Università degli Studi di Napoli “Federico II” Dottorato di Ricerca in Scienze della Terra Geologia del Sedimentario “XX Ciclo” Plio-Pleistocene large carnivores from the Italian peninsula: functional morphology and macroecology CARLO MELORO 2007 Tutor: Prof . Carmela Barbera Co-tutor: Dott . Pasquale Raia Prof . Anastassios Kotsakis Coodinatore del dottorato: Prof . Giuseppe Nardi To my family & Adele Abstract The evolutionary history of large carnivorous mammals through the Ice Age have been investigated for the Italian peninsula. No endemisms are recorded in the mainland of Italy and large carnivore species composition reflected the similar fauna of European continent. Morpho-ecological adaptation of extinct Plio-Pleistocene species have been investigated throughout temporal PaleoCommunities (9 PCOMs – spanning 3.2 to 0.3 Ma) with statistical accuracy. Trophic apparatus of extant and extinct species was investigated with a geometric morphometric analysis of mandible shape while locomotory habits were assessed using long bone indices. The mandible shape analysis performed on extant Carnivora taxa confirms their morphological differences due principally to taxonomic affiliation (family). Although, when phylogenetic history is controlled with comparative methods, significant differences still to occur among taxa with different diets and between small and large forms (threshold posed at 7 kilograms). Interestingly, both mandibular regions (corpus and ascending ramus) are informative of Carnivora ecological adaptations and they result integrated at a macroevolutionary scale. This survey allows to consider geometric morphometric as a reliable technique to apply on fossil mandibles. Feeding habits have been predicted with a good degree of accuracy in several Plio- Pleistocene large carnivores on the basis of mandible shape data. The latter data –selecting only the corpus region- have been considered also to perform a morphospace comparison between large carnivore guilds of Italian PCOMs and extant guilds representative of five mainland ecosystems worldwide. Disparity values computed for mandibular corpus shape of Plio-Pleistocene guilds did not differ significantly from extant guilds. Morphological variability in mandible shape is negatively influenced by number of species in each guild as well as number of prey confirming that ecomorph specialization does not occur at the extreme region of morphospace. Long bone proportions of Plio-Pleistocene large carnivores are grouped in the variability of extant species. Although some phenomena of morphological convergences occur among extinct and extant taxa because of similar locomotor adaptation (e.g. cursorial) and same body size constraint. These morphoecological data were also used to predict the relative adaptability of Plio-Pleistocene species to certain habitats (grassland and tropical). The macroecological analysis of presence/absence data confirms the striking relationship between the abundances of both predators and their prey thought Ice Age. On the other hand no morpho- ecological coordinate changes occurs between predators and their prey. It is noteworthy that large carnivores are overrepresented in the Italian fossil record and became rarer from Galerian to the Aurelian (also because of a possible interaction with human activities). A GIS model was then computed to compare large mammal communities toward Plio-Pleistocene in Italy. Structural changes occurred in large herbivore communities from Villafranchian through the Aurelian because of climate changes. On the other hand, the spatial structure of large carnivore communities was more affected by their prey during the Villafranchian, while in the Galerian and Aurelian there was a greater influence of uncontrolled factors like climate and human activity as well. Acknolewdegments I am indebted to several people which contribute practically and mentally to the realization of this thesis. Firstly, I express my gratitude to my friend and supervisor Pasquale Raia who always stimulated and supported my researches and my ideas even after great quarrels. His contribution was unique and I am sure that we still fighting, together. Carmela Barbera and Anastassios Kotsakis kindly show me the right pathway with their experience. I am grateful to Paul O’Higgins, Sam Cobb and Sarah Elton (HYMS, University of York) for receiving me with open arms during my period in University of York. They share with me research experiences and important opinions on my projects. I am indebted to their care during my period in UK: their humanity and friendship let me spend very good time in spite of the bad weather. It was the best growing period for myself. Matthew Collins and all the staff of Dept. of Archaeology and HYMS (University of