HOMINOID EVOLUTION AND CLIMATIC CHANGE IN EUROPE VOLUME 1 The Evolution of Neogene Terrestrial Ecosystems in Europe ppt

van der Meulen, 8 The Late Miocene small mammal succession from France, with emphasis on the Rhoˆne Valley localities Pierre Mein 140 9 Late Miocene mammals from Central Europe 10 An ove

Europe has changed greatly in terms of climate and environment in the past

20 million years Once, there were sub-tropical forests, but by the end of theMiocene, 5 million years ago, these had all gone This unique book providesevidence for the past climatic history of Europe and the Mediterranean inrelation to hominoid evolution Many diVerent lines of evidence are broughttogether including studies speciWcally on past climates and the application

of climate modelling, the reconstruction of past geographical events, andthe eVects they had on European environments and the plants and animalsliving in them Together, they form a coherent and consistent image ofenvironmental and climatic change in Europe from 18 to 1.6 million yearsago, for all those interested in mammalian and human evolution

JORGE AGUSTIis Director of the Institute of Paleontology, M Crusafont, in Sabadell,Spain He specialises in the evolution of the Neogene and Quaternary small mam-malian faunas in relation to environmental changes

LORENZO ROOKis a researcher in the Department of Earth Sciences at the University ofFlorence, working on fossil primates and carnivora and on Neogene/Quaternarybiochronology

PETER ANDREWSis a research scientist at the Natural History Museum in London,where he works on fossil primates, taphonomic and palaeoecological issues relating

to the early stages of human evolution

XXXXXX

H OM I N OI D EV OLU TI ON A N D C L I M AT I C C H AN GE I N E UR O P E

VOL UME 1

Th e E v ol u ti o n o f N e og e ne T e rr e s t r i al

E c os y s te m s i n E u r op e

XXXXXX

HOM I N O I D EVO L UT I ON AN D C LI M AT I C C H AN GE I N EUR O P E

  

Cambridge, New York, Melbourne, Madrid, Cape Town, Singapore, São Paulo Cambridge University Press

The Edinburgh Building, Cambridge  , United Kingdom

First published in print format

ISBN-13 978-0-521-64097-8 hardback

ISBN-13 978-0-511-06619-1 eBook (NetLibrary)

