shane Parker (South Australain Museum) is presently reviewing the
dwarf emu of King Island (see Parker 1984). This species, extinct soon after European
colonisation, is well-represented in a number of sites on that island.

Aboriginal middens on Hunter Island contain abundant bird and mammal material (Bowdler
1984, 1974, O'Connor 1982). The age of these middens varies from Late Pleistocene to less
than 1,000 yBP. The birds that have been identified in these deposits are a subset of the
present avifauna with the exception of a penguin, very close in size to the Rockhopper
Penguin, Eudypies chrysocome, that has been described as Tasidyptes hunteri (van Tets &
O'Connor 1983). Tasidyptes was present as recently as 760470 yBP and is apparently the only
penguin known to have become extinct in the Holocene. It is not known if it bred on Hunter
Island. The past diversity of mammals on Hunter Island was much greater than at present
(Appendix I). Bowdler (1984, 1974) states that the present mammal fauna consists of at least

VERTEBRATE FOSSIL FAUNAS FROM ISLANDS - 1349

three species, with one other probably recently extinct. Fifteen species occur in the midden
deposits. Three of these (Cercatetus nanus, Sminthopsis leucopus, Mastacomys fuscus) are
small cryptic species that might well still occur on the island but might have remained
undetected. Thus, even if these three species have survived until historic times, the mammal
fauna has experienced a decline of at least 50%, assuming that there has been no turnover of
species.

The faunal differences between the Late Pleistocene Cave Bay site and the Holocene
Stockyard site indicate that mammalian species richness had declined from fourteen in the
Pleistocene to ten in the Holocene, with the loss of the two largest herbivores (Macropus
rufogriseus, Vombatus ursinus), the largest carnivore (Dasyurus sp.) and the largest possum
(Pseudocheirus peregrinus). Aborigines were present throughout this period, at least as
seasonal visitors, and it is possible that harvesting or anthropogenic habitat alteration by fire
was the cause or contributed to this decline. Climatic change is unlikely as a cause, as all
these species have fairly wide climatic tolerances. The decline could also represent relaxation
to a new equilibrium following isolation and reduced island area, but such an interpretation is
difficult to prove (see below).

Close to Hunter Island is the small, rocky Albatross Island. Sealers and feather collectors
have left a large pile of bones of the Shy Albatross Diomedea cauta, and the Australasian
Gannet, Morus serrator, ina cave on this island. The floor of this cave seems a very likely
site for sub-fossil seabird material as well (C. Meredith, pers. obs.).

KANGAROO ISLAND

Kangaroo Island lies west of Bass Strait, at the mouth of St Vincent Gulf, 15 km off the
South Australian coast, from which it is separated by the Backstairs Passage (35 m deep) and
Investigator Strait (25m deep). It was connected to the mainland through much of the Last
Glacial Maximum, until rising sea levels opened the Backstairs Passage between 9,300 and
10,500 yBP and Investigator Strait between 8,800 and 9,900 yBP (Hope et al. 1977). Bones
of an extinct (probably soon after European colonisation) dwarf emu have come from a number
of sites on the island. Originally regarded as conspecific with Dromatus ater of King Island,
this population has recently been separated as D. baudinianus by Parker (1984). It is
interesting that these two flightless species are the only endemic bird species known from the
southeastern Australian islands.

The Seton Rock Shelter site (Hope ef al. 1977), in the south of the island, has produced a
wealth of mammal, bird and reptile remains from deposits dating from more than 16,000 yBP
to about 10,000 yBP. Two of the four stratigraphic units also contain evidence of human
occupation. Twenty-eight species of mammal were identified from this site, only seven of
which still occur on the island. Many of the extinct species are mammals of grasslands or
open vegetation, a natural habitat now lacking on the island. About 40 species of birds have
been identified, of which five waterbirds and eight landbirds are not known historically from
the island. As with the mammals, most of the extinctions amongst the landbirds were of
species typical of dry, open country. Although a number of the identifications of avian
material require re-evaluation in the light of the better comparative collections now available,
th overall picture is one of a decline in animal species associated with open vegetation and in
faunal richness over the last 10,000 years. Hope et al. (1977) suggest that this was due to a

combination of climatic change, isolation due to rising sea levels and a change in fire
frequency caused by a decline in the Aboriginal population.

1350 - MEREDITH

OTHER SOUTHEASTERN COASTAL ISLANDS

Guano deposits on Brothers Island, in St Vincent Gulf, contained a small amount of
mammal and bird material (Johns 1966; Williams 1980). The avian material was referred to
Genyornis by Rich (1979), but Patterson (1983) referred it to emu Dromaius, without
specifying which species.

Aboriginal middens on Bowen Island, off southern New South Wales, contained six species
of seabirds, all of which still occur in the area (Blackwell 1982). Blackwell's identification of
Pachyptila turtur has been altered to Pachyptila sp. in Appendix I, due to the considerable
difficulties of specific identification in this genus.

Small numbers of seabird bones and some fur seal material have been found in
archaeological deposits on Maatsuyker Island, off Tasmania (Vanderwal & Horton 1984).

WESTERN AUSTRALIAN ISLANDS

The collections of the Western Australian Museum holds a small amount of avian material
collected from dunes and caves on Dirk Hartog Island and from North Island in the Houtman
Abrolhos Archipelago, consisting of several species of seabird and some Emu eggshell. This
material suggests that it would be well worthwhile to investigate these sites further and to
search for others.

Alex Baynes (Western Australian Museum) has collected a small number of unidentified
bird bones, along with a variety of mammal bones, from Bernier Island, Shark Bay. Abundant
bird bones have recently been found in spongolite on Salisbury Island in the Recherche
Archipelago (A. Burbidge & N. McKenzie, pers. comm.), but only small amounts of material
have been collected due to the difficulties of access to the island. Guano deposits on some of
the northwestern islands are reputed to contain avian skeletal remains (R. Johnson,
pers.comm.).

OCEANIC ISLANDS

With the exception of the Fiji group and Macquarie Island, all the oceanic islands discussed
here share a similar origin, They are all remnants of a large continental block, Tasmantis
(=Lord Howe Rise, Norfolk Rise, New Caledonia and New Zealand), which was joined to
Austro-Antarclica during the Triassic and Jurassic, but which then began to split off in the Late
Cretaceous, forming the Tasman Sea (Coleman 1980). Tasmantis was split into the Lord
Howe Rise and the Norfolk Rise about 65 million years ago by the formation of the New
Caledonia Basin, and, although there has been further tectonic movement, this remains
the basic structure that exists today (Fig. 3). Despite this common tectonic origin, there has
been a great deal of diversity in the later development of these islands, with New Zealand and
New Caledonia being true continental remnants, while the Chathams are the result of
vulcanism on a submerged continental plate. Norfolk and Lord Howe Islands are volcanic
protrusions of submarine ridges, and Macquarie Island is a recently uplifted segment of oceanic
ridge. The fossil faunas of the oceanic islands discussed below are summarised in Table 1, and
localities are shown in Fig.4.

NORFOLK ISLAND

Norfolk Island (3,450ha) is an isolated basaltic outcrop of the Norfolk Rise, 1,370 km east
of Australia, the result of Late Pliocene volcanism (Jones & McDougall 1973). Most of the
island is covered by afossiliferous acidic soils, but a small coastal lowland in the southeast,

VERTEBRATE FOSSIL FAUNAS FROM ISLANDS - 1351

known as Kingston, has produced fossils from sand-dunes and beach rock. Nepean Island, one
kilometre offshore, is an erosional remnant of the Kingston lowlands and its eroding sand
capping also contains fossils. The beach rock, a massive calcarenite tentatively dated at
1,450+90 yBP (Veevers 1976), contains scarce fragments of seabird bones. Material from the
Kingston dunes is abundant and well-preserved. Four radiocarbon dates, from charcoal collected
from the top of the fossiliferous layer, cluster around 800-850 yBP. These fossiliferous sands
overlie a saprophytic clay from which dates of 6,870+30 yBP (Veevers 1976) and 4,400+90
yBP (Rich et al. 1983) have been obtained. There are no dates from the Nepean Island
deposits.

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Figure 3. Map showing the present position of the submarine ridge systems in the southwest Pacific
(ridges indicated by 2000m contour).

1352 - MEREDITH

Table 1. Islands that have produced fossil faunas, other than those detailed in Appendix I.
Unless otherwise referenced, information is from Brodkorb (1963, 1964, 1967, 1971, 1978).

St Lawrence

Aleutians

Alaskan coastal islands

Vancouver Island

Anacapa/San Nichola

Isla de Guadalupe - Hubbs & Jehl (1976)
Hawaiian Islands - Olson & James (1982a,b)
Aldabra Atoll - Harrison & Walker (1978)
Galapagos - Steadman (1986, 1981)

Easter Island - Carr (1980)

10. Henderson Island (Steadman & Olson 1985)
11. Tikopia (REFS)

12. Solomon Islands

14. Iceland

15. Newfoundland

16. Nova Scotia

17. Bermuda

18. Bahamas - Olson (1982)

19. West Indies - Olson (1978); Pregill & Olson (1981)
20. Fernando de Noronha - Olson (1981)

21. Ascension - Ashmole (1963); Olson (1977)
22. St Helena - Olson (1975a)

23. Seymour Island - Simpson (1975); Woodburne & Zinsmeister (1982)
24. Ellesmere Island

25. Great Britain

26. Jersey

27. Canary Islands

28. Balearic Islands

29. Corsica

30. Sardinia

31. Malta

32. Crete - Malatesta (1980)

33. Samos

35. Madagascar - Mahe (1972)

36. Reunion

37. Mauritius - Milne-Edwards (1874)

38. Rodriguez - Milne-Edwards (1874)

39. Amsterdam Island - Jouanin & Paulian (1960)
40. Sumatra

41. Java

42. Japan

43. Iki

44. Doiga-hama

45. Okinawa

46. New Guinea

8G 00.00 6 GN Ne SOUND

VERTEBRATE FOSSIL FAUNAS FROM ISLANDS - 1353

Figure 4. Map of the southwest Pacific region showing those oceanic islands that have vertebrate fossil
deposits: 1, New Caledonia; 2, Fiji Ids.; 3, Mangaia Is.; 4, Lord Howe Is.; 5, Norfolk Is.; 6, Kermadec Ids.; 7,
Chatham Is.; 8, New Zealand; 9, Macquarie Is.

The fauna from these sites comprises over 34 species of birds, two reptiles and two
mammals, the Pacific Rat Rattus exulans anda vespertilionid bat (Meredith 1985b). Most of
these species are known historically from the island, but six seabirds (four or five apparently
once breeding on Norfolk), a small, flightless woodhen (Rallidae), the Sub-Antarctic Snipe
(Coenocorypha prob. aucklandica), and several waders (Charadriiformes) are only recorded as
fossils. Three of these are new species: a large booby, Sula tasmani (van Tets et al. 1988), a
medium- sized gadfly petrel, Pterodroma sp., and the woodhen, Gallirallus sp. There is a
single bone of the Rockhopper Penguin Eudyptes chrysocome, a sub-Antarctic breeder that
occurs as sn occasional vagrant in the sub-tropics.

The Pacific Rat is a species commensal with Polynesian man and has been distributed
throughout the Pacific Islands by voyagers in their canoes. Although Norfolk Island was
uninhabited when first sighted by Europeans in 1774, there is archaeological evidence of a
former Polynesian presence (Specht 1984). The presence of the rat as a fossil below the 800
yBP charcoal band provides a minimum date for this early visitation (Meredith et al. 1985).

1354 - MEREDITH

Although the fossil fauna is largely similar to the island's modern fauna, the recent loss of
four or five breeding procellariiform and pelecaniform seabirds is notable. The present number
of breeding species from these orders is six. As two of the currently breeding species have only
colonised in the last few decades, this represents a decline of 50% or more. Recent declines in
seabird faunas of this magnitude are typically shown by island fossil data, but the causes of
these declines are often uncertain. Recently, archaeological studies on Norfolk Island have
located the refuse dumps of the first European colony, which contain many bird bones. A
preliminary analysis (Meredith & Varman, unpubl. data) shows that at least two of the locally
extinct seabirds that were not recorded historically survived until European colonisation
(Pterodroma pycrofti and Pterodroma new sp.). Harvesting of seabirds by these early settlers
apparently caused the extinction of some species before they could be catalogued by zoologists.
Further collecting may well reveal other species which were similarly affected.

Predation by the Pacific Rat has often been invoked as cause of extinction in seabirds on
islands, but the Norfolk Island data provide no evidence for any such predation (Meredith, in
prep.). Rat bones are absent from the lower third of the fossiliferous layer in the Kingston
dunes, but are commonly present in the upper two-thirds. There is no detectable difference
between the composition of the fauna pre- and post-rat, nor is there any evidence of tooth
marks on over 1,500 bones. Of course, the rats may have taken eggs or young birds whose
bones have not survived, but the archaeological evidence and the modern fauna show that at
least some seabirds, ranging in size from much smaller than the Pacific Rat to much larger, and
from burrow to surface nesters, have survived 800 years sympatry with the rat.

The faunal list in Appendix I contains a number of changes from the preliminary lists
presented by Rich & van Tets (1982) and Rich et al. (1983). Daption capense, in the
preliminary list, is in fact a misidentified carpometacarpus of Pterodroma_ new sp., and the
identifications of Puffinus carneipes were, in fact, all referrable to Puffinus pacificus.
Pterodroma sp. (medium) is now referred to as Pterodroma new sp., and Pterodroma sp.
(small) has been identified as Pterodroma pycrofti. Pelecanoides sp. has been deleted from the
list, as no material referrable to this genus has been located in the collections from the island.
Columba vitiensis norfolciensis, a misidentification of Hemiphaga novaeseelandiae, should
also be deleted. Plates 1-7 show a range of the fossil material from Norfolk Island.

LORD HOWE ISLAND

Lord Howe Island (1,300 ha) is a basaltic remnant of a Late Miocene shield volcano
outcropping from the Lord Howe Rise, 700 km northeast of Sydney. Subfossil avian material
has been collected from caves, sand-dunes and calcarenites on the island (Meredith, unpubl. data;
van Tets ef al. 1981, van Tets & Fullagar 1977, Bourne 1974, Lambrecht 1933). Much of this
material has yet to be analysed. So far, 13 species of birds have veen identified, mostly species
known historically from the island. Three of the seabirds known only as fossils on Norfolk
Island, P. pycrofti (listed by Rich & van Tets (1982) as Pterodroma sp. (small)), Pelagodroma
marina and Sula tasmani (van Tets et al. 1988), are similarly recorded only as fossils on Lord
Howe. A small penguin Eudyptula sp. (prob. minor) is represented by a variety of adult
bones. Eudyptula minor has been recorded several times as a vagrant to Lord Howe, but
presently only breeds in more southerly waters. Further collecting may show whether or not it
once bred on Lord Howe.

An endemic species of extinct horned turtle Meiolania platyceps occurs in Pleistocene
calcarenites (Gaffney 1983).

VERTEBRATE FOSSIL FAUNAS FROM ISLANDS - 1355

KERMADEC ISLANDS

Archaeological excavations on Raoul Island, in the Kermadec group, produced material of
seven seabirds, two landbirds, the Pacific Rat and the New Zealand Fur Seal (Anderson 1980).
Four of the seabirds are in the genus Pterodroma, a genus which presents particular difficulties
of identification, and these records need re-evaluation. For instance, the material identified as
Pterodroma magentae should be further compared with material of P. solandri, and the
supposed P. inexpectata material needs to be compared to a similar-sized extinct Pterodroma
known as a fossil from Norfolk Island and, possibly, the Chatham Islands.

NEW CALEDONIA

The fossil fauna of New Caledonia is discussed in detail in Balouet's Chapter (this volume).
It is one of the most interesting insular faunas in the southwest Pacific, and contains a number
of fossil species that are possible Gondwana vicariants, notably a crocodile Mekosuchus
inexpectatus and the large, flightless bird Sylviornis neocaledoniae, of uncertain affinities.
Fossil horned turtles have also been found on New Caledonia (Meiolania sp.), and the nearby
Loyalty Islands (Meiolania mackayi). An extinct varanid lizard Varanus sp. cf. V. indicus
from the island is believed to be most closely allied to the New Guinea varanids. A single
tooth of a large mammal, originally described as a rhinoceros, has recently been placed in the
diprotodontid marsupial genus Zygomaturus (Guerin et al. 1981). Some debate, however,
surrounds the identification and provenance of this tooth (Bertrand 1986; Rich et al. 1988), and
it appears that it is indeed a rhinoceros. Only 15 species of birds are known as fossils, a rather
low number considering the size of the present avifauna (111 species), and more will
undoubtedly be found. Most of the fossils are extant or historically known species, but there
are several extinct forms: Sylviornis neocaledoniae, Porphyrio kukweidi, Megapodius, and a
woodhen (Rallidae) probably related to Tricholimnas lafresnayanus, amongst many others
(Balouet & Olson 1989).

FIJI ISLANDS

The Fiji Islands are palacontologically quite unexplored, yet there is every reason to think
that they are likely to have a potentially very interesting fossil fauna. The only fossils so far
known are from Polynesian middens on Naiggani and Laemba islands (R.F.Baird, pers.
comm.). The two species represented are an extinct large, apparently flightless pigeon and an
extinct megapode. Middens, caves and dunes are widespread in the Fiji group, and further
exploration is likely to be most profitable (R.F.Baird, pers. comm.).

COOK ISLANDS

Avian fossils have recently been located on Mangaia in the southern Cook Islands
(Steadman 1985, 1987). They include all but two of the nine species currently found on the
island, along with nine more extinct species, once again indicating the extent of faunal change
caused by Polynesian habitat alterations.

Steadman as well as C. Williams, T. Flannery, D. Roe, M. Spriggs and D. Wickler are
currently carrying out investigations into the fossil faunas of a number of other Pacific islands
(R.F.Baird & P.V. Rich, pers. comm.).

1356 - MEREDITH

CHATHAM ISLANDS

The Chatham Islands are an archipelago comprising two main islands and some twenty
other smaller islands and reefs. They lie on the tropical convergence, about 850 km east of
New Zealand, and have been occupied by Maoris since 800-1,000 yBP.

The Chathams are part of the New Zealand continental fragment and their history dates back
to a period of subsidence and volcanism in the Late Cretaceous (Hays et al. 1970). Their
present physiography is largely the result of changing sea levels during the Pleistocene.
During periods of low sea level, the Eocene Te Whanga Limestone developed its karst
topography, and was then buried beneath the dune-bedded, quartzose Wharekauri Sands.
Fluctuations in sea level produced a series of alternating sands (both marine and dune) and peats
(representing periods of ameliorating climate), During periods of maximum transgression (sea
level during the Nukumaruan rose to at least 285 m) the land area of the Chathams was reduced
to a minute fraction of its present area, while during the Last (Otiran) Glaciation, when sea
level fell to almost 120 m below present, virtually all the islands in the group were united as
one.

Following the Flandrian Transgression (about 2 m above sea level) in the Holocene,
extensive Older Dunes formed, particularly on northern, eastern and western coasts. The
development of low barrier dunes allowed the formation of the many lagoons and low-lying
coastal lakes of Chatham Island, and the present configuration of the whole group was attained.
H.O, Forbes discovered large numbers of sub-fossil bird bones in 1892, on which he published
a series of very brief papers (Forbes 1892 a-c, 1893 a-c, 1897). His collection is in the British
Museum. Despite the efforts of Dawson (1952, 1957, 1958, 1959, 1960, 1961) and Bourne
(1967), this collection still remains inadequately studied and clearly warrants a complete and
detailed reappraisal. In addition to Forbes' collection, that of Lord Rothschild is in part also in
the British Museum, with the remainder at Tring (Andrews 1896 a-c, Rothschild 1907, Dawson
1960), while much other material is held in New Zealand (Auckland War Memorial Museum,
Canterbury Museum, National Museum, Otago University Anthropology Department).

Fossil material is known only from Chatham and Pitt islands, none of the others having
suitable sites for bone preservation. It seems that virtually all the sand-dunes from which
bones have been obtained are likely to be no older than Holocene, having developed since sea-
level stabilized at its present level 5,000-6,000 yBP (Gibb 1979; Millener 1981b). Localities
for many of the early collections are inadequately described or, indeed, not given at all, but it is
likely that most represent non-anthropogenic accumulations, although some material is ‘rom
occupation middens. Some more recent collections, particularly of seabird material, may be of
recently beach-wrecked birds.

Most identifiable dune sites are listed by Millener (1981a). Cave deposits are usually
associated with occupation debris, indicating an age of less than 800-1,000 yBP. Although
peat deposits and swamps are widespread on Chatham Island, conditions in most of them appear
to have been unsuitable for the preservation of bone, possibly due to the past frequency 0: peat
fires (Hays er al. 1970), and the acidity of the environment. A small collection of bones,
however, has recently been obtained from a dark, peaty horizon among sand-dunes on Chatham
Island (R.J. Watt, pers.comm.).

In discussing the past and present avifauna of the Chathams, there are several species whose
reported occurrence is extremely doubtful. It was reported that Maoris had said that a kiwi
(Apteryx sp.) had once been present on the Chathams (Travers 1866), but this was later denied
(Travers 1883). Gallirallus minor, only known as a fossil species, is an unsatisfactorally
defined species which falls within the size range of Gallirallus dief‘enbachii (Olson 1975b), or,
alternatively, may only represent a small form of Gallirallus australis (Oison 1975b, Millener
1981b). Strigops habroptilus and Nestor notabilis have both been reported from the Chathams

VERTEBRATE FOSSIL FAUNAS FROM ISLANDS - 1357

(T ravers 1866; Forbes 1892b & 1893c; Dawson 1959, 1960). The verbal record of S.
habroptilus was later disavowed (Travers 1883), and there is some doubt over the provenance of
the supposed fossil material of this species from the Chathams. It seems most likely that this
species was, like Gallirallus australis (see below), brought to the Chathams by the Maoris.
Dawson (1959) considered all records of fossil N. notabilis were misidentifications of N.
meridionalis.

The Weka, Gallirallus australis, is known fossil from several sites, and it is possible that it
was indigenous prior to its introduction in 1905. If so, it is most likely that this flightless
species was brought to the Chathams by the Maoris, as it is undifferentiated from the mainland
form (Olson 1975b). Its presence in Maori middens both in New Zealand and on the Chathams
indicates that it was a regular food item.

Twenty-seven procellariiform seabirds are known as fossils from the islands, of which 20
probably bred there. This compares with a present procellariiform fauna of 25 species, of
which 14 are breeding species. This is a reduction of 30% in numbers of breeding species
(Table 2). Several species recorded as fossils require further study to confirm their
identification: Diomedea chlororhynchus, Phoebetria sp., Pterodroma neglecta, Pterodroma
prob. magentae and Pterodroma cf. ultima. This last species, tentatively identified by Bourne
(1967), may be conspecific with the extinct medium-sized Pterodroma from Norfolk Island
(Meredith, pers. comm.). The pelecaniform and sphenisciform species found as fossils are the
same as the present fauna, with the exception of the Emperor Penguin Aptenodytes forsteri, an
Antarctic breeder, which is now only a rare vagrant to southern New Zealand.

Waders (Charadriiformes) are well represented in the fossil deposits, with one extinct species
present, the endemic Coenocorypha chathamica. Two skuas, Stercorarius skua and S.
longicaudus have been identified from fossil material (the latter tentatively). The status of S.
longicaudus in the southwest Pacific is very unclear (Harrison 1983), and it is unknown
historically from the Chathams.

The land and freshwater avifauna of the Chathams (both fossil and extant) is highly
endemic, with two endemic genera, nine endemic species and nine endemic subspecies, out of a
total of thirty-four breeding species and seven non-breeders (excluding exotics and Maori
introductions). Most of these species are known as sub-fossils. The 12 known only as fossils
include both endemic genera and two of the endemic species. As the passerine fossils have yet
to be fully studied, these numbers should increase.

A variety of material from stratified middens indicates that most, if not all, species of
land and freshwater birds known only as fossils survived into, but became extinct during, the
prehistoric Maori occupation. There is substantial evidence, particularly from the investigations
of Simmons (1964) and Sutton (1979, 1981), that many terrestrial birds were hunted by the
prehistoric Maori, but such birds do not form the major source of food at any site. In the
Durham area siudied by Sutton, only three landbirds were ever recorded in numbers greater than
ten individuals at any one site. Moreover, at each site the excavated midden remains probably
represent several hundred years of accumulation, The most abundant avian remains at the CHA
and the CHB sites (see Sutton 1979) were those of the Chatham Island Pigeon, Hemiphaga
novaeseelandiae chathamensis, followed by those of Dieffenbach’s Rail, Gallirallus
dieffenbachi, and Hawkin's Rail, Diaphorapteryx hawkinsi. Perhaps significantly, the first of
these species, although apparently abundant in the prehistoric period, has never been common
in European times, the second species became extinct about 1900, while the last, possibly
mentioned in Maori legend (sce White 1897) became extinct prior to European settlement. At
several sites near the Te Whanga Lagoon, bones of waterfowl, particularly of the now extinct
swan, Cygnus sumnerensis, have been found in abundance in occupation middens
(Forbes 1892ac, Sutton 1979). This evidence of exploitation may be significant; of the eight
indigenous waterfowl recorded as fossils from the Chathams, only one is still resident there.

1358 - MEREDITH
SSS_L_LL_L—LLL_L_LLLLL_— SSS —E—_———————————SS SSS S______—_

Table 2. A comparison of the procellariiform faunas of the Chatham Islands - past and present (B=breeding,
V=visitor, FB?=may have formerly bred).

RECORDED CURRENT SPECIES SUBFOSSIL STATUS
Diomedea exulans - Vv
Diomedea epomophora + B
Diomedea chlororhynchus +? -
Diomedea cauta + B
Phoebetria sp. +? -
Macronectes halli + B
Daption capense + v
Pterodroma lessonii + Vv
Pterodroma inexpectata + V(FB?)
Pterodroma ?neglecta +? >
Pterodroma ?magentae + B?
Pterodroma cf. ultima + -(FB?)
Pterodroma nigripennis + B
Pterodroma axillaris + B
Pachyptila vittata + B
Pachyptila turtur + B
Pachyptila crassirostris + B
Procellaria cinerea + V(FB?)
Procellaria aequinoctialis + V(FB?)
Puffinus carneipes + Vv
Puffinus bulleri + V(FB?)
Puffinus griseus + B
Puffinus tenuirostris + Vv
Puffinus gavia + V(FB?)
Puffinus assimilis + B
Garrodia nereis + B
Pelagodroma marina + B
Fregetta tropica - Vv
Pelecanoides urinatrix + B

This archaeological evidence indicates that reduction in numbers of land and freshwater
birds, and, for some species, extinction, may have been caused directly through exploitation by
the prehistoric Maori. However, it seems more probable (as has been argued by Millener
1981b, 1984 for the extinction of landbirds on mainland New Zealand) that a combination of
factors was involved. Foremost among them was habitat modification and the depredations of
introduced mammalian predators, followed by exploitation for food. Thirteen Chathams species
failed to survive the Polynesian period. There seems little doubt that the seven species which
have become extinct in the Chathams since 1840 succumbed to the even more drastic changes
wrought by European settlement, while several others, whose populations have been severely
diminished, and which are typically restricted to the more remote islands in the group, appear to
have little chance of longterm survival.

Archaeological investigations have also shown that the prehistoric Chatham Island Maoris
were largely dependent upon marine and littoral resources, since, due to the climate, traditional
horticulture of tropical Polynesian cultigens was impossible (Sutton 1979, 1980, 1981, 1982;
Sutton & Marshall 1980). Shellfish, fish, seabirds and marine mammals were the main food

VERTEBRATE FOSSIL FAUNAS FROM ISLANDS - 1359

resources utilised. Evidence from a large number of middens in the Durham area indicates that
sealing was by far the most important economic activity (Smith 1977).

Marine birds, particularly the fledgling young of albatrosses and petrels, were selectively
and, in some cases, intensively exploited, but, despite this, were not the major source of food
at any site. Although Sutton (1979) lists a considerable variety of seabirds from the Durham
middens, only four species were represented by more than ten individuals per site. It is notable
that the most commonly represented seabird Pterodroma sp. cf. P. magentae (over 400
individuals in sites CHA and CHB, constituting over 50% of the total bird fauna) is now the
rarest of the breeding petrels in the Chathams. The diminution in numbers of this and other
petrels on the islands can perhaps be attributed directly to prehistoric exploitation, although the
effects of habitat modification and the introduction of mammalian predators should not be
underestimated. Indeed, this last factor has been suggested as being responsible for the local
extinction of mainland breeding petrel populations in much of New Zealand (Millener 1981b).

The land and freshwater birds of the Chathams are, for the most part, closely related to New
Zealand taxa, all but nine being inseparable from, or representing endemic subspecies of,
mainland species. Diaphorapteryx seems to be a derivative from a generalised Gallirallus
philippensis-type ancestor (Andrews 1896b). Later invasions of such stock gave rise to two
other endemic rails, Gallirallus modestus and G. dieffenbachii (Olson 1975b). The true
affinities of Pachyanas have yet to be determined, but it may prove to be a derivative of some
generalised Anas-type ancestor comparable to that which gave rise to Euryanas on the New
Zealand mainland. Haliaeetus is not known from mainland New Zealand, and the endemic H.
australis of the Chathams is considered by Olson (1984) to be more similar to northern species
of the genus, particularly H. pelagicus, than to the geographically closer H. leucogaster. Olson
has suggested that the ancestor of H. australis probably colonised the Chatham Islands from the
Northern Hemisphere rather than from Australasia.

NEW ZEALAND

New Zealand has produced abundant fossil material, and this is discussed in detail by
Fordyce (this volume) and Millener (this volume). Rather than repeat this, I merely wish to
emphasise a number of points relevant to the other insular faunas discussed here.

New Zealand's vertebrate fauna, both fossil and recent, is dominated by birds. The only land
mammals present, except for those introduced by man, are two species of bats, but there is a
diverse fossil cetacean assemblage. Mesozoic, Cretaceous and Tertiary marine reptiles are
known, and at least two dinosaurs. Otherwise terrestrial reptiles are only represented by
relatively scarce Late Pleistocene and Recent material of the Tuatara Sphenodon, and a diverse
small lizard fauna, largely of extant taxa, but also including several, as yet undescribed, extinct
forms. One or more leiopelmatid frogs, larger than and not apparently referrable to any living
species, are known from many subfossil sites, particularly caves.

The avifauna, both fossil and modern, is rich in seabirds but depauperate in land and
freshwater species, with the latter group exhibiting a high degree of endemism. Flightlessness,
or reduced flying ability, is common, particularly amongst extinct species, many of which had
died out before European colonisation, The moas (Dinornithiformes), the rails (Rallidae) and,
to a lesser extent, the ducks and geese (Anseriformes) show a particularly high frequency of
flightlessness and endemism, and have undergone varying degrees of adaptive radiation.

There has been a high frequency of extinction since the Late Pleistocene (possibly only
since the Late Holocene), and this has particularly affected endemic species. The reasons for
these extinctions remain uncertain, but the various changes caused by the arrival of the Maoris
800-1,000 years ago are often invoked. Unfortunately, there is little direct evidence on these
matters, except for the extinction of the moas, where Anderson (1983) has argued convincingly

1360 - MEREDITH

on the basis of archaeological evidence for Maori hunting as the cause.

Unlike most small islands, New Zealand has produced pre-Pleistocene avian fossils.
Thirteen species of penguin (Sphenisciformes) are known from the Tertiary, eight of the genera
among which they are distributed being endemic. Manu antiquus, possibly an albatross
(Diomedeidae), is known from the Oligocene, and a false-toothed bird Pseudontornis stirtoni
(Pelecaniformes) is thought to be of Mio-Pliocene age, but could in fact be Early Pleistocene
(Fordyce 1982). The only evidence of Tertiary landbirds comprises an as yet undescribed
assemblage from Miocene lacustrine deposits, some moa footprints and some presumed kiwi
footprints.

MACQUARIE ISLAND

Macquarie Island is a small, subaerial projection of the submarine Macquarie Ridge. It lies
about 1,000 km southeast of Tasmania. Large numbers of seabirds presently breed on the
island, but, prior to the appearance of several recent adventives and introductions, only two
landbirds, Gallirallus philippensis macquariensis and Cyanoramphus novaezelandiae erythrotis,
were historically recorded.

Subfossil material of two species of penguin, Eudyptes chrysolophus schlegeli and
Aptenodytes patagonica, occurs abundantly in bone beds at Finch Creek and Bauer Bay, and
has been dated at 6,100+120 yBP and 3,980+140 yBP respectively (McEvey & Vestjens 1974).
These dates, however, may require revision in the light of the possibility of contamination by
old carbon from the seas at these latitudes. This material is indistinguishable from skeletal
material obtained from the present populations. Occasional procellariiform material is also
found in these deposits, including bones of Pterodroma lessoni, juvenile Puffinus prob.
griseus, and a small Puffinus similar to Puffinus gavia but too poorly represented to be
confidently identified (Meredith 1985a).

Vestjens (1963) collected bones of one of the two landbirds, Gallirallus philippensis
macquariensis from Eagle Cave and Aurora Cave, but considered that they were probably recent
remains left by sealers. Recent bones of a variety of seabirds are also present in Aurora Cave.
Abundant material of Pachyptila prob. desolata of unknown age, possibly recent, occurs in a
cave at Brothers Point (Meredith 1985a).

DISCUSSION

Islands have for many years been assumed to be unlikely sites for fossil deposition because
of their small size, youthful ages and, frequently, their volcanic origins. This is clearly false,
however, not only around Australia and in the southwest Pacific, but, as shown by Fig. 5 and
Table 2, throughout the world. Further finds are certain, as there remain thousands of islands
still unexplored by palaeontologists.

Insular fossil faunas share several characteristics when compared to continental faunas.
They are virtually all Late Quaternary or Recent in age. This is a reflection of a number of
factors. Many islands are relatively young geologically (mid-Late Cainozoic), limiting the
maximum possible age of any fossils found on them. Furthermore, most depositional

Figure 5. Localities of islands that have produced significant vertebrate fossil deposits, other than those
shown in Figs. 1 & 4. The numbers refer to the islands listed in Table 1. Large islands, such as Japan and
England, are not included,

VERTEBRATE FOSSIL FAUNAS FROM ISLANDS - 1361

1362 - MEREDITH

environments prior to the Late Pleistocene are likely to have been destroyed or changed by
diagenetic processes. Sand dunes built during the Last Glacial Maximum and, more recently,
since the mid-Holocene (5,000-6,000 yBP) have given rise to some of the most abundant bone
deposits (e.g. Chathams, Norfolk Island, Flinders Island, Hawaiian Islands). Some karstic
limestones which contain fossils in caves and clefts did not develop until the Pleistocene (e.g.
New Caledonia).

However, the most consistent compositional difference between insular and continental
fossil faunas is the dominance of birds, particularly seabirds. Ninety-two per cent of all
Pleistocene procellariiform fossils are from islands, and, for the marine pelecaniforms (sulids,
frigatebirds and tropic-birds), the figure is 96%. Faunas of oceanic islands are particularly
biased in this way. Those of coastal islands tend to be more similar to the adjacent mainland.
These differences largely reflect the biases of the modern faunal communities. Island faunas,
both extant and extinct, are also characterised by the frequent presence of unusual endemic
forms (e.g. flightless birds and dwarfed or gigantic forms).

The study of extant insular faunas has contributed importantly to modern biological theory.
The question arises: what can island palaeontology contribute? I see four main areas to which
fossil data may be relevant: taxonomy and phylogenetic relationships, taphonomic studies,
seabird biogeography and biogeography of insular landbirds.

TAXONOMY AND PHYLOGENETIC RELATIONSHIPS

Although the importance of fossil taxa in taxonomic and phylogenetic studies may be less
than has been generally assumed (Patterson 1981), the high level of Quaternary and ongoing
extinctions on islands means that fossil material is often of particular practical importance.
Many island taxa are known only as fossils. Even with species that are still extant, many of
these are absent or rare in osteological collections and the only material for comparative studies
that is available may be that from fossil sites.

TAPHONOMIC STUDIES

The taphonomy of insular fossil sites has so far received little attention. Such sites are
taphonomically interesting for several reasons. The small size of many islands and the nature of
their physiography often means that ihe source area for the entombed fauna can be delimited
easily and accurately, Furthermore, particularly at very recent sites, historical records of the
fauna prior to any major anthropogenic disturbance can provide an accurate estimation of the
size and composition of the source fauna, thus allowing depositional biases to be assessed.

The frequent absence of mammalian predators and carnivores on islands suggests that some
island sites could be used to provide a base line for assessing effects of such predation on
preservation, by comparing them with similar continental sites where mammalian predators are
present. Other sites may show the effects of the sudden introduction of mammalian predators
and other exotic species by man (é.g. Rattus exulans on Norfolk Island).

Penguins are among the most abundant pre-Pleistocene avian fossils in the Southern
Hemisphere, but there have been no taphonomic studies done on this group. Many islands in
the Southern Ocean offer excellent opportunities for such studies.

SEABIRD BIOGEOGRAPHY

Seabird biogeography has progressed little since the important analyses of latitudinal
zonation by Murphy (1936). One of the reasons for this has been the difficulty in detecting
patterns other than latitudinal ones in the distributions of many species, which are often either

VERTEBRATE FOSSIL FAUNAS FROM ISLANDS - 1363

patchy, with anomalous gaps, or very localised and relictual. Already, the fossil data have
begun to fill in some of these gaps and to expand the ranges of some "relics." Twenty years
ago, Bourne (1965) drew attention to the importance of fossil evidence in establishing the
true" distributions of seabirds, noting that whole extinct seabird communities had been found
on Bermuda, St Helena, St Paul, Amsterdam Island and the Chathams. Since then, several
more fossil seabirds previously unknown from these islands have been identified from St
Helena (Olson 1975a) and the Chathams (Bourne 1967, this Chapter), and others have been
found in the Bahamas (Olson 1982), Aldabra Atoll (Harrison & Walker 1978), the Mascarene
Islands (Bourne 1976), Norfolk and Lord Howe islands (this Chapter), Easter Island (Carr
1980), the Hawaiian islands (Olson & James 1982a,b) and the Cook islands (Steadman 1985).
Fossils have shown significantly larger past ranges for Pterodroma pycrofti (now relict on a
few small islands off New Zealand, subfossil on Norfolk and Lord Howe islands), the
hypoleuca" group of the gadfly petrels (Pterodroma hypoleuca, P. nigripennis and P.axillaris)
are currently restricted to the western Pacific, their extinct sister-species P. kurodai is a
Pleistocene fossil from Aldabra Atoll, Indian Ocean), Sula abbotti (now relict on Christmas
Island, Indian Ocean, subfossil on the Mascarene islands) and Pterodroma cahow (now relict on
Bermuda, subfossil on the Bahamas).

BIOGEOGRAPHY OF INSULAR LANDBIRDS

The usefulness of fossil data from islands in enabling a fuller and more accurate description
of island landbird faunas in the light of high Quaternary and post-European extinction rates is
obvious and uncontroversial. This is, however, one of the very few uncontroversial areas
remaining in island biogeography. There is now considerable argument concerning the relative
merits of dispersalist versus vicariance explanations of distribution patterns, and about the
validity and usefulness of the MacArthur-Wilson equilibrium theory of island biogeography
(e.g. Simberloff 1983). What does the fossil record have to say on these matters?

One of the main arguments used by vicariance biogeographers against dispersalist theories
is that dispersal is an historical explanation and, therefore, neither provable nor falsifiable. It
is, thus, important to consider whether palaeontological (= historical) data can prove or
disprove a dispersalist explanation. The ever-present problem of "the incompleteness of the
fossil record" means that absence of a species from the fossil record of an island can never prove
real absence from the past fauna, although it can be suggestive. For instance, the absence of
the Pukeko, Porphyrio p. melanotus, a large rail, from fossil and midden deposits in New
Zealand, when other species of similar size and habits are common to very common in such
deposits, suggests that it may have been a recent arrival in New Zealand (Millener 1981b).

The presence of a species as a fossil on an island can, under very restricted circumstances, be
positive evidence of past dispersal, if it is a species that is unknown in early historical faunal
lists (assuming these are adequately complete), but has recently recolonised by dispersal. Such
a species is the Green-winged Pigeon Chalcophaps indica on Norfolk Island (Schodde et al.
1983). It can be argued that re-dispersal following extinction has been observed to occur in
that species, and that its presence as a fossil indicates that dispersal also occurred in the past and
is not simply a modern effect associated with, say, following ships or human introductions.
Such an argument, however, is rather trivial, and in most such cases dispersal is likely to be
accepted as the most probable (although unproven) explanation anyway. So, all in all, the
palaeontologist cannot offer stronger data than the neontologist on active dispersal of species,
although they may be able to provide evidence suggestive of past absence foliowed by recent
dispersal.

Tn the second main area of controversy, the MacArthur-Wilson equilibrium theory, it has
already been claimed that fossil data from the West Indies (Pregill & Olson 1981), from Hawaii

1364 - MEREDITH

(Olson & James 1982a,b) and from the Galapagos (Steadman 1986) cast real doubts on the
value of this theory. Certainly the fossil data show that some late Quaternary island faunas
were markedly different from those of today, both in species richness and in species
composition, and that these differences may be adequately explained by climatic changes or
anthropogenic effects, without the need to invoke equilibrium theories (e.g, Hope et al. 1977,
Pregill & Olson 1981, Olson & James 1982a, b). While these data show that, when
comparing faunal richness and composition between islands, one needs to be aware of recent
prehistoric changes, the mere fact that there are changes over time in these parameters does not
necessarily invalidate the equilibrium theory.

The MacArthur-Wilson theory postulates an equilibrium between immigration and
extinction rates which determines the number of species that are found on an island (MacArthur
& Wilson 1967). Once an equilibrium state is reached, species richness should remain
constant, but community composition may change due to turnover. A relatively constant
number of species on an island is only evidence for equilibrium if rates of immigration and
extinction do not change. In each of the examples mentioned above there has clearly been an
increase in the extinction rate, and, as would be expected if the equilibrium theory applies, a
decrease in species number. This is not to say that the equilibrium theory is proved, only that
the fossil data are consistent with this aspect of it.

If there is an equilibrium, then there should be turnover and, thus, change in species
composition. There is little evidence for turnover from the fossil record, but again, due to the
incompleteness of that record, it is hard to see what would constitute valid evidence. Long,
fairly complete stratified fossil deposits are unlikely to be found on islands. Absences from the
fossil record can only be suggestive of real absences, and not proof. Only the presence of a
species in the fossil record and its current or historical absence (as discussed for dispersal) could
indicate turnover, and only providing that environmental change or anthropogenic effects can be
excluded.

The regular relationship of species diversity (especially bird species diversity) to island area
within an island chain, usually a power function, has long been used as evidence for the
equilibrium theory, on the assumption that immigration and extinction rates will also vary
regularly with island area. At present, the only fossil data relevant to this problem come from
the Hawaian islands. A recent application of the MacArthur-Wilson theory to the extant
Hawaiian avifauna (Juvik & Austring 1979) found that the relationship of both species richness
of landbirds and the species richness of the endemic honeycreepers (Drepanidini) to island area
was very close to that expected from the theory. Olson and James (1982b) strongly criticized
this on the basis that their own fossil data showed that "the recent history of the endemic
avifauna has been one of natural extinction without natural replacement", and thus modern
species richness figures could not represent a saturated equilibrium situation.

This criticism is certainly an important one, but it is not a damning one. Olson and James
themselves argue that the effects of the Holocene extinctions were common to most, if not all,
the islands, and, on each island, that the extinctions were largely confined to certain segments
of the avifauna, mainly the inhabitants of the lowland forests (these forests were almost
completely cleared by the Polynesians), the large, edible, flightless species, and the top
predators. Furthermore, Juvik and Austring found high correlations with various habitat
diversity indices as well as with area, as have a variety of studies on other islands. Could not
the modern avifaunas of Hawaii simply be equilibrial subsets of a past larger fauna that was
also in equilibrium, the modern faunal composition being the result of the systematic removal
by man on all the islands of a certain suite of species - those of the lowland forests, those that
were edible, and the large predators? Again, while raising important questions, the fossil data
remain at least consistent with equilibrium theory, and certainly do not falsify it. In fact,
perhaps the strongest criticism of the equilibrium theory that remains is the difficulty in

VERTEBRATE FOSSIL FAUNAS FROM ISLANDS - 1365

falsifying it in any way (Simberloff 1983).

Diamond (1972) introduced the term "relaxation" to describe the process of a fauna returning
to equilibrium after displacement from it, following isolation and decreased island area due to
Late Pleistocene sealevel rises. There is some fossil evidence for such a phenonemon, but it is
restricted to large mammals whose immigration rate would decline to zero after isolation,
making the equilibrium concept rather empty. Hope (1973) found that, out of a relatively
diverse assemblage of herbivorous marsupials which was present when the Bass Strait islands
were a united land bridge, the largest had died out completely and at present no island smaller
than 1.4 sq km could support one species, while two or more species required at least 6.1 sq
km. As several of the largest species (Protemnodon anak, Sthenurus occidentalis) also died out
on the mainland, their extinction cannot be counted as evidence for relaxation, nor can the
possible effects of human agency be ruled out.

