Geologica Ultraiectina

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Geologica Ultraiectina GEOLOGICA ULTRAIECTINA Mededelingen van de Faculteit Aardwetenschappen Universiteit Utrecht No. 156 THE SMALL MAMMALS FROM THE UPPER MIOCENE OF THE TERUEL-ALFAMBRA REGION (SPAIN): PALEOBIOLOGY AND PALEOCLIMATIC RECONSTRUCTIONS Cover illustration by Jaap Luteijn ISBN 90-5744-014-8 The small mammals from the upper Miocene of the Teruel-Alfambra region (Spain): paleobiology and paleoclimatic reconstructions De kleine zoogdieren van het boven Mioceen van het Teruel-Alfambra gebied (Spanje): paleobiologie en paleoklimaatsreconstructies (met een samenvatting in het Nederlands) PROEFSCHRIFT ter verkrijging van de graad van doctor aan de Universiteit Utrecht op gezag van de rector magnificus Prof. dr H. O. Voorma ingevolge het besluit van het college van dekanen in het openbaar te verdedigen op maandag 15 december 1997 des namiddags te 16.15 uur. door Jan Arie van Dam geboren op 20 maart 1959, te Krimpen aid IJssel PROMOTOR: PROF. DR J.E. MEULENKAMP CO-PROMOTOR: DR A.J. VAN DER MEULEN Research was carried out at the Research Institute for Paleoenvironments and Paleoclimate Utrecht (IPPU) of the Netherlands Research School of Sedimentary Geology (NSG). This is NSG publication number 970165. Field campaigns were partly carried out within the framework of the projects "Paleoecologia y paleoclimatologia del Neogeno continental de la fosa Calatayud-Teruel" (PB92-0013) and "Evolucion paleoambiental de sucesiones con alta resolucion bioestratignifica en el Neogeno continental: area de Daroca-Teruel (Aragon-Espana)" (PB95-0114), financed by the Direccion General de Investigacion y Tecnica, Spain. Address of the author: Faculty of Earth Sciences Utrecht University Budapestlaan 4 3584 CD Utrecht The Netherlands tel.: 31-302535178/302535122 fax: 31-302535030 E-mail: [email protected] Voor mijn ouders Voor Agnes CONTENTS Publications 8 Samenvatting (Summary in Dutch) 9 Chapter 1: Introduction and synopsis 13 Chapter 2: Mammal successions of the Teruel-Alfambra region: 17 biostratigraphy and biochronology Chapter 3: The Murinae from the upper Vallesian and lower Turolian 37 of the Teruel - Alfambra region (NE Spain) Chapter 4: Stephanodonty in fossil murids: a landmark-based 73 morphometric approach Chapter 5: Paleoclimatic reconstruction on the basis of micromammal 87 faunal compositions, with special reference to life-history aspects Chapter 6: Reconstruction of the late Miocene climate of Spain 111 using rodent paleocommunity successions: an application of end-member modelling Chapter 7: The small mammals from the upper Miocene of the 153 Teruel-Alfambra region: faunal dynamics and events Appendix A: Taxonomic remarks 171 Appendix B: Age estimates for mammal localities 175 Appendix C: A biometric analysis on fossil and extant murine molars 183 References 187 Acknowledgements 201 Curriculum vitae 204 Publications Chapter 2: will be published in an extended form as: van Dam, J. A., L. Alcala, A. Alonso Zarza, J. P. Calvo, M. Garces, and W. Krijgsman. High-resolution late Miocene mammal biochronology and paleoecology of the Teruel-Alfambra region (NE Spain). (to be submitted to the Journal a/Vertebrate Paleontology) Chapter 3: to be submitted to Palaeontographica. Chapter 4: has been published as: van Dam, J.A. 1996. Stephanodonty in fossil murids: a landmark-based morphometric approach. Pp. 449-461 in Marcus, L. F., M. Corti, A. Loy, G. J. P. Naylor and D. E. Slice, eds. Advances in Morphometries. NATO volume 284. Plenum, New York. Chapter 5: submitted to Paleobiology. Chapter 6: will be published as: van Dam, J. A. and G. J. Weltje. Reconstruction of the late Miocene climate of Spain using rodent paleocommunity successions: an application of end-member modelling. (submitted to Palaeogeography, Palaeoclimatology, Palaeoecology) Samenvatting (Summary in Dutch) Samenvatting (Summary in Dutch) Het onderzoek aan fossiele kleine zoogdieren is in de jaren 60 in een stroomversnelling geraakt, toen ontdekt werd dat grate hoeveelheden gebitselementen konden worden verkregen door grondmonsters te onderwerpen aan de techniek van het "nat zeven". Met het uitwerken van de taxonomie werd al snel duidelijk dat met name knaagdieren zeer geschikt zijn voor het bedrijven van continentale biostratigrafie, hetgeen te danken is aan de hoge diagnostische waarde van hun gebitselementen en aan hun snelle evolutie. Daarnaast worden vanaf het einde van de jaren 70 associaties van kleine zoogdieren steeds vaker gebruikt voor de reconstructies van paleomilieu en klimaat. Het onderzoek in Spaanse Neogene bekkens heeft hierbij een belangrijke ral gespeeld. De uitzonderlijke rijkdom aan zoogdierresten in de fossiele meerafzettingen van deze bekkens maken het opstellen van hoge resolutie stratigrafische opeenvolgingen mogelijk, hetgeen een