© Cambridge University Press 1999

1999

Information on this title: www.cambridge.org/9780521640978

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www.cambridge.org

C o n t en t s

1 Introduction Jorge Agustı´, Lorenzo Rook and Peter Andrews 1

PART I Palaeogeography of the circum-Mediterranean region 7

2 Mediterranean and Paratethys palaeogeography during the

3 Pliocene tephra correlations between East African hominid

localities, the Gulf of Aden, and the Arabian Sea

4 Climatic perspectives for Neogene environmental

reconstructions Eileen M O’Brien and Charles R Peters 55

5 A critical re-evaluation of the Miocene mammal units

in Western Europe: dispersal events and problems of correlation

6 Large mammals from the Vallesian of Spain

Jorge Morales, Manuel Nieto, Meike Kholer and Salvador

7 Trends in rodent assemblages from the Aragonian

(early–middle Miocene) of the Calatayud-Daroca Basin,

Aragon, Spain Remmert Daams, Albert J van der Meulen,

8 The Late Miocene small mammal succession from France,

with emphasis on the Rhoˆne Valley localities Pierre Mein 140

9 Late Miocene mammals from Central Europe

10 An overview on the Italian Miocene land mammal faunas

11 The Miocene large mammal succession in Greece

12 Chronology and mammal faunas of the Miocene Sinap

Formation, Turkey Juha Pekka Lunkka, Mikael Fortelius,

13 The Late Miocene small mammal succession in

Ukraine Valentin A Nesin and Vadim A Topachevsky 265

PART III Palaeoenvironments: non-mammalian evidence 273

14 Marine invertebrate (chieXy foraminiferal) evidence

for the palaeogeography of the Oligocene–Miocene of western

Eurasia, and consequences for terrestrial vertebrate migration

15 Palaeoclimatic implications of the energy hypothesis from

Neogene corals of the Mediterranean region Brian R Rosen 309

16 Contribution to the knowledge of Neogene climatic

changes in western and central Europe by means of non-marine

17 Sedimentary facies analysis in palaeoclimatic reconstructions.

Examples from the Upper Miocene–Pliocene successions of

south-central Tuscany (Italy) Marco Benvenuti, Mauro Papini

18 Neogene vegetation changes in West European and

West circum-Mediterranean areas Jean-Pierre Suc,

Se´verine Fauquette, Mostefa Bessedik, Adele Bertini,

Zhuo Zheng, Georges Clauzon, Danica Suballyova,

Filomena Diniz, Pierre Que´zel, Najat Feddi, Martine Clet,

the late Ezzedine Bessais, Naima Bachiri Taou Wq,

PART IV Palaeoenvironments: mammalian evidence 389

19 Shrews (Mammalia, Insectivora, Soricidae) as

paleoclimatic indicators in the European Neogene

20 Mammal turnover and global climate change in the late

Miocene terrestrial record of the Valle`s-Penede`s basin

(NE Spain) Jorge Agustı´, Lluı´s Cabrera, Miguel Garce´s and

viii

21 Palaeoenvironments of late Miocene primate localities

in Macedonia, Greece Louis de Bonis, Genevieve Bouvrain and

22 The paleoecology of the Pikermian Biome and the

savanna myth Nikos Solounias, J Michael Plavcan,

23 Vicariance biogeography and paleoecology of Eurasian

Miocene hominoid primates Peter Andrews and

ix

Department of Anatomy, Laboratory of Paleobiology, Howard University,

520 W Street, Washington DC 20059, USA

Lluı´s Cabrera

Grup de Geodinamica i Analisi de Conques, Department de EstratigraWa i

Paleontologia, Universitat de Barcelona, Campus Pedralbes, E-08028

Barcelona, Spain

Georges Clauzon

CEREGM (UMR 6635 CNRS), Europoˆle de l’Arbois, BP 80, 13545

Aix-en-Provence Cedex 04, France

Martine Clet

Morphodynamique continentale et coˆtiere, Universite´ de Caen, 24 rue des

Tilleuls, 14000 Caen, France

Nathalie Combourieu-Nebout

Pale´ontologie et Stratigraphie (ESA 7073 CNRS), case courrier 106,

Universite´ Pierre et Marie Curie, 4 place Jussieu, 75252 Paris Cedex 05,

France

Remmert Daams

Departamento de Paleontologia y UEI, Facultad de Ciencias Geolo´gicas,

Universidad Complutense y CSIC, Ciudad Universitaria, 24040 Madrid,

Spain

Louis de Bonis

Lab Ge´obiologie, Biochronologie et Pale´ontologie humaine, Faculte´ de

Sciences, Universite´ de Poitiers, 40 av du Recteur Pineau, F 86022 Poitiers

Departamento de Geologia, Universidade de Lisboa, rua Escola

Polytecnica 58, 1294 Lisboa Codex, Portugal

Burkhart Engesser

Naturhistorisches Museum, Agustinergasse 2, CH 4001 Basel, Switzerland

Daniela Esu

Dipartimento di Scienza della Terra, Universita´ ‘La Sapienza’, P le A

Moro, 5, 00185 Roma, Italy

Se´verine Fauquette

Lab de Botanique Historique et Palynologie, Inst Me´diterrane´en

d’Ecologie et de Pale´oe´col., Faculte´ des Sciences de Saint Je´roˆme, 13397

Marseille Cedex 20, France

xi

Najat Feddi

De´partment des Sciences de la Terre, Faculte´ des Sciences Semlalia,Universite´ cadi Ayyad, avenue Prince Moulay Abdellah, BP S15 Marrakech,Morocco

Mikael Fortelius

Finnish Mueseum of Natural History, University of Helsinki, PO Box 11,

0014 Helsinki, Finland

Jens Lorenz Franzen

Forschungsinstitut Senckenberg, Senckenberg-Anlage 25, D-60325

Frankfurt am Main, Germany

Miguel Garce´s

Grup de Geodinamica i Analisi de Conques, Department de EstratigraWa iPaleontologia, Universitat de Barcelona, Campus Pedralbes, E-08028Barcelona, Spain

Robert Wynn Jones

BP, Exploration Operating Co Ltd, Chertsey Road, Sunbury on ThamesMiddlesex TW16 7LN, UK

Juha Pekka Lunkka

Finnish Museum of Natural History, University of Helsinki, PO Box 11,

Henriette Meon

UFR des Sciences de la Terre, Universite´ Claude Bernard – Lyon I, 27–43

boulevard du 11 Novembre, 69622 Villeurbanne Cedex, France

Jorge Morales

Museo Nacional de Ciencias Naturales, CSIC, Jose´ Gutierrez Abascal 2,

E-28006, Madrid, Spain

Salvador Moya `-Sola`

Institute of Pakontology, M Crusafont, Escola Industrial 23, E-08201

Sabadell, Spain

Valentin A Nesin

Institute of Zoology, Ukrainian Academy of Sciences, 15 Bogdan

Khmelnitsky Str, 252030 Kiev 30, Ukraine

Manuel Nieto

Museo Nacional de Ciencias Naturales, CSIC, Jose´ Gutierrez Abascal 2,

E-28006, Madrid, Spain

Lab de Botanique Historique et Palynologie, Inst Me´diterrane´en

d’Ecologie et de Pale´oe´col., Faculte´ des Sciences de Saint Je´roˆme, 13397

Marseille Cedex 20, France

xiii

Forschungsinstitut Senckenberg, Senckenberg-Anlage 25, D-60325

Frankfurt am Main, Germany

Danica Suballyova

UFR des Sciences de la Terre, Universite´ Claude Bernard – Lyon I, 27–43

Bd du 11 Novembre 1918, 69622 Villeurbanne Cedex, France

Jean-Pierre Suc

Centre de Pale´ontologie Stratigraphique et Pale´oe´cologie (UMR 5565),Universite´ Claude Bernard – Lyon I, 27–43 Bd du 11 Novembre 1918, 69622Villeurbanne Cedex, France

Naima Bachiri Taou Wq

De´partment de Ge´ologie, Faculte´ des Sciences Ben M’Sik, Universite´Hassan II Mohammedia, BP 7955, Casablanca, Morocco

Giovanni Testa

Dipartimento di Scienze della Terre, Universita´ di Firenze, via G la Pira 4,

50121 Firenze, Italy

Vadim A Topachevsky

Institute of Zoololgy, Ukrainian Academy of Sciences, 15 Bogdan

Khmelnitsky Str, 252030 Kiev 30, Ukraine

xiv

Albert J van der Meulen

Department of Stratigraphy & Paleontology, Institute of Earth Sciences,

Budapestlaan 4, 3508 TA, Utrecht, The Netherlands

A c kn o wl ed geme nt s : T h e Eu r o p ea n

S c i e n c e F o u n d a t i o n

The European Science Foundation (ESF) acts as a catalyst for thedevelopment of science by bringing together leading scientists for fundingagencies to debate, plan and implement pan-European scientiWc andscience policy initiatives