It is on this last point, the effects of man, that palacontology has made its strongest
contribution to island biogeography so far, by emphasizing that most island faunas are likely
to have been drastically changed following the arrival of human settlers, whatever their level of
technology. The present size and composition of those faunas on islands with indigenous
human populations (or at least the size and composition prior to the arrival of European man)
cannot be assumed to be purely the result of natural ecological processes. As discussed above,
this may or may not matter in discussing particular biogeographic questions, but from now on
island biogeographers will have to be aware of the possible effects on their data of
anthropogenic changes.

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VERTEBRATE FOSSIL FAUNAS FROM ISLANDS - 1369

APPENDIX I

Australasian and southwest Pacific insular fossil faunas. An asterisk (*) after a species name
indicates an identification that is regarded by this author as uncertain.

(a) COASTAL ISLANDS.
Flinders Island.

(i) Ranga Cave (Hope 1969,1973).

BIRDS - unident. biird.

MAMMALS - Dasyuridae: Dasyurus viverrinus,
Antechinus minimus.

Peramelidae: Parameles gunnii.

Vombatidae: Vombatus ursinus.

Phalangeridae: Trichosurus vulpecula.

Petauridae: Pseudocheirus peregrinus.

Macropodidae: Potorous apicalis, Macropus
rufogriseus, Macropus giganteus, Aepyprymnus
rufescens, Thylogale billardieri.

Muridae: Rattus lutreolus, Mastacomys fuscus,
?Sminthopsis leucopus, Pseudomys cf.
novaehollandiae, Pseudomys higginsi.

Otariidae: Arctocephalus pusillus.

(ii) Palana dunes (Hope 1969,1973; Meredith, pers.obs.).
BIRDS - Spheniscidae: unident. penguin.
Procellariidae: Puffinus tenuirostris.

Pelecanidae: Pelecanus conspicillatus.

unident. birds.

MAMMALS - Dasyuridae: Dasyurus maculatus,
Sarcophilus harrisii.

Peramelidae: [soodon obesulus.

Macropodidae: Potorous apicalis.

(ii) Unspecified dune sites (Hope 1969,1973).

BIRDS - Procellariidae: Puffinus tenuirostris.

MAMMALS - Tachyglossidae: Tachyglossus
aculeatus.

Dasyuridae: ?Antechinus minimus.

Vombatidae: Vombatus ursinus.

Petauridae: Pseudocheirus peregrinus.

Macropodidae: Macropus rufogriseus, Thylogale
billardieri.

Muridae: Rattus lutreolus.

Cape Barren Island.

Unspecified dune sites (Hope 1973).

REPTILES - Scincidae: Tiliqua nigrolutea.

BIRDS - Procellariidae: Puffinus tenutrostris.

MAMMALS - Peramelidae: Jsoodon obesulus.

Phalangeridae - Trichosurus vulpecula.

Macropodidae - Thylogale billardiert.

Muridae - Rattus lutreolus.

Preservation Island.

Unspecified dune sites (Hope 1969,1973).
REPTILES - Scincidae: Tiliqua nigrolutea.
BIRDS - Procellariidae: Puffinus tenuirostris.
Anatidae: Cereopsis novaehollandiae.
unident. birds.

MAMMALS - Dasyuridae: ?Antechinus minimus.
Macropodidae: Macropus rufogriseus.
Munidae: Rattus lutreolus.
Unident. seal.
East Kangaroo Island.

Unspecified dune sites (Hope 1973).

BIRDS - Procellariidae: Puffinus tenuirostris.

Anatidae: Cereopsis novaehollandiae.
MAMMALS - Macropodidae: Thylogale billardieri.
Deal Island.

Unspecified dune sites(Hope 1973).

REPTILES - Scincidae: Tiliqua nigrolutea.

BIRDS - Procellaridae: Puffinus tenuirostris.

MAMMALS - Dasyuridae: ?Dasyurus maculatus,
7Antechinus minimus.

Vombatidae: Vombatus ursinus.

Phalangeridae: Trichosurus vulpecula.
Macropodidae: Potorous apicalis, Macropus
rufogriseus, Macropus giganteus, Thylogale
billardieri.
Muridae: Rattus lutreolus.
Erith Island.
Unspecified dune sites (Hope 1973).

REPTILES - Scincidae: Tiliqua nigrolutea.

BIRDS - Procellariidae: Puffinus tenuirostris.

unident. spp.

MAMMALS - Phalangeridae: Trichosurus vulpecula.

Macropodidae: Macropus rufogriseus, Thylogale
billardieri.

Muridae: Rattus lutreolus.

Long Island.

Unspecified dune site (Hope 1973).
MAMMALS - Muridae: Rattus lutreolus.

Three Hummock Island.
Unspecified dune site (Hope 1973).
MAMMALS - Peramelidae: [soodon obesulus.
Macropodidae: Potorous apicalis, Thylogale billardieri.
Hunter Island.

(i) Stockyard Midden (O'Connor 1982; van Tets &
O'Connor 1983).

REPTILES - Scincidae: Tiliqua sp.

BIRDS - Spheniscidae: Eudyptula minor, Tasidyptes
hunteri.

Diomedeidae: Diomedea cauta.

Procellariidae: Puffinus tenutrostris, Pachyptila sp.*

Pelecanoididae: Pelecanoides urinatrix.

Pelecanidae: Pelecanus conspicillatus.

Phalacrocoridae: Phalacrocorax fucescens.

Anatidae: Cygnus atratus, cf. Anas superciliosa,
Aythya australis, Cereopsis novaehollandiae.

Accipitridae: Haliaeetus leucogaster.

Falconidae: Falco sp.

Laridae: Larus pacificus, cf. Larus.

Psittaciformes: unident. parrot.

Cracticidae: cf. Strepera.

Corvidae: Corvus sp.*

Passeriformes: unident. passerines.

MAMMALS - Dasyuridae: Antechinus ?minimus.

Peramelidae: Isoodon obesulus.

Burramyidae: Cercartetus nanus.

Macropodidae: Potorous
billardieri.

Muridae: Hydromys chrysogaster, Rattus lutreolus,

apicalis, Thylogale

1370 - MEREDITH

Mastacomys fuscus, Sminthopsis leucopus, Pseudomys
higginsi.

Otariidae: Arctocephalus pusillus, Arctocephalus fosteri.

Phocidae: Mirounga leonina.

(ii) Cave Bay Cave (Bowdler 1974).

BIRDS - Spheniscidae: Eudyptula minor.

Procellariidae: Puffinus tenuirostris.

Psittaciformes: unident. parrot.

Corvidae: unident. raven.

unident. birds.

MAMMALS - Dasyuridae: Dasyurus sp., Antechinus
sp.

Peramelidae: Perameles gunnii, Isoodon obesulus.

Petauridae: Pseudocheirus peregrinus.

Burramyidae: Cercartetus sp.

Macropodidae: Macropus rufogriseus, Thylogale
billardieri.

Muridae: Mastacomys fuscus, unident. rat.

(iii) Muttonbird Midden (Bowdler 1974).

BIRDS - Spheniscidae: unident. penguin.
Procellariidae: Puffinus tenuirostris.
unident. bird.

MAMMALS - Dasyuridae: Antechinus sp.
Peramelidae: [soodon obesulus.
Macropodidae: Thylogale billardieri.
Muridae: Rattus sp.

(iv) Little Duck Bay (Bowdler 1974).

BIRDS - Spheniscidae: unident. penguin.
MAMMALS - Peramelidae: inident. bandicoot.
Macropodidae: Thylegale billardiert.
Muridae: Rattus sp.

Otariidae: Arctocephalus pusillus.

King Island.
(i) Egg Lagoon (Hope 1973).

MAMMALS - Diprotodontidae: Diprotodon optatum.

(ii) Settlement Lagoon (Hope 1973).

MAMMALS - Diprotodontidae: ?Nototherium sp.
Macropodidae: Protemnodon anak.

(iii) Surprise Bay (Hope 1973).

MAMMALS - Macropodidae: Sthenurus occidentalis,
Protemnodon anak.

(iv) Unspecified dune sites (Hope 1973).

REPTILES - Scincidae: ?Tiliqua nigrolutea.
BIRDS - Casuariidae: Dromaius ater.

Procellariidae: Puffinus tenuirostris.

MAMMALS - Tachyglossidae: Zaglossus harrissoni,
Tachyglossus aculeatus,

Dasyuridae: Dasyurus maculatus, Antechinus minimus.

Vombatidae: Vombatus ursinus.

Petauridae: Pseudocheirus peregrinus.

Macropodidae: Potorous apicalis, Macropus
rufogriseus, Thylogale billardieri.

Muridae: Rattus lutreolus, Pseudomys higginsi.

Unident. seal.

Kangaroo Island.

N.B. For full list of localities of Dromaius baudinianus
see Parker

(1984).

(i) Rocky River (Williams 1980).

BIRDS - Casuariidae: Dromaius baudinianus.
MAMMALS - Dasyuridae: Sarcophilus sp.
Vombatidae: unident. wombat.
Phascolarctidae: Phascolarctos cinereus.
Phalangeridae: Trichosurus vulpecula.

Diprotodontidae: Diprotodon sp., Zygomaturus
trilobus.

Macropodidae: Macropus fuliginosus, Macropus
eugenii, Protemnodon sp., Sthenurus spp.,
Sthenurus gilli.

Muridae: unident. murids.

(ii) Kelly Hill caves (Williams 1980).

BIRDS - Casuariidae: Dromaius baudinianus.

MAMMALS - Dasyuridae: Dasyurus maculatus,
Sarcophilus harrisii, Phascogale tapoatafa.

Vombatidae: Lasiorhinus sp.

Phascolarctidae: Phascolarctos cinereus.

Phalangeridae: Trichosurus vulpecula,

Petauridae: Pseudocheirus peregrinus.

Macropdidae: Macropus fuliginosus, Macropus eugenit,
Sthenurus cf. occidentalis.

(iii) Mount Taylor Cave (Williams 1980).

MAMMALS - Macropodidae: Sthenurus sp.

(iv) Emu Four Hole Cave (Williams 1980).

BIRDS - Casuariidae: Dromaius baudinianus.

MAMMALS - Tachyglossidae: Tachyglossus aculeatus.

Dasyuridae: Dasyurus viverrinnus, Sminthopsis
marina,

Peramilidae: Perameles sp., lsoodon obesulus.

Phalangeridae: Trichosurus vulpecula.

Petauridae: Pseudocheirus peregrinus.

Burramyidae: Cercartetus concinnus.

Macropodidae: Potorous platyops, Macropus
fuliginosus, Macropus eugenii.
Muridae: Rattus fuscipes, Rattus lutreolus.

(v) Fossil Cave (Williams 1980).

MAMMALS - Dasyuridae: Sarcophilus cf. harrisii.

Vombatidae: unident. wombat.

Phascolarctidae: Phascolarctos cinereus.

Macropodidae: Macropus fuliginosus, Macropus
eugenii, Sthenurus cf. brownei.

Muridae: unident. murids.

(vi) Seton Rockshelter (Williams 1980; Hope et al. 1977).

REPTILES - Elapidae: unident. elapid.

Varanidae: Varanus sp.

Scincidae: Trachydosaurus rugosus, Tiliqua nigrolutea,
cf. Egernia whitii.

Agamidae: Amphibolurus spp.

BIRDS - Procellariidae: Puffinus sp., Pachyptila cf.

salvini*.
Plataleidae: Threskiornis cf. molucca.

Anatidae: ¢f. Anseranas semipalmata, Tadorna cf.
tadornoides, Anas cf. superciliosa, Anas cf.
castanea, Malacorhynchus membranaceus.

Accipitridae: Hieraaetus morphnoides.

Falconidae: Falco berigora.

Phasianidae: Coturnix cf. pectoralis.

Tumicidae: Turnix varia, Turnix velox.

Rallidae: Gallirallus philippensis, Rallus pectoralis,
Porzana cf. fluminea, Gallinula cf. mortierii,
Gallinula cf, ventralis.

Burhinidae: Burhinus magnirostris.

Scolopacidae: Gallinago cf. hardwickii.

Laridae: Larus novaehollandiae, Sterna cf. nereis.

Columbidae: Ocyphaps lophotes.

Platycercidae: Pezoporus wallicus, Lathamus discolor.

Hirundinidae: sp. 1*, sp. 2*.

Meliphagidae: unident. honeyeaters.

Sylviidae: Cinclorhamphus cruralis.

Grallinidae: Grallina cyanoleuca.

VERTEBRATE FOSSIL FAUNAS FROM ISLANDS - 1371

Cracticidae: Gymnorhina tibicen, Strepera graculina,
Strepera versicolor.

Corvidae: Corvus sp.*

Passeriformes: unident. passerines.

MAMMALS - Dasyuridae: Dasyurus cf.
geoffroyi/viverrinus, Dasyurus maculatus,
Sarcophilus harrisii.

Peramelidae: Perameles bougainville, Isoodon
obesulus.

Vombatidae: Lasiorhinus latifrons.

Phalangeridae: Trichosurus vulpecula.

Burramyidae: Cercartetus nanus.

Macropodidae: Bettongia penicillata, Bettongia lesueur,
Potorous platyops, Macropus cf. fuliginosus,
Macropus greyi, Macropus rufogriseus, cf. Megaleia
rufa, Logorchestes leproides, Sthenurus cf. gilli.

Muridae: Hydromys chrysogaster, Rattus fuscipes,
Rattus lutreolus, Mastacomys fuscus, Pseudomys
occidentalis, Pseudomys australis/shortridgei.

Brothers Island.

Cave, western end of western island (Williams 1980).
BIRDS - Casuariidae or Dromomithidae: see text.
unident. birds.

MAMMALS - Macropodidae: Macropus sp., Sthenurus
ef. maddocki.
Otariidae: Arctocephalus sp.

Bowen Island.

Bowen Island One Midden (Blackwell 1982).

BIRDS - Spheniscidae: Eudyptula minor.

Diomedeidae: Diomedea cauta.

Procellariidae: Puffinus tenuirostris, Pachyptila sp.*

Sulidae: Morus serrator.

Phalacrocoridae: Phalacrocorax carbo.

MAMMALS - Otariidae: Arctocephalus pusillus.
Maatsuyker Island

Unspecified midden (Vanderwal & Horton 1984),

BIRDS - Spheniscidae: penguin.

Diomedeidae: Diomedea cauta.

Procellariidae: Puffinus tenuirostris, Pachyptila sp.

Phalacrocoridae: Phalacrocorax sp.

MAMMALS - Otariidae: Arctocephalus pusillus.
Houtman-Abrolhos Archipelago.

North Island (Alex Baines, pers.comm.).

BIRDS - Phalacrocoridae: Phalacrocorax sp.
unident. birds.
Dirk Hartog Island.

(i) North Point (Western Australian Museum collection).
BIRDS - Procellariidae: Puffinus sp.
Phalacrocoridae: Phalacrocorax varius.
unident. birds.

(ii) Herald Heights dunes (WAM collection).

BIRDS - Casuariidae: emu eggshell.

(iii) Cave near Herald Heights (WAM collection).

BIRDS - unident. birds.
Salisbury Island.

Spongolite, site unspecified (A.Burbidge & N.McKenzie,
pers. comm.).

BIRDS - unident. birds.
Bemier Island.
Unspecified site (A. Baines, pers.comm.).
unident. birds. ;
(b) OCEANIC ISLANDS.
Norfolk Island.
(i) Cemetery Beach dunes (Rich et al. 1983; Meredith
1985b; van Tets et al. 1988).

REPTILES - Gekkonidae: Phyllodactylus guentheri.

Scincidae: Lieolopisma lichenigera.

BIRDS - Procellariidae: Pterodroma solandri,
Pterodroma pycrofti, Pterodroma n.sp., Puffinus
pacificus, Puffinus assimlis, Pachyptila sp.,
Pelagodroma marina.

Sulidae: Sula dactylatra, Sula n.sp.

Phaethontidae: Phaethon rubricauda.

Accipitidae: Accipiter cf. fasciatus.

Rallidae: Gallirallus philippensis, Gallirallus n.sp.

Charadriidae: Pluvialis dominica, Charadrius cf.
bicinctus.

Scolopacidae: Numenius phaeops, Limosa lapponica,
Limosa haemastica, Coenocorypha prob.
aucklandica.

Laridae: Sterna fuscata.

Columbidae: Hemiphaga spadicea, Gallicolumba cf.
norfolcensis.

Psittacidae: Nestor
novaezelandiae.
Cuculidae: Eudynamis prob. taitensis.

Strigidae: Ninox cf. undulata.
Campephagidae: Lalage cf. leucopyga.

Muscicapidae: Turdus poliocephalus, Pachycephala
pectoralis.

productus, Cyanoramphus

Acanthizidae: Gerygone cf. igata.
Stumidae: Aplonis prob. fusca.
MAMMALS - Muridae: Rattus exulans.
Vespertilionidae: unident. vespertilionid bat (S.Hand,
pers.comm.).
(ii) Emily Bay (Meredith 1985b).
BIRDS - Procellariidae: Pterodroma pycrofti, Puffinus
assimilis, Pelagodroma marina.
(iii) Nepean Island (Rich et al. 1983; Meredith 1985b; van
Tets et al. 1988).
BIRDS - Spheniscidae: Eudyptes chrysocome.
Procellariidae: Pterodroma pycrofti, Pterodroma n.sp.,
Puffinus pacificus, Puffinus assimilis, Pelagodroma
marina.
Sulidae: Sula dactylatra, Sula tasmani.
Accipitidae: Accipiter cf. fasciatus.
Rallidae: Gallirallus philippensis, Gallirallus n.sp.
Charadriidae: Pluvialis dominica, Charadrius cf.
bicinctus.
Scolopacidae: Numenius phaeops, Limosa lapponica,
Calidris sp.
Laridae: Sterna fuscata, Gygis alba, Anous stolidus.
Columbidae: Hemiphaga spadicea, Chalcophaps indica.
Psittacidae: Nestor productus, Cyanoramphus
novaezelandiae.
Cuculidae: Eudynamis cf. taitensis.
Strigidae: Ninox cf. undulata.
Campephagidae: Lalage cf. leucopyga.
Muscicapidae: Turdus poliocephalus, Pachycephala
pectoralis.
Lord Howe Island.
REPTILES - Meiolaniidae: Meiolania platyceps (see
Gaffney 1983 for localities).
Unspecified sites (Rich & van Tets 1982; Meredith,
pers.obs.).
BIRDS - Spheniscidae: Eudyptula sp. (prob. minor).
Procellariidae: Pterodroma solandri, Pterodroma
pycrofti, Puffinus carneipes, Puffinus pacificus,
Puffinus assimilis, Pelagodroma marina, Fregetta
grallaria.

1372 - MEREDITH

Sulidae: Sula dactylatra, Sula n.sp.
Rallidae: Tricholimnas sylvestris, Notornis alba.
Columbidae: Columba vitiensis,

Kermadec Islands.
Raoul Island (Anderson 1980).

BIRDS - Procellariidae: Pterodroma spp. (four species),
Puffinus pacificus, Puffinus assimilis.

Pelecanoididae: Pelecanoides urinatrix.

Columbidae: unident. pigeon.

Meliphagidae: Prosthemadera novaeseelandiae.

MAMMALS - Muridae: Rattus exulans.

Otariidae: Arctocephalus pusillus.

Chatham Islands.
(Compiled by P.Millener.).

BIRDS - Spheniscidae: Aptenodytes forsteri, Eudyptula
minor chathamensis, Eudyptes sclateri.

Diomedeidae: Diomedea epomophora, Diomedea
Ichlororhynchus, Diomedea bulleri, Diomedea cauta,
Phoebetria sp.

Procellariidae: Macronectes halli, Daption capense,
Pterodroma lessonii, Pterodroma inexpecata,
Pterodroma neglecta*, Pterodroma ?magentae*,
Pterodroma cf, ultima, Pterodroma nigripennis,
Pterodroma axillaris, Pachyptila vittata, Pachyptila
turtur, Pachyptila crassirostris, Procellaria cinerea,
Procellaria aequinoctialis, Puffinus carneipes,
Puffinus bulleri, Puffinus griseus, Puffinus
tenuirostris, Puffinus gavia, Puffinus assimilis,
Garrodia nereis, Pelagodroma marina, Fregatta
tropica

Pelecanoididae: Pelecanoides urinatrix.

Sulidae: Morus serrator, Sula sp.

Phalacrocoracidae: Phalacrocorax carbo, Leucocarbo
carunculatus, Stictocarbo punctatus.

Anatidae: Cygnus sumnerensis, Tadorna variegata, Anas
superciliosa, Anas 2gibberifrons, Anas aucklandica,
Anas rhychotis, Aythya novaeseelandiae, Mergus
australis, Pachyanas chathamica.

Accipitridae: Haliaeetus australis, Circus approximans,

Falconidae: Falco novaeseelandiae.

Rallidae: Gallirallus dieffenbachii, Gallirallus modestus,
Porzana tabuensis, Porphyrio porphyrio, Fulica
chathamensis.

Haematopodidae: Haemotopus chathamensis.

Scolopacidae: Limosa lapponica, Arenaria interpres,
Coenocorypha_ chathamica, Coenocorypha
aucklandica, Calidris canutus.

Stercorariidae: Stercorarius skua, Stercorarius
Dlongicaudus,

Laridae: Larus dominicus, Larus scopulinus,
Hydroprogne caspia, Sterna striata.

Columbidae: Hemiphaga novaeseelandiae
chathamensis.

Psittacidae; Nestor meridionalis, Cyanoramphus
novaezelandiae, Cyanoramphus auriceps.

Cuculidae: Chrysococcyx lucidus.

Strigidae: Sceloglaux albifacies.
Motacillidae: Anthus novaeseelandiae.
Sylviidae: Bowdleria punctata.

Muscicapidae: Petroica macrocephala, Petroica traversi.
Meliphagidae: Prosthemadera novaeseelandiae.
Corvidae: Palaeocorax moriorum,

MAMMALS - seals.

Varanidae: Varanus cf. indicus.

CROCODILES - Eusuchia: Mekosuchus inexpectatus,

BIRDS - Incertae sedis: Sylviornis neocaledoniae.

Accipitridae: Circus approximans, Pandion haliaeetus.

Falconidae: Falco peregrinus.

Megapodiidae: Megapodius indet.

Tumicidae: Turnix varia.

Rhynochetoidae: Rhynochetus jubatus.

Rallidae: Notornis kukwiedei, Porzana tabuensis,
Tricholimnas lafrasneyanus, cf. Tricholimnas,
Gallirallus philippensis.

Columbidae: Ducula goliath, Ptilinopus greyt.

Tytonidae: Tyto longimembris, Tyto alba.

Aegothelidae: Aegotheles sayesi.

Passeriformes: indet. passerines.

MAMMALS - Rhinocerotidae: "Zygomaturus"
diahotensis*.

Muridae: Rattus exulans.

Chiroptera: indet. bats.

Fiji Islands.
Naigani and La Kemba Islands (R.F.Baird, pers.comm.).

BIRDS - Megapodiidae: megapode n.sp.
Columbidae: pigeon n.sp.

Cook Islands.
Mangaia (Steadman 1985, 1986).

REPTILES - Gekkonidae: Gehyra oceanica.

BIRDS - Procellariidae: Pterodroma sp., Puffinus sp.

Oceanitidae: Nesofregetta fuliginosa.

Phaethontidae: Phaethon rubricauda, Phaethon lepturus.

Anatidae: Anas superciliosa.

Rallidae: Porzana tabuensis, Porzana rua, Gallirallus
ripleyi.

Laridae: Gygis alba,

Columbidae: Gallicolumba sp., Ducula sp., Ptilinopus
Sp.

Psittacidae: Vint cf. kuhlii.

Alcedinidae: Halcyon mangaia.

Sylviidae: Acrocephalus kerearako.

MAMMALS - Muridae: Rattus exulans.

Pteropidae: Pteropus sp.

Macquarie Island.
(i) Finch Creek (McEvey & Vestjens 1974; Museum of

Victoria collection).

BIRDS - Spheniscidae: Eudyptes chrysolophus
schlegeli, Aptenodytes patagonica.

Procellariidae: Puffinus (small sp.), Halobaena caerulea,
Pachyptila prob. desolata,

Pelecanoididae: Pelecanoides cf. urinatrix.

Unident. birds.

(ii) Bauer Bay (McEvey & Vestjens 1974).

BIRDS - Spheniscidae: Aptenodytes patagonica,
Eudyptes chrysolophus schlegeli.

(iii) Aurora Cave (Vestjens 1963; Museum of Victoria

collection; Meredith, pers.obs.).
BIRDS - Diomedeidae: Diomedea exulans.
Procellariidae: Puffinus cf. griseus, Procellaria cinerea,
Pterodroma lessoni, Halobaena caerulea.
Stercorcariidae: Stercorcarius skua.
Anatidae: Anas sp.

Rallidae: Gallirallus philippensis macquariensis.

New Caledonia. (iv) Eagle Cave (Vestjens 1963).
For localitites, see Balouet, this volume. BIRDS - Rallidae: Gallirallus philippensis

REPTILES - Meiolaniidae: Meiolania mackayi, macquariensis.
Meiolania sp. (v) Brothers Point Cave (Meredith, pers.obs.).

BIRDS - Procellariidae: Pachyptila prob. desolata.

VERTEBRATE FOSSIL FAUNAS FROM ISLANDS - 1373

PLATES

Plate 1. Top row, left to right: A, dorsal view of the cranium and rostrum of Pterodroma pycrofti (fossil,
Norfolk Island); B, palmar view of the humerus of P. pycrofti (fossil, Norfolk Island); C, anterior view of the
tibiotarsus of P. pycrofti (fossil, Norfolk Island); D, anterior view of the tarsometatarsus of P. pycrofti (fossil,
Norfolk Island); E, lateral view of the skull of Plerodroma solandri (recent, Lord Howe Island); F, lateral view
of the rostrum of P. solandri (archaeological material, Norfolk Island); G, anterior view of the tarsometatarsus
of P. solandri (recent, Lord Howe Island); H, Anterior view of the tarsometatarsus of P. solandri (archaeological
material, Norfolk Island). Bottom row, left to right: I, dorsal view of the skull of Puffinus pacificus (recent,
Norfolk Island); J, dorsal view of the skull of P. pacificus (fossil, Norfolk Island); K, anterior view of the
tibiotarsus of P. pacificus (fossil, Norfolk Island); L, palmar view of the humerus of Puffinus assimilis (fossil,
Norfolk Island); M, palmar view of the ulna of P. assimilis (fossil, Norfolk Island); N, lateral view of the
femur of P. assimilis (fossil, Norfolk Island).

Plate 2(a). Left to right: A, palmar view of the ulna of Plerodroma sp. nov. (fossil, Norfolk Island); B,
anterior view of the tarsometatarsus of Pterodroma sp. nov. (fossil, Norfolk Island); C, anterior view of the
femur of Plerodroma sp. nov. (fossil, Norfolk Island); D, palmar view of the humerus of Pterodroma sp. nov.
(archaeological material, Norfolk Island - note charring).

Plate 2(b). Top row, left to right: F, palmar view of the humerus of Coenocorypha aucklandica
(subfossil, Auckland Island); F, palmar view of the humerus of Coenocorypha prob. C. aucklandica (fossil,
Norfolk Island), Bottom row, left to right: G, posterior view of the tibiotarsus of C. aucklandica (recent,
Auckland Island); H, posterior view of the tibiotarsus of Coenocorypha prob, C. aucklandica (fossil, Norfolk
Island); I, anterior view of the tibiotarsus of Coenocorypha prob. C. aucklandica (fossil, Norfolk Island); J,

anterior view of the tibiotarsus of C. aucklandica (subfossil, Auckland Island).

Plate 2(c). Top row, left to right: K, anterior view of the tarsometatarsus of Pelagodroma marina (fossil,

Norfolk Island); L, posterior view of the tarsometatarsus of P. marina (fossil, Norfolk Island); M, posterior view of
the femur of Fregetta grallaria (fossil, Norfolk Island); N, anterior view of partial femur of Pachyptila sp. (fossil,

Norfolk Island); P, posterior view of partial femur of Pachyptila sp. (fossil, Norfolk Island), Middle row: Q,
lateral view of the rostrum of Phaethon rubricauda (fossil, Norfolk Island). Bottom row, left to right: R, interior

view of the coracoid of Sterna fuscata (fossil, Norfolk Island); S, palmar view of the humerus of Gygis alba (fossil,
Norfolk Island).

Plate 3. Top row, from left to right: A, palmar view of the ulna of Numenius phaeops (fossil, Norfolk Island);
B, anterior view of the tibiotarsus of N, phaeops (fossil, Norfolk Island), C, anterior view of the tarsometatarsus of
Limosa lapponica (fossil, Norfolk Island); D, posterior v iew of the tarsometatarsus of

L. lapponica (fossil, Norfolk Island); E, palmar view of the humerus of Pluvialis dominica (fossil, Norfolk Island);
F, anconal view of the humerus of P. dominica (fossil, Norfolk Island); G, anterior view of the tarsometatarsus of
P. dominica (fossil, Norfolk Island). Bottom row, left to right: H, dorsal view of the cranium of Sula dactylatra
(fossil, Norfolk Island); I, dorsal view of the rostrum of S. dactylatra (fossil, Norfolk Island); J, anterior view of
the tarsometatarsus of S. dactylaira (fossil, Norfolk Island).

Plate 4. Top row, left to right: A, palmar view of partial humerus of cuculid, probably Eudynamis taitensis

(fossil, Norfolk Island); B, posterior view of partial femur of Eudynamis prob. taitensis (fossil, Norfolk Island); C,
posterior view of partial tarsometatarsus of Eudynamis prob. taitensis (fossil, Norfolk Island). Middle row, left to
right: D, palmar view of the humerus of Turdus poliocephalus (fossil, Norfolk Island); E, anterior view of the

tibiotarsus of T. poliocephalus (fossil, Norfolk Island); F, palmar view of the humerus of Gerygone olivacea (

Recent, Victoria); G, palmar view of the humerus of Gerygone prob. igata (fossil, Norfolk Island); H, interior view
of partial coracoid of Pachycephala prob, pectoralis (fossil, Norfolk Island). Bottom group, top to bottom: I,

posterior view of the tarsometatarsus of Ninox undulata (fossil, Norfolk Island); J,. anconal view of the radius of N.
undulata (fossil, Norfolk Island), K, palmar view of the humens of N. undulata (fossil, Norfolk Island).

Plate 5. Top row, left to right: A, anconal view of the humerus of Cyanorahmphus novaezelandiae (fossil,

Norfolk Island); B, posterior view of the tarsometatarsus of C. novaezelandiae (fossil, Norfolk Island); C, palmar
view of the ulna of C. novaezelandiae (fossil, Norfolk Island); D, lateral view of the rostrum of Nestor productus
(fossil, Norfolk Island); E, dorsal view of the mandible of N. productus (fossil, Norfolk Island); F, posterior view
of the tarsometatarsus of N. productus (fossil, Norfolk Island). Middle row, left to right: G, palmar view of the
humerus of Hemiphaga novaeseelandiae (fossil, Norfolk Island); H, lateral view of the sternum of H.

novaeseelandiae (fossil, Norfolk Island). Bottom row, left to right: I, interior view of the carpometacarpus of
Chalcophaps indica (fossil, Norfolk Island); J, posterior view of the tarsometatarsus of Gallicolumba jobiensis

1374 - MEREDITH

(Recent, New South Wales); K, posterior view of partial tarsometatarsus of ““~/licolumba prob. norfolciensis
(fossil, Norfolk Island); L, intemal view of the coracoid of G. jobiensis (Recent, New South Wales); M, intemal
view of partial coracoid of Gallicolumba prob. norfolciensis (fossil, Norfolk Island).

Plate 6. Top row, left to right: A, troclea for digit IV of the right tarsometatarsus of Eudyptes prob. chrysocome
(fossil, Norfolk Island); B, posterior view of the distal end of the tarsometatarsus of Accipiter fasciatus (fossil,
Norfolk Island); C, posterior view of the proximal end of the tarsometatarsus of A. fasciatus (fossil, Norfolk
Island); D, palmar view of partial ulna of Charadrius bicinctus (fossil, Norfolk Island). lMiddle row, left to right:
E, posterior view of the femur of Lalage prob leucopyga (fossil, Norfolk Island); F, posterior view of the
tibiotarsus of Lalage prob. leucopyga (fossii, Norfolk Island); G, anconal view of the ulna of Aplonis prob. fusca
(Recent, Lord Howe Island); H, anconal view of the ulna of Aplonis prob. fusca (fossil, Norfolk Island). Bottom
row, left to right: I, posterior view of the femur of a bat (fossil, Norfolk Island); J, lower jaw of Rattus exulans
(fossil, Norfolk Island); K, L, jaw bones of gecko (fossil, Norfolk Island).

Plate 7. Top row, left to right: A, palmar view of the humerus of Gallirallus philippensis (Recent, Victoria); B,
palmar view of the humerus of Gallirallus sylvestris (fossil, Lord Howe Island); C, palmar view of the humerus of
Gallirallus sp. nov. (fossil, Norfolk Island); D, anterior view of the tarsometatarsus of Tricholimnas lafrasneyanus
(Recent, New Caledonia); E, anterior view of the tarsometatarsus of G. philippensis (fossil, Norfolk Island); F,
anterior view of the tarsometatarsus of Gallirallus sp. nov. (fossil, Norfolk Island). Lower, top to bottom: G, left
side view of the pelvis of G. philippensis (Recent, Victoria); H, left side view of the pelvis of Gallirallus sp. nov.
(fossil, Norfolk Island); I, right side view of the pelvis and string of vertebrae of G. sylvestris (fossil, Lord Howe
Island).

PLATE 1 VERTEBRATE FOSSIL FAUNAS FROM ISLANDS - 1375

PLATE 2

1376 - MEREDITH

PLATE 3 VERTEBRATE FOSSIL FAUNAS FROM ISLANDS - 1377

1378 - MEREDITH PLATE 4

PLATE § VERTEBRATE FOSSIL FAUNAS FROM ISLANDS - 1379

1380.- MEREDITH PLATE 6

PLATE 7 VERTEBRATE FOSSIL FAUNAS FROM ISLANDS - 1381

1382 - MEREDITH

Quipollornis koniberi, a primitive owlet-nightjar (Aegothelidae) from Miocene-aged sediments
near Coonabarabran, New South Wales. This form, unlike its living Australian relative, was
more adapted to aerial hawking for insects rather than a life closer to the ground. (From Rich &
van Tets 1985, with permission of The Museum of Victoria).

CHAPTER 29

THE FOSSIL VERTEBRATE
RECORD OF NEW
CALEDONIA

Jean Christophe Balouet !

THtrOdtiCtion <.56 ects ccccses vee eda htereraedeceewtdtees 1384
Palaeontological Sites... eeeeeeeeeeeeeeeeee 1384
Loyalty Islands ...........ccccceeeeeeeeeee sees eeeeees 1385
Living or Recently Extinct Terrestrial Vertebrates
Of New Caledomnta...........cc.csccccsssseeeeeeesees 1386
Extinct Fossil Vertebrates of New Caledonia.... 1388
Fossil Reptiles ...........ccccseeceeeeeeeeeeeeneeenes 1388
SqQuamatar.c.::Aciseses decneaes stp neniheatcensss 1388
ChelOmia ns reccedcby sha deawteaste steele. segs 1388
Crocodilomorphs............ceceeeeeeeeeeeees 1389
FOSSIL: Bird sid vedecescs oBhecBee usenet dedstes one dations 1390
SYLVIOTNIS v..c..cpecencnngessdesteageetedtqnestes 1390
| CAS Lg COSC ee 1392
Other Fossil Birds...........cccccecseeeeeee ees 1394
Mammals. ols. iecceveinccacnteecesseg te adeeweeesseete 1395
Extant Vertebrates of New Caledonia
with a Fossil Record ............eeeeeeeeee ees 1395
Palacobiogeography.......ccccccceessreeeceeeerseeeeees 1396
Geological History of New Caledonia ........ 1396
New Caledonian Endemism and Modern
Dispersal..........cccccccceeeeeceeeceeeeeseceeees 1396
CORGIUSIONS vececnecvachee deve cess soles ods ctede bacucewe cei 1397
R@FETENCES:....ccccvckescienccctedeccdencasesccesessuesnees 1399
PU ALES ors vicleiies She cloals coticolvseiteacet seen aelelndtieades Semaine 1401

nn sneer

1 Institutut de Paleontologie, Museum national d'Histoire Naturelle, 8 rue Buffon, 75005 Paris, France.

1384 - BALOUET

INTRODUCTION

Most of the fossil vertebrates from New Caledonia are no older than a few centuries.
However, even these fossils, many discovered quite recently, differ from the living forms.
Over 15,000 bones have been collected from the 17 localities known to date. Although the
first finds were made in the middle of the last century (Filhol 1876), until recently little has
been published on this subject (Balouct 1984a, 1987). This paper is the first synthesis of all
the known information regarding the fossil vertebrate record of New Caledonia.

Intense field and laboratory research on New Caledonian fossil vertebrates began in 1979,
and it is now possible to report on the results of this work. This paper will discuss the main
palacontological sites, some of the most interesting fossils and some of the
palacobiogeographic consequences of these findings.

PALAEONTOLOGICAL SITES

Seventeen vertebrate bearing localities are known to date: two in the Mesozoic and fifteen
in the Quaternary (Fig. 1).

( Belep
20°S
Paaba © . Balabio =
Diahot ii |.Ouvea
Ct
%& |.Tiga
Pindai
Cy |.Mare
Teremba ae
22
Gilles e3
|.Page ;
Noumea
0 100km !
LY Ww lle des Pins Walpole
| |
164°E 166°E

Figure 1. Major localities producing fossil vertebrates in New Caledonia and on nearby islands.

The two Mesozoic localities are on the west coast of mainland New Caledonia. The fossil
material from both localities is poor and fragmentary. The Carnian Volcanic Series of the
Teremba Peninsula has yielded a jaw fragment and a fragment of a long bone. On Page Island
(St Vincent Bay) thoracic ribs and vertebrae were found in 1980 and 1982. All of these
fragments belong to ichthyosaurs. The fifteen Quaternary localities occur on the Loyalty
Islands, the Isle of Pines (Fig. 2) and the mainland.

FOSSIL VERTEBRATES OF NEW CALEDONIA - 1385

167°30'

Calcarenites

E= uplitted Reef Complex
= Lateric High Plain
Fossil =
Bearing Sinkholes == ma Gabbros
GOEg Serpentinites
Peridotites

Figure 2. Geology of the Isle of Pines. Main vertebrate fossil localities occur as sink hole accumulations
in the uplifted reef limestone. (After Launay in Rich et al. 1981).

LOYALTY ISLANDS

Tiga Island has produced five fragmentary bones, collected in 1954 in phosphatic deposits
(Obelliane 1958). The estimated age of these phosphates is Late Pleistocene to Holocene. The
most diagnostic fragment is a centrum of a seventh cervical vertebrae (Gaffney et al. 1984) of a
meiolaniid turtle.

On Walpole Island a few bones were collected in 1910 of an endemic terrestrial turtle
(Meiolania mackayi). These are housed in the Australian Museum in Sydney. The
sedimentary history of these deposits is probably the same as Tiga's (Recy et al. 1975), making
them Late Pleistocene to Holocene in age.

Mare Island (Menaku) has yielded four avian bones, which have been attributed to the
Rallidae and the Columbiformes. They were found in a cornice, not far from a human
cemetery, and their age is unknown.

Lifou Island has produced a partial human skeleton, found in 1980 on We Beach. The
calcareous deposits containing it could have been deposited during the last marine transgression,

1386 - BALOUET

about 2500 years ago. The specimen, that of a juvenile, is housed in the Paris Museum of
Natural History.

On the Isle of Pines a locality (Kanumera) was discovered in 1974 thanks to a local legend
(Dubois 1976, Rich et al. 1981). Four fossiliferous sink holes are now known from this area.
These contain accumulations of bones and coral reef breccia in holes that developed in the
uplifted reef (Rich et al. 1981). The bones of the largest New Caledonian bird have been dated
at 3470 + 210 yBP (Poplin et al. 1984). Over two tons of the fossiliferous breccia were
collected during the French expeditions to Kanumera in 1980 and 1981. During a cooperative
Australian-New Caledonian expedition in 1979 (Rich et al. 1981), several hundred bones were
also collected.

Mainland New Caledonia contains several localities that have yielded significant
accumulations of vertebrate fossils in the Gilles and Pindai areas as well as a number of faunal
bearing archaeological sites.

Three caves in the Gilles area have produced bones. The guano-rich deposits, mainly
accumulations left by bats and swifts, have yielded bones of reptiles (geckoes, scincids and
Varanus), bats, rats and large quantities of bird bones, including the giant bird also collected in
Kanumera (Sylviornis neocaledoniae ). Most of the small vertebrates appear to have been
accumulated by owls. In the uppermost layers, rat bones are very abundant, and marine shells
brought in by man, such as Arca and Ostrea,can been found. The history of the cave
sediments is complex, because of rock falls and water activity in the caves. Carbon dating is in
progress for the deepest, rat-bearing layers (Rattus exulans), in order to determine the timing of
the first human arrival in New Caledonia.

The Pindai caves are the richest mainland localities, Fossils were first discovered in 1983,
and to date over 10,000 bones have been collected, out of four tonnes of sediment. The
fossiliferous sediments are primarily clays, gypsum and phosphates. Bones evidently
accumulated in an underground lake, which is now empty. Carbon dating on charcoals resulted
ina 1750 £70 yBP (Gif 6341),

Archaeological sites containing Lapita pottery in the Koumac and Nessadiou (west coast)
areas have also yielded bones. These belong to reptiles (meiolaniid and the crocodile
Mekosuchus), birds (including Sylviornis) and mammals (sea-cows and bats).

LIVING OR RECENTLY EXTINCT TERRESTRIAL
VERTEBRATES OF NEW CALEDONIA

Besides introduced species, the modern fauna of New Caledonia is composed of reptiles (9
species of geckonids and 23 species of scincids), birds (111 species) and mammals (4 species of
bats). The fossil fauna (Table 1) differs somewhat from the modern fauna and includes the
following terrestrial vertebrates that are both endemic to New Caledonia and currently extinct:
reptiles (Mekosuchus inexpectatus, a eusuchian; Meiolania mackayi, a meiolaniid) and birds
(Sylviornis neocaledoniae, family incertae sedis; Megapodiidae; Porphyrio kukwiedei,
Tricholimnas cf lafresnayanus, both rails; Aegotheles savesi, an aegothelid).

Two other species, known from the New Caledonian subfossil record, are now extinct in
New Caledonia, but are still alive in other South Pacific countries: Varanus cf indicus, a
varanid which is probably specifically distinct from the New Guinea form (but for the moment
the New Caledonian fossil form will be closely allied to the New Guinean form because of the
lack of comparative material); and one owl (Ninox novaeseelandiae, a strigid).

Some of the fossil New Caledonian species became extinct very recently. This is the case

FOSSIL VERTEBRATES OF NEW CALEDONIA - 1387

rr a a EN EAR AT AE SOS

Table 1: Quatemary Fossil Vertebrates from New Caledonia

Taxon Pindai Gilles Kanumera Menaku Walpole Tiga
REPTILIA
Mekosuchus inexpectatus
Meiolania
Varanus cf indiccus
Gekonidae
Scincidae
AVES

Urodynamis taitensis
Falco peregrinus
Accipiter efficas, new species
Accipiter quartus, new species
Megapodius molistructor,
new species
Sylviornis neocaledoniae
Turnix (varia) novaecaledoniae
Ducula goliath
Drepanoptila holosericea
Columba vitiensis
Chalcophaps indica
Caloenas canacorum,
new species
Gallicolumba longitarsus,
new species
Halcyon sancta
Tyto ? letocarti, new species
Tyto alba
Ninox cf. N. novaeseelandiae
Aegotheles savesi
Collocalia spodiopygia
Collocalia esculenta
Porzana tabuensis
Gallirallus philippensis
Tricholimnas lafresnayanus
Gallinula cf. G. tenebrosa
Porphyrio kukwiedei,
new species
Porphyrio porphyrio
Rhynochetos orarius,
new species X
Charadrius mongolus -
Pluvialis dominica -
Coenocorypha? species -
Anas gracilis x 5 3
Pterodrama rostrata X

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1388 - BALOUET

with Varanus cf indicus, the victim of a car collison 20 years ago near Bouloupari. A
local legend relates that a bird which was probably Porphyrio kukwiedei was still alive
in the 1870 (Balouet 1984a, 1984b). Live specimens of Aegotheles savesi and
Tricholimnas lafresnayanus were collected in the 1900's. Tyto longimembris and
Turnix neocaledoniae were both observed by Delacour (1948), and Sylviornis
neocaledoniae and Mekosuchus inexpectatus remains have been found in the
archaeological deposits of Koumac and Nessadiou.

The oldest human artifacts in New Caledonia are dated at 3000 yBP (Frimigacci
1980). Absolute age of the Pindai and Kanumera sites, as well as archaeological
findings and ethnological relative dates, associated with remains of now extinct fauna,
indicates that the arrival of humans and their introduction of exotic fauna (e.g. Rattus
exulans) played some part in the extinguishing of a number of species (Balouet 1987).
Later European settlers and their dogs, cats, pigs and goats are responsible for further
decimation of the already depauperate fauna. This is neither the first nor the last
example of the fragility of island faunas, easily disturbed by outside influences.

