European Neogene Mammal Chronology

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European Neogene

Mammal Chronology
NATO ASI Series
Advanced Science Institutes Series

A series presenting the results of activities sponsored by the NATO Science Committee,
which aims at the dissemination of advanced scientific and technological knowledge,
with a view to strengthening links between scientific communities.

The series is published by an international board of publishers in conjunction with the


NATO Scientific Affairs Division

A Life Sciences Plenum Publishing Corporation


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c Mathematical Kluwer Academic Publishers


and Physical Sciences Dordrecht, Boston, and London
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Volume 180-European Neogene Mammal Chronology


edited by Everett H. Lindsay, Volker Fahlbusch,
and Pierre Mein

Series A: Life Sciences


European Neogene
Mammal Chronology
Edited by

Everett H. Lindsay
University of Arizona
Tucson, Arizona

Volker Fahlbusch
University of Munich
Munich, Federal Republic of Germany

and

Pierre Mein
Claude Bernard University
Lyon, France

Springer Science+ Business Media, LLC


Proceedings of a NATO Advanced Research Workshop on
European Neogene Mammal Chronology,
held May 16-20, 1988,
in Schloss Reisensburg (near GOnzburg), Federal Republic of Germany

Llbrary of Congrass Cataloglng-ln-Publlcat ton Data

European Naogane •ammal chronology 1 edited by Everett H. Lindsay,


Volkar Fahlbusch, and Piarre Mein.
p. c•. -- !NATO ASI series. Series A, Life sciences ; v.
180)
"Proceedings of a NATO Advanced Research Workshop an European
Neogene Mam1al Chronology, held May 16-20, 1988 in Schloss
Reisensburg"--T.p. versa.
"Published in cooperation with NATO Sc1ent1fic Affairs Division."
Includes bibliographical references.
ISBN 978-1-4899-2515-2 ISBN 978-1-4899-2513-8 (eBook)
DOI 10.1007/978-1-4899-2513-8
1. Mammals, Foss11--Europe--Congre sses. 2. Paleontology,
Stratigraphic--Congres ses. 3. Paleontology--Neogene- -Congresses.
4. Paleontology--Europe--C ongresse&. 5. Geology, Stratigraphic-
-Neogene--Congresses. 6. Geology--Europe--Congr esses. I. Lindsay,
Everett H. II. Fahlbusch, Volker. III. Mein, Pierre. IV. NATO
Advanced Research workshop an European Neogene Ma•mal Chronology
!1988: Schloss Reisensburg> V. North Atlantic Treaty Association.
Scientific Affairs Oivision. VI. Series.
QE881.E82 1989
569'.094--dc20 89-26648
CIP

© 1989 Springer Science+Business Media New York


Originally published by Plenum Press, New York in 1989
Softcover reprint of the hardcover 1st edition 1989
Ali rights reserved
No part of this book may be reproduced, stored in a retrieval system, or transmitted
in any form or by any means, electronic, mechanical, photocopying, microfilming,
recording, or otherwise, without written permission from the Publisher
PREFACE

During the last ZO years great progress has been achieved in our understanding of
both earth history and vertebrate evolution. The result is that climatic/tectonic
events in earth history can now be placed in a more precise and global time frame,
that permit their evaluation as abiotic causal factors which might trigger extinction
and dispersal events in vertebrate history. Great strides have also been made in
genetics and cell biology, providing new insight into phylogenetic relationships among
many vertebrates. These new data, along with data on chronologie resolution of earth
history, provide tests of previous interpretations regarding ancestral-descendant
relationships based solely on the fossil record.

It is fitting and proper that a volume on European Neogene mammal chronology


is produced at this time, to ensure that new interpretations of vertebrate evolution
and chronology are based on the most accurate and current data. Vertebrate paleon-
tologists believe that the fossil record is the only secure data for measuring the actual
course and tempo of vertebrate evolution. Knowledge of the fossil record must keep
pace with advances in other areas of science so that inferences on vertebrate evolu-
tion are accurate and meaningful.

The rich record of fossil mammals in Europe has contributed substantially to


studies of vertebrate evolution. Notable early contributions to Cenozoic geochronology
and vertebrate evolution were made by Cuvier, Geoffroy St.-Hilaire, Darwin, Huxley,
Agassiz and others. Many of the concepts that we now use in Cenozoic geochronology
and vertebrate evolution were developed in Europe during the last century.

European mammal chronology has grown from both broad synthetic studies (e.g.,
Zittel, Thenius, Kretzoi and Kurten) plus centrally focused studies (e.g., Crusafont,
Thaler, Tobien and Azzaroli). The latter of these foundation builders are included
among the contributors to this volume. We co-editors dedicate this volume to those
vertebrate paleontologists who have helped to create the foundation of European
Cenozoic mammal chronology that we have inherited.