ESF is the European association of more than 60 major national fundingagencies devoted to basic scientiWc research in over 20 countries Itrepresents all scientiWc disciplines: physical and engineering sciences, lifeand environmental sciences, medical sciences, humanities and socialsciences The Foundation assists its Member Organisations in two main ways

ways: by bringing scientists together in its scientiWc grammes, networks, exploratory workshops and Europeanresearch conferences, to work on topics of common concern;and through the joint study of issues of strategic importance

pro-in European science policy

It maintains close relations with other scientiWc institutions within andoutside Europe By its activities, the ESF adds value by cooperation andcoordination across national frontiers and endeavours, oVers expertscientiWc advice on strategic issues, and provides the European forum forfundamental science

1 Introduction

Jorge Agustı´, Lorenzo Rook and Peter Andrews

The late Neogene (the period between − 14 and − 2.4 Ma) is one of the mostinteresting phases in understand the present conWguration of terrestrialecosystems It was during this time that the change took place from themiddle Miocene dominant subtropical forests that stretched across south-ern Europe and western Asia to a more open but still wooded biotope thatnow prevails in warm–temperate areas This change in vegetation, whichstrongly aVected the composition of mammalian faunas, seems to be linked

to the rapid spread of grasses around 8–10 Ma ago Moreover, in the lateNeogene, climatic shifts and falling temperatures due to the spread of theAntarctic Ice, were followed by the Plio-Pleistocene Arctic glaciations in theNorthern Hemisphere Furthermore, during the late Neogene, importantchanges at the regional level gave rise to the present conWguration of the OldWorld land masses, and the successive drying of inland seas in the Europeanarea facilitated faunal interchange between Europe and Central Asia andAfrica At the same time, other tectonic processes like the Himalayan andthe Tibetan uplifts and the opening up of the great eastern African basinsand Red Sea, although working in opposite directions, favoured the pro-cesses of speciation and isolated evolution

All these phenomena must have had a strong inXuence on hominidevolution between 14 and 2 million years ago (Ma) The diversity of hominidspecies recorded in that period must be considered in relation to the chang-ing environmental conditions and to constrictions imposed by the existence

of signiWcant zoogeographical barriers As a consequence, at around 10 Maago Eurasia displayed a large variety of apes, with gracile forest-dwellers

(Dryopithecus, Sivapithecus), dry-adapted forms (Ankarapithecus) and large robust gorilla-like forms (Graecopithecus), and all this variety in a period

when there is a huge information gap in Africa In some ways, the EurasianMiocene radiation of apes parallels the trends observed in Africa 5 Ma laterthat led to the appearance of theWrst bipedal hominids DiVering from thePliocene hominid evolution in Africa, the changing environments did notresult in Eurasia in the emergence of bipedal apes, but in the extinction ofgeneralised morphotypes and the persistence of highly specialised forms in

relic areas (Gigantopithecus, Pongo) However, knowledge of the ecosystems

in which hominids of the late Miocene evolved is still incomplete, and verylittle integrated analysis of data has been attempted, in particular makinguse of modern techniques for palaeoenvironmental analysis such as iso-topes, pollen, micromammals

In order to focus on these issues and to foster co-operation betweenscientists dealing with them, the scientiWc network ‘Hominoid evolution

and environmental change on the Neogene of Europe’ was approved bythe European Science Foundation in 1995 One of the goals of this net-work was to create a database on Neogene Mammals of Eurasia, thatcould be used for further analysis of the ecosystems where hominoidslived The network also organised three workshops in order to analyse

diVerent aspects of the late Neogene time and their relevance to hominoidevolution

The Wrst workshop was held in Sant Feliu de Guixols (Spain), fromthe 24th to the 27th of October 1997, and had as the main topic ‘TheVallesian’ For analysing this particular interval of the late Miocene, 24scientists from 11 countries attended this Wrst workshop The workshopwas mainly devoted to analysing the faunal and environmental changesthat took place during the period known as the Vallesian stage However,this analysis was not strictly limited to that period but included also theintervals immediately preceding and succeeding the Vallesian (eg., lateAragonian and early Turolian) In this way, a number of presentationsdealt with regional mammalian successions ranging from Spain to South-

ern Asia: Spain (Agustı´ et al., Morales et al.), France (Mein), Italy (Rook &

Engesser), Central Europe (Franzen & Storch), the Aegean area (Koufos &

De Bonis), Anatolia (Fortelius et al.) and Eastern Europe (Nesin

& Topachersky) In the case of Eastern Spain, the communications ented by the Spanish team were complemented with a Weld-trip in theValle`s-Penede`s Basin, which enabled the participants to have an accurateidea of the work developed in this area

pres-The emphasis of the workshop was put on the comparison and closecorrelation of long sections bearing sequences of large and small mammallocalities, instead of the usual correlation between isolated localities lacking

an accurate geological context (for instance, in the case of karsticWssure

inWllings) In this way, comparison could be made between sections bearing

a detailed magnetostratigraphic analysis, particularly the sections of thePotwar Plateau (Pakistan), the Valle`s-Penede`s (Spain) and Sinap (Turkey).Another signiWcant topic of the workshop was the so-called ‘Mid-VallesianCrisis’, an extinction event that followed the change from the middle to thelate Miocene Results reviewed at the workshop suggested that a changesimilar to the Mid-Vallesian Crisis of Western Europe occurred at diVerenttimes in diVerent areas The change was already established in the earlyVallesian in the so-called Sub-Parathetyan Province (data from Maragheh

by Bernor), but in Western Europe did not take place until about 9.6 Ma ago(data from the Valle`s-Penede`s by Agustı´ and co-workers) On the otherhand, in Central Europe, several species indicating wet, forested conditionspersisted well into the late Vallesian (data from Franzen & Storch) Finally,

