The Ecological Context of the Early Pleistocene Hominin Dispersal to Asia

Home , Ailuropoda microta, Canis etruscus, Cercopithecoides, Chasmaporthetes, Gorgopithecus, Pachycrocuta

THE ECOLOGICAL CONTEXT OF THE EARLY PLEISTOCENE HOMININ DISPERSAL TO ASIA

by Robin Louise Teague

A.B. in Anthropology, 2001, Harvard University

A dissertation submitted to

The Faculty of The Columbian College of Arts and Sciences of The George Washington University in partial fulfillment of the requirements for the degree of Doctor of Philosophy

August 31, 2009

Dissertation directed by

Richard Potts Curator of Physical Anthropology, National Museum of Natural History, Smithsonian Institution Alison S. Brooks Professor of Anthropology

The Columbian College of Arts and Sciences of The George Washington University certifies that Robin Louise Teague has passed the Final Examination for the degree of

Doctor of Philosophy as of June 16, 2009. This is the final and approved form of the

dissertation.

THE ECOLOGICAL CONTEXT OF THE EARLY PLEISTOCENE HOMININ DISPERSAL TO ASIA

Robin Louise Teague

Dissertation Research Committee:

Richard Potts, Curator of Physical Anthropology, National Museum of

Natural History, Smithsonian Institution, Dissertation Co-Director

Alison S. Brooks, Professor of Anthropology, Co-Director

Lars Werdelin, Senior Curator, Swedish Museum of Natural History,

Committee Member

iii Acknowledgments

I would like to acknowledge a number of people who have helped me and guided me through the process of writing my dissertation. First, I would like to thank my committee: Rick Potts, Alison Brooks, Lars Werdelin, Margaret Lewis and Brian

Richmond. My advisor, Rick Potts, led me into a stimulating area of research and supported me in pursuing a large and ambitious project. He has encouraged me all through the time I have worked on this dissertation. Alison Brooks has helped me with enthusiasm, pointing out differing perspectives and opportunities. Lars Werdelin has been a source of detailed and helpful information and has always been available to answer questions and provide guidance. I am also grateful to Brian Richmond and Margaret

Lewis for many helpful comments and for making time for my dissertation.

I would also like to thank my family for their love and support during my time in graduate school and especially during the process of writing. Their encouragement was essential to my success.

I would like to thank my fellow students at GWU for many years of friendship. I would like to thank that faculty of the Hominid Paleobiology Doctoral Program and the

Department of Anthropology for their support. At the Smithsonian, I thank Jenny Clark,

Briana Pobiner and Matt Tocheri for their assistance and for making me feel welcome.

For access to mammalian skeletal collection, I thank Linda Gordon. I am also grateful to the Smithsonian Libraries for letting me check out and renew many books many times. iv Thanks go also to the many people who facilitated my access to fossils at the

Kenya National Museum and the Institute of Vertebrate Paleontology and

Paleoanthropology in Beijing, as well as many other museums. In particular, Wang Wei traveled with me, assisting with translation and helping me make the contacts necessary to study many of the fossil specimens in Chinese museums. I would also like to thank

Gao Xing, Deng Tao, Qiu Zhanxiang, Qi Guoqin, Huang Weiwen, Hou Yamei, Zhu

Rixiang, Deng Chenglong, Li Qing Kui and Wei Guangbao for helping me study fossil specimens while I was in China.

For financial support, I would like to thank the IGERT program in Hominid

Paleobiology at GWU as well as the Smithsonian National Museum of Natural History where I had a predoctoral fellowship. My years of study at GWU and at the Smithsonian

Human Origins Program have been extremely rewarding. This dissertation research was funded by NSF Grant BSC 065092.

v Abstract of Dissertation

The Ecological Context of the Early Pleistocene Hominin Dispersal to Asia

The ecological context of the first known dispersal of Homo into East Asia is investigated here using information from large mammals, and particularly from carnivores. The aims were to determine whether hominins occurred in similar ecological contexts compared with sites in East Africa, and whether carnivore guilds in East Asia and East Africa were similar in composition in terms of ecologically comparable species.

To answer these questions, dental measurements were taken on large mammalian specimens from East Asian Plio-Pleistocene sites, including hominin and non-hominin sites, and from specimens found at Olduvai Gorge and Lake Turkana in East Africa.

Dental measurements were taken to estimate body mass and hypsodonty, as well as ecomorphological characteristics in carnivores. Each large mammal species was classified as an ecotype, which is a combination of body mass, diet and substrate (i.e., terrestrial, arboreal and aquatic) characteristics. The ecotype analysis shows that East

Asian and East African fossil sites were significantly different from each other in ecological structure, with the Asian sites having a greater concentration of browsers and mixed feeders, while East African sites had more grazers. The East Asian hominin sites included varied ecological structures, implying that hominins were not tied to a single type of environment on their initial dispersal. Carnivore ecomorphological indices related to body mass and feeding adaptations, such as the amount of the dentition devoted to slicing, grinding and bone-cracking. Carnivore guilds containing sets of species with similar feeding adaptations and body mass would have presented similar opportunities for

vi scavenging and degrees of competition for hominins. The Hyaenidae differed between

Africa and Asia in features related to fourth premolar size. Omnivorous ursids were

present in Asia but not in Africa. In East Asia, there were also decreases in the number of

species of Hyaenidae and Canidae from the Late Pliocene to the early Pleistocene.

Despite this, the remaining Asian hyaenid, Pachycrocuta , would have been a formidable competitor for scavenging hominins. Overall, hominins occurred in varied ecological settings, and competed with a carnivore guild that had species with different adaptations compared with Africa.

vii Table of Contents

Acknowledgments……………………………………………………………………....iiv

Abstract of Dissertation………………………………………………………………….vi

Table of Contents…………………………………………………………………... ….viiii

List of Figures…………………………………………………………………………....ix

List of Tables……………………………………………………………………………xii

Chapter 1: Introduction…………………………………………………………………..1

Chapter 2: Background to ecological similarity analysis……………………………….11

Chapter 3: Methods to determine ecological similarity………………………………....41

Chapter 4: Results of the ecological structure analysis………………………………....94

Chapter 5: Background to carnivore ecomorphology……………………………….....129

Chapter 6: Carnivore ecomorphology methods……………………………………...... 151

Chapter 7: Carnivore ecomorphology results……………………………………….....176

Chapter 8: Discussion……………………………………………………………….....236

Bibliography…………………………………………………………………………...270

Appendices………………………………………………………………………….....292

viii List of Figures

Figure 2.1 Map of East Asian fossils……………………………………………………39

Figure 3.1 Modern Eurasian localities used for comparison………..…………………...52

Figure 3.2 Modern African localities used for comparison……………………………...53

Figure 4.1 Scatterplot of the CA of the modern faunal assemblages and ecotypes...... 99

Figure 4.2 NMDS using Euclidean distance of modern divisions……………………..100

Figure 4.3 CA scatterplot of modern faunal sites excluding rainforests…………..…..102

Figure 4.4 NMDS of modern sites excluding rainforests……………………………...104

Figure 4.5 Scatterplot of a CA of the modern sites and ancient fossil assemblages…..109

Figure 4.6 Scatterplot of a CA of the modern and ancient assemblages, axes 2 and

3………………………………………………………………………………………..110

Figure 4.7 NMDS analysis of ancient and modern sites…………………………….....112

Figure 4.8 CA scatterplot of modern and ancient faunal assemblages, excluding modern rainforests………………………………………………………………………….…...115

Figure 4.9 CA scatterplot, axes 2 and 3: Modern sites (excluding rainforests) and ancient faunas………………………………………………………………………………...... 116

Figure 4.10 NMDS plot of Plio-Pleistocene assemblages with modern sites (excluding rainforests)…………………………………………………………………………...... 120

Figure 4.11 Correspondence analysis scatterplot of Plio-Pleistocene sites……………125

Figure 7.1 Scatterplot of the CA for Canidae category scores………………….……..180

Figure 7.2 Scatterplot of NMDS analysis of Hamming distances for Canidae………..181

ix Figure 7.3 PCA Scatterplot of Canidae index values from East Africa and East

Asia…………………………………………………………………………………….182

Figure 7.4 Loadings for component 1 of the PCA of Canidae fossils……………...... 183

Figure 7.5 Loadings for component 2 of the PCA for Canidae fossils……………...... 184

Figure 7.6 CA Scatterplot of Hyaenidae category scores…………………………...... 189

Figure 7.7 NMDS scatterplot of Hyaenidae Hamming distances…………………...... 190

Figure 7.8 PCA Scatterplot of Hyaenidae index values…………………………...... 191

Figure 7.9 Component 1 PCA loadings for Hyaenidae…………………………...... 192

Figure 7.10 Component 2 PCA loadings for Hyaenidae………………………...... 193

Figure 7.11 CA Scatterplot of Felidae category scores…………………………...... 201

Figure 7.12 NMDS scatterplot of Hamming distances for Felidae…………….……....202

Figure 7.13 PCA Scatterplot of Felidae index values……………………………...... 203

Figure 7.14 PCA loadings for component 1 for Felidae…………………………...... 204

Figure 7.15 PCA loadings for component 2 for Felidae………………………...... 205

Figure 7.16 CA scatterplot of Ursidae category scores…………………………...... 208

Figure 7.17 NMDS Scatterplot of Hamming distance values for Ursidae……...... 209

Figure 7.18 PCA Scatterplot of Ursidae index values…………………………………210

Figure 7.19 Loadings for component 1 of PCA scatterplot for Ursidae……………….211

Figure 7.20 Loadings for component 2 of PCA scatterplot for Ursidae……………….212

Figure 7.21 CA Scatterplot of Mustelidae category scores…………………………….214

Figure 7.22 NMDS Scatterplot of Hamming distances for Mustelidae……………...... 215

Figure 7.23 PCA Scatterplot for index values for Mustelidae…………………...... 216

Figure 7.24 PCA of Mustelidae loadings for component 1………………………...... 217

x Figure 7.25 PCA of Mustelidae loadings for component 2……………………………218

Figure 7.26 CA Scatterplot of category scores for Herpestidae, Prionodontidae and

Viverridae………………………………………………………………………...... 221

Figure 7.27 NMDS Scatterplot of Hamming distances for Viverridae, Herpestidae and

Prionodontidae species…………………………………………………………...... 222

Figure 7.28 PCA scatterplot of index values for Viverridae, Herpestidae and

Prionodontidae…………………………………………………………………………223

Figure 7.29 Component 1 loadings for PCA of Viverridae, Herpestidae and

Prionodontidae………………………………………………………………………....224

Figure 7.30 Component 2 loadings for PCA of Viverridae, Herpestidae and

Prionodontidae…………………………………………………………………………225

Figure 7.31 Scatterplot of CA of carnivores from East Asia and East Africa…………232

Figure 7.32 Scatterplot of CA of carnivores from East Asia and East Africa, second and third axes…………………………………………………………………………...... 233

xi List of Tables

Table 3.1 Ecotype classifications…………………………………………………….....43

Table 3.2 Modern comparative localities…………………………………………….....48

Table 3.3 Plio-Pleistocene East Asian sites and their dates…………………………….56

Table 3.4 Plio-Pleistocene African species at the Turkana Basin and Olduvai………...56

Table 3.5 Plio-Pleistocene East Asian species……………………………………….....62

Table 3.6 Modern specimens of Canidae………………………………………….……75

Table 3.7 Modern specimens of Ursidae………………………………………………..76

Table 3.8 Modern samples of Mustelidae, Viverridae, Prionodontidae and Herpestidae

…………………………………………………………………………………………..76

Table 3.9 Ecological characteristics of East Asian fossil site faunas………………...... 78

Table 3.10 Summary of ecological information and ecotype assignment for African fossil species……………………………………………………………………………...……83

Table 4.1 The MEDC (mean Euclidean distance to the centroid) for each modern division…………………………………………………………………………….…….97

Table 4.2 Modern site group centroids excluding rainforests……………………...... 101

Table 4.3 Mean and maximum distance to the centroid for modern and ancient assemblages…………………………………………………………………………….108

Table 4.4 Mean and maximum distance to the centroid for modern and ancient sites excluding rainforests…………………………………………………………………...117

Table 4.5 MANOVA results for the centroids of African and Asian fossil sites from the ancient and modern CA without rainforests……………………………………………118

xii Table 4.6 Mean and maximum distance to the centroid for Plio-Pleistocene sites……123

Table 4.7 MANOVA results for the centroids of African and Asian fossil sites…...... 124

Table 6.1 Modern Felidae samples……………………………………………….……153

Table 6.2 Modern Hyaenidae samples……………………………………………...... 153

Table 6.3 Ecomorphological measurement descriptions…………………………...... 155

Table 6.4 Ecological traits and ecomorphological indices…………………………….155

Table 6.5 Index values for fossil carnivore species……………………………………158

Table 6.6 Index values for fossil carnivore species, continued……………………...... 162

Table 6.7 Category cut-off values………………………………………………...…....167

Table 6.8 Category scores for all carnivores...... 171

Table 7.1 P-values for the MANOVA of carnivore family centroids in East Asia and East

Africa…………………………………………………………………………………..226

Table 7.2 Mean distance to the centroid (MEDC) for carnivore families…………...... 227

Table 7.3 MEDC for carnivore guilds at African and Asian sites…………………...... 227

Table 7.4 MANOVA of carnivore guilds at sites in Africa and Asia………….……....228

Table A6.1 Canidae specimens measured from East Asia…………………….………292

Table A6.2 Felidae specimens measured from East Asia……………………….……..293

Table A6.3 Hyaenidae specimens measured from East Asia…………………….……295

Table A6.4 Ursidae specimens measured from East Asia……………………….….....299

Table A6.5 Mustelidae specimens measured from East Asia…………………….……302

Table A6.6 Viverridae and Prionodontidae specimens measured from East Asia…….303

Table A6.7 Canidae specimens measured from East African sites……………………303

Table A6.8 Felidae specimens measured from East African sites……………………..304

xiii Table A6.9 Hyaenidae specimens measured from East African sites…………..…...... 305

Table A6.10 Mustelidae specimens measured from East African sites…………...... 306

Table A6.11 Herpestidae specimens measured from East African sites…………...... 306

Table A6.12 Viverridae specimens measured from East African sites……………..…307

xiv Chapter 1: Introduction to the dissertation

Anthropologists have long been interested in the environments that hominins inhabited and in the way hominins interacted with the other members of the mammalian community, in particular with the large-bodied members of the order Carnivora

(carnivores). Large carnivore kills might have been scavenged by hominins for meat.

Carnivores interacted with hominins as predators, competitors, suppliers of scavengeable carcasses, and later as sources of skins and ornamentation. The circumstances surrounding the earliest currently known dispersal of hominins from Africa around 1.8

Ma are particularly interesting with regard to hominin ecology and interactions with other mammals because the localities in which the dispersing hominins have been found (the

Caucasus, continental East Asia and Java) are geographically distant from Africa and have taxonomically different faunas. The East Asian localities are in temperate and subtropical zones, which would be expected to have a different ecology compared with tropical and subtropical African sites. However, the comparative ecological contexts of these locations are not well known.

The ecology of the localities in which dispersing hominins have been found is the focus of this dissertation. The ecological context of hominin dispersal is addressed by comparing the ecological properties of mammalian species from initial dispersal sites in

East Asia and contemporaneous sites in East Africa to determine whether hominins colonized places that were ecologically similar or whether hominins were capable of adapting to different environmental settings on their initial dispersal. A subset of the study concerns carnivore feeding adaptations to determine whether the carnivore guilds were similar in East Asia and East Africa, and if not, in what ways they differed. The

implications for resource and niche availability for hominins in new environments are considered, as is the range of ecological contexts in which hominins were found in East

Asia and East Africa.

Current evidence points to an initial dispersal out of Africa around 1.8 Ma.

However, it is possible that hominins dispersed earlier and that new sites or new dates

will be found that push the date back further. Dmanisi is dated to 1.77-1.75, immediately

following the Olduvai subchron (Gabunia et al. 2000a, Vekua et al. 2002, and Rightmire

et al. 2006). The Yuanmou hominin layer is dated to ~1.7 Ma (Zhu et al. 2008), while the

oldest artifacts known from the Nihewan Basin are dated to ~1.66 Ma (Zhu et al. 2004).

Hominins in Java are dated to 1.8 to 1.6 Ma at Mojokerto (Swisher et al. 1994, 1997,

Larick et al. 2001, Huffman et al. 2006) and to ~1.6 Ma at Sangiran (Swisher et al. 1994,

Antón and Swisher 2004). These hominins were most likely members of the genus

Homo . Archaeological evidence from Africa and Asia indicates that Homo was most likely an omnivore, obtaining meat from scavenging or predation. Dmanisi, sites within the Nihewan and Yuanmou all have Oldowan stone tools.

The geographic spread of initial dispersal sites shows that hominins were able to survive in regions very distant from East Africa, with mammalian faunas consisting of many different genera and species. Dmanisi and the Nihewan Basin are located relatively far north, at about 40°N latitude, raising the possibility of that hominins had to adapt to environmental conditions quite different from the tropical and subtropical latitudes occupied by probable source populations.

Ecological Similarity of the Large Mammal Fauna

Environmental similarity has been thought to have facilitated hominin dispersal.

Dennell (2004) argued that a belt of savanna habitats across Asia and Africa provided both environmental similarity and a wide corridor of ecologically suitable habitats for hominins, resulting in dispersal into Asia. Dennell (2003, 2004) hypothesized that hominins were constrained to environments that were sufficiently similar in temperature and seasonality patterns to Africa, leading hominins to be only found south of 40°N in the early Pleistocene 1. At ‘Ubeidiya, the presence of Pelorovis was thought to imply the extension of savannas, while Kolpochoerus may have indicated the occurrence of gallery forests (Martínez-Navarro 2004). Dispersing hominins might have been part of a small group of African taxa that expanded out of Africa at that time, implying that an ecological opportunity, such as the expansion of suitable habitats, existed for certain similar species (Turner 1999). Thus, the expansion of African savannas is thought to be a facilitating factor for the initial hominin dispersal. However, there is evidence that some of these sites are not similar in their faunal structure to that of African savannas.

Belmaker (2005) found that African savanna mammals were not abundant at ‘Ubeidiya, and that overall the faunal structure was more similar to that of Mediterranean environments. Other than hominins, very few African species dispersed at this time into

Europe or Asia (Martínez-Navarro 2004), showing that it was unlikely that hominins were part of a hypothetical group of African taxa entering East Asia simultaneously. The hypotheses tested in this dissertation concern the degree to which environmental or ecological similarity was an important factor in hominins’ ability to adapt to the new

1 Pleistocene and Pliocene are used to refer to their date range prior to June 29, 2009. 3 places they colonized. Each hypothesis is tested using data from the ecological properties of large mammals.

1) Did the ecological settings and faunas associated with early Homo in East Asia

differ from those in East Africa at this time, and if so, in what ways?

2) Were there ecological differences between hominin and non-hominin sites in East

Asia?

The community or ecological structure of the mammalian fauna was evaluated using ecostructure methods. In these methods, each species is classified using a combination of ecological variables including body size, diet and substrate (i.e., terrestrial, arboreal or aquatic). Ecostructure methods have been used, for example, by

Andrews (1979, 1996), Reed (1997, 1998, 2008), Rodríguez 2004, 2006a, b), and

Mendoza (2004, 2005). Methods that classify species by ecological properties are particularly useful in comparing sites with taxonomically different faunas in order to determine how the sites were ecologically similar despite the taxonomic differences. This type of method was used by Rodríguez (2004) to determine whether ecological change was occurring along with taxonomic change at Atapuerca. Reed (1997, 1998, 2008) used ecological classifications to reconstruct the paleoenvironments of hominins at

Makapansgat, Hadar and other East and South African sites. Ecostructure methods are useful for comparison because they show which types of animals are the sources of ecological similarity or difference between taxonomically different sites. The similarities or differences between ancient East Asian and East African faunal assemblages are

described using the results of correspondence analysis and comparison of the ecotype numbers and proportions between sites. The species within the ecotypes that contribute to similarities or differences are then investigated to determine how differences in their proportions could have impacted hominins.