EXTINCT FOSSIL VERTEBRATES OF NEW CALEDONIA

FOSSIL REPTILES
Squamata (Lizards)

One hundred bones of Varanus cf inducus (Fig. 3B, Pl. 1) have been collected at Pindai, and
one vertebra has been recovered from Gilles. Identification to the order Varanoidea is possible
because the haemapophyses are in a subcentral position and are articulated (Estes 1983); to the
family Varanidae because the scapular fenestra is lacking on the scapulocoracoid (Lecuru 1968),
the ectopterygoid separates the maxillary and the suborbital fenestra (Estes 1983), and there is a
distinctive type of kinesis in the intramandibular articulation. The family Varanidae in New
Caledonia today includes two genera, the fossil genus /berovaranus and the modern genus
Varanus. The less elongated vertebrae and more transversely elongated condyles (Hoffstetter
1968) are characteristic features that allow placement of the New Caledonian fossil within the
genus Varanus.

The genus Varanus is well represented in the Australasian region (12 species in New
Guinea, 19 in Australia). Unfortunately, comparative material is rare; the species Varanus
karlschmidti is known only from the type specimen (a skin). For two other species, the only
available comparative material consists of an illustration of the skull. The New Caledonian
species, based on the material available for comparison, however, is most similar to the New
Guinean species Varanus grayi and V. indicus, which with the New Caledonian fossils share an
identical shape of the sagittal process and similar development of the posterolateral process.
The New Caledonian fossil differs from Varanus grayi in its higher and less curved dentary as
well as in its very elongate teeth.

The New Caledonian fossil Varanus differs from all species to which it could be compared
in having very elongated tecth and an elongate proximo-dorsal depression of the maxillary.
Because of the lack of a complete comparative collection, it is still not possible to determine
with certainty, however, the specific status of the New Caledonian varanid.

Chelonia (Turtles)

Cryptodire turtles are represented in the fossil assemblages by the terrestrial horned turtle,
Meiolania, which belongs to an extinct family, the Meiolaniidae (Pl. 3), restricted to the

FOSSIL VERTEBRATES OF NEW CALEDONIA - 1389

Figure 3. Mammalian and reptilian fossil vertebrates from New Caledonia: A, "Zygomaturus diahotensis";
B, Varanus cf indicus, premaxillary. Mekosuchus inexpectatus: C, premaxillary, showing the anterolateral
opening of the nares; D, procoelous vertebra; all x1.

Southern Hemisphere. The first remains of this endemic turtle, Meiolania mackayi, were
found on Walpole Island (Andrews 1922, Anderson 1925). Other specimens of meiolanids
were collected in 1954 (Obelliane 1958) from Tiga Island. A partial vertebra, identified as a
seventh cervical (Gaffney et al. 1984) is a meiolaniid. A complete meiolaniid seventh cervical
vertebra has also been found in the caves at Pindai. The New Caledonian species may be
distinct from the Lord Howe Island form.

Crocodilomorphs (Crocodilians)

The marine crocodile, Crocodylus porosus, is known in the southwest Pacific (Fiji, the
Solomons), but the discovery of Mekosuchus inexpectatus (Figs 3C,D, 4, Pl. 2) (Buffetaut
1983, Balouet & Buffetaut 1987) was totally unexpected. This crocodile is known from
Kanumera and Pindai - over three hundred bones have been collected on Pindai alone, and most
of the skeleton of this species is actually known. The first evidence of the species was a
quadratojugal from the Kanumera site, discovered in 1981 (Buffetaut 1983).

The New Caledonian crocodile is considered to be terrestrial due to the presence of
anterolaterally opening nares, strong muscular insertions on the limb bones and well developed
dorsal spines on the cervical vertebrae.

Mekosuchus exhibits a number of plesiomorphic (primitive) characteristics, such as the
dorsal insertion of the postorbital process on the quadratojugal, characteristic of mesosuchian

1390 - BALOUET

crocodiles (Balouet & Buffetaut 1987). However, the skull of this crocodile exhibits some
apomorphic (advanced) characteristics, which include: a prominent posterior development of
the pterygoids, an orbital fenestra that is limited ventrally by the maxillary and very narrow
palatine bones.

Mekosuchus belongs to the modern Eusuchia. Vertebrae are procoelous, and the choanae
open within the pterygoid. The genus differs from modern Crocodylidae in that it lacks the
quadratojugal spine. The Alligatoridae, a second family of Eusuchia, is not well defined. The
important development of the fourth maxillary tooth could be characteristic of the recent
alligatorids, but this is not a well established characteristic for all known alligatorids as of yet.

There is no synapomorphy that allows Mekosuchus to be placed in one of the two families
of Eusuchia (Crocodylidae or Alligatoridae). The Mekosuchidae is considered as a sister group
to both of these modern families (Fig. 4).

The length of Mekosuchus is estimated at 2 metres. It probably fed on molluscs based on
the morphology of its teeth (tribodont).

FOSSIL BIRDS
Sylviornis

The avifauna is the main component of both the living and the fossil fauna of New
Caledonia. One of the most unexpected elements of the fossil avifauna of New Caledonia is
the ground-dwelling Sylviornis neocaledoniae (Figs 5,6, Pl. 4-6).

Sylviornis was a neognathous bird. Its palatines are fused in a symphysis. This
characteristic, as well as the location of the ascending process of the tarsometatarsus
(MacGowan 1984), suggests that Sylviornis should be included in the Neognathae. Although
first described as a ratite (Poplin 1980), this form has recently been reinterpreted as a megapode

Mekosuchus Alligatoridae Crocodilidae

4th maxillary
tooth developed

posterior development
of pterygoids

narrow palatines

maxillary at orbit quadrotojugal spine

dorsal insertion of postorbital process

procoelous vertebrae

choanae within the pterygoid

Figure 4. Cladogram illustrating the relationship of Mekosuchus to other eusuchians.

(Poplin et al, 1984). Thanks to the Pindai discoveries (including a complete skull and
4,000 bones belonging to about 30 individuals) the skeleton of this genus is now almost
completely known. However, I have not been able to discern a single synapomorphy that
could clarify the suprageneric position of this strange form. Although a large number of
characteristics are shared with different taxa within the Galliformes, none can be seen as clearly
synapomorphic.

Autapomorphies of Sylviornis include: a palatine symphysis; an articular facet located on

FOSSIL VERTEBRATES OF NEW CALEDONIA - 1391

the posterior part of the quadrate for the posterior articular process; a massive quadratojugal,
which is S-shaped and has a strongly developed posterior process articulating with the quadrate;
a deep socket on the quadrate for the quadratojugal; a cranio-facial hinge developed along the
anterior width of the skull; the anterior part of the lacrimals bearing a rounded articular facet
for diarthrosis with the nasal bone (these two characteristics represent a new type of cranio-
facial kinesis); a basitemporal plate forming a 110° angle with the basisphenoidal rostrum; an
axis of the occipital condyle that lies parallel to the basisphenoidal rostrum; a nasal bone and
nasal process of the premaxillary that form a large hemispheric exostosis or nasal bump, which
is present on all specimens except juveniles; a nares that opens laterally and very near the jugal
insertion; maxillopalatines that are fused all along their inner surface, and an extremely reduced
sternal facet on the coracoid.

Plesiomorphic characteristics of Sylviornis include those widely distributed in the
Neognathae or present in other vertebrate groups than birds. These include: a stout
basisphenoidal rostrum bearing two well developed basipterygoid processes; a continuous
interorbital septum; temporal fossae that are limited to the lateral parts of (and not above) the
skull; large ilioischiatic fenestra (Olson 1981, Feduccia 1985); equal development of the
transverse processes of the synsacral vertebrae and parallel anterior borders of the ilia.

Some characters present in Sylviornis can also be observed in most reptiles and some
modern birds. There are no uncinate processes on the ribs. Uncinate processes are also absent
in reptiles, some megapodes and in anhimids. Uncinate processes are often considered as a
general avian characteristic. Their systematic distribution leads to two hypotheses: their lack
in Sylviornis could be retention of a character from a reptilian ancestry or they have been lost
since the derivation of this genus from some group of neognathous birds. The fused
claviculocoracoid seen in Sy/viornis also occurs in the hoatzin (Opisthocomus). This might be
interpreted in Sylviornis as a fusion related to the age of the individuals, as only 20% of the
population exhibits the fusion. In many flightless birds, fusion of elements in the pectoral
girdle is also known. The unfused, straight clavicles in Sylviornis, likewise, may be due to
the increasing ground dwelling nature of this form, no longer dependent on flight. Such
modifications are known is some species of living birds (e.g. Atrichornis, the Scrubbirds of
Australia, Rich et al. 1985). The large number of synsacral and free caudal vertebrae (20-25) is
greater than in most Neognathae (generally 11-16). In Archaeopteryx there are 22 caudal
vertebrae.

The osteological comparisons of Sylviornis to other birds leads to four possible
hypotheses about the relationships of this bird. The first, proposed by Poplin (1980) suggests
that Sylviornis was a ratite. Two of the characters mentioned above are shared only with the
moas (Dinornithiformes) - the posteromedian indentation of the sternum and the maximum
width of the skull lying in the nuchal region. Sylviornis, however, differs from the ratites
dramatically in the structure of the tarsus and the palate.

A second possibility is the relationship of Sylviornis to the galliform birds. Poplin et al.
(1984) proposed that Sylviornis shared with the galliforms more characters than with any other
avian group. None of the characters they mentioned, however, appear to be synapomorphic,
because they are also present in many other neognathous birds in addition to the galliforms.

A third possibly related group are the anseriforms. The two families in this order, the
Anhimidae and the Anatidae, strongly differ from each other, and most of the characters shared
with Sylviornis and this group (especially the Anhimidae) are also shared with the Galliformes.
It may be that this order is paraphyletic and its origin is somehow close to that of the
Galliformes. Still, there are no synapomorphic characters linking ducks and their kin with
Sylviornis . ; i.

Sylviornis also shares some features with the gruiforms, especially the Psophioidea. Here
again, however, it has not been possible to recognize a single synapomorphy that would favour
a close link with this group.

1392 - BALOUET

Because of the lack of shared synapomorphies, it is better not to assign Sylviornis to any
suprageneric group, although galliforms are presumably the most similar when all characters
are considered, Future work on definition of polarity of characters in the galliforms and
anseriforms is greatly needed, and that will perhaps lead to a better understanding of just where
Sylviornis originated,

: C
(0) 10cm \
fs :

/ /

Vigure 5, Reconstricted skeleton of Sylviorniy neocaledoniae,
Porphyrio kukwiedei

Previous to the discoveries in New Caledonia, a large Porphyrio was known only in New
Zealand (Porphyrio mantelli + Owen 1848) and on Lord Howe Island (Porphyrio albus ; van
Tets 1974), The New Zealand species was first described as a fossil from North Island, but a
small living colony of this bird was discovered about a century after the first fossil findings.
The first fossil specimens from New Caledonia were collected at Kanumera in 1980, The best

FOSSIL VERTEBRATES OF NEW CALEDONIA - 1393

specimen, a complete tibiotarsus, was collected at Pindai in 1983.
Existence and late survival of the genus Porphyrio in New Caledonia is also attested to by a
legend of the Kele tribe near Moindou. The present day chief related that his grandfather used to

hunt a large bird about 50 cm tall, very similar to the Swamp Hen (Porphyrio porphyrio) both
in shape and colour, but was much larger than the latter.

Figure 6. Sylviornis neocaledoniae. A, dorsal view of skull; B, lateral view of skull; C, anterior view of
left tarsometatarsus; D, anterior view of right tibiotarsus.

1394 - BALOUET

Based on the new material from New Caledonia, a new species, Porphyrio kukwiedei
(Balouct & Olson 1989) was established, based on the complete left tibiotarsus (holotype) and
one quadrate from Pindai, one partial tarsometatarsus from Gilles and fragments of a
uibiotarsus, one axis vertebra and one fibula from Kanumera. This species belongs in the
genus Porphyrio, because it has no tendinal canal on the hypotarsus, the proportions of the
ubiotarsus to the tarsometatarsus is 1:5 and the insertion of the extensor brevis is in contact
with the external cotyla. The distal fragment of the right tibiotarsus from Kanumera is
identical to that of the species Porphyrio mantelli (Brit. Mus. R6608). The left tibiotarsus
from Pindai has a reduced prominence for the inner ligament. Its rotular crest is nearly
horizontal, and there is a well marked depression between the rotular crest and the inner cnemial
crest. On the tarsometatarsus, the proximal inner foramen is located at the base of the main
calcaneal ridge. This character combination within Porphyrio is unique to the New
Caledonian species.

Marked differences separate Porphyrio mantelli from all the species of Porphyrio, including
a specific ectoparasite, Rallicola takahe, which differs from Rallicola lugens found in the
species of Porphyrio . Morphological differences between the Swamp Hen and the Takahe
include size (the weight of the Takahe is two to three times that of the swamp hens); the
stouter and shorter legs of Takahe; the short wings and toes in Takahe ; the shorter primaries in
Takahe in comparison with the length of the secondaries and wing coverts (Rothschild 1907).
Many of the differences noted between these two species may well be related to the degree of
flightlessness in these birds, and quite clearly a functional study of this group would be worth
pursuing.

Despite its taxonomic limbo, some interpretations regarding Sy/viornis can be made. It
was a large, flightless form (1.4 m from beak tip to pygostyle). Its weight is estimated at 30
kg. It was four-toed and had short legs in comparison to its body size (Fig. 5). The
remarkable cranio-facial hinge, similar in width to that of the Dodo, parrots, the Hoatzin and
the Takahe together with the marked development of a nuchal plate make it a very distinctive
species.

Other Fossil Birds

The fossil birds discussed above are the most abundant in the palaeontological deposits, but
seven other species, now extinct in New Caledonia, have also been recovered and are still under
study.

Bones of a large megapode, now extinct, have been collected in the Kanumera area (femur,
phalanges) and at Pindai (tarsometatarsus). This species is slightly smaller than Talegallus.
Megapodes are known from the Loyalty Islands, but the mainland bones, much larger, belong
to a new species, Megapodius molistructor (Balouet & Olson 1989).

Turnix neocaledoniae has not been observed in New Caledonia for more than 40 years. A
fossil tarsometatarsus, found at Kanumera, confirms that this species has not been recently
introduced, but has instead a lengthy history on this island.

Rhynochetus jubatus, the Kagu, is today endemic to New Caledonia. Fossil bones of a
new species, R. orarius (more than 200 specimens from Pindai) are slightly larger than the
modern species. Rhynochetus may be the sister group of the New Zealand Aptornis, and both
of these birds may possibly be related to the South American Psophioidea (Cracraft 1982).

Tricholimnas cf. lafresnayanus is an endemic rail that has not been observed except once
(Balouet 1987) for more than SO years (Delacour 1948). Fossil bones of this species have been
collected at Pindai and Kanumera (Rich e¢ a/. 1981) (Pls 7, 8). The fossil bones show some
differences from the recently extinct species (PI. 9). Another species, Tricholimnas sylvestris,
is known on Lord Howe Island. Olson (1973a,b) considered the Lord Howe species a member
of the genus Gallirallus. Olson further writes that the two species of Tricholimnas "have been

FOSSIL VERTEBRATES OF NEW CALEDONIA - 1395

derived independently and each is more closely related to some volant ancestor than to the other.
Whether this ancestor is the same cannot now be discerned."

Tyto longimembris, the Grass Owl, was mentioned by Delacour (1948), although no
specimens have apparently ever been captured or observed since. Fossil bones of an extinct
owl, Tyto letocarti (Balouet & Olson 1989) have been recovered in Gilles Cave.

Ninox novaeseelandiae, the Boobook Owl, is represented by a complete tibiotarsus
discovered in 1983 at Pindai.

Aegotheles savesi is represented by one specimen. This species is an endemic owlet
nightjar, collected at the beginning of the century. It has not been observed since, but fossil
bones of this aegothelid have been recovered from Gilles Cave (2 humeri and 1 tarsometatarsus)
(Olson, Balouet & Fisher 1987).

Other extinct bird species include two species of Accipiter (A. efficax, A. quartus), one very
large species of pigeon, Caloenas canacorum and a large species of Gallicolumba, G.
longitarsus (Balouet & Olson 1989).

MAMMALS

Although a thousand bat bones have been collected in various palaeontologically productive
deposits, little can be said about these because they are essentially identical to modern forms
(three species on the mainland and one on the Loyalty Islands).

More can be said about an isolated tooth (Fig. 3A, Pl. 10, 11), found in the 1860's in the
Diahot Valley in the north of New Caledonia. The tooth was first identified as a rhinoceros
first molar (Filhol 1876), but more recently Guerin ef al. (1981) and Guerin & Faure (1987)
reinterpreted it to be a diprotodontid premolar and renamed the species as Zygomaturus
diahotensis. Guerin et al. (1981) discussed at length the biogeographic implications of the
tooth and concluded that the Diahot tooth was evidence for interchange of this terrestrial
marsupial between New Caledonia and New Guinea and (or) Australia in the late Cainozoic.
They also mentioned that a similar case could be made for Sylviornis .

In a recent paper by Rich et al. (1987), which is well illustrated, it is clear that the Diahot
tooth is much more similar to a deciduous premolar of a Rhinocerous than to any tooth of
Zygomaturus trilobus, or for that matter any other known diprotodontid. In addition, when the
wear on the enamel of the Diahot tooth was examined, the "transverse wear relief characteristic
of forms with vertical Hunter-Schreger bands is clearly to be seen, although the morphology
suggests that the tooth surface has suffered an amount of damage since the animal was alive...
No such relief is visible on teeth of the diprotodontids, Zygomaturus ... and Diprotodon (Rich
et al . 1987). Independently, Bertrand (1986) came to the same conclusions.

Thus, the Diahot tooth appears to be that of a rhinoceros, most likely transported by human
means to New Caledonia, and has little to say about the biogeography of this island.

No other terrestrial mammals, besides those introduced by human populations, have ever
been found in New Caledonia.

EXTANT VERTEBRATES OF NEW CALEDONIA WITH A
FOSSIL RECORD

Several vertebrates that are still alive today have been found in the palaeontological sites on
New Caledonia and outlying islands. These include:

Reptilia: 3 species of geckonids, at least 3 species of scincids.
Aves: Urodynamis taitensis, Falco peregrinus, Ducula goliath,
Drepanoptila holosericea, Columba vitensis, Chalcophaps indica, Halcyon

1396 - BALOUET

sancta, Tyto alba, Collocalia spondopygia, C. esculenta, Porzana
tabuensis, Gallirallus philippensis, Gallinula cf. G. tenebrosa, Porphyrio
porphyrio, Charadrius mongolus, Pluvialis dominica, Anas gracilis,
Pterodroma rostrata. Over 20 species of passerines have also been
recognized.

Mammalia: 3 species of bats

PALAEOBIOGEOGRAPHY

One of the striking characteristics of the New Caledonian terrestrial vertebrate fauna is the
absence of mammals, excluding, of course, introduced species. Equally striking is the high
degree of endemism of the reptile fauna. Only a few species of scincids are not endemic and
these may have been introduced by humans. Unfortunately, for palaeobiogeographic
considerations, the fossil record is for the most part restricted to the Quaternary and the
Triassic.

Do the terrestrial vertebrates that are known, however, indicate vicariance? Before
proposing just how the New Caledonian fauna came to be what it is today, it is important to
know something about the geological history of this area and something about dispersal of
terrestrial vertebrates.

GEOLOGICAL HISTORY OF NEW CALEDONIA

The oldest New Caledonian rocks are of Devonian and Carboniferous age. Two hundred and
eighty million years ago, New Caledonia lay on the margin of Gondwana, near Queensland.
During the Permian and Early Triassic, New Caledonia was essentially a volcanic arc, lying
along the east of Australia. By the Late J urassic, the mainland of New Caledonia started to
emerge. By the Late Cretaceous, some 80 myBP, the neighbouring ocean basins began to
expand.

New Caledonia remained emergent until the late Eocene. After that, most rocks preserved
in New Caledonia until the Early Miocene are of ophiolitic sequences (Brothers & Black 1974).
In the Early Miocene shallow marine rocks characterise New Caledonia, indicating a marked
marine transgression at that time. Progressive uplift of these rocks sequences from Miocene to
Recent has resulted in the present day topography of this area.

NEW CALEDONIAN ENDEMISM AND MODERN DISPERSAL

The terrestrial molluscan fauna of New Caledonia is 100% endemic (Tillier & Clarke 1983),
as are the terrestrial reptiles. Out of 156 New Caledonian terrestrial vertebrate species not
obviously introduced by humans, 56 are endemic. If seabirds are excluded, as are introduced
species and other cosmopolitan species (such as Tyto alba ), the terrestrial vertebrate fauna is
about 90% endemic.

Of the fossil forms, two endemic taxa are of little interest in palaeobiogeographical
considerations: Sylviornis neocaledoniae, because its affinities are not yet clear, and
Mekosuchus inexpectatus, because its sister group is widespread.

However, other taxa are of distinct interest: these are Meiolania, Varanus, Caloenas,
Megapodius, Rhynochetus, Ninox and Aegotheles. Meiolania occurs in Australia, on Walpole
Island, Lord Howe Island, the Loyalty Islands and the mainland of New Caledonia. Although
no phylogeny has ever been proposed for that genus, Gaffney (1983) considers M. platyceps
(Lord Howe Island) and M. mackeyi (Walpole Island) more closely related to each other than
cither is to M. oweni of Australia. All are clearly related to each other, however, than to any
other group of turtles.

FOSSIL VERTEBRATES OF NEW CALEDONIA - 1397

Tricholimnas lafresnayanus has been related to the Lord Howe Island species Tricholimnas
sylvestris, and both species to the New Zealand species Gallirallus australis (Cracraft 1980).
Olson (1973a), however, disagrees and believes that all were independently derived.

The Rhynochetidae is now and in the past restricted to New Caledonia. Cracraft (1982) has
noted that Aptornis, a New Zealand fossil form, may be related to the New Caledonian
rhynochetids and together they may form a sister group to the South American eurypygids.

The aegothelids are restricted to the southwestern Pacific. Two genera are known:
Quipollornis (Australia), Aegotheles (New Guinea, New Caledonia and New Zealand).

The geckonids and scincids have been studied by Kluge (1967) and Hardy (1977), who have
reached the conclusion that the New Zealand reptiles are more closely related to those from New
Caledonia than to any other species in the world, including those of Australia. Preliminary
study of the New Caledonian Varanus suggests that the closest similarity lies with New
Guinean species.

The relationships discussed above are summarized in several cladograms presented in Fig. 7.
The biogeographic implications of these cladograms (Fig. 8) can be explained in the following
way. Many relationships of the New Caledonian biota can be explained by the land
connections during the Mesozoic (Fig. 9A). Such old links cannot explain the presence of
Varanus, however, nor the invertebrate exchanges between Tasmantis and New Guinea during
the Miocene (Stevens 1980). Miocene exchanges between Australia-New Guinea and New
Caledonia can best be explained by the availability of the Rennell Ridge (Fig. 9B), which
probably linked New Caledonia to New Guinea and Australia. The southern end of this
connection would have been destroyed by the tectonic events along the d'Entrecasteaux Fracture
Zone. Other Miocene exchanges within Tasmantis may have occurred along the Norfolk and
Lord Howe ridges. The Solomon and Vanuatu island biota cannot be explained by vicariance,
as these islands were never in contact with New Caledonia. In this case, the Australian-New
Caledonian tracks (Ardea novaehollandiae, Cyanorhamphus, Meiolania, Varanus) coincide with
the Rennell's Ridge track.

CONCLUSIONS

The fossil vertebrates recently discovered at several palacontological excavations in New
Caledonia establish a lengthy endemism to the fauna there and emphasize the importance of
recent extinctions in shaping the modern biota of New Caledonia. There are significant gaps in
the fossil record, where no terrestrial vertebrates have been found in the Late Triassic to Late
Pleistocene sediments on this island. The record of fossil vertebrates allied to marine, marsh
and montane environments is very poor.

The fossil vertebrate fauna is characterized by the absence of indigenous mammals (except
bats), the dominence of the birds and the presence of a highly endemic reptilian and avian fauna.
The tooth from Diahot, thought by some to be related to an Australian diprotodontid
marsupial, is instead a rhinoceros, and was most probably a human import.

Phylogenetic and palaeobiogeographic affinities of the fossil New Caledonian fauna are not
yet well understood. Only seven cladograms can be used in palaeobiogeographical studies of
Tasmantis, despite the fact that 131 taxa are endemic. Much is yet to be learned about the
origins of this fascinating biota.

1398 - BALOUET

South America Aust. N.C.
Eurypigidae NZ. N.C. ee
Geckonidae et
Scincidae
Rhynochetoidae
N.Z. L.H.
Tricholimnas
N.C.
Aust. NZ.

Aegothelidae ern 2.

Paradiseidae
other
Aust. LH. N.C Ss ae
Meiolania Varunus

Figure 7. Cladograms illustrating relationships of several New Caledonian terrestrial vertebrates.

New Guinea New Caledonia
Lord Howe

New Zealand
; Australia
South America

Figure 8. Geographic cladogram.

FOSSIL VERTEBRATES OF NEW CALEDONIA - 1399

Solomon Is.
Vanunta Island
Arc

New Caledonia

Australia

Antarctica’:

Figure 9. A Cretaceous palaeogeography of Australia (after Griffiths 1977); B, Miocene palaeogeography
in Tasmantis. (After Bitoune & Recy 1982, modified by Balouet 1984).

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BALOUET, J.C., 1984a. Les etranges fossiles de Nouvelle Caledonie. La Rech. 153: 390-392.

BALOUET, J.C., 1984b. Paleontologie des vertebres terrestres de Nouvelle Caledonie et Paleobiogeographie
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BALOUET, J. C., 1987. Extinction des vertébrés terrestres de Nouvelle Calédonie. Mém. Soc. géol. Fr. 150:
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BALOUET, J.C. & BUFFETAUT, E., 1987. Mekosuchus inexpectatus, n.g., n. sp., crocodilien nouveau de
I'Holocéne de Nouvelle Caledonie. C. r. Acad. Sci. 304(14): 853-856.

BALOUET, J.C. & OLSON, S., 1989. Fossil birds from late Quaternary deposits in New Caledonia. Smiths.
Contr. zool. 469.

BERTRAND, P., 1986. De l'utilisation de la structure de email dentaire en paleobiogeographie: enigmatique
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1053-1058.

BITOUNE, G. & RECY, J., 1982. Origine et evolution due bassin de Loyautes et de ses bordures apres la mise
en place de la serie ophiolitique de Nouvelle Caledonie. Cah. ORSTOM geol. Fr. 147.

BROTHERS, R.N. & BLACK, M.C., 1974. Temiary plate tectonics and high pressure metamorphism in New
Caledonia. Tectonophy. 17(4): 337-358.

BUFFETAUT, E., 1983. Sur la persistance tardive d'un crocodilien archaique dans le Pleistocene de I'lle de
Pins (Nouvelle Caledonie) et sa_ signification biogeographique. C. r. Acad. Sci. Paris 297: 89-92.

CRACRAFT, J., 1980. Biogeographic patterns of terrestrial vertebrates in the South West Pacific.
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1400 - BALOUET

CRACRAFT, J., 1982. Phylogenetic relationships and transantarctic biogeography of some gruiform birds.
Geobios 6: 393-402.

DELACOUR, J., 1948. Guide de Oiseaux de Nouvelle Caledonie et de ses dependances. Masson, Paris.

DUBOIS, M.J., 1976. Trouvailles a V'ile de Pins. Bull, Soc. Hist. Nouvelle Caledonie 32: 1-7.

ESTES, R., 1983. Sauria terrestria, Amphisbaenia. Handbuch der Paleoherpetologie, Teil 10A. Gustav
Fischer Verlag, Stuttgart.

FEDUCCIA, A., 1985. The morphological evidence for ratite monophyly: fact or fiction. Proc. XVIII Cong.
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FILHOL, H., 1876. Note sur la decouverte d'une dent de Rhinoceros fossile de la Nouvelle Caledonie. Annis.
Sci. nat. Zoologie 6(2): 1.

FRIMIGACCTI, D., 1980. Localisation ecogeographique et utilisation de l'espace de quelques sites Lapita de
Nouvelle Caledonie. J. Soc. oceanistes Paris 66 (36): 6-11.

GAFFNEY, E.S., 1983. Cranial morphology of the extinct homed turtle, Meiolania platyceps, from the
Pleistocene of Lord Howe Island. Bull. Am. Mus. nat. Hist. 175(4): 361-480.

GAFFNEY, W.S., BALOUET, J.C. & DE BROIN, F., 1984. New occurrences of extinct meiolaniid turtles in
New Caledonia. Novitates 2800: 1-6.

GRIFFITHS, J.R., 1977. Reconstruction of the Southwest Pacific margin of Gondwanaland. Nature 234: 203-
207.

GUERIN, C, WINSLOW, J.H., PIBOULE, M. & FAURE, M., 1981. Le pretendu Rhinoceros de Nouvelle
Caledonie est un marsupial (Zygomaturus diahotensis nov. sp.). Solution d'une enigme et consequences
paleogeographiques. Geobios 14: 201-217.

GUERIN, C. & FAURE, M., 1987, A propos du Zygomaturus, Diprotodontide de Nouvelle-Caledonie. C. r.
Acad. Sci. Paris 305(2): 815-817.

HARDY, G.S., 1977. The New Zealand Scincidae (Reptilia. Lacertilia), a taxonomic and zoogeographic study.
N.Z. J. Zool. 4: 221-325.

HOFFSTETER, R., 1968. Presence de Varanidae (Reptilia, Sauria) dans le Miocene de Catalogne;
considerations sur l'histoire de la Famille. Bull. Mus. Paris 40(3): 1051-1064.

KLUGE, A.G., 1967. Systematics, phylogeny and zoogeography of the lizard genus Diplodactylus. Aust. J.
Zool. 15: 1007-1108.

LECURU, S., 1968. Remarques sur le scapulocoracoide des Lacertiliens. Ann. Sci. nat. Zool. Biol. 10(4):
474-498.

MACGOWAN, C., 1984. Evolutionary relationships of ratites and carinates: evidence from ontogeny. Nature
307(5953): 733-735.

OBELLIANE, J.B., 1958. Le gisement des phosphates de Tiga. Rapport Int. Compagnie Phosphates
d'Oceanie, Paris.

OLSON, S., 1973a. A classification of the fossil Rallidae. Wilson Bull. 85(4): 381-416.

OLSON, S., 1973b. Evolution of the rails of the south Atlantic islands. Smithsonian Cont. Zool. 152: 1-53.

OLSON, S., 1981. Paleognathous carinate bird from the early Tertiary of North America. Science 214: 1236-
1237.

OLSON, S.L., BALOUET, J.C. & FISCHER, C.T. , 1987. The owlet-nightjar of New Caledonia, Aegotheles
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OWEN, R., 1848. On the remains of the gigantic and presumed wingless birds of New Zealand. Proc. zool.
Soc. Lond, 16: 1-10.

POPLIN, F., 1980. Sylviornis neocaledoniae n.g.n. sp.; Ratite eteint de la Nouvelle Caledonie. C. r. Acad.
Sci. Paris 290(D): 691-694,

POPLIN, F., MOURER-CHAUVIRE, C & EVIN, J., 1984. Position systematique et datation de Sylviornis
neocaledoniae, megapode geant de la Nouvelle Caledonie. C. r. Acad. Sci. Paris 297(II): 99-102.
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Sci. Paris 281(D): 489-492.

RICH, P. V., BALOUET, C., FLANNERY, T., FRIMIGACCI, D., LAUNAY, J & RICH, T., 1981. Kukwiede's
revenge. A view into New Caledonia's distant past. Hemisphere 26(3): 166-171.

RICH, P.V., MCEVEY, A.R & BAIRD, R.F., 1985. The scrubbirds (Africhornis) and lyrebirds (Menura) of
Australia: osteological comparison and comments on their relationships. Rec. Aust. Mus. 37: 156-
191.

RICH, T.H., FORTELIUS, M, RICH, P.V. & HOODER, D.A., 1987. The supposed Zygomaturus from New
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778.

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197,

FOSSIL VERTEBRATES OF NEW CALEDONIA - 1401

TILLIER, S. & CLARKE, B.C., 1983. Lutte biologique et destruction du patrimoine genetique: le cas des
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VAN TETS, G.F., 1974. A revision of the fossil Megapodiidae (Aves), including a description of a new
species of Progura DeVis. Trans. R. Soc. S. Aust. 98: 213-224.

PLATES

Plate 1, Varanus cf indicus, A, parietal; B, right maxillary; C, posterior part of skull, dorsal view; D, left
articular, angular and splenial, dorsal view; E, left dentary, external view; F, scapulocoracoid; G, humerus; H,
dorsal vertebra, anterior view; I, sacral vertebra, posterior view; J, axis vertebra, lateral view; K, pelvis; L,
sinkhole at Kanumera which has been breached by modem wave action; M, outcrop of raised reefal limestone
along Kanumera Bay in which sink holes have developed (L, M courtesy of P. V. Rich).

Plate 2. Mekosuchus inexpectatus. A, left premaxillary, ventral view; B, left premaxillary, dorsal view; C,
anterior tooth, lateral view; D, posterior teeth (tribodont), lateral view; E, atlas-axis vertebrae, lateral view; F,
dorsal vertebra, anterior view; G, cervical vertebra, anterior view; H, caudal vertebra, lateral view; I, right
jugal-quadratojugal, extemal view; J, right nasal, dorsal view; K, right parietal-temporal, dorsal view; L,
pterygoid, dorsal view; M, pterygoid, ventral view; N, maxillary, dorsal view, part of the ventral border of
the orbit; O, right lacrymal, dorsal view; P, frontal, dorsal view; Q, left palatine and palatine process of the
maxillary, ventral view; R, left palatine and palatine process of the maxillary, dorsal view; S, frontal,
ventral view; T, left ilia, lateral view; U, right humerus, anterior view; V, right humerus, posterior view; W,
left tibia, anterior view; X, left tibia, posterior view. (From Balouet 1984b).

Plate 3. Posterior cervical vertebrae of meiolaniid turtles. A, Pindai Cave, New Caledonia, probably a

seventh cervical; B, Lord Howe Island, Australia, a seventh cervical; C, Tiga Island, New Caledonia,

probably a seventh cervical. Views from left to right are: posterior, anterior, left lateral, ventral (anterior to

eed . Width at the base of the transverse process is for A, 19 mm; B, 32 mm; C, 25 mm (from Gaffney et al.
4).

Plate 4. Sylviornis neocaledoniae. A, B, right claviculocoracoid; C, D, left humerus; E, F, left ulna; G,
right carpometacarpus; H, I, left scapula; J, K, left radius.

Plate 5. Sylviornis neocaledoniae. A, left tibiotarsus, anterior view; B, left tibiotarsus, proximal view; C,
left tibiotarsus, posterior view; D, right proximal femur fragment, posterior view; E, right posterior femur
fragment, posterior view; F, right femoral shaft, anterior view; G, right femoral shaft, posterior view; H,
proximal femur fragment, proximal view; I, distal femur fragment, distal view; J, distal femur fragment,
anterior view; K, distal femur fragment, external view.

Plate 6. Sylviornis neocaledoniae. A, fibula, internal view; B, fibula, external view; C, left
tarsometatarsus, anterior view; D, left tarsometatarsus, posterior view; E, left tarsometatarsus, proximal view;
F, left tarsometatarsus, distal view.

Plate 7. Tricholimnas sp. from Kanumera Bay, Isle of Pines (after Rich ef al. 1981.) compared with the
extant Lord Howe Island Woodhen (Gallirallus sylvestris). Gallirallus: A, coracoid; D, carpometacarpus; F,
skull; H, mandible; J, humerus. Tricholimnas: B, C, coracoids; E, carpometacarpus; G, fragment of
maxillary; I, mandibular fragment; K, L, humeri. M, coracoid fragment of a large pigeon (Columbidae), N,
ulna fragment of unknown avian form (about x1).

Plate 8. Tricholimnas sp. from Kanumera Bay, hind limb elements, compared with the extant Lord Howe
Island Woodhen (Gallirallus sylvestris). Gallirallus : A, tibiotarsus; H, tarsometatarsus; J, femur.
Tricholimnas: B, C, D, F, tibiotarsi; E, I, pelvis fragments; G, tarsometatarsus; k, distal femur fragment;
about x 1. (After Rich et al. 1981).

Plate 9. Tricholimnas lafresnayanus, an X-ray photograph of a specimen in the Australian Museum,
Sydney, showing the reduced wings on this recently extinct species from New Caledonia; about x 0.75.

(Courtesy of W. Boles from Rich ef al. 1981).

1402 - BALOUET

Plate 10. A-C, Holotype of "Zygomaturus diahotensis" Guerin et al. 1981. Regarded as a right p3 by Guerin
et al. (1981) of a diprotodontid marsupial, it is considered by Rich et al. (1987) to be a right dM! of a
thinoceros: A, buccal view; B, occlusal view; C, lingual view; D, Eurhinoceros aff. sondaicus, left dM! , Late
Miocene of Pakistan, occlusal view. E-L, Zygomaturus trilobus from Mammoth Cave, Western Australia,
Quatemary: E, G, right P3, occlusal view, stereo pair; F, H, I-L, left p3. F, H, occlusal view, stereo pair, I, K,
buccal view, stereo pair; J, L, lingual view, stereo pair. M, N, diagrams showing dental terminology
employed with the Diahot tooth (M) and Zygomaturus (N). (From Rich ef al, 1987).

Plate 11, Light micrographs of clear resin casts of wom enamel surfaces. A, the Diahot tooth, right dM!

(detail of buccal edge of enamel at metacone); B, Zygomaturus trilobus, right P3 from Spring Creek, Victoria
(Quaternary) (detail of protoloph). (From Rich et al. 1987).

PLATE 1

FOSSIL VERTEBRATES OF

NEW CALEDONIA - 1403

PLATE 3

1404 - BALOUET

PLATE 4

FOSSIL VERTEBRATES OF NEW CALEDONIA - 1405

PLATE 5

aS ies wales,

mtn Sa

ee
erin

PLATE 6

1406 - BALOUET

PLATE 7

PLATE 8 FOSSIL VERTEBRATES OF NEW CALEDONIA - 1407

PLATE 9

1408 - BALOUET

50mm (E-L)

M Crochet

Metacone

Metaloph

Hypocone

Paracone

Parastyle

Medisinus

Protocone

Metacone

Metaloph

Hypocone

PLATE

Paracone
Parastyle

—— Protoloph

~ 2

Protocone

10

PLATE 11 FOSSIL VERTEBRATES OF NEW CALEDONIA - 1409

+ Enamel

; Dentine

Enamel —

Dentine — ‘

F

Included in this index are all those
genera of vertebrates that have a pre-
Holocene record in Australia, New Guinea,
New Zealand and the southwest Pacific. If
a Holocene history is known for those taxa
with a longer history, that is included, but
forms with only a Holocene record are not
incorporated. Families and genera yet
unnamed are mentioned only where they
significanuy extend the range of the group.

This index was compiled by Anne
Kemp, Noel Kemp, John Long, Alex
Ritchie and Sue Turmer (fish), Michael
Tyler and Anne Warren (amphibians),
Ralph Molnar (reptiles), Bob Baird, Pat
Vickers-Rich and Corrie Williams (birds),
and Timothy Flannery, Robert Bearlin,
and Thomas Rich (mammals) with the
assistance of Jennifer Monaghan and Pat
Vickers-Rich.

SUPERCLASS: AGNATHA

CLASS: PTERASPIDOMORPHI

SUBCLASS: HETEROSTRACI

ORDER: ARANDASPIFORMES
Arandaspididae. Arandaspis, M. Ord.
N.T.; Unnamed genus, M. Ord. N.T.
Family Unnamed. Porophoraspis, M.
Ord. N.T.

SUBCLASS: THELODONTI
ORDER: THELODONTIDA
Turiniidae. Turinia, E.-M. Dev.

N.S.W., N.T., Qd., S.A., Vic., W.A.;
Australiolepis, L. Dev. W.A.,
Nikoliviidae. ?Gampsolepis, E.-M.
Dev. Qd.; Nikolivia, E. Dev. N.S.W., L.
Dev. W.A.

SUPERCLASS: GNATHOSTOMATA
CLASS: PLACODERMI
SUPERORDER:
PTYCTODONTOMORPHA
ORDER: PTYCTODONTIDA
Ptyctodontidae. Crenurella,
Campbellodus, L, Dev. W.A.; Genus
indet., E. Dev. N.S.W.

SUPERORDER:
PETALICHTHYOMORPHA
ORDER: ACANTHOTHORACI.
Weejasperaspidae. Weeyasperaspis, E.
Dev. N.S.W.; Murrindalaspis, E. Dev.
N.S.W., Vic. Family uncertain.
Brindabellaspis, E. Dev, N.S.W.:

SYSTEMATIC INDEX - 1411

SYSTEMATIC, GEOGRAPHIC

AND GEOLOGIC INDEX

"Ohioaspis", body scales, E. Dev. N.S.W.,
Qd., Vic.

ORDER: PETALICHTHYIDA
Macropetalichthyidae.
Notopetalichthys, Shearsbyaspts,
Lunaspts, E. Dev. N.S.W.; Wijdeaspis,
E. Dev. N.S,.W., Vic.

SUPERORDER:
DOLICHOTHORACOMORPHA
ORDER: ARTHRODIRA
SUBORDER: ACTINOLEPIDOIDEI
INFRAORDER: ACTINOLEPIDI
Actinolepididae? Genus inde, E.-M.
Dev. N.S.W., Qd.

INFRAORDER: WUTTAGOONASPIDI
Wuttagoonaspididae. Wuttagoonaspis,
E.-M. Dev. N.S.W., Qd.

INFRAORDER: PHYLLOLEPIDI
Phyllolepididae. Austrophyllolepis,
L. Dev. N.T., Vic.; Placolepis, L. Dev.
N.S.W., N.T.; ?Phyllolepis, L. Dev.
N.S.W.

SUBORDER: PHLYCTAENIOIDEI
INFRAORDER: PHLYCTAENIL

Phlyctaeniidae. Denisonosteus, M.
Dev. N.S.W. Groenlandaspididae.
New Genus. M. Dev. N.S.W., Qd.;
Groenlandaspis, L.Dev. N.S.W., Vic.;
Holonematidae. Holonema , L. Dev.
W.A.Williamsaspididae.
Williamsaspis, E. Dev. N.S.W.

INFRAORDER: BRACHYTHORACI
Goodradigbeeonidae. Goodradigbeeon,
E. Dev. N.S.W. Burrinjucosteidae.
Burrinjucosteus, E. Dev, N.S.W.
Buchanosteidae. Buchanosteus, E. Dev.
N.S.W., Vic. Taemasosteidae.
Taemasosteus, E. Dev. N.S.W., Vic.;
Family uncertain. Areniptscts,
Errolosteus, E. Dev. N.S.W., Vic.
Homosteidae. cf. Homosteus,
Atlantidosteus, M. Dev. Qd.
cf. Plourdosteidae. Harrytoombsia,
Kimberleyichthys, Torosteus, L. Dev.
W.A. Incisoscutidae. /nctsoscutum,
L. Dev. W.A. Camuropiscidae.
Latocamurus, Camuropiscis, Rolfosteus,
Tubonasus, Fallacosteus, L. Dev. W. A.
Mylostomatidae. Kendrickichthys,

Bruntonichthys, Bullerichthys, L.Dev.
W.A. Family uncertain. Simosteus,
Pinguosteus, L. Dev. W.A.

ORDER: ANTIARCHI
SUBORDER: SINOLEPIDOIDEI
Sinolepididae. New genus, L. Dev.

N.S.W.

SUBORDER: BOTHRIOLEPIDOIDEI
Bothriolepididae. Monarolepis, M.
Dev. N.S.W.; Briagalepis, M.-L. Dev.
Vic.; Bothriolepis, M.-L. Dev, N.S.W.,
N.T., Qd., Vic., W.A.; Nawagiaspis, M.
Dev. Qd.

SUBORDER: ASTEROLEPIDOIDEI
Pterichthyodidae. Sherbonaspis, M.
Dev. N.S.W.; Wurungulepis, M. Dev. Qd.
Asterolepididae. Pambulaspis,
Remigolepis, L. Dev. N.S.W.

CLASS: CHONDRICHTHYES
SUBCLASS: ELASMOBRANCHIL
ORDER: XENACANTHIFORMES

Phoebodontidae. Phoebodus, M. Dev.-
E. Carb. N.S.W., Qd.; Thrinacodus
(Harpagodens), L. Dev.-E. Carb. Qd.,
W.A. Xenacanthidae. Antarctilamna,
M. Dev. N.S.W.; ?Xenacanthus, E. Carb.
Qd.; Pleuracanthus, E. Trias. N.S.W.

ORDER: SYMNORIIFORMES
Symmoriidae. Symmorium, L. Dev.-
E. Carb. N.S.W. Stethacanthidae.
Stethacanthus, E. Carb. N.S.W., W.A.

ORDER: EUGENEODONTIFORMES

SUPERFAMILY: EDESTOIDEA
Agassizodontidae. Helicoprion, M.
Perm. W.A.

ORDER: ORODONTIFORMES
Orodontidae. Orodus, E. Carb. W.A.

ORDER: PETALODONTIFORMES
Family uncertain. Ageleodus, E. Carb.

Qd.