2

such a change was not seen in Southern Asia (Pakistan) until 8–7.2 Ma (data

from the Potwar Plateau)

The second workshop took place in Certosa di Portignano (Siena, Italy),

devoted to the ‘Climatic and environmental change in the Neogene of

Europe’; 25 scientists from 11 countries and diVerent palaeoenviromental

disciplines had the rare opportunity to experience a real interfacing of data

from the terrestrial ecosystems, the shallow marine realm and the deep sea

AWrst set of contributions dealt with the palaeogeographic evolution of the

Tethys area and the eVects of changes in the extent of the Tethys sea on

faunal distributions and climate evolution F Ro¨gl (Vienna) presented a

sketch of the evolution of this area from the early Tertiary to the late

Miocene With the contact between the Arabian plate and the Anatolian

plate, land bridges formed between Africa and Eurasia opened and closed

throughout the middle Miocene, beginning about 19 Ma No hominoid

primates are known this early in Europe, but of particular signiWcance to

hominoid migrations was the end-Burdigalian regression of the sea at

around 16 Ma, as this coincides with the earliest evidence of hominoids in

central Europe

Contributions to climate modelling during the late Neogene were

pro-vided by P B DeMenocal & F H Brown, and by E O’Brien According to

DeMenocal & Brown, marine records of African climatic variability

docu-ment a shift toward prolonged and seasonally more arid conditions after 2.8

Ma This is linked to cold North Atlantic sea-surface temperatures

asso-ciated with onset of Arctic Ice sheets Major changes in African faunas

coincide with this climatic change suggesting that some speciation events

may have been climatically mediated E O’Brien demonstrated that a large

proportion (79%) of African woody plant species richness is accounted for

by two aspects of climate, annual rainfall and an optimised function of

energy (minimum monthly potential evapotranspiration) Finally, the

palaeobotanical approach included the point of view of the palynological

analysis (J P Suc, A Bertini, G Clauzon & D Suballyova)

Evidence for climate change was considered both from invertebrate and

vertebrate evidence Data presented by Rosen demonstrated that most coral

reefs (and their associated z-corals) occur in the Mediterranean area in three

major high sea-level phases, corresponding to the early (Aquitanian),

middle (Langhian–Serravalian) and late Miocene (Tortonian–early

Mess-inian) Small mammals were discussed by Reumer (Insectivores), Daams et

al (rodents from Central Spain) and Agustı´ et al (mainly rodents from

Eastern Spain) According to Reumer, ecological studies have shown that

environmental moisture may be the ultimate determinant of within-habitat

diversity and numerical abundance of soricids, which can be taken as a good

3

indicator of relatively warm and humid palaeoclimate Shrews (Soricidae,Insectivora) are among the most relatively sensitive mammals against cli-matic shifts because of their small size and high surface/volume ratio Threemain periods of faunal turnover, corresponding to humid and warm condi-tions, characterise the Neogene history of shrews: the early Miocene (19–20Ma), the early–late Miocene (Vallesian; 9–11 Ma), and the latest Miocene toPliocene (6 and 2.5 Ma) Climatic deterioration at around 2.7–2.3 MaWnallycaused a severe reduction of the European shrew fauna Among rodents, theanalysis of the Calatayud-Daroca succession developed by Daams and co-workers also suggests, as for the shrews, that species diversity is morerelated to relative humidity than to temperature Quantitative analysis of therodent succession developed by Agustı´ and coworkers in the Valle`s-Penede`sBasin allows the recognition of an alternation of dry and humid phasesduring the Miocene and early Pliocene times Early Miocene localities indi-cate forested, humid conditions, similar to those recorded in the earlyMiocene of Central Europe Increasingly dry conditions are recorded acrossthe late early and middle Miocene (early middle Aragonian), but a return tomore humid conditions is observed in the late Aragonian (Serravallian)times The middle/late Miocene boundary coincides with a relatively dryperiod, followed again by a humid peak in the early Vallesian (early Tor-tonian), and it is this time that coincides with the maximum abundance ofhominoid remains in Western Europe Again, as in the case of shrews, itappears that species diversity is more related to relative humidity than totemperature.

Contributions dealing with large mammal associations were those of N.Solounias, M Plavcan, L Witmer & J Quade, L de Bonis & G Koufos, L.Kordos, and R Bernor After a detailed analysis based on a variety of sources(dental and postcranial ecomorphology of bovids, palynology, isotopes),Solounias and co-workers arrived to the conclusion that the main habitat ofthe Pikermian mammals was not a savanna (‘the savanna myth’), butsclerophyllous evergreen woodland similar to today’s mixed monsoon for-est and grassland glades of north Central India Large mammals with luckyexaptations migrated into Africa from the Pikermian bioprovince A similartopic was developed by de Bonis & Koufos, who found in Greek faunasevidence of a trend towards drier conditions in the late Miocene On theother hand, the disappearance of hominoids and other forest elements inthe Pannonian Basin at 9–7 Ma was attributed by Kordos to the regression ofthe Pannonian sea rather than to a general climatic trend Finally, Bernor &Andrews discussed the patterns of hominoid immigration, dispersal andextinction Hominids entered Europe in the middle Miocene because landcrossings of the Tethys were possible at this time and the subtropical forest