Ecomorphological Similarity of the Carnivores

During the Plio-Pleistocene, hominins and carnivores had the potential to interact

while competing for carcasses. Archaeological evidence shows that Plio-Pleistocene

hominins and carnivores overlapped in their resource use, and therefore would have been

competitors (de Heinzelin 1999; Semaw et al. 2003; Dominguez-Rodrigo 2005; Semaw

2000; Potts, 1988, 2003; Bunn and Kroll 1986; Blumenschine 1995; Shipman 1986).

Hominins may have interacted with many different carnivore species in different ways.

While certain carnivores, such as saber-toothed felids, may have produced carcasses that

still contained flesh and marrow (Ewer 1954, Blumenschine 1987, Marean 1989, Arribas

and Palmqvist 1999), other carnivores, such as bone-cracking hyaenids, may have

consumed many of the carcasses on the landscape (Blumenschine 1987, Blumenschine et

al. 1994, Turner 1992). Also, carcass theft after a kill may occur depending on the body

size and grouping behaviors of the species involved (Van Valkenburgh 2001).

The prospects for hominins as hunters or scavengers during the Plio-Pleistocene would have depended in part on the group of carnivore species present and their ecological traits, such as body size and feeding adaptations. Feeding adaptations include morphological specialization for behaviors such as flesh-slicing or bone-cracking. Body

5 size is an important determinant of prey size and competitive interactions (Carbone et al.

1999, Van Valkenburgh 2001).

Feeding adaptations and body size are termed ecomorphological traits because they are based on morphological measurements that relate to the ecological characteristics of a species. These ecomorphological measurements can be used to characterize carnivores from different species according to their behaviors. For instance, ecomorphological characteristics are used to compare bone-cracking adaptations in hyenas in East Asia and East Africa, regardless of the taxonomic relationship between the species. This comparison shows differences in adaptations to bone-cracking and in the probable amounts of competition from carnivores that hominins would have faced in East

Asia and East Africa. Ecomorphological comparisons are used to determine whether specific carnivores are avatars. Avatars here are species from different regions that have similar feeding adaptations and body size (Damuth 1985). Measurements of feeding adaptations classify carnivores into dietary classes such as highly carnivorous, omnivorous, or bone-crackers. However, avatars that are similar in diet and in body size may have been different in other aspects of behavior, such as locomotion, that are not researched here.

The combination of ecological characteristics in the carnivore guild as a whole in a particular region and time would have shaped potential interactions with hominins that used meat and marrow and that may have scavenged from other animals. A guild refers to all species of a particular group that obtain and use resources in a similar manner (Root

1967). Here, the carnivore guild refers to members of the order Carnivora that are over 1 kg in body mass. Lewis (1995, 1997) hypothesized that there were ecological differences

in carnivore locomotion and prey capture behavior between East Africa and South Africa

that would have led to differences in hominin scavenging opportunities. Likewise, large numbers of bone-cracking species may have consumed many carcasses, making it more difficult for hominins to scavenge (Blumenschine 1987, Blumenschine et al. 1994, Turner

1992). This shows that the adaptations of the guild as a whole - the numbers of species with bone-cracking or flesh-slicing adaptations, as well as the numbers of omnivorous or carnivorous taxa that may have been competitors – would have been relevant to a scavenging or hunting hominin’s ability to obtain meat and bone marrow. Hominin use of marrow also occurred outside of Africa; percussion marked bones have been found dating to 1.66 Ma at Majuangou in the Nihewan Basin (Zhu et al. 2004). If, therefore, hominin behavioral potential was similar, then the composition of the carnivore guild in East Asia would have affected the possibilities for hominins living there. Carnivore adaptations were also important for dispersing hominins. Animal products have constant properties, unlike plant foods. Edible plants may vary in distribution, especially in the temperate latitudes of East Asia. In order to obtain those animal products, hominins would have had to interact with the carnivore guild. In this dissertation, similarities and differences in the carnivore guilds of East Asia and East Africa are determined in part by the presence of avatars. Guilds containing many similar avatars probably led to similar niches for carnivorous hominins because of similar relations between species. Specific differences between guilds could imply new competitors or sources of potentially scavengeable carcasses.

The structure of the East Asian carnivore guild is also relevant because hominins were a new immigrant taxon to East Asia and they were likely using resources formerly

exploited only by members of the order Carnivora. Hominins would have been entering

the guild of East Asian carnivores as a type of partially carnivorous mammal that was

unknown and unlike the others present.

Research on dispersal and immigration into new communities in general suggests that communities that are successfully colonized have suffered recent extinctions

(Vermeij 1991) or are less diverse than other communities that are saturated (Brown

1989, Ricklefs and Schluter 1993, Vermeij 1991). Immigrant taxa rarely cause replacement by competitive exclusion (Vermeij 1991), but may instead cause enrichment in the recipient community when an immigrant adds to the species diversity (Flynn et al.

1991, Vermeij 1991). Immigrant taxa may use resources in a different way compared with the incumbent taxa in order to be successfully integrated into the endemic community.

The circumstances of hominin colonization of East Asia during the early

Pleistocene with regard to the carnivore guild are investigated here. Older carnivore guilds from Pliocene faunal assemblages from Longdan, Longgupo and the Haiyan

Formation in the Yushe Basin are compared with the Pleistocene guilds from the other

East Asian sites. Comparisons of the carnivore guild prior to and after hominin arrival could show whether hominins took advantage of unrepresented roles, such as flesh- slicing, bone-crushing or group hunting, or whether hominins were likely to have enriched the carnivore guild, using resources in a different way. Character displacement in anatomical traits minimizes overlap of resource use and competition among carnivore guild members (Mooney and Cleland 2001). It may occur after the immigration of a new

species into the guild (Ricklefs and Schluter 1993). However, the possibility of character

8 displacement with regard to the East Asian carnivore guild must be evaluated in terms of both hominin immigration and general environmental change. Environmental changes may have resulted in changes in patterns of resource consumption (Sher and Hyatt 1999,

Davis et al. 2000, Shea and Chesson 2002) and provided opportunities for invaders

(Lozon and MacIsaac 1997). Environmental changes in both the Asian and African regions during the Plio-Pleistocene are discussed in chapter two. The questions asked about carnivores in this dissertation are as follows:

1) Were there carnivore avatars in East Asia and East Africa? Were the carnivore

guilds as a whole similar, with a similar distribution of ecotypes? What were the

differences in the ecological characteristics of the carnivores in these regions?

2) Were there changes in carnivore ecomorphology between the Pliocene and

Pleistocene of East Asia? Were there changes in the structure of the East Asian

carnivore guild? Prior extinctions without replacement by another carnivore

avatar may have indicated unfilled niches, whereas a lack of change in the

distribution of avatars may indicate enrichment or increased ecological diversity

of the East Asian fauna when hominins arrived. Opportunities for hominins in a

guild with unfilled niches would have differed from those in a saturated guild or

one with increasing ecological diversity.

The ecological characteristics of the fossil carnivores from East Africa and East

Asia are evaluated using ecomorphological measurements of the dentition designed to sort carnivores into feeding categories including flesh-specialists or hypercarnivores,

bone-crackers and omnivores. These measurements show quantifiable similarities and

differences between species. For each ecomorphological index, categories are created for

ecologically different subsets of measurements. These analyses show which species are

avatars, and are used to compare the guilds of East Asia and East Africa, as well as the

East Asian guilds through time.

Organization of the Dissertation

This dissertation is organized into two sections, the first concerned with questions about the overall ecological similarity of the large mammalian community, and the second with the comparative ecomorphology of the carnivore guild. Background to the question of overall mammalian ecological similarity, as well as detailed information about the sites from which the ancient faunal assemblages are drawn is contained in chapter two. Chapter three describes the methods used to analyze ecological similarity of ancient East Asian and East African faunas. The results of these analyses are given in chapter four. Background to the section on carnivore ecomorphology is in chapter five.

Chapter six describes the methods used to analyze comparative carnivore ecomorphology, and chapter seven describes the results. Discussion of all results and general conclusions are presented in chapter eight.

Chapter 2: Background to Ecological Similarity Analysis

Hominins are an integral part of the mammalian community (Foley 1987).

Mammalian remains found with hominins have been used to interpret hominin habitat preferences. The mammalian ecological structure (or community structure), which is defined as the proportions of mammals that have certain classes of adaptations, is correlated with environmental conditions, including the amount of vegetation, precipitation and the temperature range. The types of mammals present in a community may affect the animal resources available for a hominin, through scavenging or hunting.

The mammal community also reflects the vegetative community, which is also a food source for hominins. These aspects of habitat and mammalian community structure are particularly interesting when considering the ecological context of hominin dispersal to

East Asia and how those conditions compare with East Africa.

This chapter discusses the use of mammalian adaptations to make environmental determinations, with reference to ecological structure methods. The modern comparative environmental divisions used in the analysis are described. Theoretical expectations for a dispersing mammal, information about mammalian dispersals from Africa during the early Pleistocene, and environmental conditions conducive to dispersal are discussed.

Finally, information about the Plio-Pleistocene East African and East Asian research sites, with emphasis on their environmental conditions, is summarized.

Mammals as Environmental Indicators

Mammals in hominin sites have been used to determine the type of environment in which hominins lived. Functional morphology of single species or groups of species

(such as bovids or carnivores) found with hominins has been used to estimate aspects of habitat such as vegetation cover (Kappelman 1988, 1997; Lewis 1997; Spencer 1997;

Elton 2001, 2002; Vrba 1974, 1975, 1980). Abundance data for mammals that are correlated to particular habitat types (such as closed or open) reflect environmental shifts

(Bobe and Eck 2001, Bobe et al. 2002, Bobe and Behrensmeyer 2004).

Ecological structure in the mammalian community is commonly defined as the proportions of adaptations related to diet, body size, locomotion and substrate use

(Andrews 1996, Reed 1998, Rodríguez 2004). Patterns of ecological structure are based on physical factors such as climate, vegetation and precipitation, and are correlated with habitat types (Andrews et al. 1979). Communities from locations with similar physical conditions converge on a similar structure, even if the sets of species from those communities are taxonomically different. Community structure methods describe faunas by the ecological rather than taxonomic composition of the fauna. Higher temperatures and greater amounts of water lead to greater plant productivity, which in turn increases the number of herbivores (Ritchie and Olff 1999, Janis et al. 2002) and the number of species relying upon arboreal substrates. African habitats range from rainforests to deserts, with differing amounts of moisture and different temperature regimes in each.

Africa has a number of different habitats, in which different ecological structures and proportions of adaptations are found (Reed 1997, 1998, 2008; Andrews et al. 1979,

Andrews 1996). African forests have similar ecological proportions to tropical forests in

Australia, Malaya and Panama (Andrews et al. 1979). Some ecological types are

particularly useful for distinguishing habitat types. Andrews (1996) found that temperate

environments tend to have more terrestrial animals compared with tropical ones. Tropical

and non-seasonal habitats, such as evergreen forests, have more frugivorous, arboreal,

and scansorial species than environments that were drier, such as savannas (Andrews

1996). Mendoza et al. (2005) found that grazers and mixed feeders are more common in

bushland and savannas than in forests or arid environments. Mendoza et al. (2005) also

found that highly carnivorous species (including bone-cracking animals) are more

common in open environments. Within Africa, Reed (2008) found that adaptations such

as arboreality, terrestriality, frugivory, grazing and mixed feeding are most useful in

distinguishing habitats, with arboreality and frugivory signaling more vegetation cover,

or the presence of riverine gallery forests, while other adaptations signify more open

landscapes.

Ecological or community structure methods use different modern environments to

model how the ecological structure changes with habitat. Ecological structure methods

are also used to compare among ancient faunas to determine how they differ. Comparison of ancient and modern faunas described using ecological structure methods may show whether ancient faunas are analogous to modern ones. This dissertation compares Plio-

Pleistocene fossil faunas located in East Africa and East Asia. The East Asian fossil localities occur at higher latitudes compared with the East African comparative sites.

While most seasonal shifts in modern African localities concern the amount of precipitation, higher latitudes also experience temperature shifts (Reed and Rector 2007).

Construction of an ecological structure system to compare localities from distant

13 geographic locations, as well as from tropical and temperate habitats, requires the use of relatively broad environmental categories (Mendoza et al. 2005). Here, a set of fauna from modern African and Eurasian localities is classified into environmental categories using Bailey’s ecoregions (Bailey 1998), which are described in more detail below.

Biogeography and Ecological Structure :

Though ecological structures are similar in different locations under similar ecological conditions, historical factors affect the distribution of species and the proportions of ecological types, producing geographic differences in structure. For instance, Andrews (1996) found geographic effects in an ecological analysis of modern tropical evergreen forests, temperate forests, savanna woodlands, steppe, and tundra habitats. While the tropical forests were separated from the other environments, geographic substructure was evident in the separation between African and Asian tropical forests. Likewise, temperate deciduous forests in Eurasia and North America had different structures, possibly resulting from different environmental conditions unaccounted for in the study, different histories and different regional species pools.

An ecological structure study of modern localities in Eurasia and North America looked at the relative roles of convergence in mammalian communities from similar environments compared with the influence of geographic location (Rodríguez et al.

2006). Both biogeography and environmental variation played a role in the positioning these faunas in multivariate space (Rodríguez et al. 2006). Arid communities, such as deserts and steppes, were particularly convergent, perhaps because there are few ways to structure such a community with the limited available resources (Rodríguez et al. 2006).

There were significant differences between New World (Nearctic) and Old World

(Palearctic) faunal communities overall, which relate to differences in the composition of the species pools in these regions.

Another analysis compared the ecological structure of mammalian communities grouped by vegetation type in African and Asia to generate a predictive model (Mendoza et al. 2005). The discriminant function analysis grouped broadly similar vegetational communities from the two continents together showing that ecological structure can identify environmental convergence (Mendoza et al. 2005). However, some vegetation types are only found in Africa or Asia, meaning that some communities could not be compared (Mendoza et al. 2005). Evergreen forest, bushland and arid communities, which are found in both Africa and Asia, are similar in structure. Asian deciduous forests, which also contain grass areas, are similar to African wooded savannas. By grouping the habitats into relatively broad categories when comparing distant geographic localities, it is possible to see the general features of ecological structure (Mendoza et al. 2005).

Ancient Communities without Modern Analogs :

Some aspects of past ecosystems (or past ecosystems as a whole) may not have a modern analog, a phenomenon called historical non-equivalence. When analyzing the fauna of Makapansgat, Reed (1998) found that proportions of some extant taxonomic and ecological groups were very different compared with modern sites. Andrews et al. (1979) also noted some localities seemed to have a distribution of ecological types that is not represented in current African settings. In other cases, such as an Olduvai fauna from the middle of Bed I, ecological information from faunas does not match current habits of

15 close relatives, leading to an interpretation of greater structural complexity in that ancient habitat compared with the modern location (Soligo and Andrews 2004). Andrews (1996) recommended comparing ecological variables individually to determine the characteristics of ancient habitats that do not correspond to modern categories.

Environmental Profiles of Modern Comparative Sites:

Bailey’s ecoregions were used to classify modern localities. This is a hierarchical system in which the world is divided into domains and those domains are each divided into divisions. Each division may be either lowland or mountainous. Temperature and moisture patterns divide the world into tropical humid, humid temperate, polar and dry domains (Bailey 1998). Within these domains, latitude, precipitation, continental position, altitude and seasonality produce vegetation patterns that are described in the divisions. Each division has a typical series of altitudinal vegetational zones that occur when that division contains areas of mountainous terrain. Descriptions below are based on characteristics described in Bailey (1998) unless otherwise noted.

Dry Domain:

Temperate Deserts :

Temperate deserts are found in the interior of the Eurasian continent. They receive very little precipitation and have hot summer temperatures and very cold winter temperatures. The lack of water and extreme temperature range limits vegetation to woody shrubs. As altitude increases, the vegetation changes first to semi-desert woodland and then to meadow.

Temperate Steppe:

Temperate steppes typically feature grasslands with scattered scrubland and trees.

Most fauna are grazers. Winters are cold and dry, while summers are hot or warm with rainfall. If rainfall decreases, temperate steppes may become deserts. The altitudinal sequence from lower lands to higher ground for temperate steppe is: steppe, coniferous forest, tundra or in other areas, steppe, mixed forest and meadow.

Tropical-Subtropical Steppe :

Tropical-subtropical steppes are arid and hot. Precipitation occurs irregularly from year to year. The vegetation consists mainly of grass, but may include shrubs or trees.

This division may also be described as an acacia-grassland savanna. Tropical-subtropical

Steppe Mountains include a gradient that runs from steppe or semi-desert to mixed or coniferous forest to alpine meadow or steppe.

Tropical-Subtropical Desert :

These deserts are extremely arid with large variations in temperature between day and night. Very sparse vegetation includes shrubs, cacti and grass. Tropical-subtropical deserts include the Sahara, the Arabian Peninsula and the Thar. The altitudinal sequence is from lower to higher altitudes is semi-desert, shrub, open woodland and finally steppe or meadow.

Humid Temperate Domain:

Prairie :

Although the precipitation in prairies is sufficient to support grassland, it does not support trees unless the prairie is close to a wetter division. In that case, mosaics of

17 deciduous forest and grassland may occur. Mountains cause the following zones: forest- steppe, coniferous forest and meadow.

Subtropical :

The subtropical division is characterized by a climate without a dry season.

Streams contain water for most of the year. The average annual rainfall for subtropical forests of the broadleaf schlerophyllous type is 1283.7mm (Wang 1961). Precipitation is increased during the summer. The average temperature for Chinese subtropical forests is

15.9°C (Wang 1961). Subtropical divisions typically contain forests with evergreens such as oak, laurel and magnolia. The forest floor is thickly vegetated with bamboo, shrubs and herbs. At northern borders, subtropical forests may also have deciduous broadleaf trees. The subtropical mountainous zones range from mixed forests to meadows.

Hot Continental :

Hot continental vegetation includes tall broadleaf deciduous trees, with a seasonal herb layer on the ground. This is the native division type for areas in northern China and was originally found between 20 and 50°N latitude (Ching 1991). Currently, it is not well represented in China because of the large human population (Ching 1991). The hot continental division has hot summers and cool winters, without a dry season. A mountainous area will have deciduous or mixed forest, coniferous forest and finally meadows.

Humid Tropical Domain:

Savanna :

Savannas are a very variable type of division, with different subtypes of vegetation. These vegetation types include scrub woodlands (with a discontinuous

18 canopy layer) and woodland savanna, in which grassland, trees and shrubs are interspersed. Savannas are found in Africa, as well as in India and Southeast Asia.

However, the human population in the Asian areas is large and has altered the vegetation and the faunas substantially (Cole 1986). Cole (1986) notes that dry deciduous woodlands in India and Burma are similar to savanna woodlands in Africa. In Southeast

Asia, deciduous forests are found in areas with 1000-2000 mm of rain and a four to seven month dry season (Cole 1986). These Southeast Asian savannas include discontinuous canopied woodlands interspersed with denser areas of deciduous or rainforest (Cole

1986). Blasco (1983) describes this type of vegetation as an open forest with grass covering the ground. Savannas have wet and dry seasons. Streams dry out in the dry season or flood surrounding grasslands during rainy seasons. Mountainous savannas include open woodlands, deciduous forest, coniferous forest and then a steppe or meadow.

Rainforest :

Rainforests include many species of trees able to thrive in a setting with high temperatures and abundant rainfall. A subtype of rainforest (tropical deciduous) occurs in areas that have a dry season. Tropical rainforest has a continuous canopy layer of broadleaf trees. The fauna is especially rich and includes many arboreal species.

Mountainous rainforests shift from evergreen forest to meadows.

Dispersal and Biogeography :

Theoretical Expectations for Dispersal :

The dispersal of hominins into East Asia can be considered in light of biogeographic theoretical expectations. According to Vermeij’s (1991) study of historical biotic exchanges, in which species from one geographic locality colonize another location, very few species actually disperse when a geographic exchange opportunity occurs. A new colonizing species may have a competitive advantage over species in the recipient biota, especially with regard to disease or pathogen resistance (Vermeij 1991).

The study also reveals that biotic exchange often occurs primarily in one direction, leading to speculation that one ecological community is competitively superior over the other. A mediating factor in exchange asymmetry is the presence of suitable habitats for immigrant species (Ricklefs and Schluter 1993). Similarity in physical aspects of the environment is important to successful colonization (Brown 1989). Environmental and physical similarity is likely to result in similar food resources and similar types of species in the communities (Brown 1989). However, many species are able to tolerate a range of environmental conditions when there are few predator and competitor species (Brown

1989). Prior extinctions may increase a community’s vulnerability to colonization

(Vermeij 1991). Brown (1989) also noted that less diverse communities are more likely to be colonized. In particular, a deficit of predators could be an important factor.