An international workshop was held at Schloss Reisensburg in Germany on May


16-ZO, 1988, to stimulate further knowledge of European fossil mammals. This
meeting, sponsored by NATO as an Advanced Research Workshop, had the title
"European Neogene Mammal Chronology." It was attended by 47 researchers (listed at
the end of the volume) from 15 nations, including 13 European nations. Presentations
were given in English, even though English is the native tongue for only about 1Z
percent of the participants. The topics addressed at the meeting were varied, includ-
ing principles of stratigraphy, presentation of new fossil discoveries, syntheses of
climatic and biogeographic data, update of systematics for selected groups of
European mammals, and chronology of mammal faunas outside of Europe. Discussions
were friendly and lively; conflicting perspectives were addressed and discussed openly.

Contributions to this volume reflect the diversity of topics and viewpoints


presented at the Workshop. Seven chapters (e.g., 1, 6, 7, 18, 31, 35 and 37) were
initiated after the Workshop and most of the manuscripts were revised to some degree

v
after the Workshop. Note that opm1ons regarding importance and significance in
mammalian chronology are diverse; we have attempted to include all these viewpoints
in this volume, and will leave for future historians to decide which of these viewpoints
contributed the most important or significant methods and studies for advancing
mammal chronology, especially in Europe. We will claim that this volume reflects
current state-of-the-art in knowledge of European mammal chronology.

The co-editors are indebted to the North Atlantic Treaty Organization and the
National Science Foundation for supporting this Workshop. We thank the Inter-
nationales Institut fur wissenschaftliche Zusammenarbeit e.V. Schloss Reisensburg for
providing comfortable accommodations, excellent service and tasty meals during our
Workshop. A visit to the meteor crater and fossil deposits near Steinheim was led by
E.J.P. Heizmann, to whom we are grateful. An editorial board consisting of Andrews,
Fejfar, Ginsburg, Kowalski, Qiu and the co-editors critiqued the manuscripts following
the Workshop. We were assisted in the preliminary organization of the Workshop by
Mrs. Helga Fuchs, who was unable to join us during the Workshop. Ms. Gertrud
Roessner capably assisted during and after the Workshop. We are especially grateful
to Ms. JoAnn Overs for expert typing and editorial assistance in the preparation of
camera-ready manuscripts. Mr. Jim Abbott provided expert advice and skill in design
and correction of illustrations, for which we are grateful. The co-editors express their
sincere thanks to all those mentioned above, for helping to present a stimulating
Workshop and a handsome volume.

E. H. Lindsay
V. Fahlbusch
P. Mein

vi
CONTENTS

ASPECTS OF EUROPEAN MAMMAL CHRONOLOGY

The Setting . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . • . . . . . . . . . . . . . . . . . . . . . . . . . 1
E. Lindsay

European Neogene Marine/Continental Chronologie Correlations • • • • • • • • • • • • • 15


F. Steininger, R.L. Bernor, and V. Fahlbusch

A Biochronologic Subdivision of the European Paleogene Based on


Mammals - Report on Results of the Paleogene Symposium
held in Mainz in February 1987 • . • . . . • • • . . • . • • • . • . • . • • • . • • . • . . • • • • • 47
N. Schmidt-Kittler

The Ramblian and Aragonian: Limits, Subdivision, Geographical


and Temporal Extension • • • . • . . • • . . • • • . . • . . • • • • . • • . • • • • . • • • • • . • • • • 51
R. Daams and M. Freudenthal

New Neogene Rodent Assemblages from Anatolia (Turkey) 61


M. Siimengen, E. Unay, G. Sara~, H. de Bruijn,
I. Terlemez, and M. Giirbiiz

Updating of MN Zones • • . • • • . • . . . . • • • • • • • • • • • • • • • . • • • • • • • . • • . • • • • . • • • . • • 73
P. Mein

Muroid Rodent Biochronology of the Neogene and Quaternary • • • • • • • • • • • • • • • • 91


in Europe
0. Fejfar and W. Heinrich

Biozones or Mammal Units? Methods and Limits in Biochronology • • • • • • • • • • • • 119


c. Guerin
Large Mammal Dispersal Events at the Beginning of the
Late Villafranchian . • • . . • • • • • • • • . . • . • • . • . . • • • . • • . • • • • • • . • • • • . . . • . 131
F. Masini and D. Torre

REGIONAL PAPERS

Synthesis on the "Aquitanian" Lagomorph and Rodent Faunas of the


Aquitaine Basin (France) • • • • • • . • • • • • . • • . • • • • • • • . • . • • • • . • • . . • • • • • • . 139
M. Hugueney and M. Ringeade

The Faunas and Stratigraphical Subdivisions of the Orleanian


in the Loire Basin (France) . . . . . . • . . . . . . . . . . . . . . . . . . . . . . • . . . . . . • . • • 157
L. Ginsburg

vii
A Preliminary Mammal Zonation of the Upper Marine Molasse
of Switzer land . . . . • . . . . . . . . . . . . . . . . . . . • . . . . • . . . . . . . . . . . . . . . . . . . . 177
B. Engesser