4

environments were suitable By virtue of the taxonomic diversity and

re-stricted biogeographic ranges alone, Eurasian Miocene hominids show

strongly vicariant evolutionary patterns This suggests a model whereby a

founding species extends its range under favourable and speciWc

environ-mental circumstances and then becomes geographically restricted to

refugia by geographic ( = tectonic and palaeogeographic) and/or

environ-mental events A frequent byproduct of vicariance, exercised over millions

of years time, is homoplasy, and it is evident that there was a great deal of

homoplasy in Miocene hominids

The nature of the environments occupied by apes in Europe had many

structural similarities with the environments in Africa with which they are

associated at this time Palaeoecological evidence suggests that African

middle Miocene apes lived in seasonal woodlands and forests, for example

at Fort Ternan and Maboko Island The hominids at these sites were partly

terrestrial and with their large thick-enamelled teeth were adapted for

similar diets to some of the European apes The earliest European apes were

similar in being both partly terrestrial and with almost identical dietary

adaptations The community structure of the mammalian faunas in the

African and European sites were extremely similar, and by inference the

ecosystem they occupied was also similar The similarities in locomotor and

dietary adaptations of the African and European apes at this early stage

indicate further that their position in their respective ecosystems was also

very similar In one sense, therefore, these middle Miocene taxa in Europe

were not as distinct from their African relatives as taxonomic divisions and

their geographic separation may appear to indicate

Towards the end of the middle Miocene, at 13–12 Ma, the trends of

partial terrestriality and thick-enamelled frugivory continued in a group of

fossil apes assigned either to the pongine clade or to a paraphyletic group

unrelated to any living They are associated with a range of open forest to

woodland environments ranging from southeast Europe to China, and one

genus at least may be related to the orang utan At the same time, the more

arboreal and suspensory Dryopithecus emerged in association with closed

subtropical forest environments where they are sometimes found

asso-ciated with Pliopithecus and Anapithecus Similar environments and a

simi-lar, possibly heritage, adaptation for suspensory locomotion persisted in

Oreopithecus, although it has recently been argued that this fossil ape may

also have had adaptations for terrestrial bipedal mode of locomotion and

the evolutionary relationships of Oreopithecus are still unclear.

Later in the Miocene, the Alpine–Himalayan orogeny caused major

changes in land–sea relations, global climatic circulation patterns and

sea-sonality particularly in Central Asia Regression of the Paratethys likewise

5

caused a shift of habitats to greater seasonality and replacement of green subtropical forests by deciduous woodlands and, progressively in thelate Miocene, more seasonal warm temperate woodlands with progressivelymore open habitats Hominid primate distribution tracked these changesclosely during the 12–9 Ma interval, contracting their range from both westand east and Wnding temporary refuge in southeastern Europe, wherefavourable subtropical conditions persisted for a time after being lost else-where Hominids disappeared from this regionWnally during MN11, al-though they persisted until MN12 in local insular habitats in Italy and thelatest Miocene of China.

ever-6

PART I

Palaeogeography of the

circum-Mediterranean region

2 Mediterranean and Paratethys

Palaeogeography during the

Oligocene and Miocene

Fred Ro¨gl

Introduction

The Cenozoic conWguration of continents and oceans is strongly inXuenced

by plate tectonic movements Opening and closing pathways for mammalmigrations and marine exchanges are one of the triggering forces for faunalevents and evolution The impacts of a vanishing Tethys Ocean in themid-Cenozoic are not only important for the marine and continental biotas

of Eurasia and the Mediterranean, but also inXuenced the environmentalconditions worldwide The dispersal of continents in the Southern Hemi-sphere with the northward movement of the Indian and Australian conti-nents, together with the counterclockwise rotation of Africa, closed downthe Tethys Ocean Parallel to these movements the Atlantic Ocean opened.The Mesozoic oceanic circulation patterns changed to varying conditionswith decreasing temperatures in the Cenozoic Based on oxygen isotopes(Kennett, 1995), bottom water temperatures were highest in the late

Palaeocene–early Eocene (c 55 Ma) Distinct steps to colder conditions

followed around the Eocene/Oligocene boundary (33–35 Ma), in the middleMiocene (15 Ma), and in the Pliocene (3 Ma)

DiVerent palaeogeographic reconstructions over the past two decadeshave attempted to solve the development of the Cenozoic Mediterraneanand Paratethys areas In many cases these reconstructions depend on thepresent sediment distribution and do not consider palinspastic reconstruc-tions based on plate tectonic movements (e.g Vinogradov, 1967–69; Senes &

Marinescu, 1974; Steininger et al., 1985a; Hamor & Halmai, 1988; Cahuzac et

al., 1992; Popov et al., 1993) Other reconstructions include too long a time

span within one time slice to present a distinct time level within a strongly

changing environment (e.g Biju-Duval et al., 1977; Dercourt et al., 1985,

1993) The best reconstructions, based on tectonic, sedimentological andstratigraphic investigations are currently available for the Western Mediter-

ranean (Boccaletti et al., 1986, 1990) The sketches for the Neogene,

pro-duced by Ro¨gl & Steininger (1983) and Steininger & Ro¨gl (1984), were based

on plate tectonic hypotheses, sediment distribution, Eurasian mammalmigrations, and marine faunal similarities between the Mediterranean andParatethys At that time the information on faunal development and strati-graphic correlation was poor for the Eastern Paratethys Therefore a revision

of those interpretations, including the early history of the Paratethys wasdiscussed by Ro¨gl (1998a,b) The sketches presented here are based on plate

distributions at three levels from the Eocene to the late Miocene (Scotese et

al., 1988) The goal of the present palaeogeographic sketches is to explain

mammal migration possibilities in and between Eurasia and Africa Some

marine connections are still being debated (Adams, 1998), and certain

seaways proposed here are highly speculative The open questions will

require further tectonic and palaeontological studies

A postulate for any time slice is the exact stratigraphic correlation of the

diVerent basins included in the reconstructions The stratigraphic

correla-tion chart (Table 2.1) is based on the most recent time tables of Berggren et

al (1995) and Steininger et al (1996), correlated to the Paratethys (Popov et

al., 1993; Jones & Simmons, 1996; Ro¨gl, 1998b).