York) kindly support my period of visit. Several museum curators give me full confidence during my research visits: D. Hills (Natural History Museum, London); A. Kitchener (Royal Museum of Scotland, Edinburgh), M. Reilly, J. Liston (Huntherian Museum and Art Gallery, University of Glasgow); B. Sanchez, J. Morales, J. Cabarga, J. B. Rodríguez ( Museo Nacional de Ciencias Naturales, Madrid); A. Arribas (Museo Geominero, Madrid); D. Goujet, P. Tassy and C. Signe (Muséum National d’Histoire Naturelle, Paris); A. Tagliacozzo (Museo Archeologico Pigorin, Roma); R. Manni (Museo di Paleontologia Università “La Sapienza”, Roma), M. Del Re (Museo di Paleontologia, Università di Napoli). Their experience, their opinion, and their care represent an important contribution to the realization of my project. Paolo Piras is the best enjoyable landmark of my research visits in Rome. I am grateful to his hospitality and his hunger for comparative methods and science in general. We learn a lot from each other. Jim Rohlf and Andrea Cardini have been always a good support, clarifying several aspects on multivariate statistics. Their technical support was fundamental to the correct interpretation of my geometric morphometric analyses. Anna Loy kindly introduce me in the geometric morphometrics world and she always sustained me. I cannot forgive my best friends in University of Naples who let me spend good time and support me. I am particularly grateful to Francesco Carotenuto for still teaching me how differences are important in a working team. Antonello Bartiromo was and still to be a good friend of PhD adventures. The Italian community of York is another important landmark of my recent research experiences. Eline Manta, Enrico Cappellini, Elisa Giuntini, Patrizia Gremigni, Beatrice De Marchi (but also Lorenzo, Marco, Ilaria and the new Neapolitan incomers). I’ll never forgot your friendship and especially the good dinners!! A great thank you also to Olga Panagioulopu, Flora Groëning and Nhlanha for the good time in York and also in S-Block. Last but not the least are my relatives: Angelo Meloro, Grazia Landi, Maria Meloro, Gianluca Meloro. I still to learn from all of you. Adele Colamarco I am indebted to your faith and for trusting me. This research was supported by Università degli Studi di Napoli “Federico II”. Research visits at Museo Nacional de Ciencias Naturales (Madrid) and Muséum National d’Histoire Naturelle (Paris) were supported by the SYNTHESYS European Community’s Programme “Structuring the European Research Area” under. The present PhD thesis is part of a larger project: «The evolution of feeding habits in extinct European carnivores» supported under SYNTHESYS FR TAF-1680 and ES-TAF 858. Index Preface………………………………………………………………………………………………1 Chapter 1: Introduction…………………………………………………………………………...3 1.1 The order Carnivora: a synopsis…………………………………………………………..3 1.2 Aim of the thesis…………………………………………………………..........................7 1.3 Why large carnivores? …………………………………………………………............... 7 1.4 The Plio-Pleistocene of Italy: geological framework and large mammal fauna…………..9 1.5 Plio-Pleistocene large carnivores: species accounts……………………………………..12 Chapter 2: What is a large carnivore? Evidence from mandible shape……………………….40 2.1 Introduction……………………………………………………………………………….40 2.2 Materials and Methods……………………………………………………………………43 2.2.1 Sample size …………………………………………………………………….….43 2.2.2 Geometric morphometrics …………………………………………………….…..43 2.2.3 Ecological categories ……………………………………………………………..45 2.2.4 Testing morphological discontinuity …………………………………….………..47 2.3 Results…………………………………………………………………………………….47 2.3.1 Shape differences ………………………………………………………………….47 2.3.2 Ecological differences …………………………………………………….………49 2.3.3 PGLS …………………………………………………………………….………..50 2.3.4 Clustering ……………………………………………………………….………………51 2.4 Discussion…………………………………………………………………………………52 Chapter 3: Integration in mandible of carnivorous mammals: a macroevolutionary test……60 3.1 Modules of mammalian mandible………………………………………………………...60 3.2 Materials and Methods……………………………………………………………………61 3.3 Results…………………………………………………………………………………….63 3.3.1 Correlation between the corpus and the ramus region ……………………..…….63 3.3.2 Shape of the corpus mandibulae …………………………………………………..66 3.3.3 Shape of the ascending ramus …………………………………………………….68 3.4 Discussion…………………………………………………………………………………69 Chapter 4: Reconstructing feeding habits of Plio-Pleistocene large carnivores: a geometric morphometrics approach………………………………………………………………………….73 4.1 Feeding habits and jaw morphology in Carnivora…………..……………………………73 4.2 Materials and Methods……………………………………………………………………75 4.3 Complete mandibles analyses……………………………………………………………..76 4.4 Nested analyses of corpus shape………………………………………………………….82 4.4.1 Ursidae