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Community Structure Through Time: `Ubeidiya, a Lower Pleistocene Site As a Case Study

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Community Structure through Time: `Ubeidiya, a Lower Pleistocene Site as a Case Study Thesis submitted for the degree of “Doctor of Philosophy” by Miriam Belmaker Submitted to the Senate of the Hebrew University March 2006 This work was carried out under the supervision of Prof. Eitan Tchernov Prof. Ofer Bar Yosef Prof. Uzi Motro this thesis is dedicated to Eitan Tchernov, my mentor, without whom this thesis would have never happened Contents Acknowledgments xii Abstract xiv I Framework 1 1 Introduction 2 2 The Site of `Ubeidiya 9 2.1 Geology and stratigraphy of the `Ubeidiya Formation . 9 2.2 Dating of the `Ubeidiya Formation . 10 2.3 Excavation history . 15 2.4 Previous paleoecological research . 17 2.5 The presence of early hominins at `Ubeidiya . 19 II Methodology 22 3 Paleontological Methodology 23 3.1 Stratigraphy . 23 3.1.1 Assignment of specimens to stratigraphic units . 23 3.1.2 Choice of strata for analysis . 25 3.2 Taxonomy . 26 3.2.1 Identification of specimens . 26 3.2.2 Choice of taxa for analysis . 28 3.3 Quantification . 30 3.3.1 Quantification of individuals per species . 30 3.3.2 Quantification of body elements . 31 i 4 Identification of Pattern of Persistence or Change in the Large Mammalian Commu- nity throughout the `Ubeidiya Sequence 34 4.1 The statistical model . 34 4.2 Independent variables of the null hypotheses: The taphonomy of the `Ubeidiya large mammal assemblages . 37 4.2.1 Weathering . 37 4.2.2 Fluvial transport . 39 4.2.3 Agents of accumulation . 41 4.2.4 Post depositional carnivore ravaging . 47 4.2.5 Density mediated processes . 50 4.3 Independent variables of the alternative hypothesis: The environmental proxy . 51 4.4 The dependent variables: The biodiversity of the `Ubeidiya large mammal assem- blages . 53 4.4.1 Faunal turnover . 53 4.4.2 Relative frequency of taxa . 54 5 Paleoecological Reconstruction 59 III Results 62 6 The Large Mammalian Fauna of `Ubeidiya 63 6.1 Overview of the large mammal fauna of `Ubeidiya . 63 6.2 The biogeographic origins of the large mammals present in `Ubeidiya with empha- sis on African taxa . 63 7 Taphonomic Analyses of `Ubeidiya Strata 69 7.1 Weathering . 69 7.2 Fluvial transport . 70 7.3 Agents of accumulation . 73 7.3.1 Body element representation . 73 7.3.2 Density mediated processes . 74 7.3.3 Evidence for hominin involvement in the accumulation of the `Ubeidiya large mammalian assemblages . 77 7.3.4 Evidence for carnivore involvement in the accumulation of the `Ubeidiya large mammalian assemblages . 83 7.4 Evidence for post depositional carnivore ravaging . 84 7.4.1 Tooth marks . 84 7.4.2 Proportion of carnivores of total assemblage . 85 7.4.3 Proportions of ``bone cylinders'' shaft fragments . 86 7.4.4 The effect of carnivore ravaging on body part distribution . 87 7.5 Interim summary and discussion: Taphonomy and the accumulation of the large mammalian fauna of `Ubeidiya . 89 8 Identification of Patterns of Persistence or Change 98 8.1 Faunal turnover . 98 8.2 Relative frequency analysis of taxa . 100 8.2.1 Multi-species analysis . 100 8.2.2 Individual taxa analysis . 103 8.2.3 Diversity indices . 107 8.3 Interim summary and discussion : Can persistence or change in the large mam- malian community be discerned throughout the `Ubeidiya sequence? . 115 9 Paleoecological Reconstruction of `Ubeidiya 124 9.1 Multivariate analysis of the mesoherbivore community . 124 9.2 Interim summary and discussion: The paleoecological milieu of `Ubeidiya . 126 IV Discussion and Conclusions 133 10 Discussion 134 11 Conclusions 144 Bibliography 145 V Appendices 173 A Figures 174 B Tables 231 C Statistical Equations 270 D Names of Museum Collections Visited 276 E Grouping of Substrata into Strata 277 List of Figures 1 The Levant and the location of Plio-Pleistocene sites mentioned in the text . 175 2 The location of the `Ubeidiya Formation main outcrop in the central Jordan Valley . 176 3 Stratigraphy and lithology of the Fi cycle (Mallol, 2006) . 177 4 Schematic representation of the position of `Ubeidiya within the GPTS . 178 5 The plan of `Ubeidiya excavations 1960-1999 . 179 6 The stratigraphic sequence of `Ubeidiya strata . 180 7 The stratigraphic sequence of `Ubeidiya strata depicting `pooled strata’ . 181 8 Point by point rarefaction curves . 182 9 Ratio diagrams for small-medium and large mammals . 183 10 Limb bone portions . 184 11 Scattergram of correspondence analysis axes for weathering stages . 185 12 Dendrogram of correspondence analysis axes for weathering stages . 186 13 Grain shape distribution . 187 14 Scattergram of correspondence analysis axes for grain shape . 188 15 Dedrogram for correspondence analysis of grain shape . 189 16 Dedrogram of grain shape frequencies by size . 190 17 Percent MAU for Ruminantia E, epiphyseal data . 191 18 Correlation of percent MAU of Ruminantia E and BMD . 192 19 Percent MAU for Ruminantia F, epiphyseal data . 193 20 Correlation of percent MAU of Ruminantia F and BMD . 194 21 Percent MAU for Ruminantia G, epiphyseal data . 195 22 Correlation of percent MAU of Ruminantia G and BMD . 196 23 Percent MAU for Equidae, epiphyseal data . 197 24 Correlation of percent MAU of Equidae and BMD . 198 25 Percent MAU for Ruminantia E, complete limbs . 199 26 Correlation of percent MAU of Ruminantia E and FUI . 200 v 27 Percent MAU for Ruminantia F, complete limbs . 201 28 Correlation of percent MAU of Ruminantia F and FUI . 202 29 Percent MAU for Ruminantia G, complete limbs . 203 30 Correlation of percent MAU of Ruminantia G and FUI . 204 31 Percent MAU for Equidae, complete limbs . 205 32 Correlation of percent MAU of Equidae and GUI . 206 33 Cutmark on specimen UB 6227 . 207 34 PCA results for cutmark distributions . 208 35 Dendrogram of cut-mark distribution . 209 36 PCA for cut-mark frequency by body size . 210 37 Dendrogram for cut-mark frequencies by body size . 211 38 Carnivore modification by limb portion . 212 39 Percent “bone cylinders” . 213 40 Percent MAU of Ruminantia E correlated with percent MAU of hyaena ravaged bones . 214 41 Percent MAU of Ruminantia F correlated with percent MAU of hyaena ravaged bones . 215 42 Percent MAU of Ruminantia G correlated with percent MAU of hyaena ravaged bones . 216 43 Percent MAU of Equidae correlated with percent MAU of hyaena ravaged bones . 217 44 Dendrogram for body part distribution . 218 45 Scattergram for CA axes for presence-absence data . 219 46 Q and R clustering for presence-absence . 220 47 Scattergram for CA axes for NISP frequencies . 221 48 R and Q mode clustering for NISP frequencies . 222 49 Distribution of four common taxa in `Ubeidiya through time using NISP and MNI quantification . 223 50 Distribution of two groups of habitat specific taxa in `Ubeidiya using NISP quan- tification . 224 51 Species richness indices for `Ubeidiya strata . 225 52 Evenness diversity indices for `Ubeidiya strata . 226 53 Dominance diversity indices for `Ubeidiya strata . 227 54 Diversity indices for `Ubeidiya strata . 228 55 Scattergram for mesoherbivore presence-absence . 229 56 Scattergram for mesoherbivore frequencies . 230 List of Tables 1 Distribution of identified specimens from `Ubeidiya . 232 2 Large terrestrial mammal distribution among ‘Ubeidiya ‘pooled strata’ using NISP quantification . 233 3 Large terrestrial mammal distribution among ‘Ubeidiya ‘pooled strata’ using MNI quantification . 236 4 Percent of taxa of Africa biogeographic origin of total number of species across `Ubeidiya `pooled strata’ . ..
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  • Pleistocene Cave Hyenas in the Iberian Peninsula: New Insights from Los Aprendices Cave (Moncayo, Zaragoza)