Conversely, a stable community that has not suffered extinctions could be said to be saturated (Ricklefs and Schluter 1993, Vermeij 1991) and thus less susceptible to invasion. If a saturated community were invaded, the process of competitive exclusion

would cause extinctions among the incumbent species. However, Vermeij (1991) found

that colonizing species rarely caused extinctions. Instead of replacement, dispersers find a

place within the new community in a process of enrichment (Flynn et al. 1991, Vermeij

1991), perhaps because the disperser is exploiting available resources in a new way

(Ricklefs and Schluter 1993). Brown (1989) also found that dispersing species succeed in

colonization more often when they can occupy new niches compared with the native

species.

Community Structure :

The composition of local communities is determined by climate, habitat and local landscape (Ricklefs and Schluter 1993), as well as by competitive and predatory interactions between species (Morin 1999). Local community structure and composition may also be influenced by the sequence in which organisms colonize the location

(Robinson and Dickinson 1987, Robinson and Edgemon 1988, Drake 1991, Drake et al.

1993, Wilson 1992). Above the local populations are metapopulations, which are local populations in a region linked by dispersal of species between them (Morin 1999).

Species from the regional metapopulation may replace species that become extinct in the local populations depending on their dispersal ability and their habitat and landscape preference (Morin 1999). The regional species pool is an important factor in maintaining local diversity (Brown and Gibson 1983). Community structure at a local level is greatly influenced by the composition of the regional species pool (Mooney 1977, Schluter 1986,

Ricklefs 1987, 1989, Lawton 1984, Cornell and Lawton 1992, Ricklefs and Schluter

1993, Brown 1995). The species present at the regional level are determined by differences in environments as well as different interspecific interactions (Paine 1966,

1974, Lubchenco 1978, 1980, Menge 1995). Presence or absence of certain individual species may also cause differences in regional species pools (Tonn and Magnuson 1984,

Rahel 1984, McPeek 1990, Werner and McPeek 1994). Species in the regional species pool may be supplied by diversification within clades (Webb et al. 2002), or by dispersal due to linkages with other regional species sets (Ricklefs and Schluter 1993).

Diversification processes may result from adaptive radiation (Rosenzweig 1978, 1995,

Pimm 1979, Feder et al. 1988, 1990, Schluter and McPhail 1992, Schluter 1993, Rice and

Hostert 1993, Schluter and Nagel 1995, Losos et al. 1998, McPeek and Brown 2000), sexual selection (Lande 1981, 1982, Lande and Kirkpatrick 1988, West-Eberhard 1983,

Kaneshiro 1983, 1988, 1989, Seger 1985, Kaneshiro and Boake 1987, Turner and

Burrows 1995, Seehausen et al. 1997, Payne and Krakhauer 1997), the evolution of specific mate recognition systems (Paterson 1978, 1993), or chromosomal rearrangements (King 1993).

Each species has a fundamental niche of conditions in which it can survive as a viable population (Hutchinson 1957). One of the aspects of the niche is the species’ geographic range. In phylogenetic niche conservatism, the ancestral niche characteristics of a species are conserved in its descendants, leading to similarities in environmental tolerance and failure to expand into adjacent but environmentally different territory

(Weins 2004, Peterson et al. 1999, Ricklefs and Latham 1992, Ackerly 2003, Weins and

Donoghue 2004). Niche evolution, either in expansion of environmental tolerance or a shift to a new environmental specialization, would enable a species to colonize new habitats (Weins and Donoghue 2004).

The role of neutral processes in community assembly is debated. In his neutral theory of biodiversity and biogeography, Hubbell (2001) asserts that ecological communities of trophically similar sympatric species are structured by chance, historical factors and random dispersal of individuals. These communities are open to immigration

and are not at equilibrium (Hubbell 2001). This neutral view contrasts with a niche

assembly theory in which members of the community interact strongly with each other,

competition plays a strong role and the composition of the community may be deduced

from functional roles of the species (MacArthur 1970, Diamond 1975). The relative

importance of random or neutral processes compared with interactions between species

and between species and their environments has also been investigated (Kembel and

Hubbell 2006, Kelly et al. 2008, Jabot and Chave 2009).

Dispersal of African Mammals:

Plio-Pleistocene African mammals are found in Asia at the Levantine site of

‘Ubeidiya, dated to approximately 1.6 and 1.2 Ma (Belmaker 2005). Although the fauna

includes a mix of species from different realms, such as the oriental, most elements were

from the Palearctic (Tchernov 1992b). The sub-Saharan species identified from that

period are Pelorovis oldowayensis , Oryx cf. gazella , Kolpochoerus oldovaiensis , Equus

tabeti , Theropithecus cf. oswaldi , Hippopotamus gorgops , Mellivora sp., Herpestes sp. ,

Megantereon sp. and Crocuta sp. (Belmaker 2005). However, many of the lineages with

African affinity were derived from in situ evolution of previously dispersed lineages

(Tchernov 1992a, b). Other genera, such as Crocuta and Megantereon , have been found

in Pliocene Asian sites and may not have been recent immigrants or may not have been

African-derived. The earlier Pliocene site of Bethlehem shows many open country

mammals in an environment interpreted as similar to an African savanna (Tchernov

1992a, b, Turner 1999). Martínez-Navarro (2004) suggested that the presence of

Pelorovis at ‘Ubeidiya implied the extension of savannas, while Kolpochoerus indicated the occurrence of gallery forests. Belmaker (2005), however, found that African-derived species were not particularly abundant at ‘Ubeidiya and instead reconstructed the site as similar to Mediterranean environments. Most of the African species found at ‘Ubeidiya did not disperse further into East Asia. The exceptions are Theropithecus oswaldi , which is found in Europe and India, and Homo (Martínez-Navarro 2004).

Environment and Dispersal :

Environmental and geographic changes may have presented opportunities for

hominins to expand into Eurasia, by the opening of physical pathways or by the spread or

existence of environments in which hominins were able to survive. Environmental

factors, including seasonal climate, temperature range, and ecological factors, such as

food availability, would have played a role in determining whether hominins were able to

survive in new dispersal sites for the long term.

Dennell (2004) tied hominin dispersal to the presence of grassland habitats.

According to climatic reconstructions, grasslands were present in Asia at 3 Ma (Dowsett

et al. 1999). Based on information that places East Asia in tropical and subtropical

environments and describes Indonesian settings as savanna or open woodland, Dennell

(2004) concluded that hominins would have been able to disperse across Asia because of

the similarity in environments, specifically the presence of savanna-like settings across

Asia. Vegetational and faunal contrasts between different latitudes were not as strong

during the late Pliocene and early Pleistocene as they are today, especially since modern

desert barriers were not yet fully formed (Dennell 2004). The appearance of savannas and

woodlands in Asia and Africa may have been tied to potential times of dispersal, which

means that dispersal prior to the early Pleistocene is possible (Dennell and Roebroeks

2005).

East Africa

Evidence of Hominins and Dates :

East African sites have long been a source of hominin fossils, as well as those of

other mammals and have provided information about hominin anatomy, culture and

habitat. Faunas from the well-known sites in Koobi Fora, West Turkana and Olduvai

Gorge are used here to compare to the East Asian Plio-Pleistocene sites. Turkana Basin

faunas were analyzed by stratigraphic members. At Koobi Fora, these members included

the Upper Burgi, KBS, Okote and Chari. The Chari member has a very sparse fauna

(Turner et al. 1999). At West Turkana, the members Lokalalei, Kalachoro, Kaitio, Natoo,

and Nachukui were included. Olduvai Beds I and II were also analyzed. The East

Turkana sites date from 2.68 to 0.74 (1.39 if the Chari is excluded) (Feibel et al. 1989,

Brown et al. 1985). The West Turkana members date from 2.52 to 0.7 Ma (Feibel et al.

1989, Brown et al. 1985). These sites were dated by correlation with tuffs dated by K/Ar

dating. Olduvai Bed I dates to 2.03 Ma to 1.78 and Bed II from 1.78 to 1.33 Ma (Walter

et al. 1991, 1992, Tamrat et al. 1995).

Environmental History of the Turkana Basin :

Environmental evidence as a whole shows increasing aridity and variability in

East Africa after 3 Ma (deMenocal 2004). Trauth et al. (2005) found evidence of deep lakes in the periods of 2.7 to 2.5 Ma, 1.9 to 1.7 Ma and 1.1 to 0.9 Ma in East Africa, leading them to conclude that those were relatively humid periods within the overall trend toward aridification. However, those periods were highly variable in climate, with both arid and humid intervals (Owen et al. 2008). A series of temporary lakes formed in the Turkana Basin beginning at 2.0 Ma, but lacustrine facies disappeared after 1.7 Ma

(Feibel et al. 1991, Feibel 1997). Also, river channels of the Omo dried up between 1.7

Ma and 1.4 Ma (Brown and Feibel 1991).

Pedogenic carbonates show a transition between woodland to more open savanna between 3 and 1 Ma at the Turkana Basin and Olduvai Gorge (Cerling 1992, Cerling et al. 1988). More arid-adapted mammalian taxa appear between 2.5 and 1.8 Ma

(Behrensmeyer et al. 1997). Based on pollen data, Bonnefille (1995) found that the

Lower Burgi had vegetation, first signaling relative cold, and then dry climate. During the time of the upper Burgi and through the Okote, climate fluctuated between humid and arid conditions. Pedogenic carbonates sampled at Koobi Fora indicated a trend of change from closed woodlands to open woodlands or shrublands between 2 and 1.75 Ma (Quinn et al. 2007). Between 1.75 and 1.5 Ma, different types of low shrubland environments were found, which Quinn et al. (2007) interpreted as fragmentation of woodland habitats.

After 2.0 Ma, the Turkana Basin experienced a period of faunal turnover and grassland expansions (Bobe and Behrensmeyer 2004). Many grassland mammal species have first appearances during the interval of 2.0 to 1.8 Ma, including hypsodont bovids

and suids, while the species that went extinct included forest and closed habitat dwellers

(Bobe and Behrensmeyer 2004). Omo mammals associated with forest habitats decreased in abundance after 3.2 Ma, while secondary grassland taxa became more abundant after

2.5 Ma (Bobe et al. 2002).

Reed’s (1997) ecological analysis of the Upper Burgi, KBS and Okote faunas

shows environmental changes. Whereas the Burgi fauna was interpreted as a mixture of

open woodland, edaphic grasslands and riparian woodland, with frugivores/folivores,

fresh grass grazers and terrestrial/arboreal animals, the KBS had fewer arboreal taxa and

a greater proportion of grazers (Reed 1997). The KBS was interpreted as scrub woodland

or arid shrubland with grasslands. The Okote was described as edaphic grasslands, having

many grazers, but also contained arboreal animals from gallery forests (Reed 1997).

Overall, hominins in the Turkana Basin lived in different habitat types while the climate

fluctuated, but tended towards aridification.

Environments in Olduvai Gorge :

A broad and shallow lake was present during the deposition of Bed I (Hay 1976).

Faunal analyses have been used to show what types of environment were present. Butler

and Greenwood (1973) and Gentry and Gentry (1978a,b) found conditions that signaled

increased aridity at the top of Bed I. Analyses based on bovid ecomorphology found that

closed or intermediate habitats prevailed (Kappelman 1984, Plummer and Bishop 1994).

Pedogenic carbonates sampled from an interval of 1.845 to 1.785 Ma showed variation in

climatic conditions and fluctuation between woodlands with open canopies and grass, and

wooded grasslands (Sikes and Ashley 2007). The amount of C 4 grassland varied between

40-60% (Sikes and Ashley 2007). Using an ecological structure analysis, Andrews et al.

(1979) concluded that Olduvai Bed I had a similar fauna to that of the Serengeti and to

woodland-bushland communities. During the deposition of Bed II, the lake was reduced

in size and the area was aridified (Hay 1976). Based on pedogenic carbonates, Sikes

(1994) found that lower Bed II supported a riparian forest, with grassy woodland further

away. Kappelman (1997) found that bovid ecomorphology supported a range of cover

from open to heavy cover, though there were no forest species. Carnivores and hominins

have been primary accumulators for some of the assemblages in Beds I and II

(Dominguez-Rodrigo et al. 2007a, b, Egeland 2007, Egeland and Dominguez-Rodrigo

2008, Leakey 1971, Potts, 1988, Monahan 1996, Blumenschine and Masao 1991).

East Asia

Environments in East Asia :

Environments in East Asia most likely played an important role in facilitating or impeding hominin dispersal and in determining whether occupation was long-term or short-term. Evidence from East Asia, including δ18 O from deep sea cores and loess deposition analysis, indicates climate change during the Plio-Pleistocene. The δ18 O record from deep sea cores records increases in ice volume after the late Pliocene

(Shackleton et al. 1985; Shackleton et al. 1990, Shackleton et al. 1995). The northern hemisphere ice sheet increased during the intervals of 3.6 to 2.7 Ma, 2.7 to 2.1 Ma and

1.5 to 0.25 Ma (Tian et al. 2002).

Loess, carried by the winter monsoon from the northwest of China, reflects aridification of central Asia (Liu 1985). The formation of deserts in north and northwest

China is linked to the uplift of the Tibetan plateau, which blocks moisture from the Indian

Ocean (Guo et al. 2002). Loess particles deposited at specific sites are coarser during

glacial intervals due to southward migration of deserts and to increased wind intensity

(Ding et al. 2005). The summer monsoon is responsible for approximately 80% of the

moisture in the loess-desert margin area, with the desert margin moving north as the

summer monsoon strengthens and moves north (Ding et al. 2005). Changes in the sizes of

loess particles indicates that the desert margin moved south at 2.6, 1.2, 0.7 and 0.2 Ma,

corresponding decreased strength of the summer monsoon and ultimately attributable to

glaciation (Ding et al. 2005).

The mineral record provides another proxy suggesting aridification. The mineral

hematite is formed by chemical weathering. The production of hematite is decreased

during glacial periods so that the content of hematite, measured by remnant

magnetization, reflects the degree of chemical weathering and the degree of aridification.

The patterns coincide with the δ18 O record to suggest aridification and cooling throughout the Quaternary during both glacial and interglacial periods (Deng et al. 2006).

Despite the overall trends of aridification and cooling, considerable environmental fluctuation probably occurred at specific sites. The pollen record from a location on the

Chinese Loess Plateau at 35°7’ N and 107°12 E shows changes in vegetation, temperature and the moisture regime in that area (Wu et al. 2007). Between 3 and 2.6 Ma, arboreal pollen was dominant, in a climate that was mostly warm and humid. This interval was followed by a drier and cooler period from 2.6 to 1.85 Ma, which featured arid adapted plants and fewer trees. The location sampled was reconstructed as having trees on the hills and grass-filled valleys. During the 1.85 to 1.5 Ma interval, Pinus , as

well as firs and spruce trees are dominant, leading to an interpretation of a forest-steppe

with a cool and humid climate. In the next time interval (1.5 to 0.95 Ma), Pinus remains

very common, but broadleaved trees and plants thriving in temperate environments are

present, showing a warm-temperate and humid climate. The species of Pinus present is

believed to be one restricted to warm, temperate environments with more than 400 mm of

rain annually. Subsequent samples, from the interval 0.95 to 0.5 Ma, show a decrease in

tree pollen and an increase in herbs and shrubs, suggesting an open steppe with

grasslands.

Other lines of evidence suggest climatic changes in areas east of the loess plateau

in China. Using data from soil, loess, pollen and the biogeographic distribution of apes

and cercopithecids, Jablonski et al. (2000) reconstructed most of southern China as a

tropical environment suitable for pongids and hylobatids during the late Pliocene and

early Pleistocene, while parts of northern China were thought to have had a more

subtropical climate.

East Asian Focus Sites: Environment, Fauna, and Dating

Chronology of East Asian Sites :

The East Asian sites are dated by a combination of paleomagnetic stratigraphy

and biochronological inferences and comparisons with better dated localities. The

Nihewan sensu stricto is a typical fauna. Many other faunas in East Asia, particularly in

North China, have been compared with it in order to assess similarity in terms of how many species are shared and thus to infer whether the fauna comes from approximately the same time period as the classic Nihewan fauna. Past studies have also looked at the proportion of extinct taxa as a means of judging relative age (e.g., Han and Xu 1985).

Recent work has produced land mammal ages for China typified by combinations of taxa,

and correlations with European mammalian biozones (Li et al. 1984, Tedford 1995, Tong

et al. 1995, Qiu and Qiu 1995, Deng 2006). Studies in the Yushe Basin (Tedford et al.

1991, Flynn et al. 1991, 1997), at Lantian (Zhang et al. 2002) and Lingtai (Zheng and

Zhang 2001) have served to better integrate mammalian biochronology with the

paleomagnetic timescale.

Nihewan Basin :

The Nihewan Basin has produced some of the earliest evidence of hominin

presence in China. It is also known as a source of mammalian fossils, which with

environmental data, produces important evidence about the ecological context of

hominins in East Asia. The Nihewan Basin sites examined here include the

archaeological sites of Majuangou, Donggutuo and Xiaochangliang, as well as the non-

hominin fauna known as the Nihewan sensu stricto or the Xiashagou fauna. The Nihewan basin contains fluvial, lacustrine and eolian sediments (Zhu et al. 2003, Deng et al. 2008).

Xiaochangliang :

The Xiaochangliang site, located in the Nihewan Basin at 40.2°N, 114.65°E, contains artifacts and a fossil fauna (Figure 2.1). The artifact layer, which includes many small flakes, is dated by magnetostratigraphic correlation to 1.36 Ma (Zhu et al. 2001,

2003). The fauna reported by Tang et al. (1995) included typical Nihewan taxa and corresponded to an early Pleistocene age. Most remains are highly fragmented (Peterson et al. 2003). From a sample of bones, Peterson et al. (2003) found that although 8.1% had carnivore toothmarks, this percentage was too small for the fauna to have been accumulated primarily by carnivores. Shen and Chen (1999) also found carnivore,

possibly hyena, modifications on the bones. The assemblage was most likely

hydraulically transported (Peterson et al. 2003, Shen and Chen 2003). The assemblage is associated with a conglomerate.

Donggutuo :

The Donggutuo site is located at 40°2 N and 114.67° E. It contains lakeshore sediments. Schick et al. (1991) report cores and flakes. The site was dated to approximately 1.1 Ma using paleomagnetic stratigraphy (Li and Wang 1982). This was confirmed by Schick and Dong (1993), Li et al. (2002), Wang et al. (2005) and Zhu et al.

(2003). The fauna is listed in Wei (1985, 1991), and in Deng et al. (2008).

Majuangou :

Majuangou is located at 40°13.517’ N and 114°39.844’ E in the Nihewan Basin.

The four artifact layers contain cores and flakes (Zhu et al. 2004). Paleomagnetic stratigraphy was used to determine the following dates for the artifact layers: 1.32, 1.55,

1.64 and 1.66 Ma (Zhu et al. 2004). Fossil bones show percussion marks, indicative of processing for marrow (Zhu et al. 2004). Biochronologically, the fauna is typical of the

Plio-Pleistocene and is similar to that of Xiaochangliang (Tang et al. 1995). Sediments show that the artifact layers were deposited in wetlands or in lake margins. Pollen data indicates considerable variation in vegetation over time at the time (Zhu and Potts, pers. comm.)

Xiashagou or Nihewan sensu stricto:

The Nihewan fauna contains many mammal species and has come to be considered a standard north China fauna for the late Pliocene and early Pleistocene

(Lucas 2001). However, the fauna does not include any hominin specimens. This fauna

was documented by Teilhard de Chardin and Piveteau (1930) and further studied by Qiu

(2000). Deng et al. (2008) estimated the age of the Nihewan faunas to be between the

onset of the Olduvai normal and the Brunhes-Matuyama boundary. Biochronological

studies show that the fauna corresponds taxonomically to the Olivola fauna at about 1.8

Ma (Qiu 2000). About 20% of the mammals are from the Tertiary (Lucas 2001). Many of

the species are forest browsers (Lucas 2001).