absolute randvoorwaarde is om gedetailleerde fauna- en omgevingsveranderingen naar behoren te kunnen registreren. Het belangrijkste doel van dit proefschrift is de gedetailleerde reconstructie van trends en gebeurtenissen ("events") in fauna, omgeving en klimaat gedurende het laat Mioceen (Vallesien en Turolien) van Spanje. Ais onderzoeksgebied is gekozen het gebied tussen Teruel en Alfambra in het bekken van Teruel, noordoost Spanje. De keuze voor dit gebied is mede bepaald door het feit dat eerder onderzoek een solide taxonomisch en biostratigrafisch raamwerk had opgeleverd. Gedurende een reeks van veldwerkcampagnes in de jaren 90, in samenwerking met collega's van de Complutense Universiteit en het Nationaal Museum voor Natuurwetenschappen in Madrid, is het aantal vindplaatsen van fossiele kleine zoogdieren in het boven Mioceen praktisch verdubbeld. Dit heeft geresulteerd in een sequentie van 89 lokaliteiten, waarvan in totaal zo'n 20.000 gebitselementen zijn bestudeerd. De meeste van deze vindplaatsen bevinden zich in secties met meerdere fossielhoudende niveaus boven elkaar. Gedurende het onderzaek zijn vijf van deze profielen bemonsterd voor magnetostratigrafische analyse. De combinatie van de gevonden magnetische polariteitspatronen en de dateringen van zonegrenzen in andere Spaanse bekkens maakt een correlatie van de zoogdiersuccessie naar de geomagnetische polariteits tijdschaal (GPTS) mogelijk. De eerste stap na het verzamelen is de determinatie van de fossielen. Na een korte bespreking van de geologische setting van het onderzoeksgebied en de voornaamste lithologische aspecten van de fossielhoudende secties, worden in hoofdstuk 2 nieuwe range charts van de verschillende groepen kleine zoogdieren gepresenteerd. De gevonden knaagdieren, die het zwaartepunt van het onderzoek vormen, zijn zander veel problemen in reeds gedefinieerde soorten ingepast. De insekteneters en lagomorfen zijn tot op genusniveau gedetermineerd en de zeldzame vleermuizen tot op familieniveau. Het overzicht wordt gecomplementeerd met een range chart van de grote zoogdieren uit het onderzoeksgebied. Drie nieuwe lokale biostratigrafische subzones voor het boven Vallesien worden gedefinieerd op basis van muriden. Een groot deel van de lokaliteiten, 9 Samenvatting (Summary in Dutch) alsmede verschillende zone/etage grenzen worden gecorreleerd naar de numerieke tijdschaal. De resultaten voor de Masia de la Roma sectie zijn consistent met de ouderdom van 9.7 miljoenjaar geleden voor de Onder-Boven Vallesien (MN9-1O) grens, zoals die bepaald is in het Valles-Penedes bekken. De ouderdom van de Vallesien­ Turolien (MNlO-ll) grens is bepaald op 8.7 miljoenjaar geleden in de La Gloria sectie. Ouderdommen van twee andere subzone grenzen in het boven Vallesien zijn geschat op respectievelijk 9.3 en 9.0 miljoen jaar geleden. De magnetostratigrafische controle voor het midden en boven Turolien van het Teruel bekken is minder goed. Ouderdommen voor de vindplaatsen in dit traject zijn geschat door middel van biostratigrafische correlaties met secties in het Cabriel bekken. Hoofdstuk 3 beschrijft de stand van zaken voor wat betreft de taxonomie van de muriden (Muridae, muisachtigen), voor zover deze groep voorkomt in het boven Vallesien en onder Turolien van het Teruel-Alfambra gebied. De familie van de muriden de meest voorkomende groep kleine zoogdieren in het Mediterrane gebied gedurende het laat Mioceen en vroeg Plioceen. Het eerste regelmatige voorkomen van muriden in het Teruel bekken is gedateerd op ongeveer 9.6 miljoen jaar geleden. Verscheidene karakteristieken van de eerste bovenkaaksmolaar van deze vroegste muriden (Progonomys hispanicus) wijzen op een nauwe verwantschap met Progonomys cathalai, een iets grotere vorm die rond 9.3 miljoen jaar geleden in het gebied verschenen is. Met het identificeren van Parapodemus gaudryi in verschillende onder Turolien vindplaatsen is aangetoond dat de verspreiding van deze soort niet beperkt was tot zuidoost Europa. De oorsprong van de soort is niet helemaal duidelijk, maar het is aannemelijk dat Progonomys cathalai de directe voorouder was. Dit zou inhouden dat het genus Parapodemus polyphyletisch is (meerdere keren is onstaan). In hoofdstuk 4 wordt een nieuwe marrier besproken om vormvariatie in muridenkiezen te beschrijven. De methode gaat uit van van te voren gedefinieerde homologe punten ("landmarks") op het horizontaal geprojecteerde kauwvlak, waarvan de x en y coordinaten worden gemeten. De toepassing van verschillende op deze landmarks gebaseerde morphometrische technieken op een set van 250 eerste bovenkaaksmolaren van acht verschillende laat Miocene en Pliocene Spaanse associaties laat zien dat een clustering van populaties op grond van morphometrische parameters de taxonomische indeling redelijk goed, maar niet
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