    Pleistocene Cave Hyenas in the Iberian Peninsula: New Insights from Los Aprendices Cave (Moncayo, Zaragoza)

    Palaeontologia Electronica palaeo-electronica.org Pleistocene cave hyenas in the Iberian Peninsula: New insights from Los Aprendices cave (Moncayo, Zaragoza) Víctor Sauqué, Raquel Rabal-Garcés, Joan Madurell-Malaperia, Mario Gisbert, Samuel Zamora, Trinidad de Torres, José Eugenio Ortiz, and Gloria Cuenca-Bescós ABSTRACT A new Pleistocene paleontological site, Los Aprendices, located in the northwest- ern part of the Iberian Peninsula in the area of the Moncayo (Zaragoza) is presented. The layer with fossil remains has been dated by amino acid racemization to 143.8 ± 38.9 ka (earliest Late Pleistocene or latest Middle Pleistocene). Five mammal species have been identified in the assemblage: Crocuta spelaea (Goldfuss, 1823) Capra pyre- naica (Schinz, 1838), Lagomorpha indet, Arvicolidae indet and Galemys pyrenaicus (Geoffroy, 1811). The remains of C. spelaea represent a mostly complete skeleton in anatomical semi-connection. The hyena specimen represents the most complete skel- eton ever recovered in Iberia and one of the most complete remains in Europe. It has been compared anatomically and biometrically with both European cave hyenas and extant spotted hyenas. In addition, a taphonomic study has been carried out in order to understand the origin and preservation of these exceptional remains. The results sug- gest rapid burial with few scavenging modifications putatively produced by a medium sized carnivore. A review of the Pleistocene Iberian record of Crocuta spp. has been carried out, enabling us to establish one of the earliest records of C. spelaea in the recently discovered Los Aprendices cave, and also showing that the most extensive geographical distribution of this species occurred during the Late Pleistocene (MIS4- 2).