ORDER: INCERTAE SEDIS.
Family uncertain. Poecilodus, Perm.
W.A.; Holmesella, L. Carb. Qd.;
Tomodus, Carb.-Perm. Qd.:

1412 - SYSTEMATIC INDEX

Deltoptychius, Carb. Qd; Crassidonta,
Perm. W.A.; Helodus, E. Carb. Qd.,
W.A.; Ohiclepis , E. Dev. N.S.W.
Psammodontidae. Psammodus, E.
Carb. Qd.

COHORT: EUSELACHII
SUPERFAMILY:
PROTACRODONTOIDEA
Protacodontidae. Protacrodus, L.
Dev.-E. Carb. Qd.

SUPERFAMILY: CTENACANTHOIDEA
Ctenacanthidae. Cienacanthus, E.
Carb. Qd., W.A.

SUPERFAMILY: HYBODONTOIDEI
Ptychodontidae. Ptychodus, L. Cret.
W.A.

SUBCOHORT: NEOSELACHII
SUPERORDER: SQUALIMORPHII
ORDER: HEXANCHIFORMES

Family uncertain. Mcmurdodus, M.
Dev. Qd. Hexanchidae. Hexanchus, L.
Cret. N.Z., L. Palaeo. W.A., L. Palaeo.-
Eoc. Vic., M.-L. Eoc., L. Olig. 5.A.,
Rec.; Notorynchus, L. Cret.-L. Eo, N.Z.,
L. Cret. W.A., E. Mio.-E. Plio. Vic.,
Rec. Heptranchidae. Heptranchias, L.
Eoc., S.A., Rec.

ORDER: SQUALIFORMES
Squalidae. Centrophoroides,
Protosqualus, L. Cret. W.A., L. Cret.-E.
Mio. N.Z.; Scymnorhinus, M. Eo.-E.
Mio. N.Z.; Squalus, L. Cret.-Palaeo. N.Z.
Echinorhinidae. Pseudoechinorhinus,
Palaco. N.Z.

ORDER: PRISTIOPHORIFORMES
Pristiophoridae. /kamauius, L. Eo.-L.
Plio. N.Z.; Pristiophorus, E. Cret. Qd., E.
Eo.-E. Mio. N.Z., E. Mio.-E. Plio. Vic.,
E, Plio. S.A., Rec.

SUPERORDER: SQUATINOMORPHII
ORDER: SQUATINIFORMES
Squatinidae. Squatina, L, Cret. W.A.,
Palaeo. N.Z., Rec.

SUPERORDER: GALEOMORPHII

ORDER: HETERODONTIFORMES
Heterodontidae. Heterodontus, M.-L.
Eo. N-Z., L. Olig.-E. Plio. Vic., E. Mio.
Tas., E. Plio. S.A., L. Plio. Fl. Is., Rec.;
Synechodus, L. Cret.-Palaco. N.Z.

ORDER: ORECTOLOBIFORMES
Orectolobidae. Orectolobus, E.-L.
Mio. Vic., E. Plio. S.A., Rec.

ORDER: LAMNIFORMES
Odontaspididae. Carcharias, L. Palaeo.-
E. Eoc., L. Olig.-E. Plio. Vic., L. Palaeo.
W.A., M. Eoc., E.-M. Mio., Plio. S.A.,
E. Mio. Tas., ?7L. Mio. & “Tertiary”
N.S.W., Rec.; Eugomphodus, L. Cret.-E.
Plio, N.Z.; Hispidaspis, L. Cret. W.A.;
2Odontaspis, E. Mio. Tas., Rec.
Mitsukurinidae. Anomotodon, E.
Cret. Qd.; Mitsukurina, M.-L. Eoc. S.A.,
Rec.; Scapanorhynchus, E. Cret. Qd., L.
Cre W.A. Lamnidae, Carcharodoa, M.
Eo.-L. Pleist. N.Z., L. Olig.-E. Mio., E.
Plio. S.A., L. Olig.-E. Plio.. Pleist. Vic..

E. Mio. Tas., L. Plio. Fl. Is., E.-M. Mio,
W.A., Rec.; Carcharhoides, L. Olig.-E.
Mio. Vic.; /surus, Cret.-M. Mio. N.Z.,

L. Olig.-E. Plio, Vic., L. Olig.-M. Mio.,
E. Plio, S.A., E. Mio. Tas., Rec.; Lamna,
(including "Lamna" ), E. Cret. Qd., L.
Cret. W.A., Rec. Cretoxyrhinidae.
Cretolamna, Cretoxyrhina, E. Cret. Qd.,
L. Cret. W.A.; Paraisurus, E. Cret. Qd.;
?Protolamna., L. Cret. W.A. Otodon-
tidae. Otodus, E.-M. Palaeo., L. Eoc.-E.
Olig. Vic., M. Eoc. S.A. Amacoracidae.
Pseudocorax, Microcorax, E. Cret. Qd.

ORDER: CARCHARHINIFORMES
Scyliorainidae. Megascyliorhinus, L.
Olig.-E. Mio. N.Z., L. Mio.-E.Plio. Vic.;
Galeorhinus, M. Mio. Vic., E. Plio. S.A.,
Rec. Hemigaleidae. Hemiprists, E.
Mio. S.A., Rec. Carcharhinidae.
Carcharhinus, L. Olig.-E. Plio. Vic., E.
Mio. Tas., L. Plio. Fl. Is., E,-M. Mio. E.
Plio. S.A., ?L. Mio, N.S.W., Plio.-Pleist,
W.A., Rec.; Galeocerdo, L. Olig.-E.Plio.
N.Z., Vic., E.-M. Mio., E. Plio. S.A.,
Rec. Sphyrnidae. Scapanorhynchyus,
L. Cret. N.Z.; Sphyrna, E. Mio. Tas., E.-
M. Mio. Vic., E. Plio. S.A., Rec.

SUPERORDER: BATOMORPHII
ORDER: MYLIOBATIFORMES
Dasyatidae. Dasyatis, L. Olig.-E. Mio.

N.Z., ?L. Mio. N.S.W., E. Plio. S.A.,
Rec. Myliobatidae. Myliobatis, M.
Eo.-E. Mio. N.Z., M.-L. Eoc., Plio. S.A.,
L, Olig.-E. Plio. Pleist. Vic., E. Mio.
Tas., ?7L. Mio, N.S.W., E. Plio. S.A.,
Rec. Sclerorhynchidae. Onchopristis,
L. Cret. N.Z.

SUPERORDER AND ORDER:
INCERTAE SEDIS.
Palaeospinacidae. Synechodus, E. Cret.

Qd., L. Cret. W.A.

SUBCLASS: HOLOCEPHALI
ORDER: CHIMAERIDA

Callorhinchidae. Ptyktoptychion, E.
Cret. Qd. Chimaeridae. Chimaera,
Tert. N.Z.; Edaphodon, E. Cret. S.A., L.
Cret. W.A., L. Mio.-E. Plio. Vic.;
Ischyodus, L. Cret. N.Z., E. Mio. Tas., L.
Mio.-E. Plio. Vic.

CLASS: TELEOSTOMI
SUBCLASS: ACANTHODII
ORDER: ISCHNACANTHIDA
Ischnacanthidae. Rockycampacanthus,

E. Dev. Vic.; Taemasacanthus, E. Dev.
N.S.W.; Cheiracanthoides, E.-M. Dev.
N.S.W., Qd.; Gomphonchus, L. Sil. Qd.,
E.-M. Dev. Qd., N.S.W., S.A., Vic.
Poracanthodes, L, Sil. Qd.

ORDER: CLIMATIIDA
SUBORDER: CLIMATOIDEI
Families uncertain. Nostolepis, L.
Sil. Qd., E. Dev. N.S.W., Qd., Vic., M.
Dev. N.S.W.

SUBORDER: DIPLACANTHOIDEI
Culmacanthidae. Culmacanthus, L.
Dev, N.S.W., Vic. Family uncertain.
Striacanthus, L. Dev. Vic.
Gyracanthidae. Gyracanthides, E. Carb.
Od., Vic.

ORDER: ACANTHODIDA
Acanthodidae. Howittacanthus, L, Dev.
Vic.; ?Acanthodes, M. Dev. N.T., Qd, E.
Carb. Vic,

SUBCLASS: OSTEICHTHYES
INFRACLASS: ACTINOPTERYGII
ORDER: PALAEONISCIFORMES

Family uncertain. Ligulalepis, E.-M.
Dev. N.S.W., Qd.; Howqualepis, L. Dev.
Vic. Stegotrachelidae. Mimia,
Moythomasia, L. Dev. W.A.
Gonatodidae, Novogonatodus, E. Carb.
Vic. Family uncertain.
Mansfieldiscus, E. Carb. Qd., Vic.
Cryphiolepidae. Cryphiolepis, E.
Carb. Qd., Perm. W.A.

Elonichthyidae. ?Elonichthys, Perm.-
E. Trias. N.S.W. Bobasatraniidae.
Ebenaqua, L. Perm. Qd. Urosthenidae.
Urosthenes, Perm, N.S.W.
Tegeolepidae. Apateolepis, E. Trias.
N.S.W. Acrolepidae. Acrolepis, E.
Trias. N.S.W., Tas., ?Perm. W.A,;
Leptogenichthys, M. Trias. N.S.W.
Palaeoniscidae. Agecephalichthys, M.
Trias. N.S.W. Coccolepidae.
Coccolepis, L. Jur. N.S.W., E. Cret. Vic.;
Psilichthys, E. Cret. Vic.
Platysomidae. Platysomus, L. Trias.
N.S.W. Family uncertain,
Atherstonia, Belichthys, Trias. N.S.W.;
Elpisipholis, L. Trias. N.S.W.;
Leighiscus, Trias. S.A.; Megapteriscus,
Mesembroniscus, Myriolepis, Trias.
N.S.W.

SUBORDER: PHOLIDOPLEUROIDEI
Pholidopleuridae. Macroaethes, M.
Trias. N.S.W.

SUBORDER: REDFIELDOIDEI
Redfieldiidae. Beaconia, Brookvalia,
Dictyopleurichthys, Dictyopyge,
Geitonichthys, Molybdichthys,
Phlyctaentchthys, Schizurichthys, Trias.
N.S.W.; Genus indet., L. Perm. Qd.

SUBORDER: PERLOIDEI
Perleididae. Chrotichthys, Manlietta,
Pristisomus, Procheirichthys, Tripelta,
Zeuchthiscus, Trias. N.S.W.
Cleithrolepidae. Cleithrolepis, Trias.
N.S.W., Tas. Pholidophoridae.
?Thoracopterus, Trias. N.S.W.

ORDER: ?ACIPENSERIFORMES
Saurichthyidae. Saurichthys, E.-M.
Trias. N.S.W., Qd., Tas., W.A.

ORDER: ASPIDORHYNCHIFORMES
Aspidorhynchidae. Aspidorhynchus,
Belonostomus, E. Cret. Qd.

ORDER: SEMIONOTIFORMES
SUBORDER: SEMIONOTOIDEI
Semionotidae. Aetholepis, Aphnelepis,

Corungenys, Enigmatichthys, Trias.
N.S.W. Parasemionotidae.
Promecosomina, M.-L, Trias. N.S.W.
Macrosemionotidae. Uarbryichthys, L.
Jur. N.S.W.

ORDER: PHOLIDOPHORIFORMES
Archaeomaenidae. Archaeomaene,
Madariscus, L. Jur. N.S.W.; Wadeichthys,
E. Cret. Vic,

COHORT: TELEOSTEIL
SUPERORDER: LEPTOLEPOMORPHA
ORDER: LEPTOLEPIFORMES
Leptolepidae. Leprolepis, L. Jur.

N.S.W., &. Cret. Vic.

SUPERORDER: ELOPOMORPHA
ORDER: ELOPIFORMES
SUBORDER: ELOPOIDEI

Elopidae. Elops, E. Eo.-?Mio. N.Z.;
Flindersichthys, E, Cret. Qd.
Megalopidae. Megalops, L. Olig.-M.
Mio, Vic., Rec.

SUPERORDER: ALBULOIDEI
Albulidae. Prerothrissus, L. Olig. Vic.

ORDER: ANGUILLIFORMES
SUBORDER: ANGUILLOIDEI
Congridae. Genus indet. L. Eo.-M, Mio.

N.Z.; Astroconger, L, Olig. Vic., Rec.;
Conger, Mio. N.Z.; Congridarum, E, Mio.
N.Z.; Gnathophis, E. Eo.-Plio. N.Z.;
Maxwelliella, E. Eo. N.Z.; Scalanago, E.
Eo. N.Z.; Uroconger, L. Olig., Plio.
Vic., Rec. Heterenchelyidae.
Gnathophis, E. Eo.-Plio. N.Z.;
Heterenchelyidarum, E. Mio. N.Z.;
Heterenchelys, L. Olig.-M. Mio, Vic.,
Rec. Muraenesocidae, Muraenesox, M.
Mio. Vic., Rec, Ophichthidae.
?Mystriophis, M. Mio. N.Z.

SUPERORDER: CLUPEOMORPHA
ORDER: CLUPEIFORMES
Clupeidae. Genus indet., E. Mio. N.Z.;

Anchoa, E. Mio. N.Z,; Diplomystus,
Cret. N.Z.; Scombroclupea, Eo, N.Z.
Ichthyodectidae. Cooyoo,
1Cladocyclus, E. Cret. Qd.; Xiphacunus,
L. Cre. N.Z. Pachyrhizodontidae.
Pachyrhizodus, E. Cret. Qd., L. Cret.
N.Z.; ?Thrissopater, Cret. N.Z.

SUPERORDER: CLUPEODEI
Koonwarriidae. Koonwarria, E. Cret.
Vic.

SUPERORDER:
OSTEOGLOSSOMORPHA
ORDER: OSTEOGLOSSIFORMES
SUBORDER: OSTEOGLOSSOIDEI
Osteoglossidae. Phaeroides, Eoc.-Olig.

Qd.: Scleropages, Olig.-?Mio. Qd., Rec,

SUPERORDER:
PROTACANTHOPTER YGIL
ORDER: GONORHYNCHIFORMES
SUBORDER: GONORHYNCHOIDEI
Gonorhynchidae. Notogoneus, Eoc.-
Olig. Qd.

SUPERORDER: OSTARIOPHYSI
ORDER: SILURIFORMES
Plotosidae. Jandanus, ’Pleist. Qd., Rec.
Ariidae. Genus indet., L. Olig./M.

Mio. S.A.; Tachysurus, Plio. N.Z.

SUPERORDER: ”

ORDER; SALMONIFORMES
Argentinidae. Argentina, E. Mio. N.Z.;
Galaxiidae, Mio. N.Z.;
Photichthyidae. Polymetme, E. Mio.
N.Z. Gonostomatidae. Gonostoma,
E. Mio. N.Z, Aulopiidae. Aulopus, E.
Eo. N.Z. Myctophidae. Benthosema,

SYSTEMATIC INDEX -

Mio. N.Z.; Bolinichthys, M. Mao. N.Z.;
Diaphus, E. Olig.-L. Mio. N.Z.;
Diogenichthys, E. Mio. N.Z., Hygophum,
Mio, N.Z.; Lampanyctodes, E. Mio.-Plio,
N.Z.; Lampanyctus, M.-L. Mio, N.2.;
Lampichthys, L, Mio. N.2.;
Myctophidarum, L. Mio. N.Z.;
Myctophum, M. Mio. N.Z.;
Notoscopelus, E, Mio,-Plio. N.Z.;
Symbolophorus, E, Mio.-Plio, N.Z.

Sternoptychidae. Maurolicus, E. Mio.

N.Z.; Polyipnus, E, Mio. N.Z.
Chlorophthalmidae.
Chlorophthalmus, L.. Eo.-E. Mio. N.Z.
Scopelarchidae. Scopelarchus, L.. Eo.-
E. Mio. N.Z.

SUPERORDER: ?
ORDER; LOPHIIFORMES
Lophiidae. ?Lophius, E. Mio, N.Z.
Ogcocephalidae. ?Dibranchus, L. Eo.
N.Z.

SUPERORDER:

PARACANTHOPTERYGI

ORDER: GADIFORMES
Moridae. Actuartolum, L. Olig.-Mio.
N.Z.; Ausirophycis, E. Mio. N.Z.;
Lemonema, M. Mio. N.Z.; Lotella, E.
Mio. N.Z.; PAysiculus, E. Mio, N-Z.;
Raniceps, Mio. N.Z., Tripterophycts, L.
Eo, N.Z. Melanonidae. Karrerichthys,
Mio. N.Z. Bythididae. Genus indct.,
M. Mio. N. Z.; Oligopus, E. Mio. N.Z.;
Bythitidarum, E. Mio. N.Z.; Saccogaster,
Plio. N. Z. Euclichthyidae.
Euclichthys, L. Eo.-E. Mio, NZ.

SUBORDER: GADOIDEI
Gadidae. Bregmaceros, Eo. N.Z., M.
Mio. Vic., Rec.; Gadiculus, E. Mio. N.Z.;
Gadus, Mio. N.Z.; Macruronus, L. Olig.-
Plio. N.Z.; Merluccius, Mio. N.Z.;
Micromesistius, Plio, N.Z.
Merlucciidae. Merluccius, L. Olig.-M.
Mio. Vic., Rec.

SUBORDER: OPHIDOIDEI
Ophidiidae. Genus indet., M. Mio.
N.Z.; Ampheristus, E. Eo. N.Z..
Genypterus, Plio, N.Z.; Monomitopus, L.
Eo.-L. Mio, N.Z.; Neobythites, M. Mio.
N.Z.; Nolfophidion, L. Bo, N.Z.
Ophidion, L. Olig.-M, Mio. Vic., Mio.-
Plio. N.Z., Ree.; Ophidiidarum, Nio.
N.Z.: Sirembinorum, E. Mio. NZ,
Carapidae, Carapus, Olig. - M. Mio.
N.Z., M. Mio. Vic., Rec.

SUBORDER: MACROUROIDEI
Macrouridae. Bathygadus, E. Mio.
N.Z.: Coelorinchus, L. Olig.-M. Mio.
Vic., L. Olig.-Plio. N.Z., Rec.;
Lepidorhynchus, E. Mio,-Plio. N.Z.;
Macrouridarum, E, Mio. N.Z.,; Macrourus,
Mio. N.Z.; Macrurulus, Eo. N.Z.
Maorigadus, E. Eo. N.Z.; Nezumia, L.
Mio. N.Z.; Trachyrincus, Mio. N.Z.;
Ventrifossa, Mio, N.Z.

SUPERORDER: ACANTHOPTER YGII
ORDER: BERYCIFORMES
SUBORDER: BERYCOIDEI

Berycidae. Centroberyx, E. Mio. N.Z.
Caproidae. Antigonia, M. Eo, N.Z., M.
Mio, Vic.. Rec. Trachichthvidae.

1413

Egregioberyx, L. Eo. N.Z.; Hoplostethus,
E. Mio.-Plio. N.Z.; Paratrachichthys, E.-
M. Mio. N.Z.; Trachichthodes, Test.
Aust., L. Eo. N-Z.; Trachichthodes, ?Olig.
Aust. Holocentridae. Genus indet., L.
Eo. N.Z.; Adioryx, E. Eo, N.Z.
Melamphaeidae. Melamphaes, Mio.
N.Z. Monocentridae. Cleidopus, L.
Olig.-M. Mio. Vic., Rec.; Monocentris,
Plio. S.A., Rec.

ORDER: SCORPAENIFORMES

SUBORDER; SCORPAENOIDEI
Cottidae, Genus indet., L. Eo. E.Mio.
N.Z.; Cottus, Olig/Mio. N.Z.;
Cottoideorum, E. Mio. N.Z,
Cyclopteridae. Cyclopterus, Mio. N.Z.
Hoplichthyidae. Hoplichthys, E. Mio.
NZ.; Praehoplichthys, M.-L. Eo. N.Z.
Scorpaenidae. Sebastodes, M. Mio.
Vic. Trichodontidae. Trichodon, E.
Mio. N.Z.

SUBORDER: PLATYCEPHALOIDEI
Platycephalidae. Platycephalis, M.
Eo.-E. Mio, N.Z., E. Mio, Tas., M. Mio.
Vic., Rec.

ORDER: PERCIFORMES

SUBORDER: PERCOIDEI
Centropomidae. Muccullochella, Mio.
N.S.W., Pleist. Qd.; Percaletes, Eoc.-
Mio. Qd., Mio. N.Z., Rec.
Lutjanidae. Luijanus, "Mio. Qd., Rec.
Theraponidae. Genus indet., ?Olig. Qd.
Sillaginidae. Sillago, L. Olig.-Plio.
Vic., E. Mio. N.Z., Rec, Lactariidae.
Lactarius, E. Mio. Vic. Ambassidae.
Dapalis, E.-M. Eo. N.Z. Sparidae.
Dentex, E, Eo.-Mio. N.Z.; Pagellus, Mio.
NZ.; Pagrosomus, Pleist. Vic.; Sargus,
E. Mio. Vic., Mio, N.Z., Rec.;
°Chrysophrys, E. Mio, S.A.
Oplegnathidae. Oplegnathus, E. Mio.-
E. Plio. Vic.; ?Percidarum, M. Mio. Vic.,
L. Mio. N.Z. Percichthyidae. Genus
indet., L. Olig./M. Mio, S.A.
Monacanthidae. E. Plio. S.A.
Serranidae. Genus indet., M. Eo.-E.
Mio. N.Z.; Serranus, Mio. N.2.

SUBORDER: MUGILOIDEI
Sphyraenidae. Sphyraena, Pleast. Vic.,
Ree,

SUBORDER: LABROIDEI
Lubridae. Labridarum, E. Mio. N.Z.
Labrodon, E. Mio.-E. Plio. Vic., Mio,
N.Z.; Nummopalatus, E. Mio-E. Plio,
Vic., Rec.

SUBORDER: SCOMBROIDEL
Gampeilidae. Genus indet., L. Mio.
Vic. Trichiuridae. Genus indet., L.
Mio, Vic.

SUBORDER: ”
Nototheniidae. Notothenia, M.-L.
Mio. N.Z. Trachinidae. Trachinus,
Mio. N.Z. Gempylidae. Eothyrsites,
Olig. N.Z. Epigonidae. Epigonus, E.-
M. Mio. N.Z. Apogonidae.
Apogonidarum, 1. Olig.-M. Mio, N.Z.
Acropomatidae. Acropoma, Genus
indet., L. Eo. N.Z., L. Olig.-M. Mio.
NZ. Lactariidae. Lactartus, L. Olig.-
Plio. N.Z.; Neoscombrops, EB. Bo NZ.

1414 - SYSTEMATIC INDEX

Paralactarius, E. Eo.-E, Mio, N.Z.
Cierreidae. ?7Gerreidarum, E. Mio. N.Z.
Pomadasyidae. Genus indct., E. Eo.-E.
Mio. N.Z.; Pomadasyidarum, E. Mio.
N.Z. Cepolidae. Cepola, E, Mio, N.Z.
Scombridae, Genus indct., E. Mio.
N.Z. Gobiidae. Gobiidarum, E. Mio.
N.Z. Eleotrididae. Gobiomorphus, L,
Pleist, N.Z.; Paradiplospinus, E. Mio,
N.Z.

ORDER: PLEURONECTIFORMES

SUBORDER: PLEURONECTOIDEI
Bothidae. Genus indet., E. Mio, N.Z.;
Arnoglossus, L. Olig.-E. Mio. N.Z.
Pleuonectidae. Pleuronectes, M. Mio.
Vic., Rec. Soleidue. Achirus, M. Mio.
N.Z.; Solea, Mio, N.Z. Eucitharidae.
Citharus, Mio. N.Z.; Pleuronectidarum,
Mio, N.Z.

ORDER; TETRADONTIFOR MES
SUBORDER: TETRADONTOIDEL
Family Uncertain. Sparidarum, Mio.
N.Z. Diodontidae. Diodun, E.-L. Mio.
Vic., E. Plio. S.A., Rec.; Aracana, E.
Mio. Tas., Rec. Trigonadontidae.

Trigonodon, Mia. N.Z,

SUPERORDER AND ORDER ?
Mugiloididae, Genus indct., M. Eo.
N.Z.; Parapercis, L. Olig.-E, Mio. N.Z.
Uranoscopidae, Uranoscopus, M. Eo.
N.Z. Leptoscopidae, Lepltoscopus, L.
Olig.-L. Mio, N.Z. Hemerocoetidae.
Hemerocoetus, E. Mio,-Plio, N.Z.;
Krebsiella, L. Eo.-E. Mio, N.Z.; Waitakia,
L. Olig.-E, Mio. NZ.

INFRACLASS: DIPNOI
Dipnorhynchidae. Dipnorhynchus, E.
Dev. N.S.W., Vic.; Speonesydrion, E.
Dev. N.S.W. Chirodipteridae.
Chirodipterus, L, Dev. WA.
Pillurarhynachus, L, Dev. W.A
Dipteridae. Genus indet,, L. Dev. N.T,
Family uncertain, New genera, L. Dev.
Vic. Rhynchodipteridue.
Griphognathus, L. Dev. W.A,;
Soederberghia, L, Dev. N.S.W.
Holodipteridae, Holodipterus, L. Dev.
W.A. ?Ctenodontidae. Eoctenodus, L.
Dev. Vic.; Delatitia, E. Carb. Vic.
Protuceratodontidae. Ceratodus, Trias.
N.S.W., Qu., Tas., Vic., W.A., E, Cret.
N.S.W., E.-M. Plio. Qd., L, Plio.
N.S.W.; Gosfordia, Trias. N.S.W.
Neuceratodontidue. Neoceratodus, E,
Cret.-Rec. N,S.W., N-T., Qd., S.A.

INFRACLASS: CROSSOPTERYGYII
ORDER: ONYCHODONTIDA
Onychodontidae. Onychodus, Genus
indet., E. Dey, N.S.W., Qd., M. Dev.
Qu., L. Dev. W.A,

ORDER: ACTINISTIA
Covlacanthidae. Genus indet,, Trias,
Tas,, L. Jur. N.S.W.

ORDER; POROLEPIFOR MES
SUPERFAMILY: HOLOPTYCHOIDEA
Holoptychiidae. Genera indet., E. Dev,
N.5.W.; ?Glyptolepis, L. Dev. Vic.
Holoptychius, L., Dev. N S.W.

ORDER: RHIZODONTIDA
Rhizodontidae. Barameda, E. Carb.
Vic.; Genus indet., E. Carb. Qd., W.A.

ORDER: OSTEOLEPIFORMES
Canowindridae. Beelarongia, L. Dev.
Vic.; Canowindra, L. Dev. N.S.W.
Osteolepididae. Gogonasus, L. Dev.
W.A.; ?Gyroptychius, M. Dev. N.S.W.
Megalichthyidae. Megalichthys, E.
Carb. Qd. Eusthenopteridae.
Marsdenichthys, L. Dev. Vic.;
Eusthenodon, L. Dev. N.S.W.

CLASS: AMPHIBIA
SUBCLASS: LABYRINTHODONTIA

ORDER: TEMNOSPONDYLI
Family uncertain. Metaxygnathus, L.
Dev. N.S.W. Micropholidae.
Lapillopsis, E. Trias. Qd. Lydek-
kerinidae. Chomatobatrachus, E. Trias.
Tas. Capitosauridae, Parotosuchus, E.
Trias, N.S.W., Qd., Tas., W.A.;
Paracyclotosaurus, M. Trias. N.S.W.
Rhytidosteidae. Rewana, Arcadia,
Acerastea, E, Trias. Qd.; Deltusaurus,
E, Trias. Tas., W.A.; Derwentia, E. Trias.
Tas. Trematosauridae.
Erythrobatrachus, E. Trias. W.A..
Brachyopidae. Genus indet., E. Cret.
Vic.; Bothriceps, L. Perm. N.S.W.;
Blinasaurus, E. Trias. N.S.W., Tas.,
W.A.; Xenobrachyops, E. Trias, Qd.;
Notobrachyops, M. Trias. N.S.W.;
Austropelor, E, Jur. Qd.
Chigutisauridae. Keratobrachyops, E.
Trias. Qd.; Siderops, E. Jur. Qd.
Plagiosauridae. Plagiobatrachus, E.
Trias. Qd.

SUBCLASS: LISSAMPHIBIA
ORDER: ANURA

Hylidae. Australobatrachus, L. Olig./M.
Mio. S.A.; Litoria, L. Olig./M. Mio.
S.A. M. Mio. Qd. Pleist. S.A., WA,
Rec. Leptodactylidae. Limnodynastes,
L. Olig./M. Mio. Qd., S.A., M. Mio.
Qd,, Pleist. $.A., Tas., W.A., Rec.;
Crinia , L. Olig./M. Mio, Qd., Pleist.
S.A., W.A., Rec.;
Heleioporus/Neobatrachus, Geocrinia,
Pleist. §.A., W.A.; Lechriodus, Crinia,
Kyarranus, L. Olig./M. Mio. Qd.;
Neobatrachus, Plio.-Pleist. S.A., Pleist.
W.A.

SUPERCLASS; AMNIOTA

CLASS: INCERTAE SEDIS
Family uncertain. Kudnu, E. Trias.
Qd. Procolophonidae. ?Procolophon,
E. Trias. Qd.

CLASS; ANAPSIDA
SUBCLASS: TESTUDINES
SUBORDER: INCERTAE SEDIS

Fumilies uncertain.
Chelycarapookus, E. Cret. Vic.;
Cratochelone, E, Cret. Qd-
Protostegidae. Genus indet., L. Cret.
N.Z.

SUBORDER: PLEURODIRA
Chelidae, Chelodina, Mio, ?Plio. Qd.,
Pleist. Qd.; Emydura, Olig./Mio, Tas., L.
Olig./M. Mio,, S.A,, Plio, N.S.W., Qdy
Pseudemydura, Mio, Qd.

SUBORDER: CRYPTODIRA
SUPERFAMILY; BAENOIDEA
Meiolaniidae. Genus indet., L.
Olig./M. Mio. S.A.; Meiolania,
L.Olig./M. Mio. N.T., Qd., S.A., Plio.
N.S.W., Qu, Pleist. Lord Howe Is., N.
Caledonia, Qd.

SUPERFAMILY: TRIONYCHOIDEA
Carettochelyidae. ?Carettochelys,
Pleist. W.A. Trionychidae. Genus
indet. E. Tert.-Pleist. Qd., Mio. N.T.

SUPERFAMILY: CHELONIOIDEA
Family uncertain. Notochelone, E.
Cret. Qd.

CLASS: DIAPSIDA
SUBCLASS: LEPIDOSAUROMORPHA
SUPERORDER; LEPIDOSAURIA

ORDER: SPHENODONTA
Sphenodontidae. Sphenodon, L.
Pleist. N.Z., Rec.

ORDER: SQUAMATA
SUBORDER: IGUANIA
Agamidae. Amphibolurus, Pleist. S.A.,
2Qd., Rec.; Chlamydosaurus, Pleist, Qu.,

Rec.; Physignathus, L., Olig./M. Mio,
Qd., Rec.; Sulcatidens, L. Olig./M. Mio.
Qd., Rec.

SUBORDER: SCINCOMORPHA
Scincidae. Genera indet., L. Olig./M.
Plio. Qd.; Egernia, L, Olig./M. Mio.,
Pleist. S.A., Rec.; ?Sphenomorphus,
Pleist, $.A., Rec.; Tiliqua, Mio. S.A,, M.
Plio.-Pleist. Qd., S.A., Rec.;
Trachydosaurus, Plicist. S.A., Rec.

SUBORDER: ANGUIMORPHA

INFRAORDER: VARANOIDEA
Varanidae. Genus indet,, L. Olig./M.
Mio. Qd., M. Plio. S.A.; Megalania, Plio.
Qd.; Pleist. N.S.W., Qd,, S.A.; Varanus,
L. Plio.-E. Pleist. Qd., Pleist. N.
Caledonia, Rec. Mosasauridae. Genus
indet., L, Cret. W.A.; Mosasaurus, L.
Cre, N.Z.; Prognathodon, L. Cret. N.Z,;
Taniwhasaurus, L. Cret. N.Z.; Tylosaurus,
L. Cret. N.Z.

INFRAORDER: SERPENTES
Boidae. Morelia, L. Olig./Mio. N.T.,
Plio. Qd., Rec.; Montypythonoides, L.
Olig./M. Mio. Qd.; Wonambi, M. Plio.-
Pleist. S.A. Acrochordidae. Genus
indet., Plio. Qd. Elapidae. Genus
indet., L.. Olig./M. Mio. Qd.; Genus
indet., M. Plio. $.A.; Notechis,
Pseudechis, Pseudonaja, Pleist S.A.,
Rec. Typhlopidae. ?Rhamphotyphlops,
L. Olig./M. Mio, Qa.

SUPERORDER: SAUROPTERYGIA
SUPERFAMILY: PLESIOSAUROIDEA
Elasmosauridue. Genus indet., L, Cret.
N.Z.; Mauisaurus, L. Cret. N.Z.;
Tuarungisaurus, L. Cret. N.Z.;
Woolungasaurus, E. Cret, Qd., 2S.A.
Cimoliasauridae. Cimoliasaurus, E.
Cret. N.S.W.

SUPERFAMILY: PLIOSAUROIDEA
Family uncertain. Genus indet., L,

Cret. N.Z. Dolichorhynchopidae.
Dolichorhynchops, E. Cret. N.S.W.
Pliosauridae. Kronosaurus, E. Cret.

Qd

SUBCLASS: ICHTHYOPTERYGIA
Family uncertain. Genera indet., M.
Trias.-E, Cret. N.Z. Leptoptery-
giidae, Plarypterygius, E. Cret. N.T., Qd.

SUBCLASS: ARCHOSAUROMORPHA

SUPERORDER: PROTOROSAURIA
Prolacertidae. Kadimakara, E, Trias.
Qd.

SUPERORDER: ARCHOSAURIA
ORDER: PROTEROSUCHIA
Chasmatosauridae, Kalisuchus, E.
Trias. Qd.; Tasmaniosaurus, E. Trias. Tas,

ORDER: CROCODYLOTARSI
SUBORDER: CROCODYLOMORPHA
INFRAORDER: NEOSUCHIA

Family uncertain. "Crocodylus”, E.
Crec N.S.W. Crocodylidae. Baru,
Olig./Mio. Qd.; Crocodylus, Plio.-Quat
Qd., Rec.; Pallimnarchus, L. Mio. N.T.,
Plio. Pleist. Qd.; Quinkana, ?Plio., Pleist.
Qd. Mekosuchidae. Mekosuchus,
Quat N. Caledonia. Sebecosuchia or
Pristichampsinae. Plio. S.A.

ORDER: ORNITHODIRA
SUBORDER: PTEROSAURIA
Ornithocheiridae. Genus indet, E.
Cret. Qd., ?Vic. ?Azhdarchiidae.
Genus indet., L. Cret. W.A.

SUBORDER: SAURISCHIA

INFRAORDER: THEROPODA
Family uncertain. Genera indet, E.
Cret. Vic., L. Cret. N-Z.; Kakuru, E. Cret.
§.A.; Rapator, E. Cret. N.S.W.
Allosauridae. Allosaurus, E. Cret. Vic.

INFRAORDER:
SAUROPODOMORPHA
SUPERFAMILY: PROSAUROPODA
?Massospondylidae. Agrosaurus, ?L.
Trias. 7Q¢.

SUPERFAMILY: SAUROPODA
Family uncertain, Austrosaurus, E.-L.
Cret. Qd. Cetiosauridae.
Rhoetosaurus, M, Jur. Qd.

SUBORDER: ORNITHISCHIA

INFRAORDER: ORNITHOPODA
Family uncertain. Genera indet, E.
Cret. N.S.W., Qd., Vic., L. Cret. N.Z.
Hypsilophodontidae.
Atlascopcosaurus, E. Cret. Vic.;
Fulgurothertum, E. Cret. N.S.W., Vic.;
Leaellynasaura, E. Cret, Vic.

INFRAORDER: ANKYLOSAURIA
Ankylosauridae. Minmt. E. Cret. Qd.

CLASS: MAMMALIA
ORDER: MONOTREMATA
Family uncertain. Steropodon, E.
Cret., N.S.W. Ornithorhynchidae.
Obdurodon, L. Olig./M, Mio. 3.A.;
Ornithorynchus, E. Plio. N.S.W., Pleist.
Aust. Ree. Tachyglossidae.

SYSTEMATIC INDEX -

Tachyglossus, Quat. Aust, Rec.;
Zaglossus, E.-M. Mio. ?Plio-Pleist.
N.S.W,, Qd., Pleist. N.G., Rec.

SUPERCOHORT: MARSUPIALIA
Yingabalanaridae. Yingabalanara, L.
Olig./M. Mio. Qd.

ORDER: DAS YUROMORPHIA
Dasyuridae. Genus indet., L. Olig. Tas.;
Ankotarinja, L. Olig./M. Mio. S.A.;
Antechinomys, Pleist. s.cent. Aust, Rec.;
Antechinus, E. Plio., Pleist. Aust., Rec.;
Apoktesis, L. Olig./M. Mio. S.A.;
Dasycercus, Pleist. s.w. Aust, Rec.;
Dasykaluta, ?Pleist. W.A., Rec.;
Dasylurinja, L. Olig./M. Mio. S.A.;
?Dasylurinja, L. Olig,/M. Mio. S.A.;
Dasyuroides, Plio., S.A., 2E. Pleist.
N.S.W., W.A., Rec.; Dasyurus, Plio.
N.S.W,, Qd., S.A., Vic., Quat. Aust.,
Rec.; Glaucodon, L, Plio. Vic., M. Plio.
S.A. Keeuna, L. Olig/M. Mio. S.A.;
Myoictis, Plio. N.G., Rec.: Parantechinus,
Pleist. s.w. Aust., Rec.; Phascogale, 7M.
Plio. S.A. Pleist. e.s.w, Aust, Rec.;
Planigaie, E. Plio. Qd., Pleist. Aust.
except Tas., Rec.; Pseudantechinus,
?Pleist. W.A., Rec.; Sarcophilus, ?L.
Plio., N.S.W., Pleist. Aust., Rec.;
Satanellus, 2E. Plio. e.2w. Aust, N.G.,
Rec.; Sminthopsis, L. Plio. N.S.W., Qd,
IE. Pleist e.s.w. Aust., Rec.; Wakamatha,
2M, Mio. S.A. Myrmecobiidae
Myrmecobius, Pleist. s.cent. Aust, Rec.
Thylacinidae. Genera indet., L.
Olig./M. Mio. Qd.; Thylacinus,
L.Olig./M. Mio, Qd., L. Mio. N.T., Plio.
N.G., N.S.W., Qd., S.A.; Pleist. Aust.,
Rec.; Nimbacinus, L. Olig./M. Mio.
N,T., Qd.

ORDER: PERAMELEMORPHIA
Peramelidae. Chaeropus, E. Pleist.
s.cent. Aust., Rec.; /soodon, L. Plio.-
Pleist. Vic., Rec.; Perameles, E. Plio.
N.S.W., Qd., Vic., Pleist. Aust, Rec.
Thylacomyidae. /schnodon, Plio./E.
Pleist. $.A.; Macrous, Pleist. cente.s.w.
Aust, Rec.

ORDER: NOTORYCTEMORPHIA
Notoryctidae. Genus indet., L.
Olig./M, Mio. Qd.

ORDER: DIPROTODONTIA
SUBORDER: VOMBATIFORMES
INFRAORDER:

PHASCOLARCTOMORPHIA
Phascolarctidae. Koobor, Plio. Qd.;
Madakoala, L. Olig./M, Mio. S.A;
Litokoala, L. Olig./M. Mio, Qd., S.A.;
Perikoala, L. Olig./M. Mio. S.A.;
Phascolarctos, E. Plio, S.A., Pleist.
Aust., Rec.

INFRAORDER: VOMBATOMORPHIA
Diprotodontidae. Subfamily
Diprotodontinae. Bematherium, M. Mio.
Qd.; Diprotodon, Plio. N.S.W., S.A.,
Pleist, Aust.; Euowenta, E. Plio.-Pleist.
Qd., S.A., L. Plio. N.S.W., Plio.-Pleist.
Vic.: Brachalletes, Plio. Aust;
Euryzygoma, E.-M, Plio,-Pleist. Qd.;
Jlulithertum, Pleist, N.G.; Koalemus,
Plio. Aust.: Kolopsts. L. Mio. N.T.. Vic..

1415

Pho. N.G.; Kolapsoides, Plio. N.G.;
Meniscalophus, L. Plio/E. Pleist. S.A.;
Nimbadon, L. Olig./M. Mio. N.T., Qd.;
Neohelos, L. Olig./M. Mio. N.T., Qd.,
S.A,; Nototherium, Plio.-Pleist. N.G.,
N.S.W.; Plaisiodon, L. Mio. N.T.;
Pyramios, L. Mio. N.T.; Raemeotherium,
L. Olig./M. Mio. S.A.; Sthenomerus,
?Pleist, Aust. Subfamily
Zygomaturinae. Zygomaturus, L. Mio.-
Pleist, N.S.W., Qd., S.A., Tas., Vic.,
W.A., 7E. Plio. N.T., Pleist, N.G.
Palorchestidae. Ngapakaldia, L.
Olig./M. Mio. Qd., S.A.; Pitikantia, L.
Olig./M. Mio. $.A.; Palorchestes, L.
Olig./M. Mio. Qd., L. Mio. N.T., Vic.,
Plio. N.S.W., Qd., S.A. Wynyardiidae.
Namilamadeta, L. Olig/M Mio. Qd.,
S.A.; Wynyardia, L. Olig./M. Mio. Tas.;
Muramura, L. Olig/M. Mio. S.A.
Ilariidae. /laria, L. Olig./M. Mio. S.A.;
Kuterintja, L, Olig./M. Mio. S.A.
Vombatidae. Lasiorhinus, L. Plio.
N.S.W., S.A., Rec.; Phascolomys, L.
Plio, S.A., Pleist. e., s. Aust;
Phascolonus, Plio. N.S.W., S.A., Vic.,
W.A., Pleist. Aust.; Ramsayia, E. Plio.
N.S.W., Pleist. N.S.W., Qd.;
Rhizophascolonus, L. Olig./M. Mio.
5.A.; Vombatus, Plio. N,S,W., Qd.,
S.A., Vic., Pleist. Aust., Rec.; Warendja,
Pleist. S.A., Vic. Thylacoleonidae.
Thylacoleo, Plio. N.S.W., Qd., S.A.,
Vic., Pleist. Aust; Wakaleo, L. Olig./M.
Mio. N.T., Qd., S.A., Mio. N.T.;
Priscileo, L. Olig./M. Mio. S.A.

INFRAORDER: PHALANGERIDA
SUPERFAMILY: PHALANGEROIDEA
Phalangeridae, Subfamily Ailuropinae.

Ailurops, Pleist. Sulawesi. Subfamily
Phalangerinae, Tribe Trichosurini.
Trichosurus, L. Olig./M. Mio. Qd., Plio.
Vic., Rec.; Strigocuscus, L.Olig /M. Mio.
Qd., E. Plio. Vic. Tribe Phalangerini.
Phalanger, E. Plio. Vic., Rec.
Miralinidae. Miralina, L. Olig./M.
Mio. S.A. Ektopodontidae.
Ektopodon, L. Olig/M. Mio, S.A;
?Chunia, L. Olig/M. Mio. S.A.; Darcius,
E, Plio., E. Pleist. Vic.

SUPERFAMILY: MACROPODOIDEA
Potoroidae. Subfamily Bulungamayinae.
Bulungamaya, L. Olig./M. Mio. Qd.,
S.A.; Wabularoo, L. Olig./M. Mio. Qd.;
Galanarla, L. Olig./M. Mio. Qd.
Subfamily Hypsiprymnodonunae.
Hypsiprymnodon, L. Olig/M. Mio. Qd.,
Plio. Vic., Rec. Subfamily Propleopjnac.
Ekaltadeta, L. Olig./M. Mio, Qd.;
Propleopus, E, Plio.-Pleist. N.S.W., Qd.,
S.A., Vic. Subfamily Potoroinae.
Aepyprymnus, Pleist. Qd., Rec.;
Bettongia, Plio. N.S,W., S.A., Vic., Tas.,
W.A., N.S.W., Rec.; Milliyowt, E. Plio.
Vic.; Caloprymnus, Pleist. N.S.W.,
W.A., Rec.} Potorous, M. Plio. S.A., L.
Plio.-Pleist. Vic., Pleist. S.A., W.A,,
Tas., Rec.; Purtia, L. Olig./M. Mio. S.A.;
Wakiewakie, L. Olig./M. Mio. N.T., Qd.,
S.A.: Gumardee, L. Olig./M. Mio. Qd.,
S.A. Subfamily Palaeopotoroinae.
Palaeopotorous, L, Olig./M. Mio. S.A.
Macropodidae, Subfamily Balbannae.
Balbaroo, L. Olig./M. Mio. N-T,, Qd.,

1416 -

S.A. Subfamily Sthenurinae,
Lagostrophus, L.. Plio, W.A., N.S.W.,
Rec.; Proceptodon, Pleist. N.S.W., Qd.,
S.A., Vic; Sthenurus, Plio. N.S.W., Qd.
S.A., Vic., Pleist. Aust.; Troposodon,
Plio. Qd., Vic., S.A., Pleist. Aust;
Simosthenurus, L, Plio, N.S.W., Vic.,
Picist. Aust. Subfamily Macropodinae,
Bohra, Pieist, N.S.W.; Dendrolagus, E.-
M. Pliv., Pleist. N.G., N.S.W., Vic., L.
Plio. $.A., Rec.; Dorcopsis, L. Olig./M.
Mio, N.T., E. Plio. Vic., S.A., L. Plio.
N.G.; Dorcopsoides, L. Mio. N.T., E.
Plio. N.T.; Dorcopsulus, Pleist. N.G.,
Rec.; /ludronomas, L. Mio. N.T.; Kurrabi,
Plio. N.S.W., S.A, Vic.; Lagorchestes,
Picist. N.S.W., S.A,, W.A., Rec.;
Macropus, Plio, N.S.W., Qd., Vic., Rec.;
Nambaroo, L. Olig./M. Mio, Qd., S.A.;
Onychogalea, Pleist. N.S.W., W.A., Rec.;
Petrogale, Plio. N.S.W., Qd., Rec, Aust.
except Tas,; Prionotemaus, E.-M. Plio.
N.S.W., Qd., L. Plio./E. Pleist. Qd.,
§.A.; Protemnodon, E., Plio. Vic., E,
Plio-Plcist. N.G., N.S.W., Qd., S.A.,
Tas., Vic., W.A.; Setonix, PleisL. W.A.,
Rec.; Thylogale, Plio, Vic., Rec.;
Wallabia, Plio. N.S.W., Qd., Vic., Pleist.
Aust., Rec.; Watutia, Plio. N.G,

SUPERFAMILY: BURRAMYOIDEA
Burramyidae. Genus indet,, L. Olig.-E.
Mio., Tas.; Burramys, L. Olig./M. Mio.
Qd., S.A., Plio. Vic., Pleist.-Rec. Aust.;
Cercartetus, L. Olig./M. Mio., Pleist. Qd.,
Pleist,-Rec. N.S.W., S.A., W.A.