Gongwangling, Lantian :

Gongwangling is located at 34°12’ N and 109°28 E. A Homo erectus cranium was found at Gongwangling. The fossil was found in the L15 Loess Layer, which An and

Ho (1989) dated to 1.15 Ma using magnetostratigraphy. Heslop et al. (2000) estimated the age of the L15 loess at 1.22 to 1.19 Ma. The complete fauna was described by Hu and

Qi (1978). An environmental analysis of Gongwangling based on the fauna concluded that this site experienced relatively warm and moist conditions (Dong et al. 2000). The presence of forests was inferred based on a large number of forest-dwelling taxa (Dong et al. 2000). Gongwangling is located on the northern edge of the Qingling Mountains, which divide north and south China, and its fauna includes a number of typically southern Chinese forest mammals (Hu and Qi 1978). These southern taxa may indicate a warm period in which certain mammals were able to spread further north. Wang et al.

(1997) described the environment as a cold or cool dry winter with a warm, semi-humid summer based on stable isotope ratios from the last glacial-interglacial cycle.

Yuanmou :

Yuanmou is located in south China at 25°40’ N and 101°54’ E. The Yuanmou incisors and artifacts, found in a layer dated to ~ 1.7 Ma using magnetostratigraphy and

the sedimentation rate, are the earliest evidence for hominins in continental East Asia

(Zhu et al. 2008). The incisor morphology has affinities with Homo erectus and Homo

habilis (Zhu et al. 2008). The fauna contains some Pliocene survivor species, while other

species are typical of the early Pleistocene (Qian and Zhou 1991). Pollen showed the

presence of pine and other tree species, as well as herbaceous vegetation, indicating a

cool and temperate environment (Qian and Zhou 1991, Zhu et al. 2008). While many

grazing species are found in the fauna, taxa associated with other habitats, such as

bushland and forests, also occur (Qian and Zhou 1991).

Longgupo or Wushan

Longgupo is a cave site located in South China at 30.4°N, 109.1°E. It was formerly thought to be a hominin site (Huang and Fang 1991, Huang et al. 1995) but those remains are now thought to represent an ape (Wu 2000). Rocks with crude, sometimes overlapping facets were also found and considered to be stone tools by the excavation team (Huang and Fang 1991, Huang et al. 1995). The fauna from this site is used here as a non-hominin comparator site. Huang et al. (1995) estimated that the site

dates to 1.9-1.8 Ma. Biochronological analyses of the fauna based on the co-occurrence

of species showed that the site has to be Late Pliocene to early Pleistocene (Huang et al.

1995). Electron spin resonance was used to assign the ape level to the Olduvai subchron

(Huang et al. 1995). Many species in the ape zone have also been found at

Gigantopithecus Cave (Huang and Fang 1991). However, mammals from north China, as

well as the local area, are also present in the assemblage, indicating a mixture of species.

From the overall inferred habitat preferences of the species, Huang and Fang (1991)

inferred the presence of forests and a relatively moist climate. However, climate

fluctuations may have caused grasslands to develop during some phases. Pollen indicates climate change during the period when the middle hominoid zone was deposited. There was a transition from a cold and dry period with herbaceous vegetation, to a climate that

was warm and wet, with forests of evergreen trees (Huang and Fang 1991).

Mohui Cave:

Mohui Cave, located at 23°34’54” N and 107°00’08” E, is part of the Bubing

Basin, adjacent to the Bose Basin, South China. This cave is the uppermost in a sequence

of caves. The caves were formed when groundwater dissolved limestone. As the

groundwater sank to lower levels, new caves were formed, leaving the oldest caves in the

uppermost position. Flowstones from one of the younger caves were dated by U-series

analysis yielding a formation date of 350-200 ka (Wang et al. 2007). Many of the bones

were gnawed by rodents or carnivores (Wang et al. 2007). Some species typical of the

late Pliocene and early Pleistocene, Ailuropoda microta and Hesperotherium , are present in the assemblage and supported an early age assignment. The fauna is also similar in taxonomic composition to Longgupo and Gigantopithecus Cave (Wang et al. 2007), which are also dated to the late Pliocene or early Pleistocene. The ecological assessment of the fauna was aided by stable isotope analysis of some of the teeth. Results showed a closed, forest habitat (Wang et al. 2007).

Jianshi or Longgudong :

Jianshi is a multi-layered cave site located at 30°39’14.9” N, 110°04’29.1”E.

Paleomagnetic information was used to date the site. The younger layers were reported to

be from the Olduvai subchron, while older layers (including a potential hominin) were

dated to greater than 2.15 Ma. The potential hominin teeth found during recent

excavations included an upper third molar, a lower first molar and an upper third

premolar. Other teeth attributed to hominins have been obtained earlier. Zheng (2004)

concluded after metrical comparison that the specimens are similar to Meganthropus ,

Australopithecus or Pithecanthropus (based on specimens now sunk into Homo ).

However, the teeth may represent a species of non-hominin hominoid (Schwartz et al.

1995, Ciochon 2009). Most micromammalian genera come from a geographic class labeled as the “middle subtropical forest type” (Zheng et al. 2004). Environmental classifications of large mammals based on extant relatives show that many came from forested tropical or subtropical environments (Zheng et al. 2004). Pollen records showed climate differences for different layers, although the site overall is dominated by conifers from mountainous areas and by broadleaf trees (Zheng et al. 2004).

Linyi :

Linyi is a non-hominin site in Shanxi, located at 36°12’ N and 110°30’ E. The

fauna was found in sands that underlie loessic beds in a lacustrine-alluvial deposit (Tang

et al. 1983). Many of the mammalian species found at this site are typical of northern

China, with the same genera or species being found at the Nihewan Basin or at Xihoudu

(Tang et al. 1983). Due to that faunal resemblance and the fact that the fauna contains

archaic and extinct species, it is thought to date to the Middle or Late Villafranchian

period (Tang et al. 1983). Many of the species (such as Equus , Paracamelus , Gazella , and Coelodonta ) are typical of steppe faunas and are thought to have come from a grassland environment.

Longdan:

Longdan is located at 35° N and 103° E. The Longdan fauna is dated to between

2.55 and 2.16 Ma by paleomagnetic stratigraphy (Qiu et al. 2004). Based on evidence from faunal resemblance, Qiu et al. (2004) concluded that the fauna was similar to that of the late Pliocene site St. Vallier, France and dated to about 2.2 Ma. It is definitely older than that of the Nihewan Basin, since Longdan contains some primitive species or primitive forms of species (Qiu et al. 2004). Specimens from this site were obtained from private collections, so detailed information about provenance is unavailable. Collection bias may have resulted in the composition of this assemblage including many species of carnivores. Qiu et al. (2004) also noted a taphonomic bias against small animals. There is evidence that Longdan included steppe or open environments, as well as forested areas.

Climate change may have led to changes in the local habitat. Coelodonta nihowanensis and Hipparion sinense specimens were both relatively hypsodont; six of the other herbivores were hypsodont, leading Qiu et al. (2004) to infer the presence of steppe or open habitats. Six herbivore species were thought to have either lived in forested environments or to have been browsers or frugivores, supporting the inference of the presence of bushland, shrubland or forest habitats (Qiu et al. 2004).

Haiyan Formation, Yushe Basin :

The Haiyan Formation is located in the Yushe Basin in North China, at 37°5’ N and 112°59’ E. The sediments have been dated to the late Pliocene (Flynn at al. 1991).

Based on magnetostratigraphy, the Haiyan Formation corresponds to the lower

Matuyama reverse chron, and was deposited prior to the Olduvai event, with dates between 2.5 and 1.9 Ma (Flynn et al. 1991). Biochronological correlation also supports

37 this age (Qiu 1990). Micromammal faunas show considerable turnover between the preceding Mazegou Formation and the Haiyan, which has 11 first appearances (Flynn et al. 1991). Turnover also occurred at that boundary among the large mammals (Flynn et al. 1991). Seven large mammal species also made last appearances in the Haiyan

Formation (Flynn et al. 1991). This fauna is unpublished and information about it is limited.

Figure 2.1 Map of the East Asian fossil localities. The Nihewan includes the sites of

Xiaochangliang, Donggutuo and Majuangou, as well as the Nihewan sensu stricto fauna.

0 700 km

Jianshi

Summary :

This chapter introduces ecological structure methods, which describe mammalian communities using proportions of species with adaptations relating to diet, body size and substrate. Differences in ecological structure correlate to differences in climate and environment. However, ecological structure is also affected by biogeographic and historical processes, especially as these processes affect the composition of the regional species pool, from which species found in specific assemblages are drawn. In order to show how ecological structure relates to environmental differences, modern sites were classified into environmental groups based on factors such as latitude, precipitation and temperature using Bailey’s ecoregions.

Another focus of this chapter is a review of theories about the dispersal of African mammals, and Homo in particular, to new regions. Ideas considered include the simultaneous dispersal of a group of African mammals, including hominins, out of

Africa, and the idea that hominin dispersal was linked to the spread of savanna habitats.

These theories, as well as other habitat information about the Plio-Pleistocene sites will be considered in light of results presented in the following chapters of the dissertation.

Finally, the specific sites in East Asia and East Africa to be analyzed were introduced, along with general climatic and environmental conditions in the two regions.

East African sites were from Olduvai Beds I and II and East and West Turkana. East

Asian sites included both hominin and non-hominin localities. The East African Plio-

Pleistocene shows a trend toward increasing aridity and variability after 3 Ma, with habitats becoming more open, while East Asia experienced glaciation and cooling after

3.5 Ma, with decreases in precipitation after 2.6 Ma.

Chapter 3: Methods to determine ecological similarity

This research is concerned with the ecological context of the initial hominin dispersal out of Africa. Specifically it looks at the degree of ecological similarity between communities of mammals in East Asia and East Africa and compares the ecology of East

Asian communities during the Plio-Pleistocene. Ecological similarity is defined as the similarity between animals based on the ecological properties of diet, body mass and substrate use (i.e., terrestrial, arboreal or aquatic). Similarity in community or ecological structure implies communities with similar proportions of ecologically comparable animals. Ecological community structure methods are based on the principle of convergence of environmental conditions; i.e., communities from similar environments will have similar structures (Andrews 1996). Ecological structure methods are useful for determining the degree of ecological similarity between taxonomically distinct faunas.

Ecological Structure and Ecotypes:

Ecotypes were used to classify each large mammalian species ecologically (Table

3.1). An ecotype is defined here as a classification combining animals of similar diet, body mass and substrate use classes. By using ecotypes, similar species are placed in the same group or ecotype, regardless of their taxonomy. The numbers of the species in each ecotype at the sites were compared using multivariate analysis methods. The proportions of ecotypes at each site are a way of showing the community or ecological structure of the mammalian fauna. Ecological structure differences relate to environmental differences between different locations. In order to show the magnitude of differences in the proportions of ecotypes (and thus the structures) between sites, and how the

ecological structures found in ancient sites relate to environmental distinctions, the

ecological structures of modern faunas from various locations and habitats were also

analyzed by classifying their faunas as ecotypes. Similar methods have been used by

Andrews (1996), Mendoza et al. (2005), Reed (1997, 1998, 2008), and Rodríguez (2004,

2006).

The ecotype classification system has to encompass variation in the modern and

fossil mammals, use categories that could be assigned based on information available in

the fossil record and do this without using too many categories, which would lower the

power of the analysis (Mendoza 2005). Comparisons of East Asian and East African

fossil ecotypes here are limited to large mammals, defined as those over 1 kg in body

mass. Chiroptera are excluded, as in Rodríguez (2004, 2006). Similar definitions of large

mammals have been used in other studies, focusing on mammals over 500 g (Reed 1997,

1998), over 1 kg (Mendoza et al. 2005), and over 4 kg (Reed 2008). Small mammals are

not found in many assemblages collected in the early 20 th century or collected without sieving. Also, hominins were more likely to have interacted ecologically with large mammals (Foley 1987).

Table 3.1 Ecotype classifications. Ecotype descriptions and their codes are listed.

Code Description SCAR Small carnivore (< 20 kg) LCAR Large carnivore (>20 kg) INS Insectivore (feeds primarily on invertebrates) FRUG Frugivore (primarily feeds on fruits) AQ Aquatic (non-ungulate) ARB Arboreal vegetation feeder (non-frugivore) TERSM Terrestrial mixed vegetation feeder (1-20 kg) BRM Terrestrial browser (20-300 kg) GRM Terrestrial grazer (20-300 kg) MXM Terrestrial mixed feeder (20-300 kg) BRL Terrestrial browser (300-1000 kg) GRL Terrestrial grazer (300-1000 kg) MXL Terrestrial mixed feeder (300-1000 kg) MEGBR Mega browser (>1000 kg) MEGGR Mega grazer (>1000 kg) MEGMX Mega mixed-feeder (>1000 kg) OMNSM Small omnivore (1-20 kg) OMNL Large omnivore (>20 kg)

Diet Categories

Dietary categories used to classify species included grazers, browsers, mixed

vegetation feeders, omnivores, carnivores, insectivores, and frugivores. Other studies have divided grazers into fresh-grass and general grazers (Mendoza et al. 2005, Reed

1997, 1998, 2008), but the information available here is not sufficient for such divisions.

Other studies omit mixed feeder classifications of ungulates (Andrews et al. 1979,

Andrews 1996), but this loses information that is available from hypsodonty. Ungulates have also been divided into ruminants and non-ruminants, rather than dietary categories.

This eliminated the problems inherent in assigning diet to extinct animals, since ruminant adaptations are only found in certain taxonomic groups (Rodríguez 2004, 2006,

Rodríguez et al. 2006). The possible distinctions of carnivores into hypercarnivores and

bone-crushers (as in Mendoza et al. 2005, Reed 1997, 1998, 2008) were not used in this

section in order to avoid the creation of large numbers of dietary categories. Dietary

distinctions between carnivores are explored in more detail in the second half of the

dissertation. Feeders on underground storage organs such as roots and tubers have been

categorized separately in some studies (Reed 1997, 1998, 2008), but were included under

various mixed vegetation categories here. Grazers are species that feed predominantly

upon grasses. Browsers feed primarily upon leaves while frugivores feed primarily upon

fruits. Mixed feeders are defined here as eating a variety of vegetation. Omnivores are

species that take a variety of foods including vegetation as well as vertebrates or

invertebrates. Omnivores in modern faunas include suids and some carnivores. Species

subsisting primarily upon vertebrate prey are classified as carnivores.

Size Categories

Body size classes are often used to classify mammals found in fossil sites. They

were first used in African paleoecology by Brain (1974) and were modified by Bunn

(1982) to include size classes for large animals such as Hippopotamidae, Rhinocerotidae

and Elephantidae. However, using these common body mass classes would have created many different possible ecotypes, thus increasing the number of variables and decreasing the power of the analysis. Here divisions were made at 20 kg, 300 and 1000 kg. Predation patterns differ in carnivores, with those over 20 kg in body mass hunting medium and large-sized prey (Carbone et al. 1999; Kingdon 1997; Lewis 1997; Van Valkenburgh and

Koepfli 1993). Carnivores that weigh less than 20 kg tend to focus upon smaller mammals as prey or are partially omnivorous in their diet (Carbone et al. 1999).

Terrestrial herbivores were broken into classes of less than 20 kg, between 20 and 300 kg, 300 to 1000 kg and over 1000 kg. Most ungulates have a body mass greater than 20 kg. Animals over 300 kg are not normally captured by most modern carnivores, although they can be captured by lions and tigers (Lewis 1997; Pienaar 1969; Schaller 1972;

Seidensticker and McDougal 1993). Animals over 1000 kg are not regularly preyed upon by extant carnivores (see sources in Lewis 1997).

Substrate

Substrate indicates whether a mammal was terrestrial, arboreal or aquatic. The possible aerial substrate category that has been used in some studies (Andrews 1996,

Andrews et al. 1979) was not included as bats (Chiroptera) have been eliminated.

Fossorial mammals (in a subterranean substrate) used by Andrews (1996, et al. 1979),

Reed (1997, 1998, 2008) and Rodríguez (2004, 2006, et al. 2006) were also not used since these animals were too small to be included in this study. Ungulates were assumed to be primarily terrestrial in their substrate. Arboreality in primates was assigned based on functional morphological studies from the literature, the characteristic behavior of modern representatives if the species is extant, or upon characteristics of extant members of the genus or family if other information is not available. Fossil carnivore ecotypes did not include a substrate categorization because that information was not available for many of the Asian fossil species, particularly the species of felids, and the closest relatives were unknown or varied in their adaptations.

Modern Comparative Sites

Ecological structure analyses of ancient faunas use the structures of modern

faunas to interpret the amount of difference between ancient faunas, as well as their

environmental context (Andrews et al. 1979, Andrews 1996, Mendoza et al. 2005, Reed

1997, 1998, 2008, Rodríguez 2004, 2006, Rodríguez et al. 2006). Modern sites from

similar environments plot close together in a multivariate analysis. The amount of scatter

in a group of sites that are from the same environment shows how much variation is

possible within similar climatic and vegetational conditions. Comparison of modern and

ancient sites in the same analysis can be used to show whether ancient sites are analogous

to modern sites in ecological structure. Modern comparative faunal communities were

drawn from sites in Eurasia (with a focus on East Asia), Southeast Asia, the Indian

subcontinent, as well as Africa. The modern comparative localities were chosen to

include habitats found in modern East Asia and East Africa. Some of these habitats, such as subtropical forest and savanna, may have been present during the Plio-Pleistocene.

Modern localities were classified into groups with similar environments using the

Bailey system (1998). The Bailey system was chosen over other descriptive systems to

make broad geographic comparisons possible, as in Rodríguez (2006). Bailey’s

ecoregions system is an environmental classification system based upon the principle of

identifying analogous regions worldwide using factors including latitude, continental

position and altitude (Bailey 1998). This system has a broader classification of localities

than those typically used for African localities (e.g., Reed 1997, 1998, 2008), which

involve detailed vegetational gradients. African vegetational categories do not correspond

to the types of floral structure found in many Asian locations.

Latitude is a factor in the Bailey system because of its relationship to seasonality

and precipitation. Different sets of seasons (summer, winter or both) are present at

different latitudes. Lower latitudes also tend to have higher amounts of precipitation.

Temperature and precipitation combined can be used to delineate Bailey’s four domains:

humid tropical, humid temperate, polar and dry. These domains are further subdivided

based on rainfall and predominant vegetation. Within each domain, divisions are based

on regional conditions. Altitude, which does not vary regularly, is taken into account

within each division. The modern localities used for comparison in this study were

classified as at the division level (Table 3.2, Figures 3.1 and 3.2). The divisions ‘savanna’ and ‘subtropical lowland’ were common in East Asia and East Africa.

The savanna division generally occurs between 10° and 30° latitude and has wet and dry seasons (Bailey 1998). Vegetation varies, grading from arid areas without grass through grassy plains to mixtures of grass and trees (Bailey 1998). Many of the vegetational gradations used for African sites are found within the savanna division, with the stipulation that different levels of tree cover occur depending upon water availability.

Subtropical forests have changing seasons, including winter but without a dry season (Bailey 1998). The climate is typically temperate with at least 30 mm of rain

(Bailey 1998). The dominant vegetation is forest. East Asian subtropical areas have broadleaf evergreen forests with oak, laurel and magnolia trees as well as lower vegetation layers including ferns, palms, bamboo, shrubs, herbaceous plants, epiphytes

47 and lianas in the lower layers (Bailey 1998). Deciduous forests are found in northern areas of the subtropical forest division (Bailey 1998).

Faunas from nature preserves and national parks, in which wildlife and vegetation are preserved, were selected as sites for the modern comparative database. However, many more large mammals are extinct in Eurasia compared with Africa. More species of large mammals were present in historical times. This difference between Asian and

African faunas may contribute to some artificial ecological separation between the continents.