The Faunal Succession in the Bavarian Molasse Reconsidered-


Correlation of the MN5 and MN6 Faunas • • • • • • • • • • • • • • • • • • • • • • • • • • • • 181
K. Heissig

Stratigraphy of Neogene Mammals of Poland 193


K. Kowalski

The Neogene VP Sites of Czechoslovakia: A Contribution


to the Neogene Terrestric Biostratigraphy of Europe
Based on Rodents • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 211
0. Fejfar

FAUNAL DATUM PAPERS

The "Proboscidean Datum Event:" How Many Proboscideans


and How Many Events? . . . . . . . . . . . . . . . . . . . . . . . . . . . • . . . . . . . . . . . • . . . Z37
P. Tassy

The Proboscideans Data, Age, and Paleogeography: Evidence


from the Miocene of Lisbon • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 253
M. Antunes

Patterns of Old World Hipparionine Evolutionary Diversification


and Biogeographic Extension • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 263
R.L. Bernor, H. Tobien, and M. Woodburne

The Hipparions of the Lower Axios Valley {Macedonia, Greece).


Implications for the Neogene Stratigraphy and the
Evolution of Hipparions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 321
G. Koufos

The Genus Eguus in Europe • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 33 9


A. Azzaroli

BIOGEOGRAPHIC SYNTHESIS

Bioevents and Mammal Successions in the Spanish Miocene • • • • • • • • • • • • • • • • • • 357


S. Moya-Sola and J. Agustf

The Miocene Rodent Succession in Eastern Spain: A


Zoogeographical Appraisal • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 375
J. Agustf

Gundersheim-Findling, a Ruscinian Rodent Fauna of Asian


Affinities from Germany • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 405
G. Storch and 0. Fejfar

Dynamics of Old World Biogeographic Realms during the


Neogene: Implications for Biostratigraphy • • • • • • • • • • • • • • • • • • • • • • • • • • 413
M. Pickford

PALEOECOLOGICAL SYNTHESIS

Miocene Paleoecology of Pasalar, Turkey • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 443


B. Alpagut, P. Andrews, and L. Martin

viii
Taphonomic and Sedimentary Factors in the Fossil Record
of Mammals • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 461
M.A. Alvarez Sierra, M. Dfaz Molina, J .I. Lacomb a,
and N. L6pez Martmez

Relations Between Paleoclimatology and Plio-Pleistocene


Biostratigraphic Data in West European Countries , , , • , • , , •••• , , , •••• , 475
M. Bonifay

Small Mammal Taphonomy 487


P. Andrews

MAGNETOSTRATIGRAPHIC APPLICATIONS

Hipparion Datum and its Chronologie Evidence in the


Mediterranean Area . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 495
S. Sen

The Magnetic Stratigraphy of the Late Miocene Sediments


of the Gabriel Basin, Spain ••••••• , , • , •• , , , • , , •• , ••• , , , •••• , •••••• , 507
N. Opdyke, P. Mein, E. Moissenet, A. P~ez-Gonz~ez,
E. Lindsay, and M. Petko

Preliminary Magnetostratigraphic Results of Some Neogene


Mammal Localities from Anatolia (Turkey).••• , , ••• , , •••• , , • , • , , , • , , , 515
C.G. Langereis, S. Sen, M. Siimengen, and E. Unay

SEQUENCES OUTSIDE EUROPE

The Chinese Neogene Mammalian Biochronology- Its Correlation


with the European Neogene Mammalian Zonation •• , •• , ••••• , , •••• , • • 527
Z. Qiu

Key Biostratigraphic Events in the Siwalik Sequence •• , •• , ••• , , , •• , • , • • • • • • • 557


J. Barry and L. Flynn

Quo Vadis, Antemus? The Siwalik Muroid Record 573


L. Jacobs, L. Flynn, W. Downs, and J. Barry

The African Dimension in European Early Miocene Mammal Faunas ••• , • • • • • • • 587
R. Savage

Development and Application of Land Mammal Ages in


North America and Europe, A Comparison • , , •••• , ••••• , ••••••••• , , , 601
E. Lindsay and R. Tedford

NEW PERSPECTIVES

The Past, the Present, and the Future 625


V. Fahlbusch and P. Mein

Contributors . . . . . . . . . • . . • . . . . . . . . . . . • . . . . . . . . • • . . . . . . . . . . • . . . . . • . • . . • . 6Z9

Subject Index • • . . . . . . . . . . . . . • . . . . . • . . . . . . . . . . . . . . . . . . • • . . • . . . . . . . . . . . • 631

Taxonomic Index (Mammal Genera) . •• . . . •. . . • . . •. •. . . •. . . . . • . • . . . • . . •. . . 651

ix

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