SUPERFAMILY: PETAUROIDEA
Pseudocheiridae. Marlu, L. Olig./M.
Mio. §.A.; Puljara, L. Olig./M. Mio. Qd.
5.A.; Pildra, L. Olig./M. Mio. Qd., S.A.;
Pseudocheirops, L. Olig,/M. Mio. Qd,, L.
Mio, N.T., E.-M. Plio. N.S.W., Pleist.
NG.,, Rec.; Petauroides, Pleist. e. Aust,
Rec., Pseudocheirus, Pliv. Vic., Picist.
Aust. N.G., Rec.; Pseudokoala, E. Plio.
Vic. Petauridue. Subfamily
Petaurinae, Gymnobelideus, Pleist.-Rec, ¢.
Aust., N,.G.; Petaurus, E. Plio. S.A.,
Vic., Rec. Dactylopsilinac. Genus indet,
L. Olig./M. Mio. Qd. Pilkipildridae.
Pilkupildra, L. Olig./M. Mio, S.A.;
Djilgaringa, L. Olig./M. Mio. Qd.

SUPERFAMILY: TARSIPEDOIDEA
Tarsipedidae. Jarsipes, Plcist. W.A.,
Rec. Acrobatidae. Acrobates, L.
Olig./M. Mio. Qd., Pleist. Aust., Rec.

ORDER; YALKAPARIDONTIA
Yalkuparidontidae. Yalkaparidon, L.
Olig.-M. Mio. Qd.

PLACENTALIA
ORDER; CHIROPTERA
SUBORDER: MEGACHIROPTERA
Pteropodidae. Aproteles, Plcist.-Rec.
N.G.; Dobsonia, Pleist. N.G., Rec.;
Pteropus, Pleist, N.G., Rec,

SUBORDER: MICROCHIROPTERA.
Family uncertain. E. Plio. Vic.
Rhinoluphidae. Genus indct., M. Mio.
S.A, Rhinolopus, Pleist. Qd., Vic., Rec.

SYSTEMATIC INDEX

Hipposideridae. Hipposideros, M. Mio-
?Pleist. Qd.; Brachipposideros, L.
Olig./M. Mio. Qd. Megadermatidue.
Genus indct., M. Mio. Qd.; Macroderma,
L. Olig./M. Mio., Plio. Qd., E.-M. Plio.
N.S.W., Pleist. N.S.W., N.T., Qd., S.A.,
W.A., Rec. Molossidae. Genus indet.,
L, Olig./M. Mio. Qd.; Mormopterus,
Pleist. Vic., Rec,; Tadarida, Pleist. €., w.
Aust., Rec. Vespertilionidae,
Chalinolobus, Pleist. Vic., W.A., Rec.;
Eptesicus, Pleist. Vic., W.A., Rec.;
Miniopterus, Pleist. Qd., Vic., Rec.;
Nyctophilus, Pleist, Qd., Vic., W.A.,
Rec.; Pipistrellus, Pleist. Vic., W.A.,
Rec,

ORDER PRIMATES
Hominidae. /omo, L. Pleist. Aust.,
N.G.

ORDER: CARNIVORA
Otariidae. Arctocephalus, Pleist. N.Z.,
Rec.; Neophoca, M, Pleist. Vic., N.Z.,
Rec. Phocidae. Mirounga, Pleist.
N.Z.; Monarchinae indet., L. Mio-E.
Pleist.-E, Plio. Vic.; Ommatophocca, L.
Plio, N.Z., Rec.

ORDER: RODENTIA
Muridae. Subfamily Hydromyinae.
Anisomys, Pleist, N.G., Rec.; Conilurus,
Pleist. Qd., S.A., Vic., Rec.; Hydromys,
Pleist. N.S.W., Tas., Vic., W.A., Rec.;
Hyomys, Pleist. N.G., Rec.; Leggadina,
Pleist. Qd., Rec.; Leporillus, Pleist,
N.S.W., S.A., W.A., Rec.; Mallomys,
Pleist. N.G., Rec.; Mastacomys, Pleist.
N.S.W., Vic., Tas., $.A., Rec.; Melomys,
Pleist. N.G. Vic., Rec.; Mesembriomys,
?Pleist, W.A., Rec.; Notomys, Pleist.
W.A., N.S.W., S.A., Rec.; Pseudomys,
Plio. Qd., Vic., Pleist. Aust., Rec.;
Solomys, Plcist, Sol. Is.; Uromys, Pleist.
N.G., Rec.; Zyzomys, Plio. Qd., ?Picist.
N.T., Qd., W.A., Rec. Subfamily
Murinae. Rattus, L. Plio,-E. Pleist. Qd.,
Pleist. Qd., N.S.W., Vic., S.A., W.A.,
Rec.

ORDER: CETACEA
SUBORDER: INCERTAE SEDIS.
Cetotilites, E. Olig. Vic.; "Squalodon",
Olig. S.A.

SUBORDER: ARCHAEOCETI
Basilosauridae. Kekenodon, L. Olig.
N.Z.

SUBORDER: MYSTICETI,
Family uncertain. Gencra indet.,
Eo.-Olig., N.Z.; Mammalodon, L. Olig.
Vic. Cetotheriidae. Genera indet., E.
Mio.-E. Plio, Vic,, L. Mio. S.A.;
Mauicetus, L. Olig.-E. Mio, N.Z.; of.
Parietobalaena. (="Aglaocetus"), E. Mio.
S.A.; Pelocetus, E. Mio. Vic.
Balaenopteridae. Genera indet., M.
Mio.-E. Plio. Vic., L. Mio. N.Z.;
Balaenoptera, Plio, N-Z., Rec. ;
Megaptera., 2L. Plio. Tas., L. Mio.-E.
Plio. Vic., Rec, Aust. Balaenidae.
Genera indet., L. Plio. N.Z., ?Mio.-L.
Plio. S.A.; Eubalaena, Rec. Aust.; cf.
Eubalaena, L: Plio, $.A., L. Mio.-E. Plio.
Vic., Rec. Aust.; of. Morenocetus, E.-?M.
Mio, S.A,

SUBORDER: ODONTOCETI
Squaludontidae. Gencra indet., L.
Olig.-E, Mio. N.Z., E. Mio. Vic;
Austrosqualodon, L, Olig. N.Z.;
Metasqualodon, L, Olig. ¢.S.A.;
Microcetus, L, Olig.-E. Mio. N.Z.;
Parasqualodon, L. Olig. Vic.;
Prosqualodon, L, Olig.-E. Mio, N.Z., BE.
Mio. Tas.; Tangaroasaurus, E. Mio, N.Z.
Eurhinodelphidae. Genera indet., L.
Olig.-L. Mio, N.Z.; Phocaenopsis, E.
Mio. N.Z. Ziphiidae. Genera indet.,
7E Plio. Tas., M.-L. Mio, Chatham Rise;
?Hyperoodon, M.-L. Mio. N.Z.;
Mesoplodon, L. Mio.-E. Plio. Vic., Rec.;
"Ziphius", 2E Plio. Tas. Physeteridae.
Genera indet., L. Olig.-E. Mio. N.Z., E.
Mio.-E. Plio. Vic., ?7E. Plio, Tas., L.
Plio. N.Z.; Physeter, Rec.; Physetodon.,,
L. Mio-E. Plio. Vic.; Scaldicetus, M.-L.
Mio. Chatham Rise, L. Mio.-E. Plio.
Vic.; Scaptodon, ?E. Plio. Tas.
Kogiidae. ?Kogia, M.-L. Mio, Chatham
Rise. Kentriodontidae. Genera indet,
L. Olig.-E. Mio. N.Z. Rhabdosteidae.
Genus indet., L. Olig./M. Mio. S.A.
Delphinidae. Genera indet., Neogene
N.Z., L. Mio.-E. Plio. Vic., L. Plio.
N.Z.; Delphinus, L. Plio. N.Z., Quat.
Vic., Rec. ; Globicephala, Neogene N.Z.;
?Pseudorca, L. Plio. N.Z.; ?Orcinus, E.
Pleist. N.Z.; "Steno", L. Mio.-E. Plio.
Vic., E. Plio. N.Z.

ORDER; PERISSODACTYLA
Rhinoceratidae. cf. Rhinoceros, (not
Zygomaturus) Quat. N. Caledonia - not in
situ,

ORDER: PROBOSCIDEA
Mastodon, N.S.W.; Notoelephas, Qd.
(transported into Australia by trade, not
found in situ).

ORDER; SIRENIA
Halicore, Age uncertain N,G.

CLASS; AVES
ORDER: ENANTIORNITHIFORMES
Enantiornithidae. Nanantius, E. Cret,
Aust,

ORDER: SPHENISCIFORMES
Spheniscidae. Anthropodytes, E. Mio.
Vic.; Anthropornis, L, Eo. Ant.
Peninsula., $.A.; Aptenodytes, Mio,-Plio.
NZ., Rec.; Archaueospheniscus, L. Olig,
Ant. Peninsula, N.Z.; Duntroonornis, L.
Olig. N.Z.; Eudyptes, Quat. N.Z., Rec.;
Eudyptula, Quat. N.Z., Rec.; Korora, L.
Olig.-E. Mio. N.Z.; Marplesornis, Mio.-
Plio. N.Z.; Megadyptes, Quat. Chatham
Is. N.Z., Rec.; Pachydyptes, L, Eoc, N.Z.;
Palaeeudyptes, L. Eoc.-E, Tert. Ant.
Peninsula, S.A., Vic., L. Eo.-E. Mio.
N.Z.; Platydyptes, L. Olig.-E. Mio. N.Z.;
Pseudoaptenodytes, L, Mio. Vic.;
Pygoscelis, Plio. N.Z., Rec., Tasidyptes,
Quat. Tas.; Tereingaornis, M. Plio. N.Z.

ORDER: DROMORNITHIFORMES
Dromornithidae. Barawertornis, L.
Olig./M. Mio. Qd.; Bullockornis, L.
Olig./M. Mio. N.T., Qd.; Dromornis, L.
Olig./Mio. Qd. L. Mio. N.T., Plio.
N.S.W.; Genyornis, Pleist. Aust, except
N.T.; /lbandornis, L. Mio. N.T.

ORDER: CASUARIIFORMES
Casuariidae. Casuarius, Plio. N.G.,
Pleist. 2n. Aust., Rec.; Dromaius, L.
Olig./M. Mio. Qd., N.T., S.A., E.-M.
Plio, Qd., Plio, $.A., Quat. Aust., Rec.

ORDER: DINORNITHIFORMES
Anomalopterygidae. Anomalopteryx,
Quat, N.Z.; Emeus, Quat. N.Z.;
Euryupteryx, Quat. N.Z.; Megalapteryx,
Quat. N.Z.; Pachyornis, Quat. N.Z.
Dinorthithidae. Dinornis, Quat. N.Z.

ORDER: APTERYGIFORMES
Aplerygidae. Apleryx, Quut, N.Z., Rec.

ORDER: PODICIPEDIFORMES
Podicipedidue, Podiceps, Quat, N.Z,,
Ree.; Genera Indet., L. Olig./M. Mio.-
Quut. Aust.

ORDER: PROCELLARIIFORMES
Diomedeidae. Diomedea, L. Mio. Vic.,
Rec. Pelecanoididae. Pelecanoides,
Quat. Aust. N.Z., Rec. Procellariidae.
Daption, Quat. N.Z., Rec.; Macronectes,
Quat. N.Z., Rec.; Fulmarus, Quat. N.Z.,
Rece.; ?Halobaena, Quat, N.Z,, Rec.;
Pachyptila, Quat. N.Z., Rec.; Pterodroma,
Quat, N. Culedonia, N.Z., Rec.;
Procellaria, Quat. N.Z., Rec.; Puffinus,
Pleist. Aust., N.Z., L. Howe Is., Rec.
Oceanitidae. Fregeita, Quat. N.Z.,
Rec.; Garrodia, Qual. N.Z., Rec.;
Oveanites, Quat. N.Z., Ree.; Pelagodroma,
Quat. N.Z., Rec. Pelecanvididae.
Pelecunoides, Quat. N.Z., Rec.

ORDER: PELECANIFORMES
SUBORDER; PELECANI
SUPERFAMILY: PELECANOIDEA

Pelecanidae, Pelecanus, L. Olig./M.
Mio., Plio.-Pleist. $.A., E.-M. Plio. Qd.,,
Quat. N_Z., Rec. Pelagornithidae.
Neodontornis, Plio. N.Z.; ?Odentopter yx,
Pelagornis, Mio, N.Z.; Pseudoodontornis,
Tert, N.Z.

SUPERFAMILY: SULOIDEA
Sulidae. Sula, Qual. N.Z., Rec.
Anhigidae. Anhinga, L. Plio. S.A.,
Qual. S.A., Qd., Rec.
Phalacrocoracidae. Microcarbo, Plio.
Qd.; Leucocarbo, Qual. N.Z., Rec.;
Phulacrocorax, L. Olig./M, Mio.-Quat.
Aust., Quat. N.Z., Rec.; Stictocarbo,
Quat, N.Z., Rec.

ORDER: CICONIIFORMES
SUBORDER: ARDEAE
Ardeidae, Genus indet., L. Plio. S.A.;
Egretia, Quat. N.Z., Rec.; Botaurus, Quat.
NZ, Rec.; Lrobrychus, Quat. N.Z., Rec.;
Platalea, Quat. N.Z., Rec.

SUBORDER: CICONIAE
Ciconiidue. Genus indet., L. Plio.
S.A.; Ephipiorhychus, Pleist. Aust., Ree;
Ciconia, Plio.-Quat, Qd.; Xenorhychus,
Plio. n. Aust., Rec.; Palaeopelargus,
Pleist. Qd.; Threskiornithidae.

Threskiornis, Plio, Qd., Quat. Aust., Rec.

ORDER: PHOENICOPTERIFORMES
Phoenicopteridae. Genus indet., L.
Mio. N.T.; Ocyplanus, L. Plio.-Quat.

SYSTEMATIC INDEX - 1417

S.A,; Phoeniconotius, L. Olig./M. Mio.
S.A; Phoenicopterus, L. Olig./M, Mio.
S.A., Pleist. S.A,, Rec.;
Xenorhynchopsis, L. Plio.-Quat. S.A.
Palaelodidae. Palaelodus, M. Mio.-M.
Pleist. S.A.

ORDER: ANSERIFORMES
SUBORDER: ANSERES
Anatidae. Anas, Plio,-Quat. Aust., Quat.
N. Caledonia, N.Z., Rec.; Anseranas, E.-

M. Plio. Qd., Rec.; Aytha, E.-M. Plio.
Qd., Quat. Aust. N.Z., Rec.;
Archeocygnus, Quat. S.A.; Biziura, Plio.-
Quat. Aust. N.Z., Rec.; ?Cereopsis, E.-M.
Plio. Qd., Ree.; Chenopsis, Quat. S.A.;
Cnemiornis, Quat. N.Z.; Cygnus, E.-M.
Plio. Qd., Quat. Aust. N.Z., Rec.;
Dendrocygna, E.-M. Plio, Qd., Rec.;
Euryanas, Quat. N.Z.; Hymenolaimus,
Quit. N.Z., Rec.; Malacorhynchus, Quat.
Aust. N.Z., Rec.; Mergus, Quat. N.Z.,
Rec.; Oxyura, Quat. N.Z., Rec.;
Pachyanas, Quat. Chatham Is.; Tadorna,
E.-M. Plio. Qd., Quat. Aust. N.Z., Rec.

ORDER: FALCONIFORMES
SUBORDER; FALCONES
SUPERFAMILY: FALCONOIDEA
Accipitridae. Aquila, Quat. Aust., Rec.;
Hieraeetus, Quat. N.Z., Rec.; Accipiter,
Quat. N. Caledonia, N.Z., Rec.; Circus,
Quat. N.Z., Rec.; Aviceda, Quat. S.A.,

Rec.; Harpagornis, Quat. N.Z.; Hieraaetus,
Quat. Aust., Rec.; Ichthyophaga, Quat.
Chatham Is., Rec.; Necrastur, Plio. Qd.;
Haliaeetus, Pleist. Kangaroo Is., Aust.
N.Z., Rec.; Taphaetus, Quat. $.A., Qd.
Falconidae. Pliogetus, Pleist. Aust.;
Falco, Quat, Aust. N. Caledonia, N.Z.,
Rec.

ORDER: GALLIFORMES
SUBORDER: GALLI

Megapodiidae. Megapodius, Quat. N.
Caledonia, Rec.; Palaeopelargus, Pleist.
Aust; Progura, Quat. Aust; Leipoa, L.
Pleist. Aust., Rec.; Sylviornis, Quat. N.
Caledonia, Phasianidae. Cuturnix,
Quat. Aust. N.Z., Rec.

ORDER: GRUIFORMES

SUBORDER: TURNICES
Turnicidae, Turnix, Picist, Aust., N.
Caledonia, Rec.

SUBORDER: GRUES
Gruidae. Genus indet., L. Olig./M. Mio.
S.A.; Grus, L. Plio.-Quat. $.A., ne.
Aust. Rec. Rallidae. Genera indet., L.
Olig./M. Mio.-Quat. S.A.; Capellirallus,
Qual. N.Z.; Porphyrio, Quat. N.
Caledonia, N.Z., Rec.; Diaphorapteryx,
Quat. Chatham Is.; Fulica, Plio, Aust.,
Quat. Aust. Chatham Is., N.Z., Rec.;
Gallirallus, Quat. Aust. Chatham Is., N.
Caledonia, N.Z., Rec.; Gallinula, Quat.
Aust, N.Z., Rec.; Rallus, Quat. Aust.
N.Z., Rec.; Porzana, Qual. Aust., N.
Caledonia, Rec.; Tricholimnas, Quat. N.
Caledonia. Aptornithidae. Aptornis,
Quat. N.Z. Rhinochetidae.
Rhinochetos, Quat. N, Caledonia, Rec.

SUBORDER: OTIDES
Otididae, Genus indet., L. Plio. S.A,;

Ardevtis, Pleist. S.A., Rec.

ORDER; CHARADRIIFORMES
SUBORDER: CHARADRII

Charadriidae. Anarhynchus, Quat. N.Z.,
Rec.; Charadrius, Quat. N. Caledonia,
N.Z., Rec.; Calidris, Quat.-Pleist. W.A.,
S.A., Rec.;Erythrogonys, Quat. Vic.,
Rec.; Peltohyas, Quat. §.A., Rec.;
Pluvius, Quat. N, Caledonia, Rec.;
Thinornis, Quat. N.Z., Rec.; Vanellus,
Quat. Aust., Rec. Haematopodidae.
Haematopus, Quat. N.Z., Rec.
Scolopacidae. Arenaria, Quat. N.Z.,
Rec.; Coenocorypha, Quat. Chatham Is.,
N. Caledonia, Rec.; Calidris, Quat. NZ.
S.A., Rec.; Gallinago, Quat. S.A.,
Kangaroo Is., Rec.; Limosa, Quat. NZ.,
Rec.; Numenius, E.-M. Plio. Qd., Quat.
N.Z., Rec. Tringa, Quat. 8.A., Rec.;
Burhinidae. Genus indet., L. Olig./M.
Mio. S.A.; Burhinus, Quat. S.A., Rec.
Pedionomidae. Pedionomus, L. Plio.
or Quat,, Vic., Rec. Recurvirostridae.
Himantopus, Quat. N.Z., Rec.

SUBORDER: LARI
Laridae. Larus, Quat. Aust. N.Z., Rec.;
Hydroprogne, Quat. N.Z., Rec.; Sterna,
Quat. Aust. N.Z., Rec. Stercorariidae.
Stercorarius, Quat, N.Z., Rec.

ORDER: COLUMBIDIFORMES
SUBORDER: COLUMBAE
Columbidae. Genus indet., L. Olig./M.
Mio. S.A.; Caloenas, Quat. N. Caledonia,

Rec.; Chalcophaps, Quat. N. Caledonia,
Rec.; Columba, Quat. N. Caledonia, Rec.;
Drepanoptila, Quat. N. Caledonia, Rec.;
Ducula, Quat. N. Caledonia, Rec.;
Gallicolumba, Quat. N. Caledonia, Rec.;
Hemiphaga, Quat. N.Z., Rec.;
Leucosarcia, Quat. Aust., Rec.; Ocyphaps,
Quat. Aust., Rec.; Phaps, Pleist. Qd.,
S.A., Rec.; Ptilinvpus, Quat. Aust., Rec.;
?Cyanorhamphus, Qual. N.Z., Rec.

ORDER: PSITTACIFORMES
Cacatuidae. Genus indet., L. Olig./M.
Mio. Qd.; Calyptorhynchus, Quat. Aust,
Rec.; Callocephalon, Quat. Aust., Rec.;
Cacatua, Pleist. Aust., Rec. Loriidae.
Glossopsitta, Quat. Aust., Rec.;
Trichoglossus, Quat. Aust., Rec.
Platyceridae. Cyanorhamphus, Qual.
N.Z., Rec.; Psephotus, Quat. Aust., Rec.;
Melopsiltacus, Quat. Aust., Rec.;
Geopsittacus, Quat. Aust., Rec.;
Pezoporus, Quat. Aust., Rec.; Alisterus,
Quat. Aust., Rec.; Polytelis, Quat. Aust.,
Rec.; Lathamus, Quat. Aust., Rec.;
Purpureicephalus, Quat. Aust., Rec,;
Platycercus, Quat. Aust., Rec.; Barnardius,
Quat. Aust., Rec.; Northiella, Quat. Aust,,
Rec.; Neophema, Quat. Aust., Rec.
Nestoridae. Nestor, Qual. N.Z., Rec.,
Strigops, Quat. N.Z., Rec.

ORDER: CUCULIFORMES

SUBORDER: CUCULI
Cuculidae. Centropus, Quat. Aust.,
Rec.; Cuculus, Quat. Aust., Rec.;
Chrysococcyx, Quat. Aust., Rec.,
Chatham Is., Eudynamys, Quat. N.Z.,
Rec.; Urodynamis, Quat. N. Caledonia,
Rec.

SYSTEMATIC INDEX -

ORDER: STRIGIFORMES
Tytonidae. Tyto, Quat. Aust. N.Z., N.
Caledonia, Rec. Strigidae. Ninox,
Quat. Aust., N. Caledonia, N.Z., Rec.
Family uncertain. Sceloglaux, Quat.
N.Z., Rec.

ORDER: CAPRIMULGIFORMES

SUBORDER: CAPRIMULGI
Caprimulgidae. Caprimulgus, Quat.
Aust. N. Caledonia, Rec. Podargidae.
Podargus, Quat. W.A., Vic., Rec.
Aegothelidae. Aegotheles, Quat.
Aust., N. Caledonia, Rec.;
Megaegotheles, Quat. N.Z.; Quipoilornis,
Mio. N.S.W.

ORDER: APODIFORMES
SUBORDER: APODI
Apodidae. Hirundapus, Quat. Aust.,
Rec.; Collocalia, Quat. Aust., N.
Caledonia, Rec.

ORDER: CORACIIFORMES
SUBORDER: ALCEDINES
Alcedinidae. Ceyx, Quat. Aust., Rec.;
Halcyon, Quat. Aust., N. Caledonia, N.Z.,
Rec.; Dacelo, Quat. Aust, Rec.

1418

ORDER: PASSERIFORMES
Family uncertain. Genera indet., L.
Olig/M. Mio. S.A., Qd., L. Plio. S.A.
Menuridae. Genera indet., L. Olig./M.
Mio. Qd. Atrichornithidae.
Atrichornis, Quat. Aust, Rec.
Hirundinidae. Hirundo, Quat. Aust.,
Rec.; Cecropis, Quat. Aust., Rec.
Motacillidae. Anthus, Quat. Aust,
Rec. Muscicapidae. Bowdleria, Quat.
N.Z., Rec; Gerygone, Quat. N.Z., Rec.;
Mohoua, Quat. N.Z., Rec.; Oreoica, Quat.
Aust., Rec.; Pachycephala, Quat. Aust.,
Rec.; Petroica, Quat. Aust. N.Z., Rec.;
Rhipidura, Quat. N.Z., Rec.

Orthonychidae. Orthonyx, L. Olig./M.

Mio. Qd, Quat. Aust., Rec.; Cinclosoma,
Quat. Aust., Rec.; Psophodes, Quat.
Aust., Rec.; Sphenostoma, Quat. Aust.,
Rec. Timaliidae. Pomatostomus,
Quat. Aust., Rec. Sylviidae.
Cincloramphus, Quat. Aust., Rec.;
Megalurus, Quat. Aust, Rec.
Acanthisittidae. Acanthisitta, Quat.
N.Z., Rec.; Pachyplichas, Quat. N.Z.,
Rec.; Xenicus, Quat. N.Z., Rec.;
Traversia, Quat. N.Z., Rec. Maluridae.
Amytornis, Quat. Aust., Rec.; Anthus,

Quat. N.Z., Rec.; Malurus, Quat. Aust.,
Rec.; Stipiturus, Quat. Aust., Rec.
Acanthizidae. Acanthiza, Quat. Aust.,
Rec.; Daphoenositta, Quat. Aust., Rec.;
Pycnoptilus, Quat. Aust., Rec.;
Dasyornis, Quat. Aust., Rec.
Climacteridae. Climacteris, Quat.
Aust., Rec. Pardalotidae. Pardalotus,
Quat. Aust., Rec. Neosittidae.
Neositta, Quat. Aust., Rec.
Meliphagidae. Anthornis, Quat. N.Z.,
Rec.; Melithreptus, Quat. Aust., Rec.;
Notiomystis, Quat. N.Z., Rec.;
Prosthemadera, Quat. N.Z., Rec.
Ploceidae. Poephila, Quat. Aust, Rec.
Ptilonorhynchidae. Ptilonorhynchus,
Quat. Aust., Rec. Paradisaediae.
Turnagra, Quat. N.Z., Rec. Callaeidae.
Callaeas, Quat. N.Z., Rec.; Heterolocha,
Quat. N.Z.; Philesturnus, Quat. N.Z.,
Rec. Artamidae. Artamus, Quat. Aust.
Rec. Grallinidae. Grallina, Quat.
Aust, Rec. Cracticidae. Strepera,
Quat. Aust., Rec.; Gymnorhina, Quat.
Aust. Kec. Corvidae. Corvus, Quat.
Kangaroo Is. Aust., Rec.; Palaeocorax,
Quat. N.Z.

Abattoirs Bore, 517

Abderites meridonalis, 908

Aberfoyle, 372

Aboriginal legends, 2

Aboriginal middens, 304, 1348

Aborigines, 238, 820

abrasion facets, 233

abrasion, 273

abstracting joumal, 161

Acacia, 136, 140, 820, 822

acacias, 259

Acanthisitta, 1333

Acanthisittidae, 1320

Acanthiza chrysorrhoa, 840

Acanthizidae, 840

Acanthodes australis, 351, 453, 455

Acanthodes bridgei, 351

Acaathodes, 349, 352, 401, 443,
451, 452

acanthodians, 431, 432, 442-444
446, 448, 450, 451, 453, 454

Acanthodida, 350

acanthodids, 443

Acanthodii, 345

Acanthoides dublinensis, 453

acanthothoracids, 354, 367

Accipiter cirrhocephalus, 829

Accipiter fasciatus, 829

Accipiter, 1329

Accipitridae, 757, 829

accumulating agents, 271

accumulator species, 283

acellular bone, 344, 499

Acerastea wadeae, 579, 580

acetabular fossa, 596

acetate, 182

acetic acid, 174, 176, 182-186,
317, 327

acid etching, 174-188

acid neutralization, 181

acid tubs, 180

acidic environment, 272

Acidotheres, 823

Acinonyx jubatus, 304

Acrifix 92,187

Acrobates, 956

Acrobatidae, 956

acrolepid scales, 372

INDEX
Acrolepis, 586 allantois, 606
Actinistia, 368, 369, 386 Allaru Formation, 627

actinolepids, 355, 432, 444, 454

Actinopterygii, 368, 369

Aculeola, 499

Adamson, M., 11

adaptive radiation, 932

Adasaurus, 642

Adavale Basin, 120, 121

Adelaide Geos yncline, 116

Adelotus brevis, 599

Adelotus, 593

Aegotheles cristatus, 281, 761, 834

Aegotheles, 761, 835

aegothelid, 732, 811

Aegothelidae, 739, 761, 767, 834

Aepyornis, 881, 882

Aerodramus spodiopygius, 231

AESIS, 151, 153, 161, 162, 169

Aetosauna, 637

agamids, 673

Agassizodus, 442

age classes, 281

Agecephalichthys , 375

Agelcodus pectinatus, 347

agelcodonts, 441

Ageleodus, 441, 451,455

Agrosaurus macgillivrayi, 611, 644

Agrosaurus, 647, 663

Ailuropoda melanoleuca, 261

Ailuropoda, 215, 257

Ailurops ursinus, 944

air-scribe, 189, 190

Aiyenu Cave, 834

akinetic skull, 628

Alauda, 823

Albatross Island, 1349

albumin, 313, 314, 330, 332, 929

Alcedinidae, 836

Alcoou Fauna, 929

Alcoota Local Fauna, 668, 1037

Alcoota Station, 54, 66, 76, 90, 142

Alcoota, 329, 728, 732, 742-744,
756, 757, 781, 782, 1079

Aldabra Atoll, 1363

Aldinga Bay, 512

Alehvale, 670

Alisterus scapularis, 836

Allaru Mudstone, 504, 596, 507,
613, 645

Allaru Sandstone, 617

Allingham Creek, 688, 730, 752,
757

Allingham Formation, 669, 729

allochthonous assemblage, 269,
270, 271

allochthonous deposit, 273

allochthonous species, 269, 283

allopatric speciation, 1172

allopatry, 1166

Allopleuron, 713

Allosaurus fragilis, 647

Allosaurus, 612, 639, 646, 648, 664

Alphadon lulli, 904

Amadeus Basin, 117, 119, 389, 398,

434, 436, 437, 452
American Geological Institute,
160, 168
Amenidelphia, 925
amino acids, 316-318, 326
ammonia, 181
ammonites, 623
amnion, 606
amniote egg, 606
Amphiaspid Province, 400
Amphibolurus, 667, 670, 673
amphistylic, 346
Amphitheatre Cave, 290, 291-293,
810, 821, 825, 831, 837, 840,
Amphitheatre Group, 399, 447
Amsterdam Island, 1363
Amytornis textilis, 822, 839, 839
Anachronistes, 441
anacoracids, 500
analogous, 203
anamniote, 570
anapsid, 606, 607, 609
Anarhynchus, 133}
Anarosaurus, 630
Anas castanea, 828
Anas superciliosa, 828
Anas, 823
Anaspida, 343
anaspids, 431

INDEX - 1419

Anatidae, 739, 757, 768, 828

Andamooka, 633, 646

Anderson Formation, 122

Anderson, C.A., 46, 86

Andreolepis, 444

angular fossa, 224

angular process, 224

Anhinga latipes, 827

Anhinga novaehollandiae, 827

Anhinga parva, 827

Anhinga, 811

Anhingidae, 751, 827

anhydrite, 117, 122

Ankotarinja, 927

ankylosaur, 648-650, 651, 655

Anomalopteryx didiformis, 1325

Anomalopteryx oweni, 1325

Anomotodon, 507, 524

Anseriformes, 1329, 1359

Ansett, 735

Aatarctaspis, 361

Antarctic Circle, 650, 663, 664

Antarctic Peninsula, 624, 665,1175,
1177

Antarctica, 361, 362

Amtarctilamaa prisca, 346, 347, 438

Aatarctilamna, 401, 440, 448, 455

Antarctodolops, 615, 911

Aatechinus, 289

anterior cingulum, 224, 233

Aathornis, 1333

Anthracosauria, 570

anthracosauroids, 686

Anthropodytes gilli, 748, 1176

Anthropornis nordenskjoeldi, 746,
TAT, 808

Anthropornis, 615, 738, 748

Anthus novaeseelandiae, 46, 308

anti-collagen antibody, 331

antiarchs, 367

antigens (blood), 313

Aanligonia fornicata, 384

anusera, 320

antorbital fenestra, 648

ants, 238

Anura, 570, 592

Aaurognathus, 658

ANZAAS, 37, 63

1420 - INDEX

Apateolepis hamilioni, 374

Apateolepis tasmanicus, 374

apatite, 272

Apatosaurus, 642

Aplonis fusca, 1374

Apoda, 570

Apodidae, 761

aponcuroses, 651

Aptenodytes forsteri, 747, 1357

Aptenodytes patagonicus,| 326,
1360

Aptenodytes ridgeni, 1177

Apterorms (Apteromithidac), 1331

Apterygidae, 1320

Apterygiformes, 1326

Aptornis otidiformis, 1331

Aqlaocetus, 1183

Aquadere, 188

aquatic invertebrate cater, 258

Aquila audax, 287,757, 829

Aquila, 1329

aracoscelidians, 609

Araeoscelis, 607

aragonite, 873, 881

Araluen, 452

Aramac, 506

Arandaspidi formes, 343, 344

Arandaspis prionotolepis, 444, 446

Arandaspis, 343, 433, 455

arboreal animals, 256

arboreal habits, 257

Arcadia Formation 374, 470, 471,
490, 572, 574, 576-580, 583,
587, 611

Arcadia myriadens, 579, 580, 585

archacocetes, 1184

Archacoceti, 1171, 1179

Archaeochelydium, 616

Archaeocycnus lacastris, 750, 783

Archaeomene, 376

Archaaopteryx, 608

Archasospheniscus, 615, 748

Archer, M., 33, 34, 37, 61, 73, 75,
77, 79, 723

Archosauria, 636

archosauromorphs, 609, 623, 686

archosaurs, 609, 639, 655, 686

Arckaringa Basin, 120

Arctic Canada, 450

Ardeidac, 751, 827

Ardeotis australis, 760, 831

Arenipiscis westolli, 354, 356, 357

Arenoph.yne, 593, 596

Arganodus tiguidensis, 470, 484,
490

Arganodus, 472

Argyrolagidac, 911, 941

Argyrolagus scaglii, 912

arid adapted, 911

aridity, 136

Arlec Super Tool, 189

armadillos, 901

Armonca, 400

Ambhem Land, 1090, 1107, 1109

armnval of man, 1319

Artamidac, 841

Artamus cinereus, 841

Artamus cyanopterus, 841

Artamus leucorhyachus, 841, 842

Artamus minor, 341

Artamus personatus, 84)

Artamus superciliosas, 841

Artamus, 296

arthrodires, 354, 355

artiodactyls, 208, 214

Asaphus, 11

Ascot Beds, 506, 516

Ascot Formation, 517

Ashfield Shale, 374, 470

Asiatoceratodus tiguidensis, 484

Assa darlingtoni, 599

Assa, 593

asteroid impact, 1169

asteroid, 1175

asterolepidoid, 364, 455, 459

Asierolepis, 363, 400, 402

astragalo-calcaneal joint, 1111

Astroconger rostratus, 384

Astroconger, 382

Athertonia, 376

Atherurus, 286

Atlantidosteus, 355

Atlas Copco, 735

Atlascopcosaurus loadsi, 612, 688

Allascopcosamrus, 650

atmospheric carbon dioxide, 1169

Atrichornis rufescens, 819, 837,
838

Atrichornis, 296

Atnichomithidac, 767, 837

attrition facets, 233

attrition striac, 234, 240

altritional mortality, 281

Aupouri Peninsula, 1323

Aurora Cave, 1360

Austalolepis seddoni, 455

AUSTRADOC, 169

Australasian avifauna, 766

Australasian biota, 763

Australian Airlines, 735

Australian Army, 37, 70, 79

Australian Museum, 8, 13, 24, 28,
31, 86, 151, 340

Australian Museum Society, 735

Australian National University,
340, 723

Australian Research Council, 735

Australian Research Grants

Scheme, 76, 80, 81,735

AUSTRALIS, 169

Australobatrachus iliss, 596

Australobatrachus, 666

Australodelphia, 925, 929

Australolepis seddoni, 436, 447

Australolepis, 344, 434, 439

Austrobrachyops, 614

Austrolepis seddoni, 582

Austropelor wadleyi, 580, 586

Austrophyllolepis ritchiei, 361, 362

Austrophyllolepis, 316, 358, 401

Austrosamurus mckillopi, 613, 645

Austrosaurus, 612

autochthonous assemblage, 269,
271, 281

autochthonous deposit, 272

autochthonous species, 268, 281, 283

autoradiography, 328
autostylic, 391

avian camivores, 287

avian predators, 284

avian scavengers, 284
avian-accumulated deposits, 284
Avimimus, 647

Avon River Group, 364
Awa, 729

Awe Fauna, 55, 92, 98, 1044
Awe, 730, 746

Aye-Aye, 261

Ayers Rock Arkose, 116
Aythya novaeseelandiae, 1320
Aythya, 1329

Aztec Siltstone, 400

Babbage Peninsula, 473
Backstairs Passage, 1349
bacterial fermentation, 237
Baffin Island, 317

Bahamas, 1363

Bailly J.C.,3

Baird, R.F., 69, 723, 810
Balaena, 1180
Balaenoptera, 1180
Balbarinac, 951

Balcombe Bay, 340-342, 510
Baldina Creek, 329
Balgowan, 660

Baliem Valley, 1100
bamboo, 215, 257

banded iron formation (BIF),113
bandicoot, 230, 236-238
Bandringia, 441

Banks, J., 12
Baragwanathia, 447
Barambag Limestone, 452
Barameda decipiens, 389
Barameda, 388
Barawertornis tedfordi, 782
Barawertornis, 743
Barcaldine, 506

Barghoom, S., 80
Barnardius zonarious, 836
barrier, 11168, 172

Barrow Creek, 231
Bartholomai, A., 34, 73, 76
Baru, 1077, 1079

Barwick, R.E., 340, 388
basal metabolic rate, 208
Basalla, G.,2

Basilosaurus, 615

Bass Basin, 128, 129

Bass Strait islands, 722, 1346
Bass Strait, 760, 1348
Bassian Depression, 1346
Baston River Beds, 451
Batesford Limestone, 513
Batesford Quarry, 502, 1036, 1183
Batesford, 340-342, 515
Bathurst Island Formation, 611
batoids, 498, 502

Baudin, N., 2, 150

bay bars, 276

Baynes, A., 77, 1350

bears, 214

Beaumaris Local Fauna, 1037
Beaumaris nodule bed, 510

Beaumaris, 340-342, 382, 383, 516
518, 728, 748, 1179

beavers, 258

Bedacryl, 187,430

Beelarongia patrichae, 421

Beginners Luck Cave, 826

Belanorhynchus, 583

belemnites, 506

Belgian Basin, 507, 509

Belichthys, 376

Belonstomus, 382

Belvedere Formation, 434, 447,
459

Bematherium, 1079

Bernardius arnuxii, 1182

Bergmann's Rule, 664

Bermuda, 1363

Bernier Island, 1347, 1350

Berowra Creek, 659

Best, J.G., 90

Betatex, 197

Bettongia, 251, 955

bias in fossil record, 268

bibliographic services, 157

bibliographies, 156, 160

Big Sink Local Fauna, 1040

Billa Kalina Basin, 140

Billeroo Creek, 475, 483

bilophodont 209, 218, 222

bilophodont molars, 946

bilophodonty, 1094

Bindaree Road Fauna, 398

Bingara, 223,745

Bingleburra Formation, 451

binocular vision, 259

biocenose, 269

biogeographic

biogeography, 1170, 1363

biohermal limestone, 118

bioherms, 117

biamass, 257, 1133

biomass, 257

biostratigraphic subdivisions, 1323

biostromal limestone, 118

biotic events, 1167

Bird Rock, 511

Birregurra, 514

Bishanopliosaurus youngi, 633

Biziura dalautouri, 1329

Biziura lobata, 828, 1329

Black Rock Sandstone, 509, 516-
518

black shales, 338

Blackstone Formation, 611

Black water Shale, 372

Blackwater shales, 345, 347, 371

Black water, 340-342, 452

Blanche Point Formation, 738, 746

Blanche Point Mazi, 511

Blanche Point, 746

Blanchetown Clay, 483, 484, 668,
815

Blanchtown, 515

Blandowski, W., 13

bleach, 431

Blina Shale, 374, 572, 573, 578,
586, 611, 628, 470

Blina, 470-472

Blinasaurus townrowi, 574
Blinasaurus, 576
Blitzkreig effect, 1142
Blue Range Formation, 395, 398
Bluff Downs Local Fauna, 669,
942, 959, 1041
Bluff Downs, 729, 933
Boat Mountain Local Fauna, 669
Boat Mountain, 669, 729
Bob's Boulders Locality, 479
Bob’s Boulders, 481
bobasatranid, 371
Bockhara Limestone, 514
body size, 1133
body weight, 1075, 1129
Bogantungan region, 351
Bogon Gate, 447
Boguntungan, 370
Bohra paulae, 1112
Bokhara Limestone, 513
Boles, W., 723, 726
Bonaparte Basin, 358
Bonaparte Gulf Basin, 120
Bonaparte Gulf, 116, 451
bone beds, 276
bane fracture, 285
Bone Gulch Fauna, 479
Bone Gulch Local Fauna, 483, 668,
1054
Bone Gulch, 917
Bookpumong Beds, 515
Boongerooda Greensand, 349, 510
Boreopricea, 635
Borhyaemdae, 910, 929,
borhyaenids, 907, 925
Borogovia, 647
Bos taurus, 303
Bas, 969
Botaurus poiciloptilus, 1327
Bothrioceps australis, 583
Bothriolepis billilunensis, 364
Bothriolepis bindareei, 364, 365,
398
Bothriolepis canadensis, 364
Bothriolepis cullodensis, 336, 364,
365, 398
Bothriolepis fergusioni, 364
Bothriolepis gippslandiensis, 362,
363, 364, 398, 420
Bothriolepis tatongensis, 64, 398
Bothriolepis warreni, 398
Bothriolepis, 345, 358, 360, 363,
364, 397, 400-402, 442, 582
Boulia, 617, 618
Bow Local Fauna, 668, 1049
Bow, 730
Bow-wave effect, 1142
Bowdleria punctata, 1333
Bowen Basin, 122-124
Bowen Island, 1347

Boyd Volcanic Complex, 351

Brachiones, 204

Brachiosaurus brancai, 645

brachycephalic skull, 219, 1114,
1120

brachycephaly, 219

Brachyopidae, 574

brachythoracids, 355

bradyodont tooth, 347

bradyodonts, 346

Braidwood, 340-242, 360

Breccia Cave, 6,7

Bremer Basin, 511

Brigalow, 386

Brindabellaspis stensioi, 366, 367

Brisbane River, 482, 484, 490

British Museum (Natural History),
34, 35

brittle material, 206

Brogniart, A., 4, 149, 150

Broken River Formation, 434, 438,
452

Broken River, 340-342, 347, 363,
401, 434, 449, 450

Brontosaurus, 642

Brookvale Fauna, 374

Brookvale, 340-342, 375, 468, 470,
583

Brookvalia gracilis, 377, 378

Brookvalia spinosa, 377

Brookvalia, 376

Broom, R., 31

Broome Sandstone, 611, 660

Brothers Island, 329, 1347, 1350

Brown, D., 73

Brown, H. Y, L., 24, 26, 27, 33,
7122

Brown, R., 12

Browns Creek, 738, 746

browsers, 208, 249

browsing macropodines, 217

browsing niche, 257

Brunhes, 814

Bruntonichthys, 355

buccal cusps, 235

buccal pump, 368

buccal tier, 232, 234, 241

Buchan limestones, 448

Buchan, 303, 340-342, 351, 352,
391, 450, 452, 466, 745

buchanosteid, 459

Buchanosteus confertituberculatus,
353, 354, 357, 420

Buchanosteus, 356, 450, 459

Buckalow Bore, 515

Buckland, W., 3, 4,6, 11

Bugaldi, 729, 732, 761

Bulga, 449

Bulgoo Station, 340-342

Bulldog Shale, 347, 506

Bullerichthys, 355

Bullock Creek Fauna, 57,79, 1035

Bullock Creek Local Fauna, 481,
668

Bullock Creek, 142, 174, 178, 180,
182, 326, 327, 330, 331, 479,
483, 727, 729, 732, 736, 737,
740, 742-744, 756, 762, 781,
782, 1079