Birth of the Paratethys

In the late Eocene the Indian Plate collided with Eurasia The Tethys Ocean

vanished, leaving as relics the Mediterranean Sea at its western end, and to

the north the intercontinental Paratethys Sea in Eurasia (Fig 2.1) In the late

Eocene, a pelagic Globigerina marl facies developed from the western

Medi-terranean to the Alpine–Carpathian belt and the inner-Asian Transcaspian

Basin; north–south gradients in planktic faunal assemblages were evident

In the shallower parts of the basin tropical, larger foraminifera and mollusc

faunas thrived In the Eastern Paratethys the molluscs had a latitudinal

distribution, with the most tropical assemblages in Transcaucasia

(Akhal-tsikhe depression), whereas in northern Ustyurt a northern fauna of low

diversity was recorded (Krasheninnikov, 1974; Popov et al., 1993; Popov,

1994) From the eastern part of the later Paratethys, the tropical Tethys

Ocean communicated with the Polar Sea via the shallow Turgai Strait in

western Siberia during the middle–late Eocene (Vinogradov, 1967–69; Popov

et al., 1993) This marine barrier prevented a continental faunal exchange

between Asia and Europe During the Eocene, western and middle Europe

existed as an archipelago, with a distinct reduction of mammal diversity

during the Priabonian Northern Europe formed a landmass connected by

the De Geer Route (over Svalbard) with North America (Ko¨nigswald, 1981)

Around the Eocene/Oligocene boundary, continental collision and

movements along the Alpine–Himalayan tectonic belt closed oV the

Para-tethys The Turgai Strait vanished, and the continentalisation of the

Euro-pean archipelago formed new conWgurations (Ziegler, 1990) The mammal

immigration wave, coming from the east out of North America and Asia,

reached western Europe during nannoplankton zone NP 22 (32–33 Ma) and

resulted in the so-called ‘Grande Coupure’ (Stehlin, 1909; Tobien, 1987)

9

Table 2.1 Stratigraphic correlation chart of the Central and Eastern

Paratethys regional stage systems (Berggren et al., 1995; Popov et al., 1993; Ro¨gl, 1996, 1998a,b; Steininger et al., 1996).

10

[Figure 2.1]

Palaeogeographic sketch of the western end of the Tethys Ocean in the late

Eocene The western Tethys communicated with the Polar Sea via the Turgai

Strait and prevented mammal migrations between Asia and Europe (Ro¨gl,

1998a,b).

Sixteen new mammal families are mentioned by Savage (1990) New

important immigrants in mammal zone MP 21 include: Anthracotherium,

Eusmilus, Rhinocerothidae, Lagomorpha, Eomyidae, and Cricetidae This

faunal migration was made possible by the closure of the Turgai Strait and

the continuing northwestward movement of North America, closing in for a

Wrst Bering landbridge The Paratethys extended as an orogenic foredeep

and intercontinental sea from the Western Alps to Central Asia (early

Kiscel-lian/Pshekian) Marine connections existed in the far west with the

Mediter-ranean, and to the northwest along the Danish–Polish Strait with the North

Sea Basin (only in LatdorWan, NP 21)

First Paratethys isolation

Cold water inXux, dysaerobic bottom conditions, aberrant microXoras and

microfaunas, and northern molluscs characterised the early to middle

Oligocene (Baldi, 1979, 1984; Krhovsky et al., 1991; Popov et al., 1993; Rusu et

11

[Figure 2.2]

The isolated Paratethys Basin in the early Oligocene Only narrow seaways remained open in the west Salinity decreased and endemic conditions developed in the entire basin The closure of the Turgai Strait established the conditions for the Eurasian faunal exchange known as the ‘Grande Coupure’ (Ro¨gl, 1998a,b).

al., 1996) The isolation of the Paratethys culminated (Fig 2.2) during

nannoplankton zone NP 23 (Solenovian stage) with reduced salinity

condi-tions and endemic bivalves (‘Cardium lipoldi’ – Janschinella fauna) In

contrast, the Mediterranean Tethys remained open between the

Indo-Paci-Wc and Atlantic Ocean, and prevented a continental faunal exchange tween Eurasia and Africa Most of the Oligocene was characterised by acontinuous faunal exchange within Eurasia, rather than a distinct migration

be-wave, e.g the appearance of Melissiodon and Paracricetodon in MP 24, and

of Zapodidae in MP 26

Tropical marine excursion

A worldwide excursion of the tropical belt in the marine realm occurred in thelate Oligocene–early Miocene Horizons of tropical, larger foraminifera andmolluscs are reported from the Mediterranean, the Paratethys, and theMiddle East (Adams, 1973, 1976, 1983; Baldi & Senes, 1975; McGowran,

12

[Figure 2.3]

In the late Oligocene–early Miocene a transgression spread from the Middle East

to the Mediterranean and Paratethys Larger foraminifera, molluscs, and corals

fringed the eastern shelf along the Lut Block from Makran to Qum Basin, Lake

Urmia, and Transcaucasia In the Western Paratethys the Rhinegraben

connection and the seaway along the Alpine Foredeep closed In the Western

Mediterranean the Balearic Basin started to spread, with an eastward move of

the Apennines (Ro¨gl, 1998a,b).

