UNIVERSITÀ DEGLI STUDI DI NAPOLI FEDERICO II THE PLIO -HOLOCENE LARGE MAMMALS OF THE WESTERN EURASIA : MACROECOLOGICAL AND EVOLUTIONARY ANALYSES OF THE FAUNAS Dr. Francesco Carotenuto Dottorato in Scienze della Terra, Geologia del Sedimentario XXII Ciclo Tutor Prof. Carmela Barbera Cotutori Dr. Pasquale Raia, Prof. Anastassios Kotsakis 2009 To Daniela and my family “The most erroneous stories are those we think we know best - and therefore never scrutinize or question.” Stephen J. Gould Index Preface ........................................................................................................................................... IV Chapter 1 - The Eurasian Paleocommunities ................................................................................. 1 1.1 Introduction ............................................................................................................................ 1 1.1.1 The italian Paleocommunities (PCOMs) ............................................................................ 2 1.1.2 The properties of the Italian Paleocommunities: Body size and occupancy ..................... 5 1.1.3 Species turnover, diversity trends, and prey-predator relationship ................................... 6 1.2 Materials and Methods ........................................................................................................ 10 1.2.1 The data employed ........................................................................................................... 10 1.2.2 Time ordering techniques ................................................................................................. 13 1.2.2.1 Spectral ordering ........................................................................................................... 13 1.2.2.2 The Maximum Likelihood Appearance Event Ordination (MLA EO) ............................. 15 1.2.3 The Bootstrapped Cluster Analysis .................................................................................. 17 1.2.3.1 The BCA applied to the Plio-Holocene Western Eurasia LFAs .................................... 19 1.3 Results ................................................................................................................................. 20 1.3.1 The time-ordering of the Local Faunal Assemblages ...................................................... 20 1.3.2 The EA PCOM .................................................................................................................. 22 1.3.3 The geographical distribution and composition of the EA PCOMs ................................. 24 1.4. Discussion .......................................................................................................................... 31 Chapter 2 -Macroecology of the fossil mammal assemblages: occupancy, range size and phylogeny ...................................................................................................................................... 32 2.1 Introduction .......................................................................................................................... 32 2.1.1 The body size and its geographical distribution ............................................................... 33 2.1.2 Occupancy ........................................................................................................................ 34 2.1.3 The abundance-occupancy relationship .......................................................................... 35 2.1.3.1 Occupancy in paleontological studies ........................................................................... 37 2.1.4 The range size .................................................................................................................. 39 2.1.4.1 The distribution of range sizes ...................................................................................... 40 2.1.4.2 The hereditability of the range size ............................................................................... 41 2.1.5 The inclusion of phylogenetic information in comparative studies .................................. 43 2.1.5.1 The structures of a phylogenetic tree and its properties .............................................. 46 2.1.5.2 The Brow nian motion model ......................................................................................... 46 I 2.1.5.3 The Ornstein-Uhlenbeck process (OU) and the accelerating and decelerating Brow nian evolution (ACDC) ....................................................................................................... 47 2.1.5.4 Phylogenetic influence on type I and II error estimations ............................................. 48 2.1.6 Phylogenetic comparative methods ................................................................................. 50 2.1.6.1 Independent contrasts ................................................................................................... 50 2.1.6.2 Detecting the phylogenetic signal: the K statistics ........................................................ 52 2.1.6.3 Pagel’s λ ........................................................................................................................ 54 2.2 Materials and methods ........................................................................................................ 55 2.2.1 Taxa occurrences and age estimation ............................................................................. 55 2.2.2 Body size and duration ..................................................................................................... 56 2.2.3 Species level and genus level phylogenies ..................................................................... 57 2.2.4 Geographical distances betw een taxa ............................................................................. 58 2.2.5 Occupancy computation and trajectory ............................................................................ 59 2.2.6 Geographic range computation and its trajectory ............................................................ 60 2.2.7 The hereditability of the traits ........................................................................................... 61 2.3 Results ................................................................................................................................. 62 2.3.1 Occupancy and Area trajectories ..................................................................................... 62 2.3.2 Tests for phylogenetic signal and PICs correlations ........................................................ 63 2.3.3 Range position and phylogeny ......................................................................................... 64 2.4 Discussion ........................................................................................................................... 67 Chapter 3 -Macroecology of the fossil mammal assemblages: the species accumulation over space and time .............................................................................................................................. 70 3.1 Introduction .......................................................................................................................... 70 3.1.2 The species-area relationship .......................................................................................... 71 3.1.2.1 The hypotheses about the species-area relationship ................................................... 73 3.1.2.2 Combining the “area per se” and habitat heterogeneity influences in the species-area relationship: species-energy and the choros models ................................................................ 77 3.1.3 The species-time relationship .......................................................................................... 79 3.1.3.1 The factors that shape the pattern ................................................................................ 81 3.1.4 STAR model: combining the effects of both area and time in species accumulation ..... 82 3.2 Material and Methods .......................................................................................................... 85 3.2.1 Setting the data for the analyses ...................................................................................... 85 3.2.2 STR calculation and the application of the correction factors ......................................... 87 3.2.3 Comparing slopes at different time periods ..................................................................... 88 II 3.2.4 The computation of the SAR ............................................................................................ 89 3.2.5 The STAR model .............................................................................................................. 90 3.3 Results ................................................................................................................................. 91 3.3.1 The SAR and the STR ...................................................................................................... 91 3.3.2 Slopes per time ................................................................................................................. 91 3.3.3 STAR models .................................................................................................................... 92 3.4 Discussion ........................................................................................................................... 96 3.4.1 The STR and SA R models .............................................................................................