Bullockornis planei,781, 782

Bullockornis, 743

Bulmer, S., 90

Bulolo, 86, 92

Bulolo-Watut, 90

Bulungamayinae, 948

bulungamayines, 895

Bunda Plateau, 288, 817, 820, 822,
838, 839, 841

Bunga Beds, 345, 317, 351, 448

Bunga Creek, 1044

Bungil Formation, 612

Bunock Creek Group, 452

bunodont teeth, 213-215, 249, 261

bunodont, 947, 948, 956

bunolophodont molars, 257

Bunyip, 2

Bureau of Mineral Resources, 54,
340, 726

Burhinidae, 739, 760, 768, 831

Burhinus grallaria, 752, 754,760

Burhinus magnirostris, 754, 756,
831

Burke, A., 80

Burkes Cave, 304

Burnett River, 484

Burra Baldina Creek, 745

Burramyidae, 953-955

Burramyoidea, 953

Burramys parvus, 214, 258, 953,
954

Burramys wakefieldi, 258

Burrinjucosteus asymmetricus, 354

butcherbirds, 286

butchering practices, 304

Butler, H., 73

Butvar , 187, 196

Byro Group, 374

Cacatua rosicapilla, 822

Cacatua tenuirostris, 819, 835

Cacatuidae, 762, 835

caecotrophic, 237

caecum, 203-205, 209, 231, 234,
235, 237, 2A0

Caenolestes, 08

caenolestids, 924

Cahier, H., 161

Calaby, J., 766

calcaneo-cuboid joint, 1111

calcification, 273

calcified cartilage, 345, 499

calcite, 272, 873

calcium acetate solution, 430

Calidris ruficollis, 832

California Sea Otter, 261

Callaeas cinerea, 1334

Callaeatidae, 1320

Callen, R., 73

Callide Basin, 659

Callocephalon fimbriatum, 835

Callorhinchus milii, 503

Calonectris, 1327

calvarium, 472, 477

Calyptorhycnhus funereus, 835

Calyptorhynchus lathami, 822, 835

Cambelltown, 376

camels, 284

Cameron Iniet Formation, 516, 517

Camfield Beds, 479, 481, 483, 668

Camp, C.L., 58, 154

Campbell, C.,73, 74

Campbell, K. S., 33, 76, 340, 387

INDEX - 1421

Campbellodus decipiens, 366

Campbellodus, 367

Camptosaurus, 650, 651, 657

camuropiscids, 355

Camuropiscis concinaus, 359

Camuropiscis, 355

Canadian Lead, 1036

Canidae, 963

canine tusk, 621

Canis familiaris dingo, 287, 820,
963

Canis familiaris, 211, 303

Canis lupus, 305, 678

Canis, 965

Canning Basin, 117, 120-126 , 356,
364, 367, 399, 434

Canowindra grossi, 386, 387

Canowindra, 340-342

Cape Barren Island, 1347

Cape Grim beds, 513, 514

Cape Grim, 515

Cape Hillsborough Formation, 706

Cape Naturaliste, 598

Cape Otway, 688

Cape Paterson, 472, 473, 490, 624,

Cape Range Group, 513, 515

Cape River, 671

Cape Wanbrow, 1321

Cape York Peninsula, 3, 645, 944

Capellirallus karamu, 1330

Capellirallus, 1330

Capitosauridae, 574

Caprimulgidae, 761

Caprimulgiformes, 767, 1332

captorhinids, 607

captorhinomorphs, 607

Carapook, 340-342

carbon dioxide, 1169

carbonization, 273

Carcharhinus brachyurus, 502,
503, 515, 517, 530

Carcharhinus plumbeus, 503, 530

Carcharias acutissima, 511, 526

Carcharias macrotus, 510, 512,
524, 526

Carcharias tarus, 501-503, 514,
516, 518, 524, 526, 527

Carcharius marcrotus, 511,514

Carcharodon anguistidens, 515,
529

Carcharodon auriculatus, 515

Carcharodon carcharias, 517, 529

Carcharodon megalodon, 515, 517,
529

Carcharodon, 501, 530

Carcharoides tenuidens, 512

Carcharoides totuserratus, 512,
530

Carcharoids catticus, 512

Cardabia Group, 510

Cardabia Station, 509

Carduelis, 823

Carettochelys, 668

caribou, 665

Carl Creek Limestone, 55, 669,
1079

Carmichael Sandstone, 446

Camarvon Basin, 118, 119, 122-

1422 - INDEX

124, 126, 340-342, 358, 364,
374, 434, 509, 510, 513

Camarvon Gorge, 659

camassial, 255, 943

Carnivora, 963, 1171, 1178

camivore accumulators, 286

carnivores, 209, 248

carnivorous marsupials, 679

Carpentaria Basin, 124, 126

carpophagous, 258

Carretochelyidae, 708

Carribuddy Formation, 119, 399

cartilaginous endoskeletan, 499

Case, J., 80

caseodonts, 451

casichelydians, 617

cassias, 259

cassowary, 327

casting, 193, 198

casts, 193

Casuariidae, 739, 743, 766, 768,
824, 1086

Casuarina, 259, 319, 814, 1140

Casuarius bennetti, 746

Casuarius casuarius, 746, 878

Casuarius lydekkeri, 97, 98,745,
782

Casuarius unappendiculatus, 744,
7146

Casuarius, 94

catastrophic assemblage, 281

catastrophic explanations, 1168

catastrophic mortality, 281, 1133

Cathedral Cave, 329

Catomal Group, 452

cattle, 248, 250

Cave Bay Cave, 304, 817

Cave Bay, 1349

cave-dwelling species, 281

cave formation , 277

cave sediments, 1321

CD-ROMS, 169

Cecropsis ariel, 281, 838

Cedratodus palmeri, 483

Cedratodus wollastoni, 490

Celebes, 944

cellulose, 207

Central regional fauna, 1127

centres of origin, 1320

centrocoels, 628

Centrophoroides, 508, 52A

Centrophorus, 510

Centropus collassus, 819, 836, 867

Cephalaspid Province, 400

Cephalaspidomorphi, 343

cephalaspids, 431

cephalopods, 627

ceratodontid, 665

ceratodonts, 472, 478, 485

Ceratodus (Tellerodus) formosus,
468, 470

Ceratodus (Tellerodus) sturii, 471

Ceratedus avus, 470, 472, 473, 477,
484, 485

Ceratodus concinnus, 472

Ceratodus dorotheae, 484

Ceratodus formosus, 471, 477, 484,
490, 583

Ceratodus frazieri, 476

Ceratodus gypsatus, 472

Ceratodus kaupi, 476, 490

Ceratodus laticeps, 472

Ceratodus latissimus, 484

Ceratodus nargun, 476, 477

Ceratodus palmeri, 475, 484, 490

Ceratodus phillipsi, 470, 472, 490,
585

Ceratodus protopteroides, 486

Ceratodus szechuanensis, 484

Ceratodus tiguidensis, 470, 472,
490

Ceratodus wollastoni, 473-478,
480, 484

Ceratodus youngi, 484

Ceratodus, 586

ceratopsians, 648

Cerberean Cauldron, 397

Cereberan Volcanics, 398

Cercartetus caudatus, 258

Cercatetus nanus, 1303, 349

cercopithecids, 209

cerebral hemispheres, 932

Cervus, 763

Cestracion cainozoicus, 526

Cestracion longidens, 526

Cetacea, 1167, 1171, 1176, 1179

Cetacea, 1179

cetaceans, 631, 1086

cetiosaurs, 648

Cetotheriidae, 1180

Cetotolites, 1181

Ceyx azureus, 836

Chaeropus ecaudatus, 230, 238,
2A0, 241

Chaeropus, 221, 232, 234-240

Chalcophaps indica, 1363, 1374

Chambers Gorge, 308

Changpeipus bartholomaii, 611,
660

channel lag, 276

Chapman, F., 18, 151

Charadriidae, 760, 832

Charadriiformes, 760, 1331

Charadrius bicinctus, 1374

Charlotte Range, 446

charophyte algae, 585

Charters Towers, 76

Chasmatosaurus, 640

Chatham Islands, 1319, 1321, 1330-
1332, 1355, 1356, 1358, 1359,
1362, 1363

Chathams, 1350

Cheiracanthoides comptus, 451,
453

Cheiracanthoides, 351, 401, 420,
443, 455, 458, 459

Cheiracanthus, 443, 453

Cheirolepis, 370, 444

Chelidae, 704, 708

Chelodina, 670, 705

Chelonia mydas, 617

Chelonia, 1171

chelonian eggshell, 873

Chelonioidea, 708

Chelycarapookus arcuatus, 612,
617, 619, 620, 715

Chelycarapookus, 714
chemical weathering, 730
chenopods, 814

chicken, 327

Chillagoe Shelf, 119

Chiltern Hills, 474, 474
chimaerid, 349

chimaerids, 346, 349, 367
chimaeroid, 441, 451, 498, 506,

Chinchilla Fauna, 483, 933

Chinchilla Local Fauna, 1046, 1081

Chinchilla Sands, 479, 483, 490,
669, 819

Chinchilla, 329, 484, 730, 745, 752,
758, 760, 831,

Chionidae, 831

Chionis, 811

Chirodipterus australis, 393, 395,
421, 467

Chironectes minimus, 258

Chiroptera, 958

chitinous skeleton, 249

Chlamydia, 1094

Chlamydosaurus, 670, 673

Chlamydoselachus, 499, 500

choana, 445

Choanata, 369

Chomatobatrachus halei, 578

chondrichthyans, 440, 498

Chondrichthyes, 343, 345, 346, 352

chorion, 606

Chosornis praeteritus, 758

Chowilla dam, 69

Chowilla Fauna, 479, 483

Chowilla, 813

chromosome sturcture, 930

Chronozoon australe, 1178

Chrysococcyx lucidus, 1836, 332

Chrysophrys, 382

Chunia illuminata, 253, 255, 261

Chunia omega, 254

Chunia, 259, 945, 946

Ciconia nana, 752.

Ciconidae, 752, 827

Ciconiiformes, 1327

cimoliasaurs, 628

Cimoliasaurus maccoyi, 612, 1173

Cimoliasaurus planus, 631

Cimoliasaurus, 634

Cinclorahmphus cruralis, 306, 838,
839

Cinclorhamphus mathewsi,839

Ciaclosoma alesteri, 838, 842

Cinclosoma punctatum, 838

Cinclosoma cinnamomeum, 838

Circum Antarctic Current, 1169,
1185

Circus approximans, 1330

Circus eylesi, 1329

Cladocyclus sweeti, 382

cladodont sharks, 440, 449, 451,
458

Cladodus, 440

Cladorhynchus leucocephalus,

740, 752, 754, 756
Cladoselache, 440
cladoselachians, 440
Clarafield St. Kynuna, 473
Clarafield Station, 476
Clark, J.M., 641
Clarke, A., 13
Clarke, W. B., 8, 22, 722
claspers, 367, 440
classification (chondrichthyans),

500
classification (chelonian), 704
classification (fish), 414-419
classification (mammalian), 1083
classification, 166,168
claudiosaurs, 609
Claudiosaurus, 629, 630
Clayton River, 882, 888
cleithrolepids, 583
Cleithrolepis granulata, 376, 378
Cleithrolepis, 586
Clemens, W., 74, 74
Clift, W., 6
Clifton Formation, 511
Climacteridae, 281, 840
climate change, 768, 1171, 1349
climatic cooling, 907
climatic fluctuation, 1319
climatiid, 351, 443, 451, 453
Climatiida, 350
climatioids, 443
Climatius, 443
Clissold, H., 90
Clogg’s Cave, 291, 293, 295, 305,

819-820, 829, 830, 832 , 834,

836, 837, 840, 841
Cloughnan Shale, 393, 573, 576,
Cluan Formation, 572, 573, 586
Cnemiornis calcitrans, 1329
Cnemiornis gracilis, 1321, 1329
coal, 3, 120, 122, 124, 128
coarse shear, 236
coastal dunes, 276
coastal islands, 825
Cobar, 397, 434
Coccolepis australis, 316
Coccolepis woodwardi, 378, 380
coccosteomorph, 355
cochliodont, 451
Cockatoo Island, 376
Coelacanthidae, 390
coelacanths, 368, 386, 445, 446
Coelorhynchus elevatus, 384
Coelorhynchus innovatis, 382
Coenocorypha aucklandica, 1353,

1373
Coenocorypha chathamica, 1331,

1357
Coffee Hill Member, 452
Coffin Bay, 329
Coimadai Local Fauna, 1056
Colbert, E. H., 58
Coles, 735
Coli-Toro Formation, 476,
collagen, 313, 314, 316, 317, 319-

322, 324-327, 330, 331, 929
collagenase, 317, 326
collector bias, 273, 291

Collins-Rubie, K., 90, 91
Collocalia spondiopygia, 835
Collorhinchus milii, 524
colobine monkeys, 222
colon, 235

colonization, 1320

Columbia University, 69
Columbidae, 740, 762, 768, 835
Columbiformes, 1331
Combienbar River, 340-342
Comerong Volcanics, 360
common ancestor, 608
Compositae, 314

ized information services,

148, 162
Condobolin Formation, 447, 450
gas conductance, 877
conifers, 259
Conilurini, 959, 961
Conilurus albipes, 960

conodonts, 356, 430, 400, 445, 450-

452
continental drift, 765
convergent evolution, 312
Coober Pedy, 633
Cook Islands, 1355, 1363
Cooma, 391, 392
Coonabarabran, 783, 1382
Cooper Creek localities, 745
Cooper Creek, 15, 479, 723, 746,
757, 761, 811, 812, 832, 834
Coopers Creek Beds, 451
Coopers Creek Limestane, 450,
452, 458
Cooyoo australis, 382
Cophixalus ornatus, 600
Cophixalus, 593
coprolites, 286
coprophagy, 237, 239
copulation devices, 431, 432
Cora Lynn Cave, 329, 666
Coraciiformes, 1332
Coreena Formation, 473, 476, 612
cormorants, 286
coronoid process, 219, 224
Corosaurus, 630
Corvidae, 281, 842
Corvus bennetii, 842
Corvus coronoides, 303, 842
Corvus mellori, 842
Corvus orru, 842
Corvus tasmanicus, 842
Cosesaurus aviceps, 638
Cosgriff, J., 58, 59
cosmine, 368
cosmaid scales, 445
Coturnix australis, 306, 830
Coturniz chinensis, 306, 830
Coturnix japonica, 734, 735
Coturnix pectoralis, 830, 1330
Coturnix, 292, 296, 758, 829
Couman Forest Cave, 329

CRA, 735

Cracraft, J., 766, 767

crane, 185

cranial kinesis, 606, 617
Cratochelone berneyi, 613, 617,

1173

Cratochelone, 713

Cravens Peak Beds, 346, 398, 399,
434, 437, 438, 447, 448, 452,
458, 459

Crespin, I., 155

Cretolamna appendiculata, 507,
508, 524, 525

Cretoxyrhina mantelli, 507, 509,
525

Crinia georgiana, 599

Crinia signifera, 598

Crinia, 593, 668, 669

cristid obliqua, 256, 947

Croc Pot, 330

crocodiles, 287, 388, 651, 655

Crocodilia, 1171

crocodilians, 609, 610, 639, 641

Crocodilus porosus, 94

Crocodilus selaslophensis, 612, 641

crocodylids, 1086

crocodylotarsans, 609, 637

Crocod ylotarsi, 637

Crocodylus johnsoni, 671

Crocodylus porosus, 1671, 672,
677, 678, 688076, 1077

Crocodylus selaslophensis, 688

Crocodylus, 638, 667, 670

Crocuta crocuta, 678

Crossochelys, 676

crossopterygians, 432, 454

Crossoptery gii, 368, 369, 386, 444

crown height, 208

Crown Lands Building, 15

crows, 286

crushing, 205

Criana zone, 446

Cryobatrachus, 614

cryptocleidids, 634

cryptodire, 617, 714,715

CSIRO Index (CSX), 169

CSIRO, 169

ctenacanth, 440, 449, 51

ctenacanthoid, 347, 441

Ctenacanthus, 441

Ctenodus breviceps, 397

Ctenodus, 397

Ctenurella gardineri, 366, 367

Cucilidae, 836

Cuculiformes, 1332

Cuculus pallidus, 836

Cuculus pyrrhophanus, 836

Cuculus variolosus, 836

Cuddie Springs bone bed, 722

Culmacanthus stewarti, 350, 351,
420, 453, 459

Culmacanthus, 401, 402

Curramulka Local Fauna, 666,
1052

Curramulka Town Cave, 329

Curramulka, 598

Curran’s Creek Cave, 291, 295,
821, 840

currawongs, 286

curvature analysis, 384

Cuscus, 252, 259

cuscuses, 259

cuspidate, 204

Cuvier, G., 6, 8, 248

Cyanorhamphus novaezelandiae,
1360, 1373

Cyanorhamphus, 1323, 1332

cycad, 585

cycloid scales, 388, 368, 445

Cyclorana australis, 601

Cyclorana novaehollandiae, 599

Cyclorana, 593

Cygnus atratus, 1329

Cygnus sumnerensis, 1329, 1357

Cylomeia, 585

cynodonts, 622, 1084

Cynognathus, 614, 622, 895

cytotaxonomy, 594

Dacelo gigas, 307

Dacelo novaeguineae, 281, 836,
837

Dacrydium, 140

Dactylopsila trivirgata, 257

Dactylopsila, 956

Dactylopsilinae, 955, 956

Daily, B., 50, 51

Dalatias, 501, 510

Dalpiazia, 500

Daltolac SW6, 198

Dandaragan, 633

Dangarfield Formation, 451

Danitherium, 1100

Danks Trust, 735

Daption capense, 1354

Daption, 1358

Darcius dug gani, 254, 256, 946

Dare Plain, 340-342

Darling Basin, 399, 1479, 660, 689,
710, 745, 750, 752, 760, 827,
831, 835, 1081, 1112

Dartmoor, 748

Darwin, C., 8, 346

Dasyatis, 510, 517,531

Dasyornis brachypteris, 821, 823,
840

Dasyornis broadbenti, 293, 840

Dasyornis longirostris, 840

Dasyornis, 296

Dasyruidae, 925, 929, 1086

dasyurids, 257

Dasyuroidea, 1086

Dasyuroides, 929

Dasyuromorphia, 925

Dasyurus geoffroyii, #03

Dasyurus hallucatus, 291, 292, 294,

302, 303, 678, 926, 1090

Dasyurus viverinus, 303, 1084

Dasyurus, 1236, 286, 880, 927,
929, 349

databases, 148, 169

Daubentonia madagascariensis,
261

Daubentonia, 257

David Holdings, 735

Davidson, T., 149

Dawson, L., 77

de Blainville, D., 149

de Vis Symposium, 80

de Vis, C.W., 754, 722, 757, 758,
810

Deal Island, 1347

INDEX - 1423

death assemblages, 270
deathtrap caves, 279-281
deep chemical weathering, 129
Deep Creek Limestone, 434
defensive structures, 1140
deformation, 206
Deinocheirus mirificus, 647
Delatitia breviceps, 396
Delatitia, 397
Delatsaurus kimberleyensis, 580
Delphinornis, 615, 748
Delphinus delphis, 1180, 1182
deltas, 276
Deltasaurus kimberleyensis, 579,
580
Deltasaurus pustuldtus, 580, 586
Deltatheroida, 907
Deltoptychius, 442
Dempsey’s Lake, 302, 329
Denaea, 440
Dendrolagus, 216
Denisonosteus weejasperensis, 358
Denisonosteus, 355
dental adaptations, 202
dental drill, 190
dental efficiency, 205
dental formula, 901, 909
dental modifications, 203
dental morphology, 208
dental striae, 233
dental tools, 431
denticles, 431
dentine, 444, 445, 476, 500
dermal denticles, 498, 499
dermosphenotic, 471
Derwentia warreni, 579
Desmatochelyidae, 713
detergent, 431
detrital limestone, 118
Devil’s Lair, 291, 293, 295, 304,
309, 598, 668, 820, 829, 836;
837, 842
diagenesis, 273
Diaphorapteryx hawkinsi, 1330
Diaphorapteryx, 1359
diapsid, 629, 630
Diapsida, 606
diapsids, 609, 625
diastema, 250
diatomite, 783
dictionaries, 156
Dictopleurichthys, 376
Dictopyge illustrans, 376
dicynodont, 583, 621, 663
Didelphidae, 905, 907
didelphid, 924, 925, 929, 931
Didelphis virginiana, 907, 930
Didelphis, 896
Didelphodon vortax, 906
Didelphodus, 902
dict, 256
digesta, 231, 234, 237, 241
digestion, 207, 284
digestive strategy, 207, 209
dignathic heterodonty, 502
digrastic process, 224
digrastic sulcus, 224
dingo, 286, 950

1424 - INDEX

Dinmore, 659

Dinornis torosus, 1325

Dinomithidae, 1320

Dinomithiformes, 1325, 1359

Dinosaur Cove, 188, 190, 474

dinosaur stampede, 661

Diodon formosus, 383

Diomedea cauta, 1349

Diomedea chlororhynchus, 1357

Diomedea thyridata, 749

Diomedea, 1358

Diomedeidae, 749

diphycercal tail, 445, 466-468

diplacanthids, 351

diplacanthioids, 443, 453

Diplacanthus, 443

dipnoans, 445, 466

Dipnoi, 368, 369, 391, 444

Dipnorhynchus kurikae, 466

Dipnorhynachus sussmilchi, 466

Dipnorhynchus, 395, 396

Dipodidae, 911

Diprotodon australis, 1099, 1102,

Diprotodon longiceps, 1102

Diprotodon minor, 1102, 1104

Diprotodon optatum, 1935, 1075,
1101, 1104, 1105, 1107, 1136,
1137, 1159

Diprotodon, 8, 11, 2A, 28, 29, 30,
31, 38, 70, 73, 950, 951,1073,
1074, 1078, 1079, 1082, 1098,
1100, 1102-1104, 1106, 1127,
1128, 1131, 1135, 1143

diprotodant, 1086

diprotodontan, 252, 925, 929, 932

diprotodontid, 250, 724, 728

Diprotodontidae, 935

Dipterus digitatus, 393, 395

Dipterus, 395

Dirk Hartog Formation, 119

Dirk Hartog Island, 1347, 1350

discovery of gold, 12

discriminatory behaviour, 1139

dispersal routes, 1168, 1169, 1320

dispersal, 763, 765

dispersalist theories, 1363

distal caccum, 234, 236, 240, 241

distribution (non-passerine birds),
832

Ditjamanka Local Fauna, 756, 757,
933, 937, 1018

Djilgaringa gillespiei, 1069

DNA sequencing, 312, 332

DNA, 312, 332

Docodon, 897

Docodonta, 897, 922

Dog Rocks Islands, 513

Dog Rocks Local Fauna, 959, 1055,
1192,

Dog Rocks, 329

dogs, 211

dolichodephalic, 1114

Dolichorhynchops, 1173

dolines, 277, 279, 280

dolomites, 116, 139

Dondobolin Formation, 434

Doragnathus woodi, 576

Dorcopsis, 216

Dorcopsoides, 1079
dorsal ribs, 616
Dorudon, 1180, 1183
Doswellia, 637
Draper, J., 660

Dromaiidae, 811

dromaiids, 811

Dromaiinae, 768

dromaiines, 740

Dromaius ater, 315, 745, 822, 824,
1349

Dromaius baudinianus, 3, 743, 745,
782, 822, 824, 1349

Dromaius gidju, 743, 744, 782

Dromaius gracilipes, 745, 824

Dromaius novaehollandiae, 321,
323, 325, 327, 742, 745, 824

Dromaius ocypus, 744, 745

Dromaius patricius, 745, 824

Dromaius, 1350

Dromiciops, 924, 929

Dromornis australis, 781,782

Dromornis stirtoni, 781,782, 1076,
1079

Dromornis, 743, 884, 886, 888

dromomithid tracks, 738

dromomithid, 326, 811

Dromomithidae, 2, 6, 69, 722, 739,
740, 742, 766, 768, 825, 1086

drought, 1133, 1142

Drummond Basin, 403

Dry Creek Sands, 516, 517

Dry River, 670

dryosaurid, 664

dry screening, 736

Duaringa Fauna, 385, 475

Duaringa, 481, 490

Ducabrook Formation, 452

Duckworth Creek, 471, 472

ductile materials, 206

Dugan, K., 8

Dulcie Range, 340-342

Dulcie Sandstone, 399

Dundas Group, 116

dune fields, 276

dune sands, 1231

Dunkleosteus, 355, 356

Dunstan's Limestone Kiln Quarries,
302.

Duntroonornis parvas, 1177

Dusham, 1359

durodentine, 439

durophagous dentition, 355

durophagous, 346

dwarfing, 1820, 112, 1131, 1134,
1141

Dwomamor Local Fauna, 1032

Eagle Cave, 1360

ear drum, 896

car region (mammal), 896

ear region (reptile), 896

East Gondwana Province, 398

East Gondwana, 351, 362, 402

East Greenland, 388, 574

East Kangaroo Island, 1347

Eastern regional fauna, 1127

Eastmanosteus caliiaspis, 360, 355,
420, 421

Ebenaqua ritchiei, 371, 373

Echinorhinus, 499

echolocated, 1184

ecological tethering, 1142

ectotherms, 202

Edaphodon eyrensis, 349, 506

Edaphodon mirabilis, 349, 518, 531

Edaphodon sweeti, 349, 518, 531

Edaphosaurus, 607

Eden, 340-342, 389

edentates, 910

edestid shark, 347, 440, 442

Edestus, 442

EDTA, 317, 320

Egermia, 667, 673

eggs (fossil), 723, 872

egg (gas conductance), $77

egg (crystal structure), 877

egg (curvature analysis), 874

egg dimensions, 876

egg geometry, 875

egg length, 879

egg predation, 880

egg reassembly, 880

eggshell structure, 872

egg size, 878

egg surface area, 873

egg volume, 873

egg weight, 873, 876, 879

egg width, 879

eggcases, 440

eggshell, 742, 825, 872, 873, 880,
881

Egretta alba, 1327

Egypt, 512

Egyptian mummies, 313

Eight Mile Plains, 475, 675, 671

Eindjama Gorge, 668

Eklatadelta ima, 950

Ekladelta, 1079

Ektopodon serratus, 254, 946

Ektopodon stirtoni, 253, 254, 256

Ektopodon, 246, 250, 251, 252,
255, 257-259, 261, 946

Ektopodontidae, 253, 254, 256,
945, 958

ektopodontids, 258

Elanus scriptus, 288

elasmobranchs, 440

Elasmobranchiomorphii, 345

Elasmosauridae, 1173

elasmosaurids, 634

elasticity, 206

elastin, 313

electrophoresis, 312, 317, 320, 323,
327, 331

elephants, 330

Elizabeth Springs, 506

elonichthyid, 455

Elonichthyes, 370, 583

Elpisopholis, 376

Elseya, 675, 705

Emballonuridae, 958

Emberiza, 823

embrasure, 234, 236, 241

embryonic fissures, 268

Emeidae, 671, 1320

Emry, R., 73

Emu caves, 329

emu eggshell, 878

emus, 886

Emydura australis, 716

Emydura macquanii, 619, 675,711

Emydura, 667-669, 675, 705

en chevron selenes, 257

enameloid, 443, 500

enantiomithines, 737

encephalization quotient, 1137

encephalization, 1139, 1141

encyclopaedias, 156

endothermic, 665

endothermy, 665, 895

Endurance Tin Mine, 738

Enhydra lutris, 261

Enhydra, 258

Enigmatichthys, 376

Enoggera Reservoir, 468, 490

entelodonts, 1140

entoconid, 233

enzymes, 207

Eocaiman, 641

Eoctenodus, 397

eosuchian, 609, 636

ephiphyses, 285

Ephipptorhynchus asisticus, 752

Epiceratodus denticulatus, 477

Eptceratodus pattinsonae, 475

Epiceratodus, 478

epicontinental seaways, 131

Epidolops ameghinoi, 908

epiphyses, 623

epipubic bones, 1086

epoxy resins,198, 199

Equus caballus, #3

Equus, 967

Eremnochelys, 608

Ericiolacerta, 614

Ericmas Local Fauna, 667, 739,
757, 1025,

Ericmas Fauna, 480

Erith Island, 1347

Erithacus, 823

Eromanga Basin, 124-126, 129, 504

Errolosteus goodradigbeensis, 354,
356, 357

eruption pattem, 209

Erythrobatrachus
noonkanbahensis, 576, 578

Erythrogonys cinctus, 832

Erythrosuchia, 637

Esacus magnirostris, 760

ESSO, 735

estrous cycle, 904

Etadunna Formation 49, 54, 55,
140, 252, 478, 480, 482, 483,
490, 507, 596, 666, 667, 673,
716, 753, 762, 884, 931 1015,
1017, 1020,

Etheridge, R., Jr., 11, 25, 28, 151,
722

Ettrrick Mazi, 512

euantiarchs, 362

euarthrodires, 354

eubrachythoracids, 355

Eubrontes, 659
Ewcalyptus, 140, 290, 1319
Eucla Basin, 125, 128, 511
Eudolops caroloameghinoi, 908
Eudyamis taitensis, 11332, 373
Eudyptes chrysochome, 825, 1348,
1353, 1374
Eudyptes chrysolophus, 1326
Eudyptes pachyrhyachus, 1326
Eudyptes, 749
Eudyptula minor, 749, 825, 177,
1354
Eugomphodus, 501
eunotosaurs, 609
Eunotosaurus, 616
cupantotheres, 910, 924
Euparkeria, (7
Euparkeriidae, 637
Eupleurogmus creswelli, 351
Eupleurogmus, 453
Euramerica, 343, 400, 402
Eurhinosaurus, 626
Eurinella Formation, 475, 483
European cats, 1319
European man, 1365
European rats, 1319
European settlement, 357, 1358
Euryanas finschi, 1329
euryapsid, 606, 624, 625, 628
Euryapteryx, 1325
Eurydolops, 615,911
Euryzygoma dunenese, 1082, 1140
Euryzygoma, 1081, 1102
Euselachiformes, 498
eustacy, 1168
eustatic changes, 1169
eustatic, 1169
Eusthenodon, 388
eusthenopterids, 388
Eusthenopteron, 388, 445
eusuchian, 641
eutherians, 214, 905
evaponies, 112, 113, 116, 119, 120,
123, 124, 126, 128
Evergreen Formation, 572, 573,
586, 611, 633
exoskeleton, 209
extinction, 1072, 1076
extraterrestrial events, 1168, 1069
exudates, 205
Eyre Formation, 73, 140
Eyre Peninsula, 140
fabrosaurids, 650
fabrosaurs, 648, 649
facial angle, 255
faecal material, 287
faecal pellets, 237
Fairbridge, R.W., 163
Fairfield Group, 358, 449
Falco berigora, 281, 286, 288, 302,
309, 810, 829
Falco cenchoides, 281, 286, 288,
305-307, 829
Falco novaeseelandiae, 1330
Falco peregrinus, 281, 302, 829
Falco sparvarius, 305, 306
Falconer, H. 8
Falconidae, 757, 829

falconids, 286, 811

Falconiformes, 1329

Fallacosteus, 355

fan deposits, 276

fanglomerates, 117

faunal convergence, 1168

faunal divergence, 1168

feathers (fossil), 606, 737

feedback loops, 1168

feedback mechinism, 1169

Felis catus, 211, 254, 303, 822

Felis, 966

feral cats, 326

fexmentation chamber, 214

Ferugliotherium windhauseni, 897

fibred enameloid, 500

fibreoptic lamp, 230

Fifield, 459

Fiji group, 1350

Fiji Islands, 758, 1355

filter feeding, 350

fine shear, 236-238

fire ecology, 1142

Fischers Point, 515

Fisher, N.H., 86, 87, 90

Fisherman’s Cliff Loca! Fauna, 669,
1053, 1102

Fisherman's Cliff, 730, 745, 917

Fishing Point Mari, 513

fission track dating, 724

fissure fills, 276

Fitzroy Basin, 58

Fitzroy Crossing, 347

Fitzroy Graben, 124

Fitzroy River, 385

Flandrian Transgression, 1356

Flannery, T.F., 33, 34, 69, 76, 78

Flidersichthys, 382

Flinders Expedition, 3

Flindess, M., 3, 150

Flinders Island, 1183, 1347, 1362

Flinders University, 723

Floraville Local Fauna, 1670, 1046

flow-moulded vinyls, 198

flume experiments, 282

fluvial deposits, 282

folivores, 257

footpnnts, 781

Forbes, 576

Forbes, E., 149

foregut fermentation, 207-209,215

foregut, 214

foreign experts, 6, 37

forelink, 126, 224

formic acid, 176, 182

Forrest caves, 326

Forsyth’s Bank, 517, 1039

Fossil Bluff Sandstone, 513

Fossil Bluff, 514, 515

fossil localities (island vertebrates),
360-1361

fossil localities (islands), 1369-1372

fossil localities (mammalian), 920,
921, 1012

fossilization, 272

Foul Air Cave, 324

Fox Cave, 837

Fox, R., 388

foxes, 326

Freestone Cove Sandstone, 513,
515

Freestone Creek, 340-342, 351,

398, 401, 452

freeze-thaw technique, 431

Fregatta grallaria, 1373

Fregetta, 1358

Fremouw Formation, 614

Friends of the Museum of Victoria,
735

Friagilla, 823

frogs, 286

Fulbright Program, 46, 49, 61, 69,
75

Fulbright Fellowship, 87

Fulbright scholar, 69

Fulgurotherium australe, 612, 650,
688

Fulica atra, 1330

Fulica prisca, 1330

fume cupboard, 430

fume scrubber, 184

functional morphology, 202, 1167

Fumeaux Group, 1347

Fyansford Formation, 510, 513,
514

Gadus refertus, 384

Gadus, 382

Gaffney, E., 78, 80

Galapagos tortoises, 675

Galeaspid-Yunnanolepid Province,
400

Galeaspida, 343

galeaspidiforms, 433

galeaspids, 344, 431

Galeocerdo aduncus, 515,517,524,
5H

Galeocerdo cuvier, 531

Galeorhinus, 517

Galeus, 510

Galilee Basin, 123, 124

Gallicolumba jobiensis, 1374

Galliformes, 1330

Gallinago hardwicki, 832

Gallinula (Tribonyx) mortierit,
293, 810, 819,820, 821, 823,
827, 831, 1330

Gallinula tenebricosa, 831

Gallinula, 752

Gallirallus australis, 1323, 1330,
1356, 1357

Gallirallus dieffenbachi, 1130,
1256, 1357, 1359

Gallirallus hartreei, 133

Gallirallus minor, 1256, 1330

Gallirallus modestus, 1330, 1359

Gallirallus modestus, 1359

Gallirallus philippensis, 1359,
1360, 1374

Gailirallus sylvestris, 1330, 1374

Gallirallus ventralis, 1330

Gailus gallus, 303

ganoid scales, 444

ganoine, 444

Gantheaume Point, 660

Garra Formation, 434

Garrodia, 1358

INDEX | - 1425

Gascoyne River, 340-341
Geilston Bay Local Fauna, 1015
Geilston Bay, 936, 955
Geitonichthys ornatus, 378
Geitonichthys, 376
Gelada, 209
Gellibrand Clay, 514
Gellibrand Marl, 513
Gemuendina, 367
Genoa River Beds, 572, 573, 582
Genoa River, 340-342, 574
Genyornis newtoni, 741, 781, 825,
867, 878, 879, 886, 888, 1077,
1128
Genyornis, 3, 28, 270, 271, 732,
743, 820, 823, 879, 1350
GEOARCHIVE, 168
Geocrinia laevis, 599
Geocrinia, 593, 598, 667
Geological Society of America,
154, 160
Geological Society of London, 6,
153, 155, 163
Geological Survey of Victona, 17,
20
Geopsittacus occidentalis, 822, 836
Geopsittacus, 1331
GEOREF, 168
Georgetown, 452
Georgina Basin, 116, 117, 398,
399, 447
Geosystems, 162
geothermal events, 1169
Geranosaurus, 648
Gerbillinae, 203
Gerbillus, 204
Gervais, P., 8
Gerygone albofrontata, 1333
Gerygone igata, 1333, 1373
Gerygone, 1333
Giant Panda, 215, 223, 257, 261,
312
Giant Squid, 631
gigantism, 834
Gilberton, 340-342, 363
gill rakers, 443
gill skeleton, 442
gill slits, 344
Gill, E., 33, 46, 723
Gingham gap, 395
Gin Gin, 349
Gingin Chalk, 506, 508
ginkgo, 585
Gippsland Basin, 129
Gippsland Limestone, 515
giraffes, 1106
gizzard stones, 742, 825
glacial period, 812, 1319
glaciation, 113, 122, 129
glaciers, 122
Gladstone, 478, 705
Glauert, L., 46
Glen Garland, 670
Glenelg River, 841
Glenidal Formation, 572, 573, 586
Glenn Florrie, 140
global cooling, 128
Glossopsitta porpkyrocephala, 293,

1426 - INDEX

296, 836

Glossopsitta concinna, 836

Glossopsitia pusilla, 836

Glossopteris browniana, Glut, D.F.,
163

Glycol 4000, 193

glyptolepid, 390

Glyptolepis, 390, 421, 445

gnathorhizid, 484, 472

Gnathosaurus, 658

Gnathostomata, 345

Gneuda Formation, 340-342, 344,
358, 389, 393, 395, 434, 436,
447

Goat Paddock, 140

Gogo Fauna, 174

Gogo Formation, 352, 388, 391,
395, 421, 467

Gogo, 35, 36, 76, 80, 359, 364,
365, 368, 370, 390, 391, 393,
445

Gogonasus andrewsae, 387, 388

Gomphonchus, 443, 452, 453

Gondwana continents, 131

Gondwana realm, 446

Gondwana, 118, 120-123, 131, 136,
433, 445, 610, 766, 767, 959,
1318, 1325

Gondwanaland, 643, 655, 676

Gondwanan origin, 767, 769, 1320

Gondwanan, 677, 1331, 1334

Gonorhynchus, 385

Goodna, 659

Goodradig beeon australianum,
354, 356

Gore, 670

Gosford Fauna, 374

Gosford Formation, 374, 467, 470,
583

Gosford, 340-342, 345-347, 374,
467,470

Gosfordia truncata, 396, 397, 467,
468, 470, 471, 485, 583

Gosses Bluff, 340-342, 388

Goulburn, Major, 13

Gould, J., 236

Goulden's Hole Cave, 329

grabens, 276

Gracilisisuchus, 637

gracilisuchians, 609

Grallina cyanoleuca, 842

Grallinidae, 767, 842

Graminaceae, 814

Grampians Group, 572, 573, 582

Grampians Ranges, 574

Grampians, 340-342, 434

Grange Bum Formation, 252, 516,
518

Grange Bum, 183, 509, 513, 517

granite caves, 276

Grant Formation, 122

grass, 140, 227, 235-237, 249

grassland, 136, 142, 208, 259,1127,
1128

grass nodes, 237, 240

Grawan, 473, 474, 476

grazers, 208 219, 249, 950

grazing kangaroos, 236

grazing macropods, 218-219

grazing, 216

Great Artesian Basin, 36, 125,753

Great Barrier Reef, 1346

Great Buninyong Estate Mine, 1044

Green Bluff, 55

Green River Formation, 958

Green Waterhole Cave, 287, 290-
292, 821

Green Waterhole, 810, 831, 835,
838, 937

greenhouse heating, 1175

Greenland, 395, 907

Gregg, D., 77

Gregory, J. W., 15, 19, 20, 21, 33,
34, 479, 722, 723

Grenfell fauna, 402

Grenfell, 351, 362, 391, 452

grey kangaroos, 249, 310

Grice's Creek, 514

Griman Creck Formation, 473,
474, 476, 477, 490, 612, 618,
641, 671, 714,

grinding, 205

Griphognathus whitei, 393, 421,
467

Griphognathus, 391

Grippia longirostris, 626

Grippia, 625

Groberia minoprioi, 913

Groberidae, 911

grocnlandaspids, 355

Groenlandaspis, 358, %62, 401,
402, 582

Gross, W., 446

ground sloths, 1106

growth lines, 884

Gmidae, 758

Gmuiformes, 1330

Grus rubricundus, 758

Gualepis, 448, 459

guano deposits, 1350

Gulf Stream, 112

Gulgong Deep Lead, 668

Gulgong, 710

Gulo gulo, 678

Gulo luscius, 1140

gut contents, 231, 234

Gymnobelideus, 956

Gymnorhina tibicen, 309, 842

gymnosperms, 814

gynandric heterodonty, 502

Gyracanthides murrayi, 350, 351,
453

Gyracanthides, 401

agyracanthids, 351

Gyracanthus, 351

Gyroptychius australis,387

Gyroptychius, 386, 387

Hadronomus, 1079

hadrosaurs, 649

hagfish, 431

Halaeetus pelagicus, 1359

Halcyon pyrrhopygia, 836, 837

Halcyon saacta, 832, 836, 1321,
1332

Hale River, 140

Haliagetus australis, 1329, 1359

halite, 119

Hall, T. S., 15, 18, 22

Han, T., 11

Haman's Cave, 830

Hamilton Hotel, 737

Hamilton Local Fauna, 955, 956,
959, 1039

Hamilton River, 737

Hamilton, 34, 57,75, 78, 249, 254,
340-342, 513-515, 517, 1179,
1183, 1330

Hand, S., 33, 76

handbooks, 156

Harpagodens ferox, 347, 348, 450

Harpagodens, 401

Harpagornis moorei, 1321, 1329

Harpagornis, 1330

Harris, W.K., 728

Harrytoombsia, 355

harvester ants, 283

Hatchery Creck Conglomerate, 352,
355, 364, 447

Hatchery Creck Fauna, 363

Hatchery Creck Formation, 434

Hatchery Creek, 386

Hawaiian Islands, 1346, 1362, 1363

Hawkesbury Sandstone, 374, 376,
467, 470, 472, 572, 583

hawks, 286

Head, W., 75

headings, 165

heath, 1128

Heleoporus albopunctatus, 599

Heleoporus, 593, 598, 668

Helicoprion davisti, 348

Helicoprion, 347, 401

helodont, 451

helodontoids, 441

Helodus, 347, 401, 442

hemicellulose, 207

Hemiphaga novaeseelandiae, 1323,
1331, 1357, 1373,

Hemupristis elongatus, 500

Hemipristis serra, 500, 515

Henk’s Hollow Local Fauna, 1033

Henschke's Bone Dig, 305, 837

Henschke's Cave, 329, 598

Henschke's Quarry Cave, 828

Heptranchias ezoensis, 511

Heptranchias howellii, 511, 525

Heptranchias perlo, 525

Heptranchias, 499, 502

herbivores, 207-209, 250, 257

herbivorous diet, 257

herons, 286

herrerasaurs, 610, 642

Hervey Group, 395

Hervey's Range, 452

Haterenchelys regularis, 384

heterodont, 205

heterodontosaurs, 648, 649

Heterodontus cainozoicus, 513,
516, 526

Heterodontus portusjacksoni, 501,
526

Heterodontus, 501, 503, 512

heterodonty, 502

Heterolocha acutirostris,

Heteromyidae, 911

heterostracans, 431, 432, 433

Heterostraci, 343

hexacanthid, 448

hexanchid, 346

hexanchoids, 502

Hexanchus agassizi, 511, 512,525

Hexanchus gracilis, 499

Hexanchus griseus, 502

Hexanchus, 349, 499, 511

Hibbard, C., 52

Hieraaetus morphonoides, 829

Hills, E.S., 33, 46

Himalaya Mountains, 921

hind gut, 204, 214

hindgut fermenters, 207-209, 237

Hipposideridae, 958

Hirundapus caudacutus, 823, 835

Hirundinidae, 838

Hirundo neoxena, 281, 838

Hirundo rustica, 838

Hispidaspis, 508, 524

Hobart Zoo, 929

Hobson, E. C., 11

Hobson, M., 11

Hoch, E., 95

Hochstetter, 8

Holey Plains Marl Member, 514

Hollands Creek, 398

holocephalan, 347, 349, 432, 498,
503

Holoclemensia, 905

Holodipterus gogoensis, 421

Holodipterus, 395

Holonema westolli, 355, 360

Holoptychius, 390, 401, 402, 445

holosteans, 368

Homiphoca capensis, 1179

Homo sapiens, 286, 303, 304, 961,
970, 1142

homodont, 901

homologous, 203

homology, 202

homosteids, 355

honey possums, 901

honey-pot ants, 238

Honeycomb Hill Cave, 1321

Hooke, R., 148

Hookcan materials, 206

Hookina Creek, 329

Hope, J., 33, 70, 77, 1348

Hordem Vale, 515

Hom Creek Siltstone, 446

horse, 248-250, 964

Hotstuff, 187

Houtman Abrolhos Archipelago,
1347, 1350

Howittacanthus kenioni, 351, 352,
420, 453

Howgqualepis rostridens, 370, 421

Howgqualepis, 370, 401

Hughenden, 340-342, 617, 633, 656

Hulitherium tomasettii, 96-99

Hunter Island, 304, 817, 825, 1347-
1349

Hunter Siltstone, 351, 362, 388,
391, 395, 452

Hunterium Museum, 6

Huronian sequence, 113

Husst, H., 28

Hutton Sandstone, 611

Huxley, T. H., 8

hybodontid, 349

hybodontoids, 441

Hybodus, 441

Hydrilla verticillata, 482

hydrochloric acid,. 431

hydrodynamic parameters, 271

hydrodynamic sorting, 271, 482

hydrofluoric acid, 182, 430

Hydrolagus, 503

Hydromyinae, 959

Hydromyini, 959

Hydromys chrysogaster, 960

Hydromys, 258

hydroxyapatite, 436

Hylaeochampsa, 641

Hylidae, 593

Hylonomus, 607

Hymenolaimus

hyomandibular braces, 346

hypoconid, 233, 234, 237

hypoconulid, 233

hypolophid, 216, 224

Hypsilophodon, 650

hypsilophodont, 649, 657, 664, 665

Hypsiprymnodon moschatus, 1111

Hypsiprymnodon, 213-215, 948,
951,954

hypsiprymnodontines, 215

hypsodont teeth, 204, 209, 940, 943

hypsodonty, 203

Hystrix, 286

Ian Potter Foundation, 735

Ian’s Prospect, 475, 1045

Ibis (7) conditus, 756

Icaronycteris, 958

Ichthyodectidae, 382

ichthyoliths, 430

ichthyopterygians, 686

ichthyosaur, 606, 627, 631, 1173

Ichthyosaurus, 625

Ichthyostega, 574

Ichthyostegalia, 570

ichthyostegids, 576

iguanodonts, 649, 650

Haria illumidens, 940, 941, 1069
Ilaria, 90

Mlariidac, 939

ilariids, 932

Ilbandornis lawsoni, 743, 781, 782
Ilbandornis woodburnei, 781, 782
immunoassays, 317
immunoblotting, 319-321, 328
immunofluorescence, 313
immunoreactive, 320, 323, 326
Incisoscutum ritchiei, 355, 359