1979a,b; Popov et al., 1993; McCall et al., 1994) The Mediterranean–Indian

Ocean gateway was open between the Anatolian and Arabian Plates (Fig 2.3)

A tropical-subtropical southeastern connection for the Paratethys newly

opened across the Iranian plate (Transcaucasia–Qom Basin–Makran) In the

south, the rotation of Arabia opened the graben of the Red Sea in the late

Oligocene (Jones & Racey, 1994) The Central Paratethys was connected to

the Mediterranean along fault structures in the Alps and between the Alps

and Dinarides The marine straits in the western Alpine foredeep and the

Rhinegrabenwere closed intermittently The Alpine foredeep re-opened for a

seaway to the Rhoˆne Basin in the early Burdigalian (Eggenburgian)

In the late Oligocene (mammal zone MP 28–30) new forms appeared in

Europe successively: Heterocricetodon, Rhizospalax, and the second

in-vasion of Lagomorpha In the early Miocene (Aquitanian), important new

forms include the suoids Palaeochoerus and Hyotherium in the European

mammal zone MN 1 and Bunolistriodon in Dera Bughti (Pakistan) In the

lower Burdigalian (MN 3) the North American immigrant Anchitherium

reached Europe (Thenius, 1972; Steininger et al., 1985b; Made, 1990) A

13

[Figure 2.4]

The Gomphotherium Landbridge The Eastern Mediterranean seaway closed,

and a landbridge opened between the Anatolian and Arabian/African Plates, enabling a remarkable faunal exchange The Eastern Paratethys became entirely isolated, with reduced salinity and endemic faunas The Mediterranean communicated with the Atlantic Ocean and fed the Western and Central Paratethys The renewed seaway in the Alpine Foredeep was again connected

Eurasian–African faunal exchange was still impossible, and no relationexisted to the Wrst East African mammal fauna of Meswa Bridge (23.5Ma)

The Gomphotherium Landbridge

During the middle Burdigalian (upper Eggenburgian–Ottnangian), strongmovements of the Savic tectonic phase changed the palaeogeographic pat-terns in the circum-Mediterranean (Fig 2.4) The rotation of AfricaWnallyclosed the gap to Eurasia The Arabian peninsula collided with the Anatolian

Plate The ‘Gomphotherium Landbridge’ was established Continental faunal

exchanges in both directions started in MN 4, around 19 Ma Successivearrivals of gomphotheres, deinotheres, and primates are observed in Eurasia,

while rhinos, tayassuids, or Amphicyon arrived in Africa The Wrst

Gom-photherium are recorded in the Siwaliks (Kamlial Formation) at 18 Ma,

14

[Figure 2.5]

Indo-Pacific recurrence For a short time the Mediterranean–Indo-Pacific seaway

reopened around the early–middle Miocene boundary From Eastern Anatolia a

new transgression flooded the Paratethys The intramountain basins and the

Carpathian foredeep in the Central Paratethys were covered by

tropical–subtropical waters A connection is proposed along the Rhodopes and

Pontides, south of the Black Sea Plate The Eastern Paratethys stayed in reduced

and in Poland in the coal mine Belchatow C also at this time (Thenius, 1979;

Steininger et al., 1985b; Goldsmith et al., 1988; Barry & Flynn, 1989; Pickford,

1989; Kowalski & Kubiak, 1993; Fortelius et al., 1996; Bernor et al., 1996).

The Paratethys Sea was divided in two separate realms The Eastern

Paratethys formed an enclosed basin with endemisms under reduced

salinity (Kotsakhurian stage) The Western and Central Paratethys remained

connected to the Mediterranean and via the Rhinegraben, to the North Sea

Cooler conditions were observed in the marine faunas By the end of

Ot-tnangian the Alpine foredeep became dry land The Mediterranean itself

was open to the Atlantic Ocean

Re-opening of the Indo-Pacific gateway

The exact process by which the seaway between the Indian Ocean and the

Mediterranean reopened is still being discussed and remains controversial,

15

[Figure 2.6]

The Paratethys closure The end of open marine connections transformed the Paratethys environment into an endemic bioprovince under reduced salinity conditions Since early Oligocene times, the most uniform biofacies was observed from the Vienna to the Caspian Basin in the Sarmatian Intermittent narrow marine straits opened from the Mediterranean to the Eastern Paratethys along the East Anatolian fault zone The Persian Gulf was characterised by evaporitic and continental sedimentation of the Fars and Gachsaran Formations

based on the interpretation of the distribution of larger foraminifera (Adams

et al., 1983; Adams, 1998) At least for the lower Langhian, such a short

connection along the Bitlis Zone between the Anatolian and Arabian Plates

is required (Fig 2.5) Another connection must have existed in easternAnatolia linking the Paratethys with the Indian Ocean for an early Badeniantransgression During the time of open seaways the Eurasian/African mam-mal migrations were interrupted, which seems to be demonstrated by

migration waves, e.g arrivals of primates: Pliopithecus in MN 5,

Griphopithecus and Plesiopliopithecus in MN 6, and Dryopithecus in MN 8

(Andrews et al., 1996; Begun, 1996).