Table 3.2 Modern comparative localities. The numbers in the first column are the codes to the localities and can be used to find the sites on the maps, Figures 3.1 and 3.2. The numbers in the final column, R, are the references, listed at the end of the table. Abbreviations: Temp. = temperate; Eur. = Europe Locality Cont. Country Lat. Long. Domain Division R. 1 Ailaoshan Asia China 23°36’- 100°54’- Humid Subtropical 1 24°44’ 101°3’E Temp. Mountains N 2 Alaungdaw Asia Myanmar 22°08’- 94°15’- Humid Rainforest 2 Kathapa 22°42’ 94°27’ E Tropical National Park N 3 Amboseli Africa Kenya 02°33’ - 37°06’- Humid Savanna 3 02°45’ 37°24’ E Tropical S 4 Amudarya Asia Turkmenistan 41°N 61°47’ E Dry Temperate 3 Zapovednik Desert 5 Astrakhanskiy Asia Russia 45°28' - 47°49' - Dry Temperate 3 46°22'N 49°07'E Desert 6 Baimaxueshan Asia China 27°46'- 98°57'- Dry Tropical/ 1 28°36'N 99°20'E Subtropical Steppe Mountains 7 Boucle du Africa Mali 13°10' - 08°25' - Dry Tropical/ 3 Baoule 14°30'N 09°50'W Subtropical Steppe 8 Changbai Asia China 41°41' - 127°43' - Humid Hot 3 42°51'N 128°16' Temp. Continenta E l Mountains 9 Changshanerhai Asia China 25°36’- 99°57'- Humid Subtropical 1 26° N 100°17' Temp. Regime E Mountains 10 Chernyje Zemli Asia Russia 46°05'N 42°20'E Dry Temperate 3

Locality Cont. Country Lat. Long. Domain Division R. Biosphere Steppe Reserve 11 Daurskiy Asia Russia 49°55' - 115°05' - Dry - Temperate 3 50°14'N 115°98' Humid Steppe - E Temp. Prairie 12 Daweishan Asia China 22°58' 113°.51' Humid Savanna 3 N E Tropical 13 Dinghushan Asia China 23°10' - 112°31' - Humid Subtropical 3 23°11'N 112°34' Temp. E 14 Doi Chiang Dao Asia Thailand 19°22' 98°58' E Humid Savanna 3 N Tropical 15 Doi Inthanon Asia Thailand 18°35'N 98°29'E Humid Savanna 3 Tropical 16 Doi Pha Chang Asia Thailand 19°9' N 100°30' Humid Savanna 3 E Tropical 17 Eravikulam Asia India 10°05' - 77°- Humid Savanna 3 10°20' 77°10' E Tropical N 18 Fanjingshan Asia China 27°46' - 108°45' - Humid Subtropical 3 28°01'N 108°48' Temp. E 19 Fujian Wuyi Asia China 27°40' 117°45' Humid Subtropical 3 N E Temp. 20 Gaoligong Asia China 24°55' - 98°7' - Humid Savanna 1,4 Mountain 28°22' 98°49' E Temp. N Humid Rainforest Tropical Mountains Dry Tropical/ Subtropical Steppe Mountains Humid Subtropical Temp. Regime Mountains 21 Gombe Africa Tanzania 4°40' S 29°38' E Humid Savanna 5 Tropical 22 Great Gobi Asia Mongolia 43°- 45° 94°-98° Dry Temperate 3 N E Desert Temperate Steppe Humid Prairie Temp. Regime Mountains 23 Great Himalayan Asia India 31°44' 77°33' E Humid Savanna 3 National Park N Tropical 24 Indravati Tiger Asia India 18°51- 80°16 - Humid Savanna 3 Reserve 19°24 N 80°44 E Tropical 25 Jiuzhaigou Asia China 32°54'- 103°46'- Humid Subtropical 2 33°19'N 104°4'E Temp. Regime Mountains 26 Kalahari Africa South Africa 25°40' S 20°19' E Dry Tropical/ 3 Gemsbok Subtropical Steppe

Locality Cont. Country Lat. Long. Domain Division R. 27 Kanha Tiger Asia India 22°13'- 89°32' - Humid Savanna 3 Reserve 22°27 N 89°45' E Tropical 28 Kaplankyr Asia Turkmenistan 40°56 N 57°02 E Dry Temperate 3 Desert 29 Kodry Eur. Moldova 47°47' 28°19' E Humid Prairie 3 Zapovednik N Temp. 30 Kudremukh Asia India 13°01' - 75°00' - Humid Rainforest 3 National Park 13°29' 75°25' E Tropical N 31 Lake Manyara Africa Tanzania 3°30' S 35°50' E Dry Tropical/ 3 Subtropical Steppe 32 Lake Nakuru Africa Kenya 0°24’ S 36°05’ E Humid Savanna 3, Tropical 6 Dry Tropical/ Subtropical Steppe 33 Ma Sa-Kog Ma Asia Thailand 18°45'; 98°45; Humid Savanna 3 Reserve 18°49- 98°47- Tropical 56' N 58' E 34 Mamili Africa Namibia 18°14' S 23°39 E Humid Savanna 3 Tropical 35 Manas National Asia India 26°30' 91°51' E Humid Rainforest 2, Park N Tropical Rainforest 3 Mountains 36 Maolan Asia China 25°09' - 107°52' - Humid Subtropical 3 Biosphere 25°20'N 108°05' Temp. Reserve E 37 Mare aux Africa Burkina Faso 11°30' - 04°05' - Humid Rainforest 3 Hippopotames 11°45'N 04°12'W Tropical 38 Medog Nature Asia China 29°15' 95°30' E Humid Rainforest 1 Reserve N Tropical Mountains 39 Moremi Africa Botswana 19°16' S 23°12' E Humid Savanna 3 Tropical 40 Mount Kenya Africa Kenya 0°10' S 37°20' E Humid Savanna 3 Tropical 41 Nouable-Ndoki Africa Republic of 2°15' N 16°24' E Humid Rainforest 3, the Congo Tropical 7 42 Nujiang Asia China 28°02' 98°35' E Dry Tropical/ 3 N Subtropical Steppe Mountains 43 Palava Eur. Czech 48°37’ - 16°36’ - Humid Prairie 3 Republic 48°53’ 17°05’ E Temp. N 44 Queen Elizabeth Africa Uganda 0°06' - 29°47' - Humid Savanna 3 0°46' N 30°11' E Tropical 45 Repetek Asia Turkmenistan 38°34'N 63°11'E Dry Temperate 3 Desert 46 Richtersveld Africa South Africa 28°17' S 17°8' E Dry Tropical/ 3 National Park Subtropical Steppe 47 Royal Chitwan Asia Nepal 27°30 N 84°20' E Humid Savanna

Locality Cont. Country Lat. Long. Domain Division R. Tropical 48 Sakaerat Asia Thailand 14°26 - 101°50 - Humid Savanna 3 Environmental 14°32 N 101°57 Tropical Research Station E 49 Serengeti Africa Tanzania 01°30' - 34°00' - Humid Savanna 3 Ngorongoro 03°20'S 35°15'E Tropical Dry Tropical/ Subtropical Steppe 50 Srebarna Eur. Bulgaria 44°05 N 27°07 E Dry Temperate 3 Steppe 51 Syunt- Asia Turkmenistan 38°30' 55°30' E Dry Tropical/ 3 Khasardagh N Subtropical Desert 52 Tai Forest Africa Ivory Coast 05°15' - 07°25' - Humid Rainforest 3 06°07'N 07°54'W Tropical 53 Tianmushan Asia China 30°18' - 119°23' - Humid Subtropical 3 Biosphere 30°25'N 119°29' Temp. Reserve E 54 Tsavo National Africa Kenya 2°46' - 38°8' - Humid Savanna 3 Park 3°19' S 38°46' E Tropical 55 Tsentral'no- Asia Russia 51°00'N 36°40'E Humid Prairie 3 chernozemny Temp. 56 Voronezshkiyi Asia Russia 51°51' - 39°21' - Humid Prairie 3 52°02'N 39°47'E Temp. 57 Wolong Asia China 30°45' - 102°52' - Humid Subtropical 3 31°25'N 103°25' Temp. Regime E Mountains 58 Xishuangbanna Asia China 21°10' - 100°16' - Humid Savanna 3 22°24'N 101°55' Tropical E References:

1. China Species Information Service

(http://www.chinabiodiversity.com/search/english/readd.shtm )

2. World database on protected areas (http://www.wdpa.org/MultiSelect.aspx )

3. Man and Biosphere Faunal Database Bioinventory website (MAB)

(http://www.ice.ucdavis.edu/bioinventory/bioinventory.html )

4. UNESCO World Heritage List (http://whc.unesco.org/en/list )

5. McGrew et al. 1996

6. Kenya Wildlife Service (http://www.kws.org/ )

7. Hernandez-Fernandez and Vrba 2006

Figure 3.1 Modern Eurasian localities u sed for comparison. Sites are numbered as in table 3.1: 1:Ailaoshan; 2:Alaungdaw Kathapa NP; 4:Amudarya Zapovednik; 5:Astrakhanskiy; 6:Baimaxueshan; 8:Changbaishan; 9:Changshanerhai; 10:Chernyje Zemli Biosphere Reserve; 11:Daurskiy: 12:Daweishan; 13:Dinghu shan; 14:Doi Chiang Dao; 15:Doi Inthanon; 16:Doi Pha Chang; 17:Eravikulam; 18:Fanjingshan; 19:Fujian Wuyi; 20:Gaoligong Mountain; 22:Great Gobi; 23:Great Himalayan National Park; 24:Indravati Tiger Reserve; 25:Jiuzhaigou; 27:Kanha Tiger Reserve; 28:Kaplank yr; 29:Kodry Zapovednik; 30:Kudremukh National Park; 33:Ma Sa -Kog Ma Reserve; 35:Manas National Park; 36:Maolan Biosphere Reserve; 38:Medog Nature Reserve; 42:Nujiang; 43:Palava; 45:Repetek; 47:Royal Chitwan; 48:Sakaerat Environmental Research Station; 50: Srebarna; 51:Syunt -Khasardagh; 53:Tianmushan Biosphere Reserve; 55:Tsentral'no-chernozemny; 56:Voronezshkiyi; 57:Wolong; 58:Xishuangbanna

Figure 3.2 Modern African localities used for comparison. Site numbers are as in table 3.2. 3:Amboseli; 7:Boucle du Baoule; 21:Gombe; 26:Kalahari Gemsbok; 31:Lake Manyara; 32:Lake Nakuru; 34:Mamili; 37:Mare aux Hippopotames; 39:Moremi; 40:Mount Kenya; 41:Nouable-Ndoki; 44:Queen Elizabeth; 46:Richtersveld National Park; 49:Serengeti Ngorongoro; 52:Tai Forest; 54:Tsavo National Park

Ecotype Assignment

Modern Species

The ecological attributes of modern species were assigned based on information from the literature, especially Nowak (1999) and Kingdon (1997). If a range of body

53 masses was given in the literature, then the average body mass was used to represent the species. Dietary and substrate categories were assigned based on the definitions above.

Ancient Species

The ancient dataset included Plio-Pleistocene species from sites in East Asia

(Table 3.3) and African species from the Turkana Basin and Olduvai Gorge. Species lists for Koobi Fora and West Turkana (divided by members) and Olduvai Gorge Beds I and

II were compiled based on Turner et al. (1999). Modifications to the list of carnivores present were based on Werdelin and Lewis (2005) and Werdelin (personal communication). Modifications to the primates listed were based on Frost (2001) and

Frost and Delson (2002). Species lists for the East Asian sites were derived from the primary papers and monographs describing the sites, as well as later published additions that take into account taxonomic revisions and additional specimens found later. See

Tables 3.4 and 3.5 for the species lists from sites in East Africa and East Asia. Since it was often difficult to track down specific specimens attributed to a species at the aff. or cf. level for East African specimens, data from the fully identified specimens were used for the aff. and cf. specimens of the same potential species. The species lists include only taxa identifiable at the species or genus level due to the difficulty of assigning ecotypes to other specimens identified only at higher levels, such as that of the family. In cases where taxa were identified at the genus level and other members of that genus were present at the site or member, only one occurrence (genus and species) was recorded unless information existed specifying this as a different taxon.

Data collected from specimens and information from the literature were used to assign ancient species to ecotypes. For East Asian fossil sites, all available craniodental specimens were measured. Some specimens have been lost or were unavailable to researchers. For the East African fossil sample, fossil specimens were selected based on completeness and the amount of dental wear. Relatively unworn teeth in which the entire crown was visible were favored. Molars were favored over premolars, as molar lengths are more accurate for body mass estimation (Janis 1990). For East African species, specimens were measured from each member in which the species was present whenever possible. The measurements included maximum length, width and crown height of each cheek tooth. Additional measurements were taken as necessary to compute body mass

(e.g., Delson et al. 2000).

Sample size is a possible confounding factor for these comparisons. The three sites from the Nihewan Basin (Xiaochangliang, Donggutuo and Majuangou) all have very small samples of large mammals. Each mammal present, therefore, makes up a larger proportion of the fauna. The Nihewan fauna sensu stricto may be considered as a proxy for these sites. The Nihewan s.s . fauna was deposited during the period when the hominin sites in the basin were deposited, so it may have sampled fauna from the same region.

The Chari member from the Turkana Basin also has a small sample size. Sites with small sample sizes may not show some of the ecological diversity that sites with larger faunas do.

Table 3.3 Plio-Pleistocene East Asian sites and their dates.

Sites Dates Reference Hominin Sites Xiaochangliang, Nihewan 1.36 Ma Zhu et al. 2001, Zhu et al. 2003 Basin Donggutuo, Nihewan Basin 1.1 Ma Li and Wang 1982; Schick and Dong 1993; Li et al. 2002; Wang et al. 2005 Majuangou, Nihewan Basin 1.32-1.66 Ma Zhu et al. 2004 Yuanmou Member 4 ~1.7 Ma Li et al. 1976; Zhu et al. 2008 Gongwangling, Lantian 1.15 Ma or An and Ho 1989; Heslop et al. 2000 1.22-1.19 Ma Non-Hominin Sites Longdan 2.55 – 2.16 Ma Qiu et al. 2004 Jianshi ~2.15 Ma- Zheng et al. 2004 ~1.78 Ma Nihewan Basin sensu stricto 2-0.78 or ~1.8 Deng et al. 2008; Qiu et al. 2000 Ma Longgupo, Wushan ~1.9- ~1.8 Ma Huang et al. 1995 Linyi Late Pliocene Tang et al. 1983 or Early Pleistocene Mohui Cave Late Pliocene Wang et al. 2007 or Early Pleistocene

Table 3.4 Plio-Pleistocene African species at the Turkana Basin and Olduvai Beds I and II. Ecotype codes are as in Table 3.1. The Turkana sites are listed in order of decreasing age, starting with West Turkana, and then East Turkana (Koobi Fora). Abbreviations: Hippo: Hippopotamidae; Rhino: Rhinocerotidae; Cerco: Cercopithecidae; Deinother.: Deinotheriidae; Elephant.: Elephantidae Olduvai West Turkana East Turkana

Species Family ET Olduvai I Olduvai II Lokalalei Kalochoro Kaitio Natoo Nariokotome Upper Burgi KBS Okote Chari Carnivora Canis sp. Canidae SCAR X Canis mesomelas Canidae SCAR X Canis cf. mesomelas Canidae SCAR X X X X Canis africanus Canidae LCAR X

Olduvai West Turkana East Turkana

Species Family ET Olduvai I Olduvai II Lokalalei Kalochoro Kaitio Natoo Nariokotome Upper Burgi KBS Okote Chari Protocyon recki Canidae INS X Vulpes cf. zerda Canidae OMNSM X X Acinonyx jubatus Felidae LCAR X Acinonyx sp. Felidae LCAR X X X Caracal sp. Felidae SCAR X Cf. Caracal Felidae SCAR Dinofelis aronoki Felidae LCAR X X Dinofelis petteri Felidae LCAR Dinofelis piveteaui Felidae LCAR X Dinofelis sp. D Felidae LCAR X X Homotherium sp. Felidae LCAR X X X X X Megantereon sp. Felidae LCAR X X X X Megantereon whitei Felidae LCAR X Panthera cf. leo Felidae LCAR X X Panthera leo Felidae LCAR X X X Panthera pardus Felidae LCAR X X X Panthera pardus ? Felidae LCAR X Atilax paludinosus Herpestidae AQ X Herpestes or Herpestidae SCAR X Galerella primitivus Chasmaporthetes Hyaenidae LCAR X nitidula Crocuta crocuta Hyaenidae LCAR X X X X Crocuta dietrichi Hyaenidae LCAR X X Crocuta sp. Hyaenidae LCAR X X X Crocuta sp. nov Hyaenidae LCAR Crocuta ultra Hyaenidae LCAR X X X X Cf Hyaena sp. Hyaenidae LCAR X X X X X Hyaena cf. Hyaenidae LCAR X makapani Hyaena makapani Hyaenidae LCAR X Hyaena hyaena Hyaenidae LCAR X X Hyaena cf. hyaena Hyaenidae LCAR Aonyxini Mustelidae AQ X Lutra maculicollis Mustelidae AQ X X Cf. Torolutra Mustelidae AQ X Enhydriodon Mustelidae AQ X Lutrinae Mustelidae AQ X Mellivora Benfield Mustelidae SCAR X Mellivora sp. Mustelidae SCAR X Torolutra cf. Mustelidae AQ X X Ougandensis Torolutra sp.? Mustelidae AQ X Ursidae indet. Ursidae OMNL Pseudocivetta Viverridae LCAR X X X X

Olduvai West Turkana East Turkana

Species Family ET Olduvai I Olduvai II Lokalalei Kalochoro Kaitio Natoo Nariokotome Upper Burgi KBS Okote Chari ingens Artiodactyla Aepyceros Bovidae MXM X X X X X X X X X melampus Aepyceros Bovidae MXM X shungurae Alcelaphus sp. Bovidae GRM X Antidorcas recki Bovidae GRM X X X X X X X Antidorcas sp. Bovidae TERSM X Antilopini sp. Bovidae BRM X X Antilopini sp. B Bovidae MXM X Beatragus cf. Bovidae GRM X X X antiquus Beatragus hunteri Bovidae GRM X Beatragus sp. Bovidae GRM X Beatragus/ Bovidae GRM X X Parmularius Caprini A Bovidae MXM X X Caprini C Bovidae MXM X Caprini sp. Bovidae MXM X X Connochaetes Bovidae GRM X africanus Connochaetes sp. Bovidae GRM X X Connochaetes sp. / Bovidae GRM X X Connochaetes gentryi Connochaetes Bovidae GRM X X taurinus Damalavus cf. Bovidae GRM X makapani Damaliscus agelaius Bovidae GRM X Damaliscus eppsi Bovidae GRM X Damaliscus niro Bovidae GRM X Damaliscus sp. Bovidae GRM X X Damaliscus sp. nov. Bovidae GRM X Gazella janenschi Bovidae MXM X X Gazella Bovidae MXM X X X praethomsoni Gazella sp. Bovidae MXM X X X X Hippotragus gigas Bovidae MXL X X X X X Kobus ancystrocera Bovidae GRM X X X Kobus Bovidae GRM X X X X X X ellipsiprymnus Kobus kob Bovidae GRM X X X X X X Kobus cf. leche Bovidae GRM X X

Olduvai West Turkana East Turkana

Species Family ET Olduvai I Olduvai II Lokalalei Kalochoro Kaitio Natoo Nariokotome Upper Burgi KBS Okote Chari Kobus leche Bovidae GRM X Kobus sigmoidalis Bovidae GRM X X X X X X Kobus sp. Bovidae GRM X Kobus sp. C Bovidae GRM X Madoqua Bovidae TERSM X Megalotragus Bovidae GRM X X X X Megalotragus isaaci Bovidae GRL X X X Megalotragus Bovidae GRL X X X X X kattwinkeli Menelikia leakeyi Bovidae MXM Menelikia lyrocera Bovidae MXM X X X X X Menelikia sp. Bovidae MXM X Neotragini indet. Bovidae TERSM X Oryx sp. Bovidae MXM X X Parmularius Bovidae GRM X Parmularius Bovidae GRM X altidens Parmularius Bovidae GRM X X X angusticornis Parmularius cf. Bovidae GRM X altidens Parmularius eppsi Bovidae GRM X Parmularius Bovidae GRM X rugosus Parmularius sp. new Bovidae GRM X X Pelorovis Bovidae GRL x X X oldowayensis Pelorovis sp. Bovidae GRL X X X X Pelorovis Bovidae GRL X X X X turkanensis Rabaticeras sp. Bovidae TERSM X Redunca sp. Bovidae GRM X X X Sigmoceros Bovidae GRM X Syncerus acoelotus Bovidae GRL X X Syncerus caffer Bovidae GRL X Syncerus sp. Bovidae GRL X Taurotragus arkelli Bovidae BRL X Tragelaphini sp. Bovidae MXM X X X Tragelaphus nakuae Bovidae BRL X X X X Tragelaphus Bovidae BRM X X X scriptus Tragelaphus aff. Bovidae BRM X scriptus Tragelaphus cf. Bovidae BRM X scriptus