Index to Scientific Reviews,162

indexing joumals, 161

Indomalaysian origin, 769

inferior dental foramen, 224

inflected angle, 1086

Ingersoll-Rand, 735

Inglis, G., 73

Ingram Trust, 735

Inoceramus, 516

insectivores, 209, 249, 257, 958

insectovorous diet, 258

insectovory, 257

insular fossil faunas, 1360

intelligence, 1138

intercentrum, 571

interglacial periods, 313

Investigator Strait, 1349

invisible college, 157

Ips wich-Moreton Basin, 124

Inian Jaya, 446, 812, 1100

Inshtown, 745

irruptive behaviour, 293

ischnacanthid, 351, 443, 451, 453

Ischnacanthida, 350

Ischnacanthus, 443

Ischnodon, 930

Ischyodus dolloi, 518, 531

Ischyodus mortoni, 515

Ischyodus newtoni, 349

Ischyodus, 508

Ishua Group, 113

island faunas, 1346

island area, 1364

island biogeography, 1363

Isoodon obesulus, 303, 930

Isoodon, 232, 234

isotope istry, 126

Isurus benedeni, 501, 514, 515,
527

Tsurus desori, 512, 514, 517, 527,
528

Isurus escheri, 517

Isurus hastalis, 510, 516,517. 528

Isurus oxyrinchus, 502, 514,517,
524, 527

Isurus paucus, 501, 512,528

Isurus planus, 501, 512,514, 528

Isurus retroflexus, 501,514,517,
5A, 529

Jablonski, D., 163

Jack limestone, 452

Jack's Lookout, 340-342

James Cook University, 723,

James Ross Island, 614

Jameson, R., 6

Jandakot beds, 517

Jan Juc Formation, 511, 512

Jan Juc, 340-342

Jemalong Gap, 340-342, 391, 402

Jemmys Point, 340-342

Jemmys Point Formation, 516,
517

Jesse Limestone, 448

Jones, R., 80, 340

Jubilee Dam, 671

Jukes, W., 12

Julia Creek, 617

Junee Group, 117

Kadimakara australiensis, 611, 635

Kadimakara, 583, 636

Kagu, 1331

Kakuru kujani, 612, 646

Kakuru, 647, 648

Kalisuchus rewanensis, 621, 640

Kalisuchus, 583, 639, 641

Kangaroo Island, 347, 723,745,
824, 827, 831, 839, 840, 882,
1302, 1349

Kangaroo Well Local Fauna, 668,
1027

Kangaroo Well, 54, 668

Kannemeyeria, 614, 620

Kanunka Fauna, 483

Kanunka Local Fauna, 667, 1051,
1102

Karawarren Limestone, 512

karotypic diversity, 1141

karst topography, 1356

karstic limestones, 1362

karstic settings, 276, 277

karyotypological evidence, 1096

Katapini Formation, 479, 490

Katipiri Sands, 80, 483, 667

katoporids, 439

Keast, A., 766

Keeuna, 927

Keilor, 514

Kekenodon onamaia, 1181, 1182

Kelloggs, 735

Kemp, A., 76, 340

Kemp, N. 77, 343

Kempfield, 434

Kendrickchthys, 355

Keratobrachyops australis, 581,
585

Kermadec islands, 1355

kerosene, 191

keywords, 165

Kimberley district, 929, 1127

Kimberleyichthys, 355

King George Island, 615

King Island Emu, 3

King Island Rise, 1346

King Island, 3, 324, 329, 723,
745, 824, 1090, 1128, 1346-
1348

kingfishers, 286

Kingoria, 614, 621

Kings Creek, 324

Kingston, 1351

Kirrak, 474, 477

Knobby Sandstone, 364

Knocklofty Formation, 340-342,
374, 390, 470, 472, 580, 586

Knocklofty Sandstone, 572, 573,
574

Knowledge Creek, 660

Knowles, P., 73

Kockatea Shale, 572, 573, 580, 586

Kolopsis torus, 934, 1080

Kolopsoides, 97

Konig, C.,4

Koobor jimbarratti, 933

Koobor notabilis, 933

Koobor, 934, 940

Koonalda Cave, 291, 294, 295, 306,

INDEX - 1427

308, 309, 325, 820-822, 837,
841
Koonwarra, 340-342, 377, 380,
382, 477, 726, 729, 737, 781
Koorwarria manifrons, 380
Korora oliveri, 1177
Kow Swamp, 961
Krefft, G., 8, 13, 20, 236
Krefft, J.L.G., 25, 27, 28
Kronosaurus queenslandicus, 613,
632, 633, 1172, 1173
Kronosaurus, 613, 631, 634,1173
Krui River Loca] Fauna, 668, 1050
Kudnampirra, 834
Kudnu mackinlayi, 585, 611, 623,
624
Kudau, 583
Kuhneotheriwn praecursoris, 898,
902
Kummel, B., 164
Kuterintja, 940
Kutjamarpu Fauna, 482
Kutjamarpu Loca] Fauna, 481, 667,
739, 750, 934, 941, 945,
946,1025
Kutjitara Formation, 757
Kyancutta Museum, 14
Kyarranus kundagungan, 599
Kyarranus, 593, 598, 669
La Meseta Formation, 615
labyrinthodonts, 570, 625
Lachlan Fold Belt, 117, 118, 121
Lactarius tunulatus, 384
lactation period, 905
Laemba, 1355
Lagorchestes hirsutus, 237
Lagostrophus, 953
lagosuchians, 610
Lagosuchidae, 637
Laguna Umayo Local Fauna, 910
Laidlaw Volcanics, 447, 452
Lake Bonney, 515
Lake Bungunnia, 813-815
Lake Callabonna, 2,18, 28- 31,
36, 70, 73, 324, 329, 722,
742, 745, 781, 1102
Lake Colongulac, 324
Lake Eyre Basin, 15, 33, 37, 49,
50, 57, 74, 80, 140, 254, 1015
Lake Eyre localities, 812
Lake Eyre region, 21
Lake Eyre Sub-basin, 27, 79, 140,
142, 726, 739, 753, 757
Lake Eyre, 142, 249, 340-342, 474
Lake Frome, 69, 71, 73, 140, 480,
1183
Lake George, 129, 814, 816, 817
Lake Kanunka, 54, 56, 58, 329,
480, 667, 726, 730, 745, 751,
757, 758, 760
Lake Kittakittaooloo, 745
Lake Menindee, 220, 745, 883,
1142
Lake Mungo, 813, 961
Lake Namba, 326, 330, 331, 481
Lake Ngapakaldi, 57, 59, 62-64,
2A9, 251, 329, 481, 726, 739,
757, 758, 782

1428 - INDEX

Lake Palankarinna, 48, 49, 52, 53,
60, 61, 73, 75, 76, 249, 253,
32A, 329, 330, 480, 592, 596,
671, 726, 728, 730, 732, 745,
753, 756-758, 760, 762, 783,
937

Lake Pinpa, 249, 252, 480, 726,
746, 757, 760, 762

Lake Pitikanta, 726

Lake Tandou, 745

Lake Tarkarooloo, 75, 249, 252

Lake Tinko, 481

Lake Tyers, 1043

Lake Tyrrell, 813, 815

Lake Yanda, 481, 596, 726, 757

Lakes Entrance, 515

Lalage leucopyga, 1374

Lamarck, J. B., 3, 149, 150

Lambdodus, 441

Lamna arcuata, 307

Lamna cattica, 516

Lamnaa nasus, 527

Lamaa venusta, 524

Lemna, 347

Lamniformes, 502, 508

lampreys, 431

Lancefield, 305, 324, 745, 825,
1109, 1142

Lancefield South, 324

land plants, 120

land-sea distribution, 1169

landbridge, 1346, 1347

Lang, J. D.,6

Lansdowne, 671

Laornis, 08

Lapillopsis nanus, 577, 578

large colon, 237

larger foraminifera, 129

Laridae, 760, 831

Larus bulleri, 1320

Larus novaehollandiae, 832

Lasiorhinus angustidens, 1096

Lasiorhinus kreffti, 1096

Lasiorhinus medius, 1096

Lasiorhinus, 940, 943

last glacial maximum, 1346

last glacial, 820

Last Minute Site, 481

lateral line canals, 570

latex peel, 196, 197

latex rubber, 196, 197

Lathamus discolor, 836

Latimeria chalumnae, 445

Latimenia, 386

latipinnate, 625

Latocamurus coulthardi, 460, 421

Laura Basin, 124, 126

Laurasia, 131, 136, 648

Laurel Formation, 347

lava caves, 276

Law of Succession, 8

Lawson Quarry, 52, 53, 60, 61, 745

Lawson, P.F., 32, 47, 48, 50, 53,
773, 811

Lawson-Daily Quarry, 475

Leaellynasaura amicagraphica, 688

Leaeltynasaura, 650

Leaf Locality, 62-64, 251, 329, 782

leaf-cating habit, 257

Leakeyornis aethiopicus, 753

leaves, 214

Lechriodus intergerivus, 598

Lechroidus fletcheri, 599

Lechriodus, 593, 669

Leguninosae, 259

Lehman, 77

Leichhardt, F.W. L., 8

Leidy, J., 631

Leigh Creck Coal measures, 576

Leigh Creek, 124, 340-342, 376

Leighiscus, 376

Leiopelma, 601

Leipoa, 829

Lemley, R-E., 76, 79

Leonardo da Vinci, 148

lepidomorium, 440

lepidosaur, 624

lepidosauromorphs, 609, 623, 686

lepidosaurs, 609, 622, 623, 636

Lepidosiren paradoxa, 466, 486

Lepidosiren, 391

lepidotrichia, 444

lepospondyls, 570

Leptocleidus superstes, 633

Leptocleidus, 634, 1173

Leptodactylidae, 593

Leptogenichthys, 374

Leptolepis crassicaudata, 377

Leptolepis gregarious, 376

Leptolepis koonwarmi, 377, 380

Leptolepis lowei, 376

Leptolepis talbragaensis, 376

Leptophloeum, 121

Lepus, 971

Lea, M., 452

Leucocarbo, 751

Leucosarcia melanoleuca, 835

Leucosarcia proevisa, 757, 835

Library of Congress Subject Index,
159

Lichenostomus virescens,839

Lidicker, W., 90, 91

Lightning Ridge Local Fauna, 1014

Lightning Ridge, 391, 473, 474,
618, 633, 641, 646, 651, 663,
664, 688, 894

lignin, 207

lignites, 128, 139

Ligulalepis toombsi, 368, 370, 421,
449, 451, 455

Ligulalepis, 401, 444

limestone caves, 280, 282

limestone, 116, 117

limiting factors, 1170

Limnodynastes archeri, 596

Limnodynastes convexiusculus, 594

Limnodynastes dorsalis, 598

Limnodynastes peroni, 599

Limnodynastes tasmaniensis, 598

Limnodynastes, 593, 667, 669

Limosa lapponica, 1373

lingual tier, 232, 234

Linnean Society, 149, 153

Linquornis, 767

lissamphibians, 570

Lissemys, 714

Lithophaps ulnaris, 835

Litokoala kautjamarpensis, 934

Litokoala, 933

Litoria adelaidensis, 598

Litoria caerulea, 594, 596, 599

Litoria cyclorhunchus, 598

Litoria eucnemis, 599

Litoria ewingi, 598

Litoria infrafrenata, 592.

Litoria lesueuri, 599

Litoria moorei, 598

Litoria splendida, 594

Litoria, 593, 667-669

Liverpool subgroup, 374

living fossils, 444

lizards, 610

Lloyd, A., 54

localities (New Zealand fossil

birds), 1328

Locktite Superglue, 187

loess, 814

loganiids, 439

London Clay, 20, 509, 762

Long Island, 1347

Long Swamp, 481

Long, J.A., 33, 34, 69, 76, 77, 340

longipinnate, 625

Longisquama, 637

Longman, H., 46

Longreach, 474

lophodont, 948

lophodonty, 203

Lophortyx, 823

Lophosteus, 444

Lord Howe Island, 78, 80, 675,
676, 688, 708-710, 716, 748,
762, 1350, 1354

Lord Howe Rise, 1318, 1350, 1354

Loriidae, 835

Lome, 340-342

Los Alamitos Formation, 910

Lotosauria, 637

Lovelle Downes Station, 473, 476

low-crowned molars, 250

low-melt metals, 198

Loxton Sands, 516-518

Loyalty Group, 676

Lunaspis, 366

Lundelius E.L., 33, 34, 50, 52, 57,
75-78

lungfish eggs, 466

lungfish nests, 466

lungfish, 432, 451, 454, 665

Litjanus, 385

Lycopsis longirostrus, 909

Lydekker's Line, 763

Lydekker, R., 8,722

Lyell, C., 24

Lystrosaurus Zone, 578, 583

Lystrosaurus, 614, 621

M-27 Cave, 303

M-29 Cave, 303

Maatsuyker Island, 1347

Mabel Cave, 291, 292-294, 303,
309, 823, 835, 837, 839-841

MacArthur-Wilson equilibrium

theory, 1363, 1364

Machaeracanthus, 451, 453, 459

Machairodontinae, 910

Mackunda Formation, 473, 476,
504, 612

Macleay, A., 12, 13, 24

Macleay, W.S., 12

Macquarie Island, 1810, 350, 1360

Macquarie Island, 810

Macroderma gigas, 286, 305

Macronectes, 1358

Macropodinae, 215-217, 946, 951

macropodines, 215

Macropodaidea, 946

macropodoids, 213, 947

Macropus agilis, 1114, 1134

Macropus atlas, 1112

Macropus birdselli, 1112

Macropus cooperi, 1076, 1112,
1116

Macropus ferragus, 1112, 1116,
1125

Macropus fuliginosus, 1116

Macropus giganteus, 1219, 964111,
1116, 1135

Macropus gouldi, 1112

Macropus ospranter, 112A

Macropus parryi, 1112

Macropus piltonensis, 1112

Macropus robustus, 1112, 1116,
1134

Macropus rufogriscus, 230, 303,
1349

Macropus siva, 1112, 1134.

Macropus titan, 1076, 1112, 1114,
1116, 1118, 1124, 1134

Macropus, 218, 255, 951, 9521074,
1075, 1120, 1123, 1125, 1127,
1128

Macrotis, 929

macrovertebrate assemblage, 283

Madagascar, 386, 629

Madakoala, 900, 933

Madariscus, 376

Madtsoia, 674

Madura Cave, 291

Madura Cave, 291, 293, 295, 303,
325, 820-822, 832, 836, 837

Magadan mammoth, 314, 315

megafaunal extinctions, 820

magnetic reversals, 1169, 1175

magpies, 286

Mahoney, J.A., 33

Main Cave, 1090

Main Site, 481

Main, A.R., 50

mainland islands, 1346

Mair’s Cave, 287, 839

Malacorhynchus membranaceus,
740, 1329

Malacorhynchus scarletti, 1329

Malay Archipelago, 763

Malkuni Fauna, 479, 483

Mallee Ridge, 814

Maluridae, 839

Malurus cyaneus, 303

Malurus, 839

Mamenchisaurus, 198

mammae, 895

mammalian camivores, 302

mammalian predators, 284
mammalian scavengers, 284, 302
mammalian-accumulated deposits,
284
Mammalodon colliveri, 1181
Mammalon Hill, 252, 253, 254,
255, 329, 480
mammillary cores, 873, 880
mammnillary Surface, 873
Mammoth Cave, 304, 1088
mammoth, 330
Mampuwordu Sand, 52, 60, 475,
479, 483, 667
man, 1334
Manabla Sandstone, 354
Manarrina Hill, 474
Manarrina, 473
Mandagery Sandstone, 386
mandibular foramen, 224
Maadrillus, 210
Mandu Calcarenite, 513, 515
Mangaia, 1355
Mangere Island, 1330
Manilietta crassa, 375, 378
Manlietta, 376
Mannum Formation, 513
Mannum, 513
Mansfield Basin, 340-342
Mansfield Group, 351, 397
Mansfield, 340, 370, 388, 389, 421,
451,452
Mansfieldiscus sweeti, 371, 421
Mansfieldiscus, 370
Manu antiquus, 1327, 1360
manual dexterity, 256
Maoris, 320, 1356, 1357
Marburg Sandstone, 572, 573, 586
Marcus, L.F., 50
Maree Formation, 611, 646
marine mammals, 1167
marine vertebrates, 1166
Marsdenichthys longioccipitus, 421
Marsdenichthys, 401
Marshal] Plan, 47
Marshall, L., 69
marsupial lion, 210
marsupial radiation, 894
Marsupialia, 924
marsupials, 896
Martins Well Limestone, 450, 452,
459
Mary River, 484
Mascarene Islands, 1363
masseteric crest, 224
masseteric foramen, 224, 935
masseteric fossa, 224
masseteric processes, 1140
Massospondylus, 645
Mastacomys fuscus, 1349
Masters., G., 12
mastication, 203, 206
mastodont, 8, 330
Matuyama/Bruhnes magnetic
boundary, 813, 814, 816
Maude Formation, 512
Mauicetus brevicollis, 1182
Mauicetus lophocephalus, 1182
Mauicetus parki, 1181, 1182

Mauicetus waitakiensis, 1182

Mauisaurus haasti, 1174

Mawby, J., 54, 723

maxillolabialis muscles, 1102

Maxwellton, 645

Mayr, E., 763

McCoy, F., 8, 13, 15, 17, 22, 150,
151

McDonalds Bank, 518

McDonalds, 735

McEachem’s Cave, 324-326, 723,
745, 834, 837

McEvey, A.R., 723, 810

McFadden, B., 80

Mcmurdodus featherensis, 347

Mcmurdodus whitei, 346, 458

Mcmurdodus, 401, 440, 448

McNamara, G., 723,

McNamara, K., 340

mechanised screen 192

Meckels cartilage, 896

megachiropterans, 958

Megadermatidae, 958

Megadyptes antipodes, 1326

Megadyptes, 749

Megaegotheles, 834

megafauna (Australian), 1126

megafauna (measurements), 1150-
1158

megafauna, 82, 215, 1072, 1073,
1076, 1083, 1129, 1131, 1132,
1142, 1143

megafaunal communities, 1111,
1128

Megalania prisca, 673,678, 741,
927, 1076, 1128, 1133

Megalania, 24, 668, 69, 671, 677,
1074, 1079, 1080

Megalanocosaurus, 637

Megalapteryx benhami, 1325

Megalapteryx didinus, 313, 315

Megalapteryx, 1321

Megalichthys, 388, 401, 421, 445,
451,455

Megalops lissa, 384

Megalops, 382

Megalosauropus broomensis, 611,
660

Megalurus timoriensis, 839

Megapleuron zangleri, 484

Megapodiidae, 758, 766-768, 829

megapodiids, 1086

Megapodius reinwardt, 758 824,
826, 828

Megapodius, 758, 1355

Megaptera, 1180

Megapteriscus, 376

Megascyliorhinus cooperi, 517

Megistolotis, 593, 594, 595

Meiolania mackayi, 713, 1355

Meiolania oweni, 675, 676, 689,
710

Meiolania platyceps, 676, 688, 710,
712, 716, 1354

Meiolania, 78, 80, 667, 669

Meiolaniidae, 708, 709

Mekosuchus inexpectatus,1355

Melboume Punch, 13

Melboume Univessity, 479, 722

Meliornis, 1333

Meliphaga, 1333

Meliphagidae, 841

Melithreptus, 841

Mellinjene Limestone, 399

Melody’s Maze, 481

Melomys cervinipes, 960

Melopsittacus undulatus, 820, 836

Menaspis, 442

menaspoids, 442

Meniscoessus robustus, 252, 898

mental foramen, 224

Menuridae, 762, 767, 837

Meredith, C.W., 69, 723, 810

Mereenie Sandstone, 119

Mergus australis, 1320, 1329

Merino Formation, 714

Merino Group, 612

Meriones, 204, 205

Merluccius fimbriatis, 384

Merluccius, 382

Menmtilees, D., 34

Merniman’s Creek, 514

Merrimbula Group, 452

Mesembroniscus, 376

mesenosaurs, 609

Mesenosaurus romeri, 638

mesodentine, 442

Mesodmus, 451

Mesonychidae, 1179

Mesoplodon longirostris, 1180

Mesoplodon, 1180

mesosaurs, 609, 613, 686

mesosuchians, 641

metabolic requirements, 205

metacones, 234

metaconid, 233

metacristid, 240

metaloph, 216

Metapteryx bifrons, 824

Metaxygnathus denticulus, 576

methacrylate glue, 182

Microbiotheria, 924

Microcetus hectori, 1182

microchiropteran, 958

Microcorax, 507, 525

microfloral analysis, 822

microfossils (vertebrates), 430

micrographs, 231

Microhylidae, 593, 598

microphagous, 346

microphagy, 350

Micropholis stowi, 578

microstriations, 304

Microtinae, 203

microvertebrate assemblages, 268

microwear anal ysis,214

microwear, 210, 215

midlink, 216, 224

Midway Point, 470

mihirung eggshell, 878

mihirung, 879, 886

Milankovitch parameters, 1169

Miles, R. 33

Miller, A. H., 47, 53, 57, 58, 90,
91, 723, 811

Miller, V., 47

INDEX - 1429

Milne-Edwrds, H., 149

Milvus migrans, 287

Mimia toombsi, 368

Mineral Hill Volcanics, 434, 447

Mineral Hill, 434

Mining and Metallurgy, Institute of,
20

mining boom, 11

mining of gold, 11

Minmi paravertebra, 612, 651, 655

Miami, 613, 656, 663, 664, 689

Miolania, 1128

Miralina, 900, 945

Miralinidae, 945

Miria Formation, 509

Miria Marl, 510

Miro, 1333

Mirrabooka Formation, 447

Mitchell, T. L., 4-6, 5, 9, 722

Mithaka Waterhole, 340-342

mitochondrial DNA, 314

Mitsukurina maslinensis, 511,514

Mixophyes fasciolatus, 599

mixosaurs, 625

Mixosaurus timorensis, 625

moa, 323, 327

Moa-hunters, 1325

Mobil Oil, 735

Mogorafugwa, 1045

Mohoua albicilla, 1333

Mohoua ochrocephala, 1333

molar mophotype, 209

molar morphology, 230, 232

molar occlusion, 255

molar structure, 203

molasse, 276

Mole Creek, 745

Molecap Greensand, 349, 506, 508

molecular clocks, 312

molecular fingerprinting, 332

molecular phylogenies, 332

Molnar, R., 33, 69, 76

Molong High, 446, 447

Molossidae, 958

Molybdichthys junior, 375, 378

Molybdichthys, 376

Monarolepis verrucosa, 363

Monarolepis, 364, 400

Monash Univesity, 59, 69, 70, 188,
7123

monitor lizards, 286

monochloracetic, 430

Monodia, 204

monognathic heterodonty, 502

monophyletic taxa, 1167

Monotremata, 922

monotremes, 252, 622, 894, 896,
1143

Montagu, 1099

Montypythonoides riversleighensis,
675

Montypythonoides, 669

Moorna Formation, 483, 669

Moorna Station, 479

Mootwingee, 340-342

Moray eels, 391

Morelia antiquua, 668, 674, 675

Morelia, 670

1430 - INDEX

Morgan Limestone, 383

Morgan, 329, 513-515

Momington, 515

Morobe Gold Field, 86

Morison Formation, 316

Morrison, P., 75

Morus serrator, 1349

Morwell Local Fauna, 1057

Morwell, 730, 783, 830

mosasaur, 623, 624, 665, 1173

Mosasauridae, 1171

Mosasaurus mokora, 1174

mould supports, 197

moulding,193

moulds,193,198, 199

Mountain Limestone, 4

Mountain Zebra, 315

Mourer-Chauvire, C., 767

Mowbray Swamp, 745

Mowilith 144,187,188

Mowital B30, 430

Mowotil 7001, 144, 182

Moythomasia durgaringa, 368

Moythomasia, 444, 455

Mt Charlette, 340-342

Mt Deerina, 340-342

Mt Frome Limestone, 434, 448,
459

Mt Frome, 434

Mt Gambier Limestone, 515

Mt Gambier, 24, 511, 515, 747

Mt Grenfell Station, 340-342

Mt Howitt Fauna, 394

Mt Howitt, 338, 340-342, 351, 361,
362, 370, 390, 393, 395, 397,
401, 402, 421, 452

Mt Ida Formation, 452

Mt Jack, 340-342

Mt Kent Conglomerate, 398

Mt Knowles Limestone, 434

Mt Knowles, 434

Mt Kosciusko, 819

Mt Morgan, 633, 634, 660

Mt Tambo, 340-342

Mulga Downs Formation, 447

Mulga Downs Group, 354, 368,
390, 397, 398, 399, 434, 452

Muller’s Line, 763

multituberculates, 250, 252, 898,
910, 922, 945

Mulvihill, J., 160

mummification, 313

mummy, 315

Munyarai sequence, 399, 434

Munyarai Formation, 434

Munyurai, 437

Muraenesox, 382

Muramura williamsi, 937

Murda-Meloola, 434

Murgon, 452

murid rodents, 136

Mundae, 763, 959, 961

Mumdel HLS., 24

Murray Basin, 126, 128, 512, 515

Murray Bndge, 513

Murray Group, 517

Murray River, 69

Mumay, P., 33, 69, 77, 726

Murray-Darling catchment, 831

Murray-Darling plains, 236

Murray’s Line, 763

Murrindalaspis bairdi, 420

Murrindalaspis, 367, 450

Murrumbidgee Group, 354, 391,
448

Museum of Victoria, 8, 13, 15, 16
69, 70, 151, 188, 236, 340

Mus musculus, 128, 303, 307,
308,138

mustelids, 1319

Mustelus, 518

Muttaburrasaurus langdoni, 650,
652

Muttaburrasaurus, 612, 613, 651,
653, 654, 657, 663

Myer Foundation, 735

Myliobatis, 349, 501,511, 512,
518,524

Mylostomatidae, 355

Myobatrachus gouldii, 600

Myobatrachus, 593

myodomes, 368

Myosaurus, 614

Myriolepis clarkei, 376

Mymmecobiidae, 927

myrmecobiids, 929

Myrmecobius fasciatus, 927

Myrtlevale Formation, 452

mysticestes, 1181

Mysticeti, 1171

Milllerian ducts, 902

N’Dahla Member, 399, 434, 436,
437

N.S.W. Geol. Sur., 154, 155

Naiggani, 1355

Namba Fm., 140

Namba Formation, 73, 252, 140,
475, 478, 480-483, 475, 490,
596, 762, 931, 1183

Namilamadeta snideri, 938, 940

Namoi Formation, 451

Nanantius eos, 737

Nanantius, 738

Nanarup Limestone, 511

Nangetty Formation, 123

Napier Formation, 367

Napperby, 140

Narabeen Group, 374

Naracoorte Cave, 667

Naracoorte, 76, 745

nares, 466

Narrabeen Group, 467, 470, 572,
573, 583

Narrien Range, 388, 451

nasohypophysial foramen, 343

nasomaxillolabialis muscles, 1106

National Geographic Society, 75,
80, 81, 151, 735

native cats, 286

Natural Trap Cave, 291
290, 292

Neales River, 612

near-polar dinsoaurs, 664

Necrolestes patagonensis, 913

Nelson Bay, 329

Neobatrachus centralis, 600

Neobatrachus pictus, 598

Neobatrachus, 593, 598, 666, 668

neoceratodont, 484, 485, 490

Neoceratodus djelleh, 475, 478,
480, 483, 490

Neoceratodus eyrensis, 479, 480,
481, 483, 490

Neoceratodus forsteri, 391, 466-
468, 471, 474, 477, 479, 480,
483-486, 490

Neoceratodus gregoryi, 475, 477-
481, 483, 484, 490

Neoceratodus nar gun, 474, 476,
477, 480, 482, 483, 490

Neoceratodus, 445, 481

Neohelos tirarensis, 1079, 1080

Neophema chrysostoma, 836

Neophema elegans, 836

Neophema splendida, 836

Neophoca cinerea, 1178

Neoplatycephalus bassensis, 382

neoselachian sharks, 448,452, 498,
500

Neoselachii, 500

Neositta chrysoptera, 840

Neosittidae, 840

Neotoma, 286

Nepean Island, 1351

nest environment, 879

Nestor meridionalis, 1323, 1332,
1357

Nestor notabilis, 1332, 1356

Nestor productus, 1332, 1373

nettosuchians, 639

neural arch, 570

Neville’s Nirvanah, 783

New Caledonia, 676, 708, 713,

758, 835, 318, 1331, 1346, 1350,
1355, 1362

New Guinea, 726, 730

New Ireland, 835

New World, 735

New Zealand avifauna, 1318

New Zealand, 1350

Newcastle Coal Measures, 371,

572, 573, 576, 583

Newcastle, 340-342

Newsome, A., 90

Ngama Local Fauna, 667, 757, 769,
1019

Ngapakaldi Fauna, 80, 480, 596,

598, 958, 1016, 1018

Ngapakaldi Local Fauna, 666, 739,
753, 757, 760

Ngapakaldi Quarry, 57, 59

Negapakaldia tedfordi, 59, 936, 937,
939

Neapakaldia, 44, 1078, 1106

Nikolivia, 439

nikoliviid, 439, 447, 448

Nimbacinus dicksoni, 927

Nimbadon, 1079

Ningbing Limestone, 358

Ninox conaivens, 834

Ninox novaeseelandiae, 281, 289,
307, 1332

Ninox undulata, 1373

Niolamia, 676

nitrocellulose membrane, 328

Noah’s Ark, 133, 766

Noakes, L.C., 86

nocturnal habits, 256

Nodocosta, 453

Nolan, J., 28

nomina dubia, 1181

non-reefal limestones, 116

nonruminant artiodactyls, 208

Nooraleeba Local Fauna, 1029

Nora Formation, 446

Norfolk Island, 1723, 762, 810,
811, 332, 1350, 1354, 1355,
1357, 1362, 1363

Norfolk Rise, 1350

Normanton Formation, 613

Normanville, 329

North Cape, 1327

Northem region fauna, 1127

nostolepid, 459

Nostolepis costata, 450, 453, 455

Nostolepis striata, 350, 453, 455

Nostolepis, 420, 443, 447, 450,
459

Notaden melanoscaphus, 600

Notaden, 593

notarium, 657

Notechis, 667

Nothofagus johnstonii, 738

Nothofagus, 140, 142

nothosaurs, 609, 629

nothosaurs, 629

Notidanoides muensteri, 499

Notiomystis, 1333

Notobrachyops picketti, 583

Notochelone costata, 613, 617, 618,
713, 1173

Notochelone, 619, 627, 663

Notogoneus parvus, 385

Notomys, 911

Notoryctemorphia, 931

Notoryctes typhlops, 932, 956

Notoryctidae, 931

Notorynchus aptiensis, 508

Notorynchus cepadianus, 516, 525,
526

Notorynchus primigenius,

Notorynchus, 499, 502, 524

notosaurs, 628

Nototheirum tasmanicum, 1098

Nototherinae, 935

Nototherium , 8

Notothertseum inerme, 1104, 1106,
1143

Nototherium mitchelli, 1098

Nototherium victoriae, 1098

Nototherium watutense, 86, 87,89

Nototherium, 1073, 1128

Novogonatodus kazanisavae, 371

Novogonatodus, 370

Nullarbor Plain, 157, 291, 309,
340-342

Numenius phoeops, 1373

Nummopalatus depressus, 383

nutritional demands, 208

Nyctmene major, 1069

Nyctimystes sweifeli, 5099

Nyctimystes, 593

525

Nyroca effodiata, 757, 835
018/016 ratios, 130
Oakley Creek, 329
Oakover beds, 75
Obdurodon insignis, 258, 923, 924
Obdurodon, 71
occlusal cycle, 235, 238
occlusal design, 208
occlusal maps, 231
occlusal mechanics, 218
occlusal surface, 255
occlusal wear facets, 255
occlusion, 230, 232
occupation middens, 1321
oceanic islands, 1346
oceanic nutrients, 1175
Ocyphaps lophotes, 822
Ocyplanus proeses, 753, 754, 755,
756, 832
Odantaspis acutissima, 516
Odobenidae, 1178
Odontaspis, 349, 501, 502, 527
odontocetes, 1181, 1184
Odontoceti, 1171
odontode, 440
Officer Basin, 122, 125, 399, 434
Ohioaspis , 442, 448, 450
Ohiolepis, 438, 448, 455
Olanga Calcilutite, 509
Old Beach, 470
oligotrophic ecosystem, 1131
Olney Formation, 511
Olney No. | Bore, 511
Olson, S.L., 723, 766, 811
Ommataphoca rossi, 1179
omnivores, 208
omnivorous diet, 249
Onchodus, 445
onychodontid, 391, 455
Onychodontiformes, 369, 386
Onychodus, 390, 391, 445,
oolites, 126
oolitic limestone, 116, 117, 119,
122
Oplegnathus manni, 383
Orange District, 447
orangutan, 332
orbital cavity, 255
orbits, 256
Orbost, 340-342, 515
Orcinus, 1182
Ord Basin, 116
orectoloboid, 500
Orectolobus gippslandicus, 514,
516
Orectolobus, 526
Oriental Biogeographic Realm, 136
Oriental biota, 763
Oriental Region, 598
Origma solitaria, 281
Orlov, A., 164
omithischians, 610, 642, 648, 649,
658
Ornitholestes orntiholestoides, 646
omithopods, 648, 650, 661, 662
Omithorhynchidae, 922
omithorhynchids, 1084
Ornithorhynchus anatinus, 252,

458

923,924

Ornithorhynchus, 258

omithosuchians, 609, 637, 639

Omithosuchidae, 637

omithosuchids, 610, 686

orodonts, 451

Orodus, 347, 442,451

Orthacanthus, 440

orthacodontids, 500

orthodentine, 444

Orthodonta, 500

Orthonychidae, 762, 838

Orthonyx hypsilophus, 819, 838

Orthonyx spauldingi, 838

Orthonyx temminchi, 838

Orthonyx, 819

Orthopteryx, 615

Orvikuina, 444

Oryctolagus cuniculus, 303, 822

Osphranter, 1128

Osteichthyes, 345, 352, 367, 369,
498

osteocalcin, 316, 317, 323-325

osteodentine, 500

Osteodonta, 500

osteolepids, 388

Osteolepiformes, 368, 369, 386,
454, 445

osteostracans, 431

Osteostraci, 343

ostrich, 327

ostrich eggs, 878

Otariidae, 1171, 1178

Othnielia, 650

Otibanda Formation, 55, 90, 91, 98

otic capsule, 615

Otididae, 760, 831

Otodus obliquus, 511, 518,527

Otodus, 349

otoliths, 368, 382, 384

otters, 258

Otway Basin, 126, 128, 510, 513

Otway Group, 377, 380, 474, 476,
586, 612, 617, 646, 658, 714,
737

Otway Ranges, 660, 738, 841

Otway region, 340-342

Overkill model, 1142

Overland Comer, 814

oviraptorosaurs, 621

Ovis aries, 303

Ovis, 822, 968

Owen, R., 6, 8, 10, 27, 28, 149,
150, 722, 1143

owls, 286

Oxley Group, 669

oxygen isotopes, 112, 113

Oxynotus, 510

Oxyura australis, 1329

P.VA., 188, 192

Pachyanas chathamica, 1329

Pachyanus, 1359

Pachycephala pecioralis, 1373

pachycephalosaurs, 648-650

Pachydyptes simpsoni, 738, 747,

Pachydyptes, 748

Pachymylus, 503

Pachyornis australis, 1325

Pachyornis elephantopus, 318
pachyosteomorphs, 355
Pachyplichas jagmi, 1333
Pachyplichas yaldwyni, 1333
Pachyptila belcheri, 734, 735
Pachyptila desolata, 1360
Pachyptila turtar, 1350
Pachyptila, 1350, 1358
Pachyrhizodontidae, 382
Pachyrhizodus marathonensis, 381
Pacific Rat, 1354
Padaeosaurus, 614
Paine Quarry, 68
Palaecorax moriorum, 1334
Palaeeudyptes macraei, 1176
Palaeeudyptes marplesi, 1177
Palaeeudyptes, 615, 738, 748, 1176
palaelodid, 783
Palaelodidae, 739, 768, 819, 834
Palaelodus gracilipes, 7S7
Palaelodus, 756, 757, 783, 834
Palaeloduscrassipes, 783
palacobiogeography, 765
palaeoclimate, 1166
palaeogeography, 1166
Palaeohypotodus, 507
Palaconiscidae, 444
palaconiscoid, 452, 368, 370, 372,
451
Palaeoniscus antipodeus, 376
Palaeoniscus crassus, 376
Palaeoniscus feistmanteli, 376
Palaeoniscus randsi, 370
Palaeoniscus, 375
Palaeopsittacus georgei, 762
Palaeortyx, 767
palacosol, 252
palacospinacids, 500
Palaeospinax, 507
Palaeudyptes, 1177
Palankarinna Fauna, 483
Palankarinna Local Fauna, 667,
745, 1050
Palankarinna South Local Fauna,
1017
Palecanus erythrorhynchos, 751
paliguanid, 609, 623
Pallimnarchus pollens, 671, 672,
678
Pallimnarchus, 668-671, 677, 1077
Palorchestes azeal, 1078, 1081,
1103, 1106, 1107, 1164
Palorchestes novaculacephalus,
1106
Palorchestes painei, 1078, 1079,
1081
Palorchestes parvus, 1081, 1164
Palorchestes, 1073, 1077, 1083,
1116
Palorchestidae, 935
palpebral cartilage, 648
palynomorphs, 400
Pambula River site, 364, 389
Pambula, 452
Pambulaspis cobahdrensis, 364
Pambulaspis, 402
Panama Isthmus, 1169
Pancake Site, 330

INDEX - 1431

pandas, 215, 215

Panderichthyidae, 369

Pangaca, 587

pantadactyl limb, 570

Panthera leo, 678, 1075, 1138,
1139

Panthera pardus, 304

Panthera spelaeus, 1075

Papio, 209, 210

Paraceratodus germaini, 467

paracones, 234

paraconid, 233

paracristid,233

Paracyclotosaurus david, 574, 578,

583
Paraisurus macrorhiza, 507
parapatric species, 840
parapsid, 606, 624
paratheres, 910
Pardalotidae, 841
parental care, 466
parieasaurs, 613, 686
Paris Basin, 509
Parker, S., 723, 1348
Parksosaurus, 664
Parotodus, 501
Parotosuchus aliciae, 574, 578,
584, 585
Parotosuchus brookvalensis, 578,
583
Parotosuchus gunganj, 574, 584
Parotosuchus, 576, 577
Passer, 823
Passeriformes, 762, 768, 911
Patagonia peregrina, 911,914
Patagonia, 512
Patagoniidae, 911
Patene, 910
Patterson, C., 723
Paulik’s Bore, 517
pavement teeth, 501
Peak Downs, 730, 781
Pearce, F., 73
peat bogs, 276
peat, 121, 139
peat-forming swamps, 142
Pebble Point Formation, 510
pectin, 207
Pediomyidae, 905
pediomyids, 907
Pediomys cooki, 905
Pedionomidae, 766-768, 830
Pedionomus torquatus, 760, 783,
820
Pedionomus, 810, 811
pedogenic carbonate, 880
Peel’s Bore, 511
Pelagodroma marina, 1354, 1373
Pelagodroma, 1358
Pelecanidae, 739, 750, 768, 827
Pelecaniformes, 1327
Pelecanoididae, 826
Pelecanoides urinatrix, 826
Pelecanoides, 1354, 1358
Pelecanus conspicillatus, 750
Pelecanus grandiceps, 750
Pelecanus minor, 748
Pelecanus cadimurka, 749, 750,

1432 - INDEX

751, 783, 827
Pelecanus conspicallatus, 782, 749-
751, 783, 827
Pelecanus crispus, 751
Pelecanus grandiceps, 783
Pelecanus novaezealandiae, 749,
751
Pelecanus onocrotalus, 751
Pelecanus philippensis, 751
Pelecanus proavus, 749, 750
Pelecanus refescens, 751
Pelecanus tirarensis, 749-751, 783
Pelecanus validipes, 750, 751, 783
Pelecanus, 811
peleothyrids, 609
pellets, 287, 288
Pelodryas, 594
Peltobatrachus pustulatus, 580
Peltohyas australis, 832
pelycosaur, 620, 638
Penola, 24
Pentland, J. B., 6
Pentland, W., 8
penultimate interglacial, 819
Peppermint Grove Limestone,516
pepsin digestion, 317
Peramelemorpha, 929
Perameles gunnii, 303
Perameles, 234, 931
Peramelidae, 230
peramelids, 236, 929-932
Perameloidea, 239, 1086
Percalates antiquus, 385
pericarp, 258
Perikoala palankarinnica, 933
Perikoala, 933
periodical literature, 159
perissodactyls, 208, 664
permafrost, 276
Pernjara Group, 399
Perth Basin, 122-124, 126, 506
petalichthyids, 354, 365
petalodont, 442, 451
Petalodus, 442
Petaurinae, 955, 956
Petauroidea, 955
Petaurus breviceps, 956
Petaurus, 956
petrifaction, 276
Petroica macrocephala, 1333
Petroica, 1333
Pezoporus wallicus, 296, 836
Phaerodus, 384
Phaeton rubicauda, 1373
Phalacrocoracidae, 739, 751, 827
Phalacrocorax gregorii, 827
Phalanger orientalis, 261
Phalanger, 252, 257, 944
Phalangerida, 944, 1086
Phalangeridac, 256, 944
Phalangeriformes, 947
phalangeroid, 252, 255
Phalangeroidea, 944
Phaps chalcoptera, 294, 835
Phaps elegans, 835
Phaps histrionica, 835
Phascolarctidae, 932, 945
phascolarctids, 945

Phascolarctomorphia, 932

Phascolarctos cinerus, 933, 934,
1094

Phascolarctos maris, 934

Phascolarctos stirtoni, 1094, 1134,
1143

Phascolomys magnus, 1094, 1096-
1098

Phascolomys medius, 1094, 1096-
1098, 1143

Phascolomys, 1128

Phascolonus gigas, 942, 943, 1075,
1094, 1096-1099, 1128, 1136,
1137

Phascolonus medius, 1094

Phascolonus, 1073, 1074, 1077,
1128

Phascolotherium. bucklandi, 897

Phasianidae, 767, 829

Philesturnus caruaculatus, 1334

Phillip Island, 326

Philoria frosti, 600

Philoria, 593

Phlyctaenichthys pectinatus, 377,
378

Phlyctaenichthys, 376

phlyctaenioid, 355

Phoberodon, 1182

Phocaenopsis mantelli, 1182

Phocarctos hookeri, 1178

phocid, 1172

Phocidae, 1171

Phocoidea, 1178

Phoebetria, 1357, 1358

phoebodont, 449, 455

Phoebodus austaliensis, 449

Phoebodus, 347, 348, 401, 440

Phoeniconaias gracilis, 755, 756

Phoeniconaias minor, 752-756

Phoeniconotius eyrensis, 753, 755

Phoenicopteridac, 739, 753, 768,
811, 832

phoenicopterids, 811

Phoenicopterus novaehollandiae,
753, 755

Phoenicopterus ruber , 752-756,
832

phosphatic microfossils, 430

phreatic caves, 277

phyllolepid, 336, 354

Phyllolepis, 358, 362, 401, 582

phylogeny (Australian mammals),
962

phylogeny (turtles), 706, 707
Physeter macrocephalus, 1183
Physeteridae, 1180
Physetodon baileyi, 1180
Physignathus, 669, 673
phytoliths, 249

