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Perseverance of Pikas in the Miocene

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Perseverance of Pikas in the Miocene Perseverance of pikas in the Miocene Kees Hordijk GEOLOGICA ULTRAIECTINA Mededelingen van de Faculteit Geowetenschappen departement Aardwetenschappen Universiteit Utrecht No. 333 Members of the dissertation committee: Prof.dr. Poppe L. de Boer Sedimentology, Department of Earth Sciences Utrecht University, Utrecht, the Netherlands Prof.dr. Henk Brinkhuis Biomarine Sciences, Department of Biology Utrecht University, Utrecht, the Netherlands Prof.dr. Andy F. Lotter Paleoecology, Department of Biology Utrecht University, Utrecht, the Netherlands Prof.dr. Jack J. Middelburg Biogeochemistry, Department of Earth Sciences Utrecht University, Utrecht, the Netherlands The research presented in this thesis was funded by Faculty of Geosciences, Utrecht University Research was carried out at the Stratigraphy & Paleontology group, department of Earth Sciences, Utrecht University, Budapestlaan 4, 3584 CD, Utrecht, the Netherlands. (http://www.uu.nl/geo/ stratpal) ISBN 978-90-5744-194-3 Coverdesign: Ivo Duijnstee Graphic design: Geomedia, Faculteit Geowetenschappen, Universiteit Utrecht Printed by: Wöhrmann Print Service, Zutphen Perseverance of pikas in the Miocene Interplay of climate and competition in the evolution of Spanish Ochotonidae (Lagomorpha, Mammalia) Overleving van fluithazen in het Mioceen De rol van klimaat en competitie in de evolutie van Spaanse Ochotonidae (Lagomorpha, Mammalia) (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. J.C. Stoof, ingevolge het besluit van het college voor promoties in het openbaar te verdedigen op vrijdag 26 november 2010 des middags te 2.30 uur door Kees Hordijk geboren op 17 december 1975 te Warnsveld Promotor: Prof.dr. J.W.F. Reumer Prof.dr. G.J. van der Zwaan met dank aan Albert Contents 1. Introduction, summary and background 9 2. Systematics of resident species of Lagopsis and Prolagus (Ochotonidae, 17 Lagomorpha, Mammalia) from the late early and middle Miocene of north- eastern Central Spain Kees Hordijk & Albert J. van der Meulen – to be submitted 3. Systematics of immigrant species of Prolagus (Ochotonidae, Lagomorpha, 87 Mammalia) from the late early and middle Miocene of north-eastern Central Spain Kees Hordijk & Albert J. van der Meulen – to be submitted 4. Speciation, evolution and competition in pikas (Ochotonidae, Lagomorpha, 143 Mammalia) from the early and middle Miocene of Southwestern Europe Kees Hordijk, Ivo A.P. Duijnstee & Albert J. van der Meulen – to be submitted 5. Persistence and abundance of Miocene pikas (Ochotonidae, Lagomorpha, 175 Mammalia) in small-mammal communities from north-eastern Central Spain Kees Hordijk, Albert van der Meulen & Pablo Peláez-Campomanes – to be submitted 6. Impact of middle Miocene climate change on small-mammal evolution and 193 community composition in north-eastern Central Spain Kees Hordijk, Albert van der Meulen & Pablo Peláez-Campomanes – to be submitted References 205 Introductie & samenvatting (in Dutch) 225 Acknowledgements 229 Curriculum Vitae 231 Bibliography 232 7 8 chapter 1 Introduction, summary and background 1.1 Introduction Pikas represent a family of small-sized herbivores that is closely related to the family of rabbits and hares. Although they are currently extinct in Europe, pikas are a common component in small- mammal fossil assemblages of Miocene age in which they are often abundantly present. In spite of their regular occurrence, they are a relatively sparsely studied mammal group, in particular in comparison to rodents. Teeth represent the most important fossils of small mammals, because of their excellent preservation potential in the geologic record and their high diagnostic potential in taxonomy, which is often distinctive at the species level. In addition to its phylogenetic value, tooth shape is strongly related to diet and may provide information on past environmental conditions. The sedimentary succession near the village of Villafeliche in the Calatayud-Montalbán Basin (Figure 1.1) is well known for its exceptionally rich fossil mammal record spanning the Early and Middle Miocene interval and has been magnetostratigraphically dated (Krijgsman et al., 1994, 1996; Daams et al., 1999a). Extensive research has been focused on Neogene mammal paleontology in the Daroca-Villafeliche area for the last five decades, which has resulted in a database consisting of several tens of thousands of small-mammal teeth collected from over 100 mammal localities spanning the interval between ~17 and 10 Ma. The time interval covered by the mammal succession is characterized by major shifts and distinctive episodes in global climate, such as the Miocene Climate Optimum, which represents the