Olduvai West Turkana East Turkana

Species Family ET Olduvai I Olduvai II Lokalalei Kalochoro Kaitio Natoo Nariokotome Upper Burgi KBS Okote Chari Tragelaphus sp. Bovidae MXM X Tragelaphus Bovidae BRM X X X X X X X X strepsiceros Ugandax sp. Bovidae GRM X X Camelus sp. Camelidae MEGMX X X Giraffa stillei Giraffidae BRL X X X X Giraffa jumae Giraffidae BRL X X X X Giraffa pygmaea Giraffidae BRM X X X Giraffa sp. Giraffidae BRL X X Sivatherium Giraffidae MEGBR X X X X X maurusium Hexaprotodon Hippo. MEGGR X X X X X X aethiopicus Hexaprotodon Hippo. MEGGR X X X X karumensis Hexaprotodon Hippo. MEGGR X protamphibius Hippopotamus Hippo. MEGGR amphibius Hippopotamus Hippo. MEGGR X X X X X X X gorgops Kolpochoerus Suidae GRL X X X X X X X X X X limnetes Metridiochoerus Suidae GRL X X X X X X X andrewsi Metridiochoerus Suidae GRL X X X X X X X compactus Metridiochoerus Suidae GRL X X X X X hopwoodi Metridiochoerus Suidae GRL X X X X X X modestus Notochoerus scotti Suidae GRL X X X X X Potamochoerus Suidae OMNL X X porcus Perissodactyla Equinae indet. Equidae GRL X Equus cf. burchelli Equidae GRM X Equus burchelli Equidae GRM X Equus cf. grevyi Equidae GRL X Equus koobiforensis Equidae GRL X X Equus sp. Equidae GRM X X X Equus sp. Equidae GRL X X X X Equus cf. tabeti Equidae GRL X Hipparion Equidae GRM X X cornelianum Hipparion cf. Equidae GRM X X 60

Olduvai West Turkana East Turkana

Species Family ET Olduvai I Olduvai II Lokalalei Kalochoro Kaitio Natoo Nariokotome Upper Burgi KBS Okote Chari ethiopicum Hipparion Equidae GRM X X X X ethiopicum Hipparion Equidae GRL X hasumense Hipparion lybicum Equidae GRM X Hipparion sp. B Equidae GRM X X Ceratotherium Rhino. MEGGR X X X X X X X simum Diceros bicornis Rhino. MEGBR X X X X X X Primates Cercocebus/Lophoce Cerco. FRUG X X X X bus Cercopithecoides Cerco. MXM X X X kimeui Cercopithecoides Cerco. TERSM X williamsi Cercopithecus sp. Cerco. TERSM X Colobinae indet. Cerco. ARB X X X Colobus Cerco. FRUG X X X freedmanensis Colobus sp/ cf. Cerco. FRUG X X X X Procolobus Gorgopithecus Cerco. MXM X X major Lophocebus Cerco. FRUG X Papio sp. /Parapapio Cerco. ARB X X X A/Parapapio C Papionini indet. Cerco. TERSM X Paracolobus mutiwa Cerco. ARB X Parapapio Cerco. ARB X X Parapapio B Cerco. ARB X X X Rhinocolobus Cerco. ARB X X X turkanensis Theropithecus Cerco. GRM X X X X X X X X X oswaldi Homo erectus Hominidae OMNL X X X Homo ergaster Hominidae OMNL X X X X Homo habilis Hominidae OMNL X X X X X Homo rudolfensis Hominidae OMNL X X X Paranthropus boisei Hominidae MXM X X X X X X X X Proboscidea Deinotherium bozasi Deinother. MEGBR X X x X Elephas recki Elephant. MEGGR X X X X X X X X Loxodonta adaurora Elephant. MEGGR X

Olduvai West Turkana East Turkana

Species Family ET Olduvai I Olduvai II Lokalalei Kalochoro Kaitio Natoo Nariokotome Upper Burgi KBS Okote Chari Loxodonta sp. Elephant. MEGGR X

Table 3.5 Plio-Pleistocene East Asian species. Ecotype codes are as in Table 3.1. Northern China sites are on the left, with South China sites (starting with Yuanmou) on the right. Abbreviations: Chalico: Chalicotheriidae; Rhino: Rhinocerotidae; Cerco: Cercopithecidae; Gompho: Gomphotheriidae; Pliohyrac.: Pliohyracidae North China South China Non- Non- Hominin Hominin Sites Hominin Sites Sites

Species Family ET Longdan Linyi Nihewan Xiaochangliang Donggutuo Majuangou Gongwangling Yuanmou Jianshi Longgupo Mohui Carnivora Canis Canidae LCAR X brevicephalus Canis chihliensis Canidae LCAR X Canis Canidae LCAR X longdanensis Canis sp. Canidae LCAR X X Canis teilhardi Canidae LCAR X Canis variabilis Canidae OMNSM X Cuon dubius Canidae LCAR X X X Nyctereutes cf. Canidae OMNSM X X sinensis Sinicuon cf. Canidae LCAR X dubius Vulpes Canidae OMNSM X chikushanensis Vulpes sp. Canidae OMNSM X Felis microta Felidae SCAR X Felis sp. Felidae SCAR X X Felis teilhardi Felidae LCAR X X X Homotherium cf. Felidae LCAR X X crenatidens Homotherium Felidae LCAR X crendatidens Homotherium sp. Felidae LCAR X

North China South China Non- Non- Hominin Hominin Sites Hominin Sites Sites

Species Family ET Longdan Linyi Nihewan Xiaochangliang Donggutuo Majuangou Gongwangling Yuanmou Jianshi Longgupo Mohui Lynx shansius Felidae LCAR X X Megantereon Felidae LCAR X X X X nihowanensis Megantereon sp. Felidae LCAR X X Panthera cf. Felidae LCAR X pardus Panthera pardus Felidae LCAR X X Panthera sp. Felidae LCAR X Panthera cf. tigris Felidae LCAR X Panthera sp. (cf. Felidae LCAR X palaeosinensis) Panthera tigris Felidae LCAR X Panthera Felidae LCAR X palaeosinensis Sivapanthera Felidae LCAR X linxiaensis Sivapanthera Felidae LCAR X X X X pleistocaenicus Chasmaporthetes Hyaenidae LCAR X cf. ossifragus Chasmaporthetes Hyaenidae LCAR X progressus Crocuta Hyaenidae LCAR X honanensis Pachycrocuta Hyaenidae LCAR X X X X X X X X brevirostris Pachycrocuta sp. Hyaenidae LCAR X Hyaena sp. Hyaenidae LCAR X Arctonyx cf. Mustelidae OMNSM X minor Arctonyx collaris Mustelidae OMNSM X Arctonyx sp. Mustelidae OMNSM X Eirictis robusta Mustelidae OMNSM X Lutra licenti Mustelidae AQ X Lutra sp. Mustelidae AQ X Martes sp. Mustelidae OMNSM X X Martes sp. 1 Mustelidae SCAR X Martes sp. 2 Mustelidae OMNSM X Meles cf. chiai Mustelidae OMNSM X Meles cf. leucurus Mustelidae OMNL X Meles chiai Mustelidae OMNSM X Meles teilhardi Mustelidae OMNSM X

North China South China Non- Non- Hominin Hominin Sites Hominin Sites Sites

Species Family ET Longdan Linyi Nihewan Xiaochangliang Donggutuo Majuangou Gongwangling Yuanmou Jianshi Longgupo Mohui Ailuropoda Ursidae BRM X melanoleuca Ailuropoda Ursidae BRM X X microta Ailuropoda Ursidae BRM X wulingshanensis Ursus aff. Ursidae OMNL X thibetanus Ursus cf. etruscus Ursidae OMNL X X Ursus sp. Ursidae OMNL X X Ursus thibetanus Ursidae OMNL X Megaviverra Viverridae LCAR X pleistocaenica Prionodon sp. Viverridae SCAR X Viverra sp. Viverridae SCAR X Viverricula Viverridae SCAR X malaccensis Artiodactyla Antilope sp. Bovidae TERSM X Bibos sp. Bovidae MXL X Bison Bovidae MXL X palaeosinensis Bison sp. Bovidae MXL X X Bos (Bibos) sp. Bovidae MXL X Bos sp. Bovidae MXL X Bubalus sp. Bovidae MXL X Budorcas sp. Bovidae MXM X Capricornis sp. Bovidae MXM X Capricornis Bovidae MXM X jianshiensis Capricornis Bovidae MXM X sumatraensis Gazella cf. blacki Bovidae MXM X Gazella sinensis Bovidae MXM X Gazella sp. Bovidae MXM X X X X X Gazella Bovidae MXM X subgutturosa Hemibos gracilis Bovidae GRL X Leptobos Bovidae GRL X X brevicornis Leptobos sp. Bovidae GRL X Megalovis Bovidae MXL X X

North China South China Non- Non- Hominin Hominin Sites Hominin Sites Sites

Species Family ET Longdan Linyi Nihewan Xiaochangliang Donggutuo Majuangou Gongwangling Yuanmou Jianshi Longgupo Mohui guangxiensis Megalovis Bovidae MXL X piveteaui Ovis Bovidae GRM X shantungensis Spiroceros wongi Bovidae MXM X Spirocerus cf. Bovidae MXM X wongi Spirocerus peii Bovidae MXM X Paracamelus Camelidae MEGMX X gigas Paracamelus sp. Camelidae MEGMX X Axis cf. rugosus Cervidae GRM X Axis rugosus Cervidae GRM X Axis shansius Cervidae GRM X Axis sp. Cervidae BRM X Cervavitus Cervidae BRM X ultimus Cervocerus Cervidae BRM X ultimus Cervulus cf. Cervidae MXM X sinensis Cervus (Rusa) cf. Cervidae MXM X unicolor Cervus cf. fengqii Cervidae BRM X Cervus fengqii Cervidae BRM X Cervus stehlini Cervidae BRM X Cervus sp. Cervidae BRM X X X X X X Elaphodus Cervidae MXM X cephalaphus Elaphurus Cervidae GRM X bifurcatus Eostylocerus Cervidae BRM X longchuanensis Eucladoceros Cervidae BRM X boulei Euctenoceros sp. Cervidae BRM X Metacervulus Cervidae MXM X X capreolinus Metacervulus cf. Cervidae MXM X attenuatus Muntiacus Cervidae MXM X X

North China South China Non- Non- Hominin Hominin Sites Hominin Sites Sites

Species Family ET Longdan Linyi Nihewan Xiaochangliang Donggutuo Majuangou Gongwangling Yuanmou Jianshi Longgupo Mohui lacustris Muntiacus nanus Cervidae MXM X Muntiacus sp. Cervidae MXM X X Nipponicervus Cervidae BRM X longdanensis Paracervulus Cervidae BRM X attenuatus Procapreolus Cervidae BRM X stenosis Pseudaxis grayi Cervidae MXM X Pseudodama Cervidae MXM X elegans Rusa sp. Cervidae BRM X Rusa yunnanensis Cervidae BRM X X X Sinomegaceros Cervidae BRL X konwanlinensis Moschus Moschidae TERSM X moschiferus Dicoryphochoeru Suidae BRM X s ultimus Sus cf. lydekkeri Suidae OMNL X Sus liuchengensis Suidae BRM X Sus lydekkeri Suidae OMNL X Sus peii Suidae OMNL X X X Sus sp. (Linyi) Suidae OMNL X Sus sp. Suidae BRL X (Yuanmou) Sus xiaozhu Suidae BRM X X X Dorcabune Tragulidae BRM X liuchengense Perissodactyla Hesperotherium Chalico. BRL X X X X X sp. Hesperotherium Chalico. BRL X aff. sinensis Hesperotherium Chalico. BRL X sinense Equus aff. Equidae GRL X yunnanensis Equus Equidae GRL X eisenmannae Equus Equidae GRL X X X X X X

North China South China Non- Non- Hominin Hominin Sites Hominin Sites Sites

Species Family ET Longdan Linyi Nihewan Xiaochangliang Donggutuo Majuangou Gongwangling Yuanmou Jianshi Longgupo Mohui sanmeniensis Equus sp. Equidae GRL X Equus Equidae GRL X yunnanensis Hipparion Equidae GRM (Proboscidippario X n) sinense Hipparion sp. Equidae GRM X X Proboscidippario Equidae GRL X X X n sinensis Coelodonta Rhino. MEGGR X X X antiquitatis Coelodonta Rhino. MEGGR X X X nihowanensis Dicerorhinus cf. Rhino. MEGBR X merckii Dicerorhinus Rhino. MEGBR X lantianensis Dicerorhinus sp. Rhino. MEGBR X Elasmotherium Rhino. MEGGR X sp. Rhinoceros cf. Rhino. MEGBR X sinensis Rhinoceros Rhino. MEGBR X X X X sinensis Megatapirus Tapiridae BRL X augustus Tapirus Tapiridae BRL X X X sanyuanensis Tapirus sinensis Tapiridae MXL X X Primates Colobinae indet. Cerco. ARB X Macaca cf. Cerco. FRUG X anderssoni Macaca sp. Cerco. FRUG X X Macaca sp. 1 Cerco. FRUG X Macaca sp. 2 Cerco. FRUG X Megamacaca Cerco. ARB X lantianensis Paradolichopithe Cerco. FRUG X cus gansuensis Procynocephalus Cerco. FRUG X

North China South China Non- Non- Hominin Hominin Sites Hominin Sites Sites

Species Family ET Longdan Linyi Nihewan Xiaochangliang Donggutuo Majuangou Gongwangling Yuanmou Jianshi Longgupo Mohui cf. wimani Rhinopithecus sp. Cerco. ARB X Gigantopithecus Hominidae MXM X X X blacki Hominidae indet. Hominidae MXM X X X Homo sp. Hominidae OMNL X X X X X Proboscidea Elephas sp. Elephantidae MEGGR X Mammuthus Elephantidae MEGGR X trogontherii Palaeoloxodon Elephantidae MEGMX X namadicus Palaeoloxodon Elephantidae MEGMX X X sp. Paleoloxodon Elephantidae MEGMX X tokunagai Sinomastodon sp. Gompho. MEGBR X X Sinomastodon Gompho. MEGBR X yangziensis Stegodon cf. Elephantidae MEGBR X zdanskyi Stegodon Elephantidae MEGBR X elephantoides Stegodon Elephantidae MEGBR X orientalis Stegodon Elephantidae MEGBR X X X preorientalis Stegodon Elephantidae MEGBR X wushanensis Hyracoidea Postschizotherium Pliohyrac. MEGMX X chardini Lagomorpha Sericolagus Leporidae TERSM X brachypus Rodentia Hystrix Hystricidae TERSM X kiangsenensis Hystrix magna Hystricidae TERSM X X Hystrix Hystricidae TERSM X X subcristata Petaurista sp. Sciuridae ARB X

North China South China Non- Non- Hominin Hominin Sites Hominin Sites Sites

Species Family ET Longdan Linyi Nihewan Xiaochangliang Donggutuo Majuangou Gongwangling Yuanmou Jianshi Longgupo Mohui Trogontherium Castoridae AQ X Erinaceomorpha Erinaceus cf. Erinaceidae OMNSM X dealbatus

Hypsodonty

Hypsodonty was used to assess diet in extinct species. High hypsodonty indices

are correlated with diets high in abrasive materials (Janis 1988, Solounias and Dawson-

Saunders 1988, Pérez-Barbería et al. 2001). Greater crown height helps teeth withstand

abrasion by phytoliths and grit (McNaughton et al. 1985, Janis 1988, 1989, 1995,

MacFadden, 1997). Hypsodonty was measured as crown height divided by the maximum

tooth width. This measurement was made on all available molars. Hypsodonty estimates

from worn teeth were regarded as minima. The maximum hypsodonty estimates,

hypsodonty estimates from lower third molars, and the range of hypsodonty indices for

all unworn teeth were all considered and compared to the values of extant species in

order to assign dietary categories. Extant species’ hypsodonty values were from Mendoza

et al. (2002) and Mendoza and Palmqvist (2008). Hypsodonty values of extinct species

were compared with the full modern range, within their family, and within their genus.

Different families and genera may have different patterns in the relationship between

hypsodonty and diet. When the data from hypsodonty were not sufficient to distinguish

between dietary categories, or when hypsodonty was not available, information from

functional morphology (Spencer 1997) and isotopic studies (Cerling et al. 1999, Kingston

and Harrison 2007, van der Merwe et al. 2008, Zazzo et al. 2000, Harris and Cerling

2002, Harris et al. 2008) was used to assign diet.

Body Mass

Body mass was estimated using regression equations on dental dimensions from

Damuth (1990), Janis (1990), Van Valkenburgh (1990) and Delson et al. (2000). Unless otherwise specified by the regression equations, maximum length and width of molars and premolars were used. Janis’ (1990) equations for ungulates specified the use of occlusal length and width, while Delson et al.’s (2000) primate equations included separate predictions for anterior and posterior molar width. If a range of body mass estimates fell within one of the size groups, that size group was assigned. When the estimates spanned several size categories without a clear mode in one category, prediction errors were used to choose the most reliable regression equations from which to take body mass results. When living, each species would have had a range of body mass, possibly including sexual dimorphism. In choosing a value for each fossil species, some of this variation has been lost. However, the body mass ranges in the ecotype classification system are large, and each species would have had a range of body masses within its category.

Though body mass was based upon dental measurements, teeth are not weight- bearing elements, and dental dimensions are not as accurate at estimating body mass compared with postcrania (Damuth and MacFadden 1990). Regression equations for dental estimators typically have percent standard errors greater than 30% (Damuth and

MacFadden 1990). Postcranial fossil materials were not available for the Chinese fossils

studied here. Despite estimation errors for regression equations using teeth, dental

materials continue to be used in many studies because of the paucity of identifiable

postcranial remains compared with the good preservation and identifiability of dental

remains. The size and shape of teeth are under selection for properties that are not

directly related to body size. In herbivores, tooth size and shape relates to diet and

digestive physiology (Janis 1990, Damuth 1990). Damuth found that grazers have

relatively narrow molars compared to browsers, while Perissodactyla have relatively

larger teeth compared with Artiodactyla (Damuth 1990). Groups of ungulates with

selenodont and non-selenodont teeth also have different relationships between dental size and body mass (Damuth 1990, Fortelius 1990, Janis 1990).

There are various sources of error in using dental remains to estimate body mass.

Damuth (1990) distinguishes between imprecision or random error in the estimates, and systematic bias. Fossil species may differ from extant species in the relationship between the dimensions of the estimator dimension and the predicted body mass despite a high correlation in the living species used to create the regression equations (Damuth 1990,

Conroy 1987, Gingerich et al. 1982). The modern samples on which the regression is based may also introduce error (Damuth and MacFadden 1990). A larger modern sample should be a more accurate reflection of the values of the predictor dimensions for each species, which is reflected in studies with mean values having lower prediction errors

(Janis 1990, Damuth 1990). If a larger sample were measured, or if each specimen had a record of living body mass, the regression equations would be more accurate (Van

Valkenburgh 1990). There is also much variation in living body mass between members of the same species in the Carnivora (Van Valkenburgh 1990).

The predictor dimensions vary in the accuracy with which they predict body mass

(percent prediction error and percent standard error), but high accuracy dimensions, such

as head-body length, are seldom available for fossil samples (Damuth 1990). Occlusal

and maximal dimensions did not seem to be significantly different in predication

accuracy once sample sizes were taken into account (Damuth 1990). Janis (1990) found

that molar lengths were the best dental dimensions for estimating body mass in ungulates.

In particular, lower molar row length was found to be accurate even for herbivores with

enlarged individual molars (Janis 1990).

Carnivore body mass was estimated using regression equations based on the

length of the first lower molar, which was available for many of the fossil specimens,

unlike other estimator dimensions (Van Valkenburgh 1990). Lower first molar length was

chosen by Van Valkenburgh (1990) as a dental measurement that might be related to

body mass because the other teeth have different functions and levels of development in

various species, while the M 1 functions as a flesh-slicing blade (Van Valkenburgh 1990,

Van Valkenburgh 1989, Van Valkenburgh and Ruff 1987).