Final closure between Eurasia and Africa

The Serravallian regression coincided with the re-establishment of the

‘Gomphotherium Landbridge’ Evaporitic sedimentation and

continen-talisation took place in the area around the Persian Gulf (Jones & Racey,

16

1994) TheWnal closure of the circum-equatorial oceanic current system

caused worldwide cooling and an increased accumulation of the East

Ant-arctic ice sheet around 15 Ma (Kennett, 1995) Short-lived connections

between the Indo-PaciWc, the Mediterranean and the Eastern Paratethys

(Fig 2.6) existed in eastern Anatolia in the upper middle Miocene in the

Konkian/late Badenian and Sarmatian, but also in the Pliocene (Nevesskaya

et al., 1984; Chepalyga, 1995; Iljina, 1995) These channels may have brieXy

acted as barriers for mammal exchanges The regression of the sea from the

Greek mainland and the Aegean landmass at the end of the Burdigalian

prevented marine connections with the Paratethys in the area of the

Dar-danelles The earliest rodent faunas are reported from the Aegean in

Or-leanian times (Sen, 1982) Beginning with the Tortonian transgression, an

Aegean seaway opened to the Black Sea Basin (Ro¨gl & Steininger, 1983;

Marunteanu & Papaianopol, 1995)

Since the Sarmatian the open ocean connections of the Paratethys were

interrupted The gigantic inland sea of the Paratethys turned into

continu-ously shrinking basins A reduced salinity and the alkaline chemistry of the

aquatic realm led to strong endemisms and caused the stenohaline

organ-isms in the Sarmatian sea to disappear (Pisera, 1996) The environments of

the Eastern Paratethys Sea had higher salinities from the Sarmatian up to

the Maeotian (late Miocene) and extended to the west to the Dacian Basin,

whereas the enclosed Pannonian Basin turned to nearly freshwater

condi-tions (Pannonian stage) The connecting facies between the Pannonian and

Euxinian basins is expressed by the local Malvensian stage in the Eastern

Carpathians and Dacian realm (Motas & Marinescu, 1969; Marinescu, 1985;

Papaianopol et al., 1995) During MN 6 an increasing faunal exchange

oc-curred between Eurasia and Africa (Bernor et al., 1996; Fortelius et al., 1996).

The appearance of ‘Hipparion’ is one of the last Miocene immigration

events from North America A strong sea level drop around 11 Ma (Haq et al.,

1988) once again opened the Bering landbridge In a rapid wave the

three-toed horse spread from China to the Eastern Paratethys and western Europe

(Bernor et al., 1989) During the Turolian (late Miocene, MN 11–13) in

southeastern Europe and southwestern Asia, the ‘Pikermian’ faunal

prov-ince extended with a radiation in hipparionine horses and ruminants

Addi-tionally, Asian ‘steppe elements’ such as Cricetus and Pliospalax appeared

in Europe (Steininger et al., 1985b; Bernor, 1984; Bernor et al., 1996; Gentry &

Heizmann, 1996) Finally, new but limited faunal routes with Africa were

opened by the Messinian salinity crisis in the late Turolian (MN 13)

Import-ant representatives include Protatera, Hippopotamus, and Macaca (Agustı´,

1996) Modern conditions were established in the circum-Mediterranean by

the Pliocene marine transgression coming from the Atlantic Ocean

17

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22

3 Pliocene tephra correlations between East African hominid localities, the Gulf of Aden, and the Arabian Sea

Peter B deMenocal and Francis H Brown

Introduction

The Pliocene–Pleistocene chronology of hominid and other vertebrate ution in East Africa is largely constrained by isotopic dating and regionalintercorrelation of volcanic ash layers Some eruptions were of suYcientmagnitude or duration that their widespread tephra dispersal deWnes aseries of dated marker horizons throughout the fossil-bearing sedimentary

evol-deposits of Tanzania, Uganda, Kenya, and Ethiopia (Brown, 1982; Feibel et

al., 1989; Haileab & Brown, 1992, 1994; Pickford et al., 1991; WoldeGabriel et al., 1994) Although many of the larger eruptive events have been dated

directly the ages of many tephra are only constrained by indirect graphic interpolation between dated levels The geochemical compositions

strati-of volcanic glasses from each eruption are unique, providing a deWnitivemeans to establish broad tephrostratigraphic correlations linking the re-gional climatic, tectonic, and biologic histories of this distinctive archive ofEarth history

This same tephrostratigraphic approach has been used to extend the EastAfrican tephra correlations into the continuous and well-dated marine

sediment record of regional and global paleoclimate variability (Brown et al., 1992; Sarna-Wojcicki et al., 1985) These authors identiWed several mega-scopic volcanic ash layers within Deep-Sea Drilling Project (DSDP) sites fromthe Gulf of Aden, nearly 1000 km northeast of hominid localities in Ethiopiaand Kenya Major element chemistries of volcanic glass shards extractedfrom these marine sediments were used to establish precise tephrostrati-graphic correlations into the fossil-bearing East African sedimentary se-quences Moreover, controversy concerning the ages of speciWc eruptiveevents which then deWned key temporaljunctures in early hominidevolutioncould be tested using the independent marine sediment chronostratigraphic

framework (Brown et al., 1992; Sarna-Wojcicki et al., 1985) A more

compre-hensive eVort to establish more Pliocene–Pleistocene tephra correlationsbetween terrestrial and marine sequences was hampered by incomplete andoften disturbed core recovery at these early DSDP sites in the Gulf of Aden.Subsequent advances in deep-sea drilling technology and the return ofscientiWc ocean drilling to the Arabian Sea present a new opportunity toestablish tephrostratigraphic links between East African terrestrial and ad-jacent marine sedimentary sequences Leg 117 of the Ocean Drilling Pro-gram drilled twelve sites oV the Omani margin and Arabian Sea (Fig 3.1;