pleistocene mammals in europe

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PleistoceneMammalsof EUfopePleistoceneMammalsof EuropeBjorn Kurten~ ~ AldineTransactionU A Division of TransactionPublishers '-t;", NewBrunswick (U.S.A.) andLondon (U.K.)-===Firstpaperback printing 2007Copyright 1968 by BjornKurten.All rights reserved under International and Pan-AmericanCopyrightConventions. No part of this book may be reproduced or transmitted in any formor byanymeans, electronicor mechanical, includingphotocopy, recording,oranyinformationstorageandretrieval system, withoutprior permissioninwritingfromthe publisher. All inquiries should be addressedto AldineTrans-action, ADivision of TransactionPublishers, Rutgers-TheStateUniversityof New Jersey, 35Berrue Circle, Piscataway,NewJersey08854-8042. www.transactionpub.comThis bookisprintedonacid-freepaper that meets theAmericanNationalStandard for Permanence of Paperfor Printed Library Materials.Library of Congress Catalog Number: 2007020579ISBN: 978-0-202-30953-8Printed in the UnitedStatesof AmericaLibrary of Congress Cataloging-in-Publication DataPleistocene mammals of Europe.p.cm.Orginially published: London: Weidenfeld &Nicolson; Chicago: AldinePub. Co., 1968..Includes bibliographical references and index.ISBN 978-0-202-30953-8 (pbk.:acid-free paper)-ISBN978-0-202-30946-0 (pbk.: acid-free paper)1. Mammals, Fossil-Europe. 2. Paleontology-Pleistocene.3. Paleontology-Europe. I. Title.QE881.K8 2007569.094--dc222007020579ContentsForewordPartOne Faunal SequenceinEuropeviiI SETTINGTHE ST AGE 32 THEVILLAFRANCHIAN, PRELUDE TO THE ICE AGE 83 CHRONOLOGYOF THE ICE AGE 184 THE AGE OF INTERGLACIALS 225 THE AGE OF GLACIATIONS 28Part Two Pleistocene Mammal Species6 INTRODUCINGTHE PLEISTOCENEMAMMALS7 ORDER INSECTIVORA8 ORDERCHIROPTERA9 ORDER PRIMATES10 ORDERCARNIVORAI I ORDERPROBOSCIDEA12 ORDERPERISSODACTYLA13 ORDERARTIODACTYLA14 ORDERRODENTIA15 ORDERLAGOMORPHAPart Three TheChangingFauna16 THE SPECIES PROBLEMINTHEQUATERNARY17 SIZE ANDNUMBERS18 ORIGINATIONOF SPECIES19 FAUNAL TURNOVER20 ANIMALGEOGRAPHY21 MANANDTHE FAUNA3942515862130139153191226237244253260265270vCONTENTSAppendix StratigraphicRange ofSpeciesReferencesIndex27528634ForewordTHE mammals of the Pleistocene are but now being discovered by theevolutionists. Here we get the sought-for tie-in between zoology andpalaeontology:many Pleistocene mammals are still inexistence andmay bestudied as living beings against the background of their ownfossil history, extending back formillennia through geological time.But thePleistocene of Europe anditsmammalian faunais of in-teresttomany othersbeside thezoologist, the evolutionist, andthepalaeontologist. It is of exceptional importance tothe student of hu-man origins and of Stone Agearchaeology. Among others who mayprofit from a study of this topic may be mentioned the geologist, thepalaeoclimatologist andthepalaeogeographer, andindeedanyonewhotakes an interest in natural historyandwishes to re-enter acolourful past.The first version of this book wasa semipopular paperback intheSwedish Aldus series. The present edition is completely rewritten andgreatly expanded, butIhave retained the non-technical approach tomake the story accessible toreaders with varying backgrounds. Thefirst part of the book is an outline of the Pleistocene history of Europe,with its climatic changes and succession of mammalian faunas. In thesecond partarelistedall thespecies of Mammalia knownfromthePleistoceneandPostglacial of Europe, withtheevolution, rangeintimeandspace, andmodeof lifeset downfor eachspecies, asfarasknown. Thefinal part isan evaluationof thestoryintermsofevolution and palaeogeography.The species is the pivot of the present treatment. The species is, ofcourse, the most important taxonomic category in the study of evolu-tion [195], and towrite a comprehensive zoology of all the species ofanentiregeological epoch struck measanexciting andchallengingproject. In most palaeontological texts the genus is the basic category,not the species. Withincreasingprecisionofstudythe species isgraduallytakingover this role, andthegreatest progress is beingFOREWORDmade in the case of the Quaternary, for which we have a moredetailed record than forthe more distant past.Originally the study had been planned to cover only the Carnivora,but it was soonfoundnecessarytointroducematerialontheothermammalian orders; in the end equal emphasis was given to all of them.It should be noted,however, that much of what issaid about carni-vores here isbased on original research, while this is not the case asregards the other orders of mammals.Thanks toa three-year travel and research grant from the Univer-sity of Helsinki I havehadopportunitytostudymost of thelargercollections of European Pleistocene mammals on thespot. Much ofthe work wasdone during two long sojourns inEngland in1961 and1962, during which Dr K. A. Joysey (Cambridge) and Dr A. J.Sutcliffe (London) provided most valuable assistance and criticism.Ihavealsoprofitedgreatlyfromstimulatingdiscussionswith DrK.Kowalski (Krakow). Dr