Van Valkenburgh (1990) found that the regression equation for all carnivores

using M 1 has a large standard error, but that the equations using family subsets for

Canidae, Felidae, Mustelidae and Ursidae are more accurate. For Felidae, M1 length is the most accurate predictor dimension (Van Valkenburgh 1990). There are also different allometric relationships between body size and first molar length for families with canids and ursids scaling negatively and felids and mustelids scaling positively (Van

Valkenburgh 1990). Regression equations for other families, such as Hyaenidae and

Viverridae, were not available. Many of these scaling differences relate to behavioral

differences between the families (Van Valkenburgh 1990).

Substrate

Substrate categories were assigned using information from the literature.

Ungulates were assumed to be primarily terrestrial. Functional morphological and

anatomical studies were used to assign substrates to extinct primates (Leakey and Leakey

1976, Jablonski 1993, Fleagle 1999, Frost 2001, Frost and Delson 2002).

The Order Carnivora: Meat-eating Carnivores, Omnivores, Herbivores or Insectivores:

Members of the order Carnivora have the potential to be included in several

dietary groups: flesh-eating carnivores, omnivores that consume a mixture of vegetation

and animal food, herbivores ( Ailuropoda ), and insectivores. Dietary classifications were made based on ecomorphological indices that were applied within families. These ecomorphological indices are based upon dental properties that relate to activities such as flesh-slicing, grinding of vegetable matter, or bone crushing (Van Valkenburgh 1988,

1989, Van Valkenburgh and Koepfli 1993, Sacco and Van Valkenburgh 2004, Friscia et al. 2007). Members of Felidae and Hyaenidae were assumed to be carnivores in this part of the study (Ewer 1973, Werdelin and Solounias 1991, Friscia et al. 2007).

Measurements from modern carnivore specimens were used to show how index values relate to dietary differences in species with known diets. The modern carnivore ratios were averaged and compared with values for fossil species using univariate and bivariate plots. The dietary categorization of fossil specimens was determined by similarity of

73 index values to those of modern species of known dietary class. Body mass was estimated using regression equations for the length of the lower first molar (Van

Valkenburgh 1990). (See discussion of body mass estimation for carnivores above.)

Canidae:

Indices proven to distinguish carnivory and omnivory in canids were used. Fossil canids were compared with modern canids that have been classified into dietary categories (Table 3.6) using Van Valkenburgh and Koepfli’s (1993) classifications that were based upon a combination of diet, specifically the percentage of vertebrate foods, and relative prey size compared with predator body size. Highly carnivorous canids consume over 70% vertebrate foods and moderately carnivorous ones eat 50-70% (Van

Valkenburgh and Koepfli 1993). Indices that were used to compare canids included relative blade length (RBL), relative grinding area in the upper and lower dentition (RGA and RUMGA), and the width of the fourth premolar (P4WL). These indices are described in more detail in chapter 6.

Ursidae:

Ursid dental remains are found in East Asian sites. Though the fossils may be related to modern Ursus thibetanus or Ailuropoda melanoleuca , they were compared with the dental adaptations of a selection of other modern ursids as well (Table 3.7). Patterns specific to the family Ursidae can help distinguish carnivorous, omnivorous and herbivorous bears (Sacco and Van Valkenburgh 2004). Carnivorous bears can be distinguished from omnivores and herbivores by the amount of their dentition devoted to grinding (RGA and RUMGA) and the relative size of their second molars (UM21), which

is also related to grinding. Carnivorous bears have less area in the dentition devoted to

grinding, in both upper and lower teeth, while herbivorous and omnivorous bears have

more area (Sacco and Van Valkenburgh 2004). Omnivorous bears could be distinguished

from the herbivorous Ailuropoda by the relative size of their grinding areas, which were not as big as in herbivores, but larger than in carnivores.

Viverridae, Mustelidae, Prionodontidae and Herpestidae:

The dietary adaptations of fossil Viverridae, Herpestidae, Prionodontidae and

Mustelidae were tested using ecomorphological indices devised by Friscia et al. (2007).

Modern specimens used are listed in Table 3.8. Fossils were also compared with the

values of the indices of a larger sample of modern taxa listed in Friscia et al. (2007).

Among the indices used were relative grinding area (RGA, RUMGA), relative size of the

upper second molar (UM21) and relative blade size (RBL). Omnivores tend to have

larger areas devoted to grinding. Insectivores tend to have larger upper second molars

compared with omnivores (Friscia et al. 2007).

Table 3.6 Modern specimens of Canidae. References: 1. Van Valkenburgh and Koepfli 1993 2. Friscia et al. 2007 Species Classifications Specimens Sex Canis adustus Omnivorous (1) USNM 182343 Male USNM 182348 Male USNM 181488 Female Canis aureus Moderately carnivorous (1) USNM 321951 Male USNM 290135 Male USNM 321952 Female USNM 290136 Male Canis latrans Moderately carnivorous (1) USNM 288195 Female USNM 064756 Female USNM 064755 Male Canis lupus Highly carnivorous (1) USNM 228267 Male USNM 150421 Female USNM 289933 Male Canis mesomelas Moderately carnivorous (1) USNM 182132 Female

Species Classifications Specimens Sex USNM 182133 Male Canis simensis Highly carnivorous (1) USNM 259450 Male Cerdocyon thous Omnivorous (1); USNM 443506 Male Carnivorous (2) USNM 443678 Female Cuon alpinus Highly carnivorous (1) USNM 083522 Male USNM 258648 Male USNM 196976 Unknown Lycaon pictus Highly carnivorous (1) USNM 181512 Male USNM 181511 Female Nyctereutes procyonoides Omnivorous(1); USNM 255532 Female omnivorous/hard-object (2) USNM 255530 Male Speothos venaticus Highly carnivorous (1); USNM 270368 Male carnivorous (2) USNM 521045 Female Urocyon cinerargenteus Omnivorous (1)/carnivorous (2) USNM 024489 Male USNM 058394 Female Vulpes bengalensis Omnivorous (1)/carnivorous (2) USNM 327159 Male USNM 257970 Female Vulpes chama Omnivorous (1)/carnivorous (2) USNM 469835 Female USNM 469834 Male Vulpes velox Highly carnivorous (1) USNM 126725 Male /carnivorous (2) USNM 188005 Female Vulpes vulpes Moderately carnivorous (1); USNM 265300 Female carnivorous (2) USNM 101488 Male

Table 3.7 Modern specimens of Ursidae. Dietary classes are after Sacco and Van Valkenburgh (2004). Species Dietary Class Specimens Sex Ailuropoda melanoleuca Herbivore USNM 579891 Female Helarctos malayanus Omnivore USNM 123139 Female USNM 198713 Male Tremarctos ornatus Omnivore USNM 194309 Male USNM 271418 Female Ursus americanus Omnivore USNM 210248 Male USNM 210298 Female Ursus arctos Carnivore USNM 233002 Male USNM 233012 Unknown Ursus maritimus Carnivore USNM 200770 Female USNM 228307 Male Ursus thibetanus Omnivore USNM 240670 Female USNM 258646 Male

Table 3.8: Mustelidae, Viverridae, Prionodontidae and Herpestidae modern samples. References: 1. Friscia et al. 2007 2. Nowak 1999 Species Family Diet Specimens Sex Atilax paludinosus Herpestidae Omnivore (2) USNM 342097 Female USNM 463357 Male Crossarchus obscurus Herpestidae Insectivore (2) USNM 481994 Female USNM 481996 Male Herpestes edwardsi Herpestidae Carnivore (1) USNM 328552 Female USNM 328553 Male

Species Family Diet Specimens Sex Herpestes ichneumon Herpestidae Carnivore (1) USNM 326263 Male USNM 326265 Female USNM 326271 Male Ichneumia albicauda Herpestidae Insectivore (1) USNM 380458 Male USNM 381467 Female Arctonyx collaris Mustelidae Omnivore (2) USNM 241159 Male USNM 259015 Female Eira barbara Mustelidae Carnivore (1) USNM 051274 Male USNM 051325 Female Enhydra lutris Mustelidae Omnivore (1) USNM 527191 Male USNM 527203 Female Gulo gulo Mustelidae Carnivore (1) USNM 089511 Female USNM 275161 Male Lutra lutra Mustelidae Omnivore (2) USNM 154157 Male USNM 333178 Female Lutra maculicollis Mustelidae Omnivore (2) USNM 220396 Unknown USNM 429136 Female Lutrogale perspicillata Mustelidae Omnivore (1) USNM 083252 Female USNM 240483 Male Meles meles Mustelidae Omnivore (1) USNM 152163 Unknown USNM 199041 Male Mellivora capensis Mustelidae Carnivore (1) USNM 270224 Male USNM 296107 Male Mustela altaica Mustelidae Carnivore (2) USNM 062110 Male USNM 198473 Female Mustela putorius Mustelidae Carnivore (1) USNM 115213 Male USNM 123629 Female Arctictis binturong Viverridae Omnivore (1) USNM 143618 Male Civettictis civetta Viverridae Carnivore (1) USNM 267187 Male USNM 270480 Female Paguma larvata Viverridae Omnivore (1) USNM 254640 Female USNM 258554 Female Paradoxurus hermaphroditus Viverridae Omnivore (1) USNM 356590 Male USNM 356998 Female Viverra zibetha Viverridae Carnivore (1) USNM 258341 Female USNM 238736 Male Viverricula indica Viverridae Carnivore (1) USNM 258266 Male USNM 294958 Female Prionodon linsang Prionodontidae Carnivore (2) USNM 144109 Male USNM 481276 Female

Species for which Data were not Available

Data were not available for some species due to specimen loss or unavailability, damage, or specimen unsuitability, when fossils did not include craniodental materials.

When listed specimens were not found in the collections, measurements recorded in monographs were used to estimate ecotypes when possible. Information from the

literature, such as functional morphological or isotopic studies, and inferences based

upon taxonomic affinity were also used when specimen measurements were not

available.

Analysis

Ecotype distributions at ancient sites were analyzed using multivariate methods.

Information about body mass, diet and substrate were used to put each species into an ecotype category (Tables 3.9 and 3.10).

Table 3.9 Ecological characteristics of East Asian fossil site faunas. The following references were used when information from the literature was used for some or all of the ecological categories, or when this information was found to support the analysis. References: 1. Nowak 1999 2. Garutt et al. 1970, Guthrie 1990; Lavarev 1977 in Boeskorov 2001; 3. Qiu et al. 2004; 4. Fleagle 1999; 5. Janis and Fortelius 1988; 6. Noskova 2001; 7. Wei and Lister 2005; 8. Dong 2006; 9. McHenry 1992a, b; 10. Van der Geer and Sondaar 2002; 11. Pan and Jablonski 1987; 12. Raia 2007; 13. Palmqvist et al. 2008; 14. Wang et al. 2007; 15. Ciochon 2009, Schwartz et al. 1995; 16. Sukumar 2003, Haynes 1991; 17. Schwartz et al. 1995 *Based on analysis of other fossil members of the genus/family

Species Family Body Diet Substrate Ecotype Refs. Mass (kg) Ailuropoda melanoleuca Ursidae 93.9 Browser Terrestrial BRM Ailuropoda microta Ursidae 86.9 Browser Terrestrial BRM Ailuropoda Ursidae 89.8-101 Browser Terrestrial BRM wulingshanensis Antilope sp. Bovidae 18.2 Grazer Terrestrial TERSM Arctonyx cf. minor Mustelidae 8.49 Omnivore Terrestrial OMNSM Arctonyx collaris Mustelidae 8.49 Omnivore Terrestrial OMNSM Arctonyx sp. Mustelidae 8.49 Omnivore Terrestrial OMNSM Axis cf. rugosus Cervidae 36-50 Grazer Terrestrial GRM 1 Axis rugosus Cervidae 36-50 Grazer Terrestrial GRM 1 Axis shansius Cervidae 36-50 Grazer Terrestrial GRM 1 Axis sp. Cervidae 188 Browser Terrestrial BRM Bibos sp. Bovidae 335.6- Mixed Terrestrial MXL 590.3 Bison palaeosinensis Bovidae 411.03 Mixed Terrestrial MXL Bison sp. Bovidae 583.9 Mixed Terrestrial MXL Bos (Bibos) sp. Bovidae 244-419 Mixed Terrestrial MXL

Species Family Body Diet Substrate Ecotype Refs. Mass (kg) Bos sp. Bovidae 325-446 Mixed Terrestrial MXL Bubalus sp. Bovidae 700-1200 Mixed Terrestrial MXL 1 Budorcas sp. Bovidae 173-538.6 Mixed Terrestrial MXM Canis brevicephalus Canidae 29.6 Carnivore Terrestrial LCAR Canis chihliensis Canidae 19.8-22.8 Carnivore Terrestrial LCAR Canis longdanensis Canidae 18.9-21.6 Carnivore Terrestrial LCAR Canis sp. Canidae Carnivore Terrestrial LCAR Canis teilhardi Canidae 20.5-26.8 Carnivore Terrestrial LCAR Canis variabilis Canidae 17-21 Omnivore Terrestrial OMNSM Capricornis jianshiensis Bovidae 88.8 - Mixed Terrestrial MXM 231.5 Capricornis sp. Bovidae 127-200 Mixed Terrestrial MXM Capricornis sumatraensis Bovidae 147.6 Mixed Terrestrial MXM qinlingensis Cervavitus ultimus Cervidae 91.7 Browser Terrestrial BRM Cervocerus ultimus Cervidae 44-62 Browser Terrestrial BRM Cervulus cf. sinensis Cervidae 22.5 Mixed Terrestrial MXM 1 Cervus (Rusa) cf. Cervidae 105-312.7 Mixed Terrestrial MXM unicolor Cervus cf. fengqii Cervidae 77.5-172.4 Browser Terrestrial BRM Cervus fengqii Cervidae 63-82 Browser Terrestrial BRM Cervus stehlini Cervidae Browser Terrestrial BRM 1 * Cervus sp. Cervidae 41-60 Browser Terrestrial BRM 1 * Chasmaporthetes cf. Hyaenidae 90.9 Carnivore Terrestrial LCAR ossifragus Chasmaporthetes Hyaenidae 60.5-76 Carnivore Terrestrial LCAR progressus Coelodonta antiquitatis Rhinocerotidae 1648-4052 Grazer Terrestrial MEGGR 2 Coelodonta nihowanensis Rhinocerotidae 1648-4052 Grazer Terrestrial MEGGR 3 Coelodonta sp. Rhinocerotidae 1484.23 Grazer Terrestrial MEGGR 2 Colobinae indet. Cercopithecidae 37-40 Browser Arboreal ARB 4 Crocuta honanensis Hyaenidae 69.6-83.5 Carnivore Terrestrial LCAR Cuon dubius Canidae 28.2 Carnivore Terrestrial LCAR Dicerorhinus cf. merckii Rhinocerotidae 1152 Browser Terrestrial MEGBR * Dicerorhinus Rhinocerotidae 2377-2684 Browser Terrestrial MEGBR * lantianensis Dicerorhinus sp. Rhinocerotidae 874.9- Browser Terrestrial MEGBR 2067.9 Dicoryphochoerus Suidae 88.9-93.4 Browser Terrestrial BRM ultimus Dorcabune liuchengense Tragulidae 67-88 Browser Terrestrial BRM Eirictis robusta Mustelidae 13 Omnivore Terrestrial OMNSM Elaphodus cephalaphus Cervidae 20-22 Mixed Terrestrial MXM Elaphurus bifurcatus Cervidae 159-214 Grazer Terrestrial GRM 1 Elasmotherium sp. Rhinocerotidae 5000 Grazer Terrestrial MEGGR 6 Elephas sp. Elephantidae Grazer Terrestrial MEGGR Eostylocerus Cervidae Browser Terrestrial BRM longchuanensis Equus aff. yunnanensis Equidae 278.6- Grazer Terrestrial GRL 705.3

Species Family Body Diet Substrate Ecotype Refs. Mass (kg) Equus eisenmannae Equidae 378.6- Grazer Terrestrial GRL 803.4 Equus sanmeniensis Equidae 277-811 Grazer Terrestrial GRL Equus sp. Equidae 301-727.4 Grazer Terrestrial GRL Equus yunnanensis Equidae 273-619 Grazer Terrestrial GRL Erinaceus cf. dealbatus Erinaceidae 1 Omnivore Terrestrial OMNSM 1 Eucladoceros boulei Cervidae 256 Browser Terrestrial BRM 8 Euctenoceros sp. Cervidae Browser Terrestrial BRM * Felis microta Felidae 7 Carnivore SCAR Felis sp. Felidae 12.49-23.6 Carnivore SCAR Felis teilhardi Felidae 21.6-47 Carnivore LCAR Gazella cf. blacki Bovidae 29-47.6 Mixed Terrestrial MXM 1 Gazella sinensis Bovidae 33-54.8 Mixed Terrestrial MXM 1 Gazella sp. Bovidae 29-54.8 Mixed Terrestrial MXM 1 Gazella subgutturosa Bovidae 29-54.8 Mixed Terrestrial MXM 1 Gigantopithecus blacki Hominidae 225 Browser Terrestrial MXM 4 Hemibos gracilis Bovidae 389.9 Grazer Terrestrial GRL Hesperotherium sp. Chalicotheriidae 340.6- Browser Terrestrial BRL 575.1 Hesperotherium aff. Chalicotheriidae 193.7- Browser Terrestrial BRL sinensis 1723 Hesperotherium sinense Chalicotheriidae 887.6 Browser Terrestrial BRL Hipparion Equidae 284 Grazer Terrestrial GRM * (Proboscidipparion) sinense Hipparion sp. Equidae 284 Grazer Terrestrial GRM Hominidae indet. Hominidae 26-76 kg Herbivore MXM 4, 15 Homotherium cf. Felidae 143.6- Carnivore Terrestrial LCAR crenatidens 249.2 Homotherium Felidae 143.6- Carnivore Terrestrial LCAR crenatidens 249.2 Homotherium sp. Felidae 143.6- Carnivore Terrestrial LCAR 249.2 Homo sp. Hominidae 52-62 Omnivore Terrestrial OMNL 9 “Hyaena” sp. (probably Hyaenidae 88.4-150.6 Carnivore Terrestrial LCAR Pachycrocuta ) Hystrix kiangsenensis Hystricidae 10-30 Herbivore Terrestrial TERSM 1 Hystrix magna Hystricidae 10-30 Herbivore Terrestrial TERSM 1 Hystrix subcristata Hystricidae 10-30 Herbivore Terrestrial TERSM 1 Leptobos brevicornis Bovidae 181-435.7 Grazer Terrestrial GRL Leptobos sp. Bovidae 181-1016 Grazer Terrestrial GRL Lutra licenti Mustelidae 6.8 Omnivore Aquatic AQ Lutra sp. Mustelidae 6.8 Omnivore Aquatic AQ * Lynx shansius Felidae 27.1-30.1 Carnivore LCAR Macaca cf. anderssoni Cercopithecidae 25.4 Frugivore FRUG 4 Macaca sp. Cercopithecidae 13.9-15.5 Frugivore FRUG 4 Macaca sp. 1 Cercopithecidae Frugivore FRUG 4 Macaca sp. 2 Cercopithecidae 17 Frugivore FRUG 4 Mammuthus trogontherii Elephantidae Grazer Terrestrial MEGGR 7 Martes sp. Mustelidae 1 Omnivore OMNSM