Phytosauria, 637

Pidinga Formation, 140
Pigfooted Bandicoot, 230
Pilbara region, 75, 305
Pilkipildridae, 956

Piltdown Man, 332
pinnepeds, 1086

Pinnipedia, 1178

Pinpa Fauna, 1021

Pinpa Formation, 480

Pinpa Local Fauna, 739, 750, 757

Pinpa, 938

Pioneer, 781

pisolitic limestones, 116

pistosaurs, 631

pitfall caves, 279-281, 283, 291,
326

Pitikantia, 1078

Pitt Island, 1330, 1356

Placentalia, 958

placentals, 896

Placodermi, 345, 430, 442, 444,
448, 454, 498

Placodonta, 606

placodonts, 609, 624

placaid scale, 441, 448

placoid structures, 439

Placolepis budawangensis, 361

plagiaulacoid premolars, 214

plagiaulacoid, 258, 947

Plagiobatrachus australis, 580

Plaisiodon centralis, 1079

planation surfaces, 276

Plane, M., 55, 61, 73, 75, 76, 726

Planigale, 289

plaster jacketing, 192

Plaster of Paris, 192, 197, 198

plastic glues, 187

plasticine, 197

Platalea leucorrhoa, 827

Platalea subtenuis, 752, 827

plate tectonics, 765

Platecarpus, 624

Platycephalus bassensis, 383

Platycephalus petilis, 384

Platycephalus, 382, 383

Platycercidae, 836

Platycercus caledonicus, 309

Platycercus elegans, 281, 836

Platycercus eximius, 309, 836

Platycercus icterotus, 836

Platydyptes amiesi, 1177

Platydyptes marplesi, 1177

Platyosphys, 1181

Platyplectron, 668

Platypterygius australis, 613, 626-
628, 1173

Platypterygius, 611, 1173

platypus, 258

platysomids, 444

Platysomus, 376

Plectropterna, 659, 660

Pledge, N., 33, 73, 76, 77, 340

Plesiocetus dyticus, 1182

plesiosaur, 633

Plesiosauria, 1171, 1173

Plesiosauroidea, 1171

plesiosauroid, 609, 628, 630

plesiosaurs, 628-630, 665, 1173

Plesiosaurus macrospondylus, 631

Plesiosaurus rostratus, 629

Plesiosaurus sutherlandi, 631

Plesiosaurus, 630

pleuracanth, 346, 349

Pleuracanthus parvidens, 347

pleurocentra, 570, 571

pleurocoels, 643

pleurodires, 617, 619
Pleurodus, 442

Pleuronectes, 382
Plicatolamna, 507

Plioaetus, 757

Ptiosauroidea, 1171
pliosauroids, 609, 628, 631
Ploceidac, 841

plourdosteids, 355
Plourdosteus, 355

Pluvialis dominica, 1373
Podargidae, 761, 767, 834
Podargus strigoides, #7, 834
Podargus, 761

Podiceps cristatus, 1326
Podiceps rufopectus, 1326
Podicipedidae, 746, 825
Podicipediformes, 1326
Poephila guttata, 841
Poggendorf, J.C., 154

Point Addis Limestone, 511
Point Lewis, 474

Point McDonnell, 515

polar ice, 1169

polar latitudes, 726

Polda Basin coals, 124

pollen assemblages, 817
polyacrylamide gel, 323, 326
polybutylmethacrylate, 175
polycotylid, 632
Polydolopidae, 910, 911
polyesters, 198
Polyglyphanodon, 607
Polynesian dogs, 1319
Polynesian man, 1353
Polynesian Period, 1358
Polynesian Rat, 1319
Polynesians, 1364
polyprotodont, 927, 1086
Polyrhizodus, 442
polyurethane foam, 175, 187, 198
polyvinyl acetate, 188, 430
Poposauria, 637

population irruptions, 296
population structure, 1142
Poracanthodes, 443, 453
pore-canals, 874

pore-canal system, 368
Porolepiformes, 368, 369, 386, 454
porolepiforms, 445
Porolepis, 445
Porophoraspis crenulata, 446
Porophoraspis, 343, 433
Porphyrio kukweidi, 1355
Porphyrio melanotus, 1321, 1363
porpoises, 901

Port Augusta, 888

Port Campbell Limestone, 513
Port Campbell, 513

Port Jackson Shark, 514

Port Phillip district 11

Port Pirie, 329

Port Willunga Formation, 512
Portland Local Fauna, 917, 946
Portland, 191, 253, 517
Porzana fluminea, 830
Porzana tabuensis, 831
Porzana, 1331

Postmetacrista, 236
postparacrista, 236
postalveolar fossa, 224
postalveolar process, 224
postcrista, 255, 257
posthypocristid, 233
postmetacrista, 234, 240
posttemporal fenestra, 608, 615
Potoroidae, 215, 947
potoroines, 215
Potorus tridactylus, 303
Pragian Limestone, 452
Pratt-Winter, S., 24
Precambrian, 113
Precipice Sandstone, 611, 659
precrista, 255, 257
predator diversity, 1133
predator species, 286
predator, 285, 287
preferential preservation, 272
prehypocrista, 233
prehypoctistid, 233
premetacrista, 236
preparacrista, 234, 236, 240
prepubis, 648
Preservation Island, 1347
prey species, 283
Priacodon, 897
primary journal, 159
primary publications, 157-159
primates, 256, 259, 958, 961
Princetown, 510, 511
Prionotemaus, 250, 1112
Priscileo pitakantensis, 1081
Priscileo, 1944, 079, 1109
Pristiophorus lanceolatus, 513,
516, 517, 526
Pristiophorus tumidens, 506, 524
Pristiophorus, 500
Procelariidae, 825
Procellaria, 1327
Procellaniformes, 1327
proceratodontid, 484
Proceratodus carlinvillensis, 471
Procheirichthys ferox, 375
Procheirichthys, 376
Procolophon trigoniceps, 616
Procolophon, 608, 614
procolophonians, 609, 613, 686
procolophonid, 583, 663
Procoptodon goliah, 219, 223, 224,
953, 1114, 1118, 1121, 1123-
1125, 1128, 1143, 1159
Procoptodon pusio, 218, 223, 1112,
1114, 1116, 1125
Procoptodon rapha, 219, 223, 953,
1114, 1118, 1125
Procoptodon texasensis, 1114, 11 25
Procoptodon, 1128, 215, 219, 250,
950, 073, 1074, 1077, 1127,
1128, 1133
procumbent incisors, 217
proganochelydians, 617
Proganochelys, 608
Prognathodon waiparaensis, 1174
Progura gallinacea, 826, 828, 829,
867
Progura naracoortensis, 758, 824,

826, 828, 829

Progura, 758, 819, 820

Prolacerta broomi, 614, 635

Prolacerta, 636

prolacertids, 635

prolacertiform, 638

Prolacertiformes, 609, 623, 634,

635, 638, 663, 686

Promecosomina beaconensis, 376

Promecosomina formosa, 376

Propalorchestes novaculacephalus,
1078

Propalorchestes, 1079

Properamedlidae, 931

Propleopinae, 948

propleopines, 215

Propleopus chillagoensis, 1111

Propleopus oscillans, 200, 213,
949, 1111, 1112, 1114, 1116,
1133

Propleopus, 213, 214, 880, 945,
951, 1213, 1073, 1074, 1076,
1079, 1128

prosauropods, 643, 644

Prosqualodon davidis, 1180-1183

Prosqualodon hamiltoni, 1182

Prosqualodon marplesi, 1182

Prosqualodon, 1180, 1185

protacrodonts, 449

Protacrodus, 348, 440, 455

Proteaceae, 140

Protemnodon anak, 1112, 1116,
1118, 1365

Protemnodon brehus, 1112, 1116,
1127

Protemnodon buloensis, 98

Protemnodon otibandus, 93, 94,
1114, 1124, 1134

Protemnodon roechus, 1112, 1116,
1118

Protemnodon, 90, 250, 950-952,
1073, 1174, 1176, 1179, 1114,
1120, 1135

Proterochampsidae, 637

Proterosuchia, 637

proterosuchian, 609, 638, 639

protocone, 233, 234, 236, 237, 907

protoconid, 947

Protogonacanthus, 351

Protolamna, 508, 525

protoloph, 216

protolophid, 216, 224

Protopterus protopteroides, 486

Protopterns, 391, 466, 467

protorosaurs, 609, 623, 635

protoroth yrids, 607

Protosauria, 623

Protosqualus, 508, 524

Protungulatum community, 907
proventriculus, 284

proximal caecum, 240, 241
proximal colon, 240, 241
psammodont, 451

Psammomys, 204, 205
Psephotus pulcherrhinus, 82.2

Psephotus varius, 820, 836
Pseudaptenodytes macraei, 748
Pseudaptenodytes, 748
Pseudapteryx gracilis, 1326
Pseudechis, 667
Pseudemydura, 705
Pueudocheiridae, 955
peeudocheirids, 257, 916, 945, 947
pseudocheirines, 932, 945
Pseudocheirus peregrinus, 955,
1349
Pseudocorax australis, 507
Pseudocorax, 518
Pseudodoniornis stirtoni, 1360
Pseudodontorns, 1327
pscudofossil, 617
Pseudomys vandycki, 959
Pseudonaja,.673
Pseudophryne bibroni, 00
Pseudophryne guentheri, 598
Pseudophryne, 593
Pseudorca, 1182
Pseudosuchia, 637
pseudosuchians, 609
peeudovagina, 902, 903
Psilichthys selwyni, 377
Psittacidae, 768
Psittaciformes, 740, 762, 1331
Psophodes nigrogularis, 839
Psophodes olivaceus, 839
Psuedomys, 961
psychrosphere, 1185
Pteranodon, 658
Pteraspidomorphi, 343
pterodactyloids, 657
Pterodactylus, 658
Pterodaustro, 658
Pterodroma axillaris, 1363
Pterodroma cahow, 1363
Pterodroma hypoleuca, 1327, 1363
Pterodroma inexpectata, 1355
Pterodroma magentae, 1355, 1357,
1359
Pterodroma nigripennis, 1363
Pterodroma pacificus, 1373
Pterodroma pycrofti, 1354, 1373
Pterodroma solandri, 1355, 1373
Pterodroma ultima, 1357
Pterodroma, 1353, 1354, 1358,
1360
Pteropodidae, 958
pterosaur, 688, 610, 657, 659, 686
Ptilinopus, 835
Ptillorhoa, 838
ptilodontid, 898
Prilonorhynchidae, 841
Ptilonorhynchus violaceus, 821
ptychoceratodont, 484
ptyctodontid, 354, 367, 442
Ptykoptychion tayyo, 506”
Ptykoptychion, 347
Pucapristis, 500
Puebla Formation, 514
Puffinus assimilis, 1373
Piffinus carneipes, 1354
Puffinus gavia, 1360
Puffinus tenuirostris, 326, 330,
825, 1347

INDEX - 1433

Piffinus, 1358
Pulbeena Swamp, 817
Pulchera Waterhole, 506
pulp cavity, 440
puncture crushing, 205, 256, 258
Pureni Local Fauna, 1098
Pureni, 95-97, 746, 782
Purgatorius ceratopsis, 961
Purpureicephalus spurius, 836
Putjamarpa Rock Shelter, 304
Pycnoptilus floccosus, 840
Pycnoptilus, 819
Pyctnoctenion, 388
Pygoscelis tyreei, 1177
Pyramids Cave, 291, 295, 309, 819,
837, 838, 840
Pyramios alcootaensis, 935, 1079,
1082, 1102
Python amethystina, 674
pythons, 286
Quagga, 314, 315, 332
Quanbun Local Fauna, 671, 1048
Qantas, 735
Queanbeyan, 447
Queen Victoria Museum, 14
Queensland Museum, 14, 31, 69,
150, 340, 633
Quercy, 758, 767
Quetzalcoatlus, 657
Quilty, P., 151, 155
Quinkan Gallery, 3
Quinkana fortirostrum, 672, 678,
688
Quinkana, 669, 670, 674, 1077
Quipollornis koniberi, 761, 783,
1382
Quipollornis, 729
rabbits, 250
Rackham’s Roost Local Fauna, 670,
959, 961, 1042
radiocarbon dates, 849, 1321, 1351
radiocarbon dating, 881
radioimmunoassay, 317-320, 325,
327, 331, 332
radiometric dating, 724, 917, 929
radotinid, 450, 459
Ragless, F.B., 28
rainfall, 818
rainforest, 129
Rallidae, 739, 760, 768, 830, 1359
Rallus pectoralis, 830
Rallus philippensis, 830
Ramsayia curvirostris, 1094, 1096-
1098
Ramsayia lemleyi, 942
Ramsayia, 943, 1073, 1128
Rana papua, 600
Rancho la Brea, 316
Rancholabrean megafauna, 1131
Rando’s Bore, 517
Ranidae, 598
Ranidella parinsignifera, 600
Ranidella, 667, 668
Ranken, G., 4, 722
Rannuaspid Province, 400
Rapator ornitholestoides, 612, 646,
647
Rapator, 6A8

1434 - INDEX

rat kangaroos, 251

rates of evolution, 677

ratite, 872

Rattus exulans, 353, 1362, 1373

Rattus lutreolus, 303, 960

Rattus, 959, 961

Raup, D., 164

Ray, C., 73

Raymond Formation, 352, 371

Raymond, 452

rays, 498

Razorback beds, 611

Receptaculites limestones, 466

recrystallisation, 881

ted bed facies, 338

ted beds, 116, 124, 126

Redbank Plains Group, 477, 478,

Redbank Plains Series, 669, 671

Redbank Plains, 340-342, 385, 477,
675, 729

Redcliff Mountain, 340-342

redfieldiids, 444

reef limestones, 112

reefs, 122, 126, 467

Reefton Beds, 448, 451

reference books, 156

regional fauna, 1127

regression, 1168, 1175

regurgitation pellets, 287

relative ages, 916

relaxation (island biogeography),
1364

Remigolepis, 358, 363, 364, 401,
402, 420, 582

reproduction, 1141

reptilian eggs, 873

research monographs, 156

restoration of fossil animals, 248

retroarticular process, 571

review articles, 162

Revultex, 197

Rewan Formation, 470, 471, 576,
578

Rewan, 80, 620, 635

Rewana quadricuneata, 580

reworked deposit, 285

reworking, 272

Rhabdosteidae, 182, 1180, 1183

rhabdosteids, 1185

thamphorhynchoids, 657

Rhamphotyphlops, 669

rhea, 327

thenanid, 367, 455, 459

Rheobatrachus silus, 600

Rheobatrachus, 593

Rheodytes, 705

Rhigosaurus, 614

Rhina, 510

Rhinolophidae, 958

Rhinolophus ferrumequinum, 1069

Rhipidistia, 386

thipidistians, 445, 466

Rhipidura, 1333

Rhizodontida, 369, 386

thizodontids, 445

Rhizodontiformes, 388, 454

Rhizoduz, 445

Rhizophascolonus crowcrofti, 941,

1096

Rhoetosaurus brownei, 611, 645

Rhoetosaurus, 648, 663

thomboid scales, 444

Rhombomys, 204, 205

Rhondda Colliery, 659

Rhynchocephalia, 623

thynchosaurs, 609

Rhynochetos jubatus, 1331

thyodacite, 398

thyolite, 398

Rich, P., 73, 76, 77, 723, 767, 811

Rich, T.H., 34, @, 70, 73, 74, 76-
79, 340, 723

Richmond, 633, 689

Ride, W.D.L. 33, 59

Riedel, W., 50

rift valley slumps, 276

Ringtail Site, 481

Riojasuchus, 638

riparian woodland, 1127

Ritchie, A., 33, 34, 76-78, 80, 340,
346

Rivernook, 511

Riversleigh District, 1027

Riversleigh Local Fauna, 1028

Riversleigh Society, 79

Riversleigh, 37, 55, 69, 79, 80,
142, 249, 253, 254, 330,479,
481. 483, 490, 598, 601, 669,
671, 675, 705, 710, 727-728,
7%, 737, 739, 743, 752, 760-
762, 781, 782, 945, 956, 958
1127

RNA, 312-315

Robe, 511

Robertson, G., 151

Robinson, J., 631

Rochow, K., 90

rock paintings, 1109

Rockhampton, 451

Rockhampton Group, 451

rockshelters, 277

Rockycampacanthus milesi, 420,
454

Rockycampacanthus, 351, 401

Rodentia, 959

rodents, 258

Roe Plain, 836

Rohon, J., 446

Rolfosteus canningensis, 359, 360

Rolfosteus, 355

Rolleston, 471

Rolling Downs Group, 347, 377,

380, 504, 506

Rolling Downs, 382

root damage (to bones), 286

Rosella Plains, 670

rotary grinders, 189, 190

Royal Society of London, 22, 149,
154

Ruess, D.J., 153

ruminants, 207-209

rumination, 208

Rundle Formation, 478, 490, 705

Rundle Oil Shale, 669, 671

Rundle, 477, 688

Sacabambaspis, 446, 455

Sacrophilus, 1074

Safeway, 735

sagenodont lungfish, 470, 471, 484

Sagenodus laticeps, 472

salinity changes, 1175

Salisbury Island, 1347, 1350

Salt Creek, 329

saltation, 1111

sampling effect, 664

sand flow lines, 817

Sandover River, 54

sandstone caves, 276, 277

Santa Marta Formation, 614

Sarcophilus harrisii, 284, 302, 303,
678, 880,1090, 1093, 1134,
1140

Sarcophilus laniarius, 302, 1086,
1090, 1134, 1143

Sarcophilus, 211, 285, 302, 305,
826, 927, 1076, 1092

Sarcopterygii, 369, 386, 444, 466

Sarjeant, W.A.S., 154

Saurichthys gracilis, 374

Saurichthys parvidens, 374

Saurichthys, 583, 585, 586

saurischians, 642

Sauropelta, 655

sauropodomorphs, 610, 642, 643

sauropods, 643, 644, 662

Sauroptery gia, 606

sauropterygians, 609, 624, 628,
629, 686

saurostemids, 609

savanna grassland, 1080, 1083

savanna(h) woodland, 822, 1127

Scaldicetus lodgei, 1180

Scaldicetus macgeei, 1180

scale species, 445

Scapanorhyachus subulatus, 507,
524

Scapanorhynchus, 347, 508

Scaptodon lodderi, 1180

Scarlett, R., 723

Scaumenacia, 396

scavenger accumulators, 286

scavenger, 285, 287

scelidosaurs, 648

Sceloglaux albifacies, 1332

Schizurichthys pulcher, 375

Schizurichthys, 376

schlepp effect, 271

Schodde, R., 766, 767

scincids, 286

Sclater’s Line, 763

Sclater, P.L., 763

Scleromochlus, 637

Scleropages, 385

sclerophyll forest, 1127

Sclerorhynchidae, 500

Scolopacidae, 760, 832

Scotchtown Cave, 745

Scot River egg, 881

Scott River, 834

screen washing, 190

scrub, 1127

Scylliorhinus, 510

sea caves, 276

Sea Cow, 330

sea level, 816

seafloor spreading, 127, 765

sealevel changes, 1169

Seaspray Group, 514

Seaton Rock Shelter, 302, 823, 827,
828, 831, 836, 837, 839, 840,

secondary publications, 156
sectioning, 231

Sedgwick, A., 22

sediment settling tank, 184
sedimentation rate, 1168
sedimentation rate, 1168

selachians, 441, 498

selenes, 250

selenodont molars, 932, 1094

selenodont teeth, 257

selenodont, 947

Selwyn, A. R.C., 12

Semionotus, 376

septarian concretion, 617

Serengeti-type kills, 1133

Sereno, P., 80

serological evidence, 1096, 1114

serum albumin, 316, 594

serum proteins, 332

Serventy, D., 766

Set Site, 329

Setonix, 951

sexual dimorphism, 288, 502, 829,
1141

Seymour Island, 614, 738, 746,
747, 1177

Seymour, 748

shagreen, 436

Shale Oil Site (Rundle), 478

shark scales, 439, 440, 451

shark spines, 440

Shark Tooth Hill, 509

sharks, 430-432, 446, 448, 451,
453, 454, 498

Sharpey’s fibres, 443

Shearbyaspis, 367

shearing crests, 237

Sheehy, E.P., 163

sheep, 214, 964

shelf carbonates, 121

shell thickness, 876

shell fragments, 877

shell membrane, 880

shell microstructure, 872

shell porosity, 872, 877, 885

shell structure, 878, 879

shell thickness, 876, 879

Shell, 735

shelly facies, 117

Sherbonaspis hillsi, 363

Sherbonaspis, 401

shrublands, 208

Siderops kehli, 580, 581

sieving boxes, 273

silcrete, 139, 182

silica crystals, 207

silicification, 273

silicone mould, 197
Sillago pliocaenica, 384
Sillago, 382
Silverband Formation, 434, 447-
449, 452, 582,
Silverdale Formation, 447
Simosteus, 355
Simos thenurus, 215, 218, 219, 250,
1073, 1074, 1076, 1114, 1133
Simpson Desert, 475, 507,1078
Simpson, G.G., 723
Sinacanthus, 453
sinolepidoids, 362
Sinolepis, 402
Sirenia, 1171, 1178
Skamolepts fragilis, 448
Skamolepis, 346, 438, 455
Skartopus australis, 660
skates, 498
skeletal reconstructions
(megafauna), 1129, 1130
skeletal reconstructions, 248
Skull Cave, 291, 295, 298, 668,
820, 821, 837
Smeaton, 1057
Sminthopsis leucopus, 1349
Sminthopsis, 289, 899
Smith, D., 735
Smith's, W., 149
Snake Dam, 882-886, 888
Snobs Creek Volcanics, 398
Snow Hill Group, 614
Snowy Mountains, 818
Snowy Plains Formation, 398
Soederberghia, 393, 582
solar radiation, 1169
Somersby Falls, 467, 470
sorting, 273, 275
South Australian Museum, 340, 723
South Blue Range, 340-342, 386
South China Province, 343
South Mt Cameron, 738
Southeast Pacific Rise, 1318
Southern Alps, 955
Southwest Pacific islands, 1346
South westem regional fauna, 1127,
1128
Sowerby, J., 4
speciation patterns, 1131
species diversity, 1364
species-groups, 594
speleo, 276
speleophiles, 834, 838
Spenostoma occidentalis, 839
Speonesydrion iani, 392, 395
Speonesydrion, 396, 466
Spheniscidae, 825
Sphenisciformes, 1171, 1175, 1326
Sphenodon, 1359
sphenodontians, 609
Sphenomorphus, 667
Sphenophryne robusta, 600
Sphenophryne, 593
Sphyrna, 515, 517,531

Spring Creek, 305, 748

Springer, M., 80

springs, 276

Springsure, 340-342, 633, 671

Spurr's resin, 231

Squalodon serratus, 1181, 1183

Squalus, 501, 510

squamatans, 609, 623

Squatina, 508

St Helena, 1363

St Paul, 1363

St Peters, 472

St John, O., 446

St Peters fauna, 374

St Peters, 340-342, 470, 583

St Vincent Basin, 511, 512, 516

Stairway Sandstone, 343, 446

Stanley, G.A.V., 86

Staphylococcus aureus, 319

Steadman, D., 766

stegosaurs, 648, 649

steinkem, 714

Steller’s Sea Cow, 332

Steno cudmorei, 1180, 1183

steppe limestones, 276

Stercorarius longicaudus, 1357

Stercorarius skua, 1357

stereoscopic vision, 256-258

Sterna fuscata, 1373

Steropodon galmani, 922-924

Steropodon, 1084

stethacanthid, 440, 451, 455, 458

Stethacanthus thomasi, 347, 348

Stethacanthus, 440, 441,451

Stetosaurus, 633

Sthenurinae, 215, 217, 1116

sthenurine kangaroos, 213, 219

Sthenurus (Simosthenurus)

brownmei, 1118-1122

Sthenurus (Simosthenurus) gilli,

1119, 1120, 1122

Sthenurus (Simosthenurus)

maddoch, 1119, 1120, 1122, 1128

Sthenurus (Simosthenurus)

occidentalis, 938, 1118, 1120,1121,
1128, 1135, 1365

Sthenurus (Simosthenurus)

orientalis, 1118, 1120, 1122, 1123

Sthenurus (Simosthenurus), 1128

Sthenurus andersoni, 1112, 1125

Sthenurus atlas, 220, 1112, 1124,
1125, 1128

Sthenurus oreas, 1112, 1122, 1123

Sthenurus pales, 1112, 1122, 1123

Sthenurus tindalei, 220, 1122, 1125,
1128

Sthenurus, 215, 218, 219, 251, 950,
951, 953, 1073, 1074, 1076,
1079, 1114

Stictonetia naevosa, 740

Stipiturus malachurus, 840

Stipiturus, 840

Stirling, E. C,, 18, 28,722

Stirton Quarry, 54, 56, 58

Stirton, R. A., 20, 32, 33, 37, 47,
56, 62, 64, 75, 90, 91, 250,
252, 723, 735, 756, 811

Stockyard Site, 1349

stomach acid pH, 287, 288

stomach acid, 284

stomach contents, 627

stomach, 240, 241

stomatosuchians, 639

strain, 206

Strathalbyn, 513

Strathdownie Cave, 324, 1143

stratigraphic correlation, 1167

Strepera graculina, 842

Strepera versicolor, 842

Strepsodus decipiens, 388

Strepsodus, 445

Streptopelia, 823

Striacanthus sicaeformis, 453

Striation pattem, 210

stride length, 662

Strigidac, 761, 834

Strigiformes, 1332

Strigocuscus reidi, 945

Strigocuscus, 944, 954

Strigops habroptilus, 1331, 1356,
1357

Striped Possum, 257

Strongs Cave, 516

Struniiformes, 368

Strunius, 390, 391,445

Struthio camelus, 1325

Struthiomimus altus, 647

Swzelecki Creek, 51

Strzelecki Formation, 474, 477

Strzelecki Group, 377, 472, 473,
490, 572, 573, 580, 586, 612,
617, 646, 714, 737, 765

Sturzelecki, 477

Sturnmus vulgaris, 306, 309

Sturmus, 823

Sturt, C., 236

Stutchbury, S., 8

stylar shelf, 232, 910

stylid cusps, 256

subfossil, 1321

subholostean, 374

Sula abbotti, 1363

Sula dactylatra, 1373

Sula tasmani, 1353, 1354

Sumatra, 384

Sumba, 763

Sunlands Local Fauna, 1934, 038

Sunlands, 329

supernova, 1175

Suprasec 5005, 198

supraspinatus fossa, 1120

Surat Basin, 124, 125

Surcaudalus, 347

Sus, 763, 970

swallow hole, 282

swamp alluvium, 1321

sweepstakes dispersal, 1321, 1320,
1334

swim-bladder, 345

Sydney Basin, 123, 124, 345, 467,
576, 583

Sydney University, 151

Sylviidae, 839

Sylviornis neocaledoniae,

Sylviornis, 758

symbionts, 207, 208

Symmorium, 440

1355

INDEX - 1435

symphseal teeth, 502
synapsid, 609, 620, 686
Synechodus validus, 508
Synechodus, 507, 525
Synocryl, 175, 182, 187
Synodontaspis acutissima, 512
Synodontaspis taurus, 512
Synodontaspis, 501
synthetic community, 1129
Tachyglossidse, 922
tachyglossids, 1084
Tachyglossus aculeatus, 303, 1084,
1086, 1088-1090
Tadorna, 1329
Tacmas Group, 458, 459
Taemas, 351, 352, 356, 367, 391,
392, 395, 450, 452, 466
Tacmas-Wee Jasper, 340-342, 354,
391, 434
Taemasacanthus erroli, 453
Taemasacanthus, 351, 401, 420
Taemasosteus novaustrocambricus,
354, 356, 357
Taemasosteus erroli, 420
Taeniolabis, 250
Taggerty Fauna, 398,
Taggerty, 340-342, 395, 397, 398
Talbragar, 340-342, 376, 382
Talingaboolba Formation,
Talki Limestone, 513
talonid basin, 907
talonid, 233, 901
Talyawalka, 1052
Tambar Springs, 329
Tambo Series, 382
Tamworth Trough, 122
Tandalgoo Formation, 399
Tandalgoo Red Beds, 434, 452
Tandanus, 385
tanins, 235
Taniwhasaurus oweni, 1174
taphonomic accumulator, 289, 290
taphonomic classification, 275
taphonomic history, 275, 285
taphonomic input, 275
taphonomic studies, 1362
taphonomy, 268, 286, 880
tapirs, 1106
tar pits, 276, 316
Tara Creek, 670, 705, 1043
Tarkarooloo Local Fauna, 481,
667, 939, 941, 1023
Tarkarooloo Sub-basin, 140, 142,
254, 726, 732, 746, 757
Tarkarooloo, 762
Taroona, 668
tarsal pattern, 929
Tarsipedidae, 956
Tarsipedoidea, 956
Tarsipes spencerae, 956
Tasidyptes hunteri, 749, 825, 1348
Tasman Sea, 1318
Tasmanian Devil, 284
Tasmanian devils, 286
Tasmanian Journal, 22,23
Tasmanian Museum, 14, 69, 343
Tasmanian regional fauna, 1127
Tasmanian Wolf, 332

447

1436 - INDEX

Tasmaniosaurus triassicus, 639,
640

Tasmaniosaurus, 586, 641

Tasmantis, 1350

Tate, R., 11, 20, 25, 151

Tatera, 204, 205

Tatong Fauna, 398

Tatong, 340-342

Taudactylus ditrnus, 0

Taudactylus, 593

Te Whanga lagoon, 1329

Te Whanga Limestone, 1356

Tea Tree Cave, 670

Teddy Mount Formation, 449, 458

Tedford locality, 49, 252, 330

Tedford, R.H., 20, 32, 50, 53, 62,
73, 74, 80, 723, 811

Tegeolepidae, 374

Teichert, C., 356

Telemon Formation, 452

lteleostean, 368, 374

Teleosteomi, 345

telestomes, 442, 450

Tellerodus, 471

TEM, 232

Temnodontosaurus, 625

Temnospondyli, 570, 571

temnospondyls, 640

temporal fenestrae, 606, 607

terrestrial communities, 268

Tertiary mammalian genera, 1008-
1011

tertiary publications, 156, 157, 163,
164

tesserae, 499

testudines, 609, 615, 675

Testudinidae, 704, 713

tetrabromocthane, 431

tetrapod trackway, 574

Tetrapoda, 369

tetrapods, 368, 570

tetraradiate squamosal, 635

Texas Cave, 670

textbooks, 156

thalattosuchians, 639

thanatocoenosis, 586

The Crater, 470

Thecodontosaurus minor, 645

thecodonts, 636, 637, 639, 641, 663

Thelodinti, 343

thelodont scales, 439, 445

thelodont, 344, 397, 401, 430, 432-
434, 436, 437, 439, 446-448,
459

Thelodus, 434, 439

therapsid, 620, 622, 648

therians, 1084

Therizinosaurus cheloniformis, 647

Theropithecus, 209, 210, 223

theropod, 660, 610, 643, 644, 661,
662

Thescelosaurus, 638, 664

Thickthom, 31

thin sections, 877

Thinoconidae, 761, 766

Thinornis, 1331

Thotnpson, A. M., 20, 151

Thorlindah, 745

Three Hummock Island, 1347

Threskiornis spinicollis, 756

Threskiornis mollusca, 827

Threskiornis spinicolliz, 752, 754,

Threskiomithidae, 752, 827

Thrinacodus (Harpagodens) ferox,
347, 348, 441, 451, 455, 458

Thrinacodus, 440, 450

Thrinaxodon, 614, 622, 898

Thulbom, R.A., 33, 76, 660-662,
645

Thursius, 388

thylacine, 286, 314, 332, 907

Thylacinidac, 927, 1086

Thylacinus cynocephalus, 286, 678,
880, 927-929, 950, 1079,1086,
1090, 1092, 1093, 1133, 1136,
1143

Thylacinus major, 1093

Thylacinus potens, 929, 1079

Thylacinus rostralis, 1093

Tiylacinus spelaeus, 1093

Thylacinus, 90, 94, 211, 895, 1074

Thylacoleo carnifex, 212, 286, 304,
310, 678, 943, 950, 1081,
1108, 1109, 1133, 1136, 1138,
1139, 1143, 1159

Thylacoleo crassidens, 1081

Thylacoleo, 27, 210, 213, 214, 305,
944, 945, 1073, 1074, 1076,
1107, 1137

Thylacoleonidae, 943, 944

Thylacomyidae, 930

Thylacosmilus atrox, 911

Thylacosmilus, 1107

Thylogale billardieri, 230, 303,
1135

tibia-astragular joint, 1111

Tiga Island, 676

Tiliqua, 667, 668, 670, 673

tillites, 112

Timbu Tasong, 91

Timor, 763

Tingamarra Local Fauna, 925,927,
1014

Tinodon bellus, 901

Tirari Formation, 49, 54, 56, 80

Tiran Local Fauna, 1079

Titanopteryx, 657

titanothere, 1100

Tiupampea Loca] Fauna, 910

Toescatter Locality, 480

Toko Syncline, 346, 434

Toko Syncline, 447

Tom O's Quarry, 321, 324, 330,
732

Tomasettii, B. Fr., 97

tongue, 250

Toolebuc Formation, 504, 506,
507, 613, 617, 627, 737,

Toolinna Limestone, 511

Toomba Range, 340-342

Toombosteus denisoni, 354, 357

tooth form , 205

tooth function, 205

tooth morphology, 207, 215, 257

tooth nomenclature, 899, 900

tooth ultrastructure, 500

tooth wear, 210

toothwhorls, 443

topographic relief, 1319

Torquay Basin, 513

Torquay, 511, 512, 514, 515

Torres Strait, 86, 1346

Torresian, 812

Tower Hill, 324

Town Well Cave, 1052

trabecular dentine, 500

Trachichthyodes, 382

Trachydosaurus, 668, 669

trachytic pumices, 276

tracks (dinosaur), 659

tracks (omithopod), 660

trackways (bird), 742

trackways (dinosaur), 659

Transantarctic Mountains, 614, 622

transgression, 1168, 1319

translations, 164

Traversia lyalli, 1333

Traversia, 1333

treatise, 156, 157, 164

Triakis, 510

Triazeugacanthus, 351

tribosphenic therian, 902

tribosphenic, 931

Triceratops community, 907

Trichoglossus haematonotus, 836

Tricholimnas lafrasneyanus, 1374

Trichosurus dicksoni, 945

Trichosurus vulpecula, 303, 944,
1084, 1138

Trichosurus, 954

triconodontians, 896, 897, 922

triconodonts, 910

Trigonia, 3

trigonid, 233, 901

Trigonotreta stokesii, 4

Trinacromerum leucoscopelus, 632

Trinacromerum osborni, 632

Trinacromerum, 613, 633

Tringa glareola, 832

Trionychidae, 676, 708, 714

Tristychius, 441

Trochocyathus Bed, 511

Trochocyathus, 525, 526

Troposodon kenti, 56, 216,1125

Troposodon minor, 215, 1112,
1116, 1125

Troposodon, 128, 215, 250, 1114

Trundle Beds, 448, 450, 452, 459

Trundle Group, 434

Trundle, 452

Trundle-Kadungle, 434

Tsintosaurus, 198

tubercular, 213

Tubonasus, 355

tuffs, 276

Tulki Limestone, 515

Tullamore, 458

tumbler experiements, 732

Tumblong Oolite, 434

Tumbunan, 812

Turdus merula, 738

Turdus poliocephalus, 1373

Turdus, 823

Turena, 614

Turinia australiensis, 344, 434,
4%, 437, 447, 459

Turinia fuscina, 434, 447

Turinia hutkensis, 344, 434, 437

Turinia pagei, 434, 459

Turinia polita, 434, 447

Turinia, 439

turiniids, 439, 447

Turnagra capensis.,1333

Tumbull, W., 33, 34, 50, 57, 75,
78

Tumer Brook, 302

Tumer, S., 33, 34, 76, 340

Tumicidae, 758, 830

Turnix pyrrhothorax, 830

Turnix varia, 252, 296, 326, 830

Turnix velox, 292-294, 296, 830

Turnix, 325, 758

turtles, 286, 665, 686

Tutur, 823

Tuva, 400

Tyers district, 450

Tyers, 458

Tylosaurus haumuriensis, 1174

tympanic process, 1086

type localities, 1167

Tyrannosauropus, 660

Tyrannosaurus, 637, 642

Tyto alba, 286, 289-292, 295, 296,
307-309, 325, 834, 836, 1332

Tyto longimembris, 289

Tyto novaehollandiae, 281, 286,
287, 289, 295, 309, 761, 834

Tyto tenebricosa, 285, 287, 289,
290, 306-309

Tytonidae, 761, 834

tytonids, 811

U.S. Geological Survey, 160

Uabryichthys latus, 376, 380

Ulrich,159

ultrasound, 431

ungulates, 208

uniformitarian approach, 1168,
1170

University of Adelaide, 20, 48

University of California (Berkeley),
49, 69, 73, 154, 723, 735

University of Califomia-South

Australian Museum Expedition,
723

University of Chicago, 50

University of Melbourne, 14, 15,
16

University of Sydney, 20

University of Texas, 75

Uperoleia, 593, 599, 600

Upper Murray Cliffs, 340-342

Upper Site Local Fauna, 1030

Upper Swan River, 304

Uranolophus, 395, 396

Uroconger, 382

Urodela, 570

urogenital tract, 903

Uromyini, 959

Urosthenes australis, 371

Urosthenes, 372, 583

Ursus arctos, 678

Ursus, 215

uruguaysuchians, 639
Utah Mining, 735,
Utting Calcarenite, 345, 451, 452
vadose caves, 277, 278
vagility, 1170
vagina, 902
Vallisneria spiralis, 482
van Tets, G.F., 723, 811, 827
varanids, 136, 673, 1086, 1086
Varanus giganteus, 678
Varanus indicus, 1355
Varanus komodoensis, 673, 1076
Varanus, 667-670, 895
varves, 665
Vega Island, 614
ventriculus, 284
vertebrate accumulated assemblage,
83
Vespertilionidae, 958
vibro-engraver, 189, 190
vicariance, 763, 765, 1171, 1172,
1363
WVicecomodora Marambio Island,
614, 615, 624
Vickers, M., 75, 77
Victoria Cave, 80, 598, 842
Victoria Fossil Cave, 811, 829-832,
836, 837
Victoria Land, 122
Victorian Chamber of Mines, 20
viking funeral ship, 133, 766
Vinalak 63-513, 187, 196
Viverra civetta, 678
volcanism, 1169
volcanic caves, 277, 280
Vombatidae, 939, 940
Vombatiformes, 932, 1086
Vombatomorphia, 935
Vombatus hacketti, 1095, 1096
Vombatus ursinus, 303, 1084, 1094,
1096, 1098, 1099, 1136, 1137,
1349
Vombatus, 940, 943
von Buch, L., 3
von Waldheim, f., 149
vulcanism, 1169, 1175
Vulpes vulpes, 305, 822
Wade, M., 76, 340, 661
Wade, Rev. R.T., 46
Wadeichthys oxyops, 380
Wadikali Local Fauna, 1024
Waikerie, 329
Wairuna Formation, 119
Waite Formation, 668, 728
Wakaleo alcootaensis, 1079,
Wakaleo vanderleueri, 944, 1079,

Waldman, M., 61

Walford, AJ., 163

Walgett, 340-342, 473, 474
Walgettosuchus woodward, 646
Wallabia bicolor, 216-218, 1134
Wallabia indra, 1134

Wallabia rifogrisea, 93
Wallace’s Line, 136, 765, 767

Wallace, A.R., 1346

Walloon Group, 611

Wallumbilla Formation, 126, 504,
506, 613, 633

Walpole Island, 676, 708, 713

Walsh River, 633

Wanneroo, 302

Waratah Bay, 452

Warendja wakefieldei, 943, 1094

Warra Station, 737

Warren, A., 76

Warren, J., 58, 59, 73, 76

Warrumbungles, 340-342

washing box, 191

water rat, 258, 286

waterhole tethering, 1079

Watut Valley, 86, 91,95, 98

Watut-Bulolo, 91

Waum Ponds Limestone, 512
Waum Ponds, 340-342, 511
wear facet formation, 218

wear facets, 230

weathered profiles, 129
weathering, 272, 273

Wobdb's Caves, 302

Weber’s Line, 136, 763

Wee Jasper, 121, 458, 459, 466
Weejasperaspis gavini, 366
Weekes Cave, 811, 827, 839, 841,

842
Weetalibah, 329
Wellington caves, 6-8, 324, 329,
669, 722
Wellington Valley, 4, 5, 36
Wellington, 434, 512,

Wells, R., 33, 73, 74, 76, 77, 80, 97

Welsh Borderland, 459

Wendover Site, 315

West Indies, 1346

Westem Australian Museum, 340

Wester blotting, 323

Wester Mining, 735

Westoll-lines, 445

Westralichthys uwagadensis, 356

wetscreening, 191, 275, 736

Whalers Bluff Formation, 517

White Cliffs, 473, 632

White Sands Basin, 480,470, 472

Whitelaw, M., 80

Wianamatta Group, 374, 572, 573,
583

Wianamatta Series, 375

Wijdeaspis warrooensis, 366

Wild, J.,70

Wilkinson, R., 77

Williams, D.L.G., 80, 723, 886

Williams, P., 97

Williamsaspis bedfordi, 354, 357,

Willis, P.M.A., 671

Wilsons Promontory, 1346, 1347

Wimanornis, 615, 748

Winduck Group, 399

Winteracese, 140

Winton Formation, 73, 473, 474,
612, 618, 645, 661, 662, 688

Winton, 474, 618, 661

Wintonopus latomorum, 660

Wipejira Formation, 478, 481-484,
667,716

Witton Bluff, 746

Witwatersrand Group, 113

Wombeyan caves, 745

Wombeyan Quarry Cave, 324

Wonambi naracoortensis, 674,
675,1128

Wonambi, 666, 668, 677

Wonthaggi, 646

Woodard Quarry, 48, 53, 329

Woodard, G.D., 48, 55, 90

Woodbume M.O., 33, 53, 54, 63,
68, 73, 74, 80, 723

Woods, J.E.T., 24,722

Woodward, H., 149

Wooley, P., 95

Wollaston, T.C., 474

Woolungasaurus glendowerensis,
613, 631, 632, 688, 1173

Woolungasaurus, 634, 663, 1173

Worange Point, 452

Worthen, A.H., 446

Warrumbungle Ranges, 386

Wuttagoona Station, 340-342

Wuttagoonaspid-Phyllolepid
Prov ince, 400

Wuttagoonaspis fauna, 344

Wuttagoonaspis fletcheri, 354, 357,
358

Wuttagoonaspis scales, 455

Wuttagoonaspis, 355, 397-399, 401

Wyandotte Creek, 324, 831

Wyandotte Station, 710

Wyloo group, 113

Wynyard, 1036

Wynyardia bassiana, 937, 938

Wynyardia, 46

Wynyardiidae, 937, 939

X-ray diffraction, 881

xenacanthid, 499, 583

xenacanths, 440, 451

Xenacanthus, 346, 440, 455

Xenicus gilviventris, 1333

Xenicus longipes, 1333

Xenicus, 1333

Xenobrachyops allos, 574, 576, 585

xenoliths, 113

Xenorhynchopsis minor, 753, 754

Xenorhynchopsis tibalis, 752-754,
832

Xenorhynchopsis, 819

Xenorhynchus asiaticus, 752, 754,
827

Xenorhynchus minor, 832

Xenorhynchus, 752

Xiphactinus australis, 382

Xiphactinus, 381

Yalkaparidon coheni, 956

Yalkaparidontia, 956

Yalkaparidontidae, 956

Yanda Local Fauna, 481, 667, 739,
757, 1022

Yapok, 258

Yarra Yarra Creek Group, 443,
447, 450, 452

Yarrol Trough, 122

INDEX - 1437

Yellow Drum Sandstone, 358

yolk sack, 606

Young, G., 33, 76, 340, 355, 630

Youngina, 630

younginiforms, 609

Youngolepis, 369, 386

yunnanolepidoids, 362

Zaglossus briuijnii, 1086, 1088,
1090

Zaglossus hacketti, 1086, 1089

Zaglossus ramsayi, 1086, 1088-
1090

Zaglossus robusta, 192A, 086, 1090

Zaglossus, 69, 1073, 1074, 1076

Zeitz, A-H.C., 722

zeuglodons, 1180

Zietz, A-H.C., 18, 28

Ziphius, 1180

ziphodont crocodiles, 639, 669-671,
677, 679, 927

zoogeography, 1170, 1171

Zoological Record, 149, 155

zygomatic arch, 255

Zygomaturinae, 935

Zygomaturus keani, 1098

Zygomaturus trilobus, 934, 1073,
1074, 1079, 1080, 1098, 1100,
1101, 1103, 1106, 1107, 1127,
1136, 1137, 1140, 1143, 1159

Zygomaturus, 197, 1081, 1102,
1135, 1355

Zyzomys, 961

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