M. Crusafont Pairo (Barcelona) kindlychecked the material on Spanish fossils and provided important newinformation. Many other colleagues have contributed by showing methe collections intheir care, and giving generously of their time andadvice. Among theseIwish particularly to mention Drs K. D. Adam(Stuttgart), c;:. Arambourg (Paris), A. Azzaroli (Florence), H. Bohlken(Kiel), M. Degerbel (Copenhagen), K. Ehrenberg (Vienna), E. W.Guenther (Kiel), D.A.Hooijer (Leiden), }.Hiirzeler (Basle), V.Jaanusson (Uppsala), H. D. Kahlke (Weimar), F.E. Koby (Basle),J.P. Lehman(Paris), K. P. Oakley(London), H. E. P. Spencer (I ps-wich), D. Starck (Frankfurt), L. Thaler (Montpellier), E. Thenius(Vienna), H. Tobien(Mainz),J. F. deVillaltaComella(Barcelona),R. West (Cambridge), P. Woldstedt (Bonn) andH. Zapfe(Vienna).Mrs Sonia Cole read a preliminary draft of the text and made valuablesuggestions for improving it. To allthesepersons and institutionsIwish toexpress my sincere gratitude. Finally, Iwish todedicate thisbook to three inspiring teachers: Pontus Palmgren (Helsingfors),Birger Bohlin (Uppsala) and George Gaylord Simpson (Cambridge,Massachusetts) .Helsingfors BJORN!ZURTENviiiPart OneFaunal SequenceinEuropeChapter1Setting the StageAHUNDREDMILLIONyearsago, duringthereign of thedinosaurs,theearthwas amoist, warmplanetunderatropical sun. Much ofEuropewas floodedby theChalkSea, teemingwithreptilianlife,while ponderous monsters moved slowly over the endless beaches andplains of the low-lying land. This was the crest of one of the great heatwaves in geological time. It lasted more than 200 million years.To find the trough preceding it we have to move back into Permiantimes,250 million yearsago and more;there once again wefindthepresence of continental glaciers, a sight which wastobecomeso fa-miliar duringtheperiod withwhichweareconcerned, thePleisto-cene. After Permian times the temperature curve ascended graduallyand the face of the globe became more uniform. Mountains were ero-ded and asthey wereplaned down thelandscape became more andmoremonotonous. Thesealevel roseandthecontinental marginswere flooded. The pace of geological processes had become impercep-tible, as though in a world left over to entropy.Afterthispeak thetemperaturecurvebegantotrenddownwardagain. The sea, which had spread halfway across the continents, verygradually, and with manyhalts andreversals,began torecede. Thecoastal lagoons dried up and new land waslaid bare. Once more theinner forcesof the earth ground into gear and new mountainswerebuilt up. The poles cooled off more rapidly than the tropics and onceagain the earth was girded by distinctive climatic belts. The giant rep-tilesbecame scarce and finally none were left. The mammals, whichhadledatimidlifeinthebackground, developedlargerandmorevaried forms. ThustheCretaceous passed into the Tertiary Period,65 million years ago.Still the temperature fell. Thisis the basic theme throughout theTertiary, the background against which we must view the long, com-plicated evolutionary history of the mammals: over millions of years3FAUNAL SEQUENCEINEUROPEthe temperature fell, the searetreated and the mountain chains rose.These tendencies fluctuated in strength and direction, but they werealways there. Out of the great Mediterranean sea that extended fromEurope through Asia inthe beginning of the Tertiary, known as theTethys Sea, therearoseamountainous archipelagowhichbecameconsolidated and grew higher. Eventually the firstsnow glittered onits highest peaks andthe Alps, the Himalayas andtheAmericanCordillera were born.In Europe the climate was subtropical and as late as 15million yearsago lush jungles covered most of the continent. But as the temperaturecontinued to fall, about 10 million years ago in the early Pliocene therecameasharplymarkedshift. Theclimatebecamedrier andgreatgrasslands, savannas and steppes spread over the continent.To uswho observe it inthe foreshortenedperspective of the geo-logical time scale, the shift appears dramatically sudden; but it musthave gone on formany thousands of years and to the living beings ofthe time it would not have been noticeable. It is easyenough forustoobserve the descent of the temperature curve - by ingenious methodsit can actually be directly measured [74]. Yet the change in the face ofthe earth at that time was slow in comparison with what wasto followduring the Ice Age: this was like an explosion, a total revolution in thetempo of geological events. The world we now liveinis the world oftheIce Age. Even if wedisregardtheinfluence of man it isa worldwhich is dynamically changing at anabnormal rate; itis ina state offlux unequalled since the Permian Ice Age more than 250 million yearsago.Measuredbyoureverydaystandardseventhistempomayseemslow. Yet we are nowable within a single generationto observesuch sub-phases of the climatic evolution as the evident ameliorationduringthefirst half of thiscentury. Thisistypicalof theIceAge.Temperature fluctuations of a kind requiring milli

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