Species Family Body Diet Substrate Ecotype Refs. Mass (kg) Martes sp. 1 Mustelidae 1.36 Carnivore SCAR Martes sp. 2 Mustelidae 12.1-13 Omnivore OMNSM Megalovis guangxiensis Bovidae 381.7- Mixed Terrestrial MXL 511.5 Megalovis piveteaui Bovidae 546.6 Mixed Terrestrial MXL Megamacaca Cercopithecidae 29.9 Vegetation Arboreal ARB 1 lantianensis (Rhinopithecus ) Megantereon Felidae 74.1-114.7 Carnivore Terrestrial LCAR nihowanensis Megantereon sp. Felidae 74-114.7 Carnivore Terrestrial LCAR Megatapirus augustus Tapiridae 431.7 Browser Terrestrial BRL Megaviverra Viverridae 30-45 Carnivore LCAR pleistocaenica Meles cf. chiai Mustelidae 9-20 Omnivore Terrestrial OMNSM Meles cf. leucurus Mustelidae 27.6 Omnivore Terrestrial OMNL Meles chiai Mustelidae 13.7-14.3 Omnivore Terrestrial OMNSM Meles teilhardi Mustelidae 16.6 Omnivore Terrestrial OMNSM Metacervulus Cervidae 16.3-49 Mixed Terrestrial MXM capreolinus Metacervulus cf. Cervidae Terrestrial MXM * attenuatus Moschus moschiferus Moschidae 9.5 Mixed Terrestrial TERSM Muntiacus lacustris Cervidae Mixed Terrestrial MXM * Muntiacus nanus Cervidae Mixed Terrestrial MXM * Muntiacus sp. Cervidae 12.6-45.6 Mixed Terrestrial MXM Nipponicervus Cervidae 206.3 Browser Terrestrial BRM longdanensis Nyctereutes cf. sinensis Canidae 4-10 Omnivore Terrestrial OMNSM Nyctereutes sinensis Canidae 4-10 Omnivore Terrestrial OMNSM Ovis shantungensis Bovidae 139 Grazer Terrestrial GRM 1 Pachycrocuta Hyaenidae 88.4-150.6 Carnivore Terrestrial LCAR brevirostris Pachycrocuta sp. Hyaenidae 88.4-150.6 Carnivore Terrestrial LCAR Palaeoloxodon Elephantidae Mixed Terrestrial MEGMX 16 namadicus Palaeoloxodon sp. Elephantidae Mixed Terrestrial MEGMX 16 Paleoloxodon tokunagai Elephantidae Mixed Terrestrial MEGMX 16 Panthera cf. pardus Felidae 71-108 Carnivore LCAR Panthera pardus Felidae 53.-120.1 Carnivore LCAR Panthera sp. Felidae Carnivore LCAR Panthera cf. tigris Felidae Carnivore LCAR Panthera palaeosinensis Felidae Carnivore LCAR Panthera sp. (cf. Felidae Carnivore LCAR palaeosinensis) Panthera tigris Felidae Carnivore LCAR Paracamelus gigas Camelidae 1560.7- Mixed Terrestrial MEGMX 3568.4 Paracamelus sp. Camelidae 1560.7- Mixed Terrestrial MEGMX 3568.4 Paracervulus attenuatus Cervidae Terrestrial BRM 1* 81

Species Family Body Diet Substrate Ecotype Refs. Mass (kg) Paradolichopithecus Cercopithecidae 27-31.3 Frugivore Terrestrial FRUG 10 gansuensis Petaurista sp. Sciuridae 1-2.5 Vegetation Arboreal ARB 1 Postschizotherium Pliohyracidae 1248 Vegetation Terrestrial MEGMX 17 chardini Prionodon sp. Prionodontidae 1.1 Carnivore SCAR Proboscidipparion Equidae 662.3- Grazer Terrestrial GRL sinensis 823.69 Procapreolus stenosis Cervidae 73.8 Browser Terrestrial BRM Procynocephalus cf. Cercopithecidae 37 Frugivore Terrestrial FRUG 11 wimani Pseudaxis grayi Cervidae 51-158.7 Mixed Terrestrial MXM Pseudodama elegans Cervidae 88-111 Mixed Terrestrial MXM 12,13 Rhinoceros cf. sinensis Rhinocerotidae 1192-2876 Browser Terrestrial MEGBR 14. Rhinoceros sinensis Rhinocerotidae 487.9- Browser Terrestrial 4364 Rhinoceros sp. Rhinocerotidae Browser Terrestrial Rhinopithecus sp. Cercopithecidae 17-21.5 Vegetation Arboreal ARB 1 Rusa sp. Cervidae 36-260 Browser Terrestrial BRM 1 Rusa yunnanensis Cervidae 212-314.9 Browser Terrestrial BRM Sericolagus brachypus Leporidae 1 Vegetation Terrestrial TERSM 1 Sinicuon cf. dubius Canidae 28.2 Carnivore Terrestrial LCAR Sinomastodon sp. Gomphotheriidae Browser Terrestrial MEGBR Sinomastodon Gomphotheriidae Browser Terrestrial MEGBR yangziensis Sinomegaceros Cervidae 343.3 Browser Terrestrial BRL konwanlinensis Sivapanthera linxiaensis Felidae 60-103.5 Carnivore Terrestrial LCAR Sivapanthera Felidae 122.2 Carnivore Terrestrial LCAR pleistocaenicus Spiroceros wongi Bovidae 196.6-254 Mixed Terrestrial MXM Spirocerus cf. wongi Bovidae 105.4- Mixed Terrestrial MXM 207.6 Spirocerus peii Bovidae 55.9-196 Mixed Terrestrial MXM Stegodon cf. zdanskyi Elephantidae Browser Terrestrial MEGBR Stegodon elephantoides Elephantidae Browser Terrestrial MEGBR Stegodon orientalis Elephantidae Browser Terrestrial MEGBR Stegodon preorientalis Elephantidae Browser Terrestrial MEGBR Stegodon sp. Elephantidae Browser Terrestrial MEGBR Stegodon wushanensis Elephantidae Browser Terrestrial MEGBR Sus cf. lydekkeri Suidae Omnivore Terrestrial OMNL 1 Sus liuchengensis Suidae 104-350.8 Browser Terrestrial BRM Sus lydekkeri Suidae 85.8 Omnivore Terrestrial OMNL 1 Sus peii Suidae 53.9-671 Omnivore Terrestrial OMNL 1 Sus sp. Suidae Omnivore Terrestrial OMNL * Sus sp. (Yuanmou) Suidae 594-792 Browser Terrestrial BRL Sus xiaozhu Suidae 10.5-99 Browser Terrestrial BRM Tapirus sanyuanensis Tapiridae 150.8-598 browser Terrestrial BRL Tapirus sinensis Tapiridae 412.7-545 Mixed Terrestrial MXL Trogontherium sp. Castoridae Aquatic AQ 1

Species Family Body Diet Substrate Ecotype Refs. Mass (kg) Ursus aff. thibetanus Ursidae 78 Omnivore Terrestrial OMNL Ursus cf. etruscus Ursidae 65-150 Omnivore Terrestrial OMNL Ursus sp. Ursidae 75.7-79.7 Omnivore Terrestrial OMNL Ursus thibetanus Ursidae 78 Omnivore Terrestrial OMNL Viverra sp. Viverridae 11.5 Carnivore Terrestrial SCAR Viverricula malaccensis Viverridae 3.4 Carnivore Terrestrial SCAR 1 Vulpes chikushanensis Canidae 7.17 Omnivore Terrestrial OMNSM Vulpes sp. Canidae Omnivore Terrestrial OMNSM *

Table 3.10 Summary of ecological information and ecotype assignment for African fossil species. Abbreviations: Rhino.: Rhinocerotidae; Cerco.: Cercopithecidae; Hippo.: Hippopotamidae. The following references were used when information from the literature was used for some or all of the ecological categories, or when this information was found to support the analysis. References: 1. Nowak 1999; 2. Harris et al. 1988; 3. Spencer 1997; 4. Lee- Thorpe and van der Merwe 1994; 5. Van der Merwe et al. 2003; 6: Fleagle 1999; 7: Benefit 2000; 8. Frost and Delson 2002; 9. Leakey 1982; 10. Fourie et al. 2008; 11. Elton 2001; 12: Leakey 1976; 13. Bobe and Behrensmeyer 2004; 14. Gentry and Gentry 1978a, b 15. Kingdon 1997; 16. Christiansen 2004; 17. Cerling et al. 1999; 18. Eisenmann 1983; 19. Kingston and Harrison 2007; 20. Harrison and Su 2004; 22. Harris et al. 2008; 23. Harris 1991; 24. Zazzo 2000; 25. Sponheimer and Lee-Thorpe 1999; 26. Ungar et al. 2006; 27. Blumenschine and Masao 1991; 28. McHenry 1992a, b; 29. van der Merwe 2008; 30. Harris and Cerling 2002; 31. Gagnon and Chew 1999; 32. Dunbar 1992; 33. Werdelin and Lewis 2005

Species Family Body Diet Substrate Ecotype Refs. Mass (kg) Acinonyx jubatus Felidae 21-72 Carnivore Terrestrial LCAR 1 Acinonyx sp. Felidae 21-72 Carnivore Terrestrial LCAR 1 Aepyceros melampus Bovidae 36.7- Mixed Terrestrial MXM 82 Aepyceros shungurae Bovidae 52- Mixed Terrestrial MXM 2 80.1 Alcelaphus sp. Bovidae 100- Grazer Terrestrial GRM 225 Antidorcas recki Bovidae 23.5- Mixed Terrestrial GRM 3; 4; 62.5 5 Antidorcas sp. Bovidae 7.6 Mixed Terrestrial TERSM Antilopini sp. Bovidae 28-52 Browser Terrestrial BRM 1 Antilopini sp. B Bovidae 278.7 Mixed Terrestrial MXM Aonyxini Mustelidae Aquatic AQ Atilax paludinosus Herpestidae 4.2-4.5 Omnivore/ Aquatic AQ 1 Insectivore Beatragus cf. Bovidae 187.3- Grazer Terrestrial GRM

Species Family Body Diet Substrate Ecotype Refs. Mass (kg) antiquus 223.8 Beatragus hunteri Bovidae Terrestrial GRM Beatragus sp. Bovidae Terrestrial GRM Beatragus/Parmulari Bovidae 89- Grazer Terrestrial GRM us 205.3 Camelus sp. Camelidae 1407. Mixed Terrestrial MEGMX 1 5- 1647 Canis cf. mesomelas Canidae 10.5- Carnivore Terrestrial SCAR 10.7 Canis mesomelas Canidae 11.6- Carnivore Terrestrial SCAR 13.3 Canis sp. Canidae Carnivore Terrestrial SCAR Canis ? Canidae Carnivore Terrestrial SCAR Canis africanus Canidae 21.9- Carnivore Terrestrial LCAR 24.8 Caprini A Bovidae Terrestrial MXM Caprini C Bovidae Terrestrial MXM Caprini sp. Bovidae 40.5- Mixed Terrestrial MXM 72.2 Ceratotherium simum Rhino. 2592 Grazer Terrestrial MEGGR 4018 Cercocebus/Lophoce Cerco. 15.5- Frugivore Arboreal? FRUG 6; 7 bus 21.2 Cercopithecoides Cerco. 20.9- Mixed Terrestrial MXM 8; 9 kimeui 32.6 Cercopithecoides sp. Cerco. 7.2 Mixed Terrestrial TERSM 8 Cercopithecoides Cerco. 18.1- Mixed Terrestrial TERSM 7; 10; williamsi 21.6 11 Cercopithecus sp. Cerco. 3.5 Browser Terrestrial TERSM cf. Lophocebus Cerco. 14.4 Frugivore Arboreal? FRUG 1 Chasmaporthetes Hyaenidae 86.2 Carnivore Terrestrial LCAR nitidula Colobinae indet. Cerco. 9.8- Browser Arboreal/ ARB 1; 12; 33.9 Terrestrial Colobus Cerco. 9.9- Frugivore Arboreal/ FRUG 7 freedmanensis 10.9 Terrestrial Colobus sp/cf. Cerco. Frugivore Arboreal/ FRUG 7 Procolobus Terrestrial Connochaetes Bovidae 181 Grazer Terrestrial GRM africanus Connochaetes sp. Bovidae Terrestrial GRM Connochaetes sp./ Bovidae 209.4- Grazer Terrestrial GRM Connochaeates 317 gentryi Connochaetes Bovidae 140- Grazer Terrestrial GRM 1 taurinus 290 Crocuta dietrichi Hyaenidae 92- Carnivore Terrestrial LCAR 108.8 Crocuta dietrichi? Hyaenidae Carnivore Terrestrial LCAR Crocuta ultra Hyaenidae 97.3- Carnivore Terrestrial LCAR 84

Species Family Body Diet Substrate Ecotype Refs. Mass (kg) 102.2 Crocuta cf. ultra Hyaenidae 97.3- Carnivore Terrestrial LCAR 102.2 Crocuta? Hyaenidae Carnivore Terrestrial LCAR Crocuta sp. Hyaenidae Carnivore Terrestrial LCAR Crocuta sp. nov. Hyaenidae 194.9 Carnivore Terrestrial LCAR Crocuta crocuta Hyaenidae 82.8- Carnivore Terrestrial LCAR 102.9 Damalavus cf. Bovidae Terrestrial GRM makapani Damaliscus agelaius Bovidae 137.9 Grazer Terrestrial GRM Damaliscus eppsi Bovidae 288.9- Grazer Terrestrial GRM 13 305.2 Damaliscus niro Bovidae 68- Grazer Terrestrial GRM 14, 155 15 Damaliscus sp. Bovidae Terrestrial GRM Damaliscus sp. nov Bovidae 67.9 Grazer Terrestrial GRM 13 Deinotherium bozasi Deinotheriidae 12000 Browser Terrestrial MEGBR 16, - 17 17000 Diceros bicornis Rhino. 2374. browser Terrestrial MEGBR 1- 3827 Cf. Dinofelis Felidae Carnivore LCAR Dinofelis aronoki Felidae 191.7 Carnivore LCAR Dinofelis petteri Felidae Carnivore LCAR Dinofelis piveteaui Felidae 130.4 Carnivore LCAR

Dinofelis sp. D Felidae Carnivore LCAR Dinofelis sp. Felidae Carnivore LCAR Elephas recki Elephantidae 3870- Grazer Terrestrial MEGGR 16, 10497 17 Enhydriodon Mustelidae 72.1 Omnivore Aquatic AQ Enhydriodon? Mustelidae 72.1 Omnivore Aquatic AQ Equinae indet. Equidae Grazer Terrestrial GRL Equus cf. burchelli Equidae 175- Grazer Terrestrial GRM 15 385 Equus burchelli Equidae 175- Grazer Terrestrial GRM 15 385 Equus cf. grevyi Equidae 385.2- Grazer Terrestrial GRL 665.8 Equus koobiforensis Equidae 546.2 Grazer Terrestrial GRL Equus sp. (KF) Equidae 291.3- Grazer Terrestrial GRM 18 303.3 Equus sp. (WT) Equidae 299- Grazer Terrestrial GRL 2 677.7 Equus cf. tabeti Equidae 244.5- Grazer Terrestrial GRL 502 Cf. Caracal Felidae 14.1 Carnivore Terrestrial SCAR 1 Caracal sp. Felidae 14.1 Carnivore Terrestrial SCAR 1

Species Family Body Diet Substrate Ecotype Refs. Mass (kg) Gazella janenschi Bovidae 20-50 Mixed Terrestrial MXM 19; 20 Gazella Bovidae 10.3- Mixed Terrestrial MXM 2 praethomsoni 56.9 Gazella sp. Bovidae 13- Mixed Terrestrial MXM 125.6 Giraffa jumae Giraffidae 705- Browser Terrestrial BRL 863 Giraffa pygmaea Giraffidae 114.1- Browser Terrestrial BRM 318 Giraffa sp. Giraffidae 310.5 Browser Terrestrial BRL 2 Giraffa stillei Giraffidae 338.8- browser Terrestrial BRL 865.2 Gorgopithecus major Cerco. 41 Mixed Terrestrial MXM 6 Herpestes or Herpestidae 1.4 Carnivore Terrestrial SCAR 1,20 Galerella primitivus Hexaprotodon Hippo. 438.3- Grazer Terrestrial/ MEGGR 22 aethiopicus 1553. Aquatic 6

Hexaprotodon Hippo. 1536. Grazer Terrestrial/ MEGGR 22 karumensis 4- Aquatic 4596. 3

Hexaprotodon Hippo. 1795. Grazer Terrestrial/ MEGGR 22, protamphibius 6 Aquatic 23 Hipparion Equidae 277 Grazer Terrestrial GRM cornelianum Hipparion cf. Equidae 95.1- Grazer Terrestrial GRM ethiopicum 286.8 Hipparion Equidae 231.1- Grazer Terrestrial GRM ethiopicum 231.9 Hipparion hasumense Equidae Grazer Terrestrial GRL 24 Hipparion lybicum Equidae Grazer Terrestrial GRM 25; Hipparion sp. B Equidae 130- Grazer Terrestrial GRM 188.5 Hippopotamus Hippo. 1000- Grazer Terrestrial/ MEGGR amphibius 4500 Aquatic Hippopotamus Hippo. 1333- Grazer Terrestrial/ MEGGR 22 gorgops 6350. Aquatic 9 Hippotragus gigas Bovidae 322.2 Mixed Terrestrial MXL Homo erectus Hominidae Omnivore Terrestrial OMNL 26, 27 Homo ergaster Hominidae 52-62 Omnivore Terrestrial OMNL 28 Homo habilis Hominidae 32-52 Omnivore Terrestrial OMNL 28, 29, 26 Homo rudolfensis Hominidae Omnivore Terrestrial OMNL 26 Homo sp. Hominidae Omnivore Terrestrial OMNL

Species Family Body Diet Substrate Ecotype Refs. Mass (kg) Homotherium sp. Felidae 374- Carnivore Terrestrial LCAR 388 Homotherium? Felidae Carnivore Terrestrial LCAR Hyaena sp. Hyaenidae 44.5- Carnivore Terrestrial LCAR 64.9 Hyaena cf. makapani Hyaenidae Carnivore Terrestrial LCAR Hyaena makapani Hyaenidae 32.2 Carnivore Terrestrial LCAR Hyaena sp.? Hyaenidae 29.2 Carnivore Terrestrial LCAR Hyaena hyaena Hyaenidae Carnivore Terrestrial LCAR Hyaena cf. hyaena Hyaenidae Carnivore Terrestrial LCAR Kobus ancystrocera Bovidae Grazer Terrestrial GRM Kobus ellipsiprymnus Bovidae 160- Grazer Terrestrial GRM 15 300 Kobus kob Bovidae 64.6- Grazer Terrestrial GRM 214.2 Kobus cf. leche Bovidae 60- Grazer Terrestrial GRM 15 130 Kobus leche Bovidae 60- Grazer Terrestrial GRM 15 130 Kobus sigmoidalis Bovidae 108.3- Grazer Terrestrial GRM 3 149.7 Kobus sp. Bovidae Grazer Terrestrial GRM Kobus sp. C Bovidae Grazer Terrestrial GRM Kolpochoerus Suidae 93.3- Grazer Terrestrial GRL 30 limnetes 655.6 Lophocebus Cerco. 12.7 Frugivore Arboreal FRUG 6 Loxodonta adaurora Elephantidae Grazer Terrestrial MEGGR 17 Loxodonta sp. Elephantidae Grazer Terrestrial MEGGR 17 Lutra maculicollis Mustelidae 5-14 Carnivore Aquatic AQ 1 Lutrinae Mustelidae omnivore Aquatic AQ Madoqua Bovidae 4.3-5 Browser Terrestrial TERSM Megalotragus Bovidae 242 Grazer Terrestrial GRM Megalotragus isaaci Bovidae 301.1- Grazer Terrestrial GRL 847.7 Megalotragus Bovidae 588.4 Grazer Terrestrial GRL 14 kattwinkeli Megantereon sp. Felidae 37.6 Carnivore LCAR Megantereon whitei Felidae 37.6 Carnivore LCAR Megantereon? Felidae Carnivore LCAR Mellivora benfieldi Mustelidae 6.12 Carnivore Terrestrial SCAR Mellivora sp. Mustelidae Carnivore Terrestrial SCAR Menelikia leakeyi Bovidae 169.1 Mixed Terrestrial MXM 3 Menelikia lyrocera Bovidae 91- Mixed Terrestrial MXM 3 163.6 Menelikia sp. Bovidae Mixed Terrestrial MXM 3 Metridiochoerus Suidae 167- Grazer Terrestrial GRL 30 andrewsi 524.6 Metridiochoerus Suidae 472 Grazer Terrestrial GRL compactus Metridiochoerus Suidae 714.1- Grazer Terrestrial GRL

Species Family Body Diet Substrate Ecotype Refs. Mass (kg) hopwoodi 1761. 1 Metridiochoerus Suidae 337.4- Grazer Terrestrial GRL modestus 570.1 Neotragini indet. Bovidae 3-21 Browser Terrestrial TERSM 31 Notochoerus scotti Suidae 603.7- Grazer Terrestrial GRL 30 1275 Oryx sp. Bovidae 127.3- Mixed Terrestrial MXM 231.3 Panthera (lion-sized) Felidae Carnivore Terrestrial LCAR Panthera (lion- Felidae Carnivore Terrestrial LCAR sized)? Panthera cf. leo Felidae Carnivore Terrestrial LCAR