warmest phase of the Neogene, and the Middle Miocene Climate Transition, which encompasses one of the major cooling phases in the Cenozoic (Flower & Kennett, 1994; Zachos et al., 2001). The combination of a rich and dense fossil record with good time control provide ideal conditions for studying the biostratigraphy, evolutionary trends and changes in the community structure of small mammals under the influence of major changes in climate. In recent years, studies focusing on the rodents have yielded significant new insights in small-mammal community dynamics (van der Meulen et al., 2005; van Dam et al., 2006). The role of the pikas in this context has so far remained unexplored. The taxonomy of fossil pikas is not very straightforward, which is mostly due to mosaic evolution, convergent and parallel trends and generally conservative evolutionary change, while having considerable intra-specific and ontogenetic variation. The comprehensive work of López- Martínez (1989) comprises a firm taxonomic framework for Spanish Miocene ochotonids and provides a wealth of information on the evolution of the group, which serves as the main starting point in the analysis of the several thousand ochotonid tooth elements from the Daroca- Villafeliche succession. Introduction, summary and background 9 The focus of this thesis is fourfold, comprising: 1) A detailed analysis and description of the extensive new ochotonid material from the Daroca- Villafeliche succession and re-evaluation of the existing taxonomy and biostratigraphy (Chapters 2 & 3); 2) Analysis of the main evolutionary trends in the lineages with special attention to speciation and competition (Chapter 4); 3) Analysis of the role of pikas in the small-mammal primary consumer communities (e.g. rodents and pikas) in the succession and analysis of the community structure in terms of community membership (“residents” and “transients”) and persistence-abundance patterns (Chapter 5); 4) Establishing the impact and effect of long-term climate changes on the small-mammal community composition and evolution of individual lineages by comparison to marine stable isotope records that serve as a proxy for changes in global climate (Chapter 6). Figure 1.1: Location of the Calatayud−Montalbán Basin and synthetic geology of the surrounding region. The square box denotes the position of the studied succession near the village of Villafeliche. (Modified after García- Paredes et al., 2010). 10 Chapter 1 1.2 Summary The high resolution mammal record in the Calatayud-Montalbán Basin provides an ideal setting for verifying and specifying the existing taxonomy and biostratigraphy of middle Miocene ochotonids. Chapter 2 deals with the systematics of the two ochotonid lineages that were already present in the region during a significant part of the early Miocene: the Lagopsis penai – L. verus and the Prolagus lopezmartinezae n. sp. – P. tobieni lineage. The definition of the chronospecies has been refined both in terms of morphological characteristics and in the extent of their temporal ranges. The anagenetic transition in theLagopsis lineage has been shifted to a level ~400 kyr older. The presence of P. lopezmartinezae marks the onset of the endemic evolution of this lineage and represents a different stage of evolution relative to P. tobieni. Chapter 3 is focussed on the systematics of the two immigrant lineages in the area: the Prolagus vargasensis n. sp. – P. major and the well known Prolagus oeningensis lineages. Similar to Chapter 2, the morphological definition and temporal ranges of the chronospecies have been refined. It is demonstrated that P. vargasensis arrived ~2 Myr earlier than P. oeningensis into the area and that P. major evolved from P. vargasensis in the early late Aragonian, thereby rejecting earlier hypotheses on the phylogenetic origin of P. major. In Chapter 4 the intraspecific variation and interspecific disparity between the four lineages described in Chapters 2 & 3 is demonstrated by multivariate analyses of size and shape characteristics of the P3. On the basis of our results we hypothesize that competition in combination with climate likely played a role in the evolutionary and co-existence patterns of the ochotonid lineages present in the Daroca-Villafeliche area. The first appearance of the endemic P. vargasensis n. sp. is inferred to be the result of parapatric speciation from a P. cf. oeningenensis stock from a more northern region. We deduce from our modelled disparity pattern between the morphologically similar P. vargasensis-P. major and P. oeningensis lineages that apart from climate, biotic mechanisms such as competitive exclusion of neighbouring allopatric species and limiting similarity between co-occurring species are likely to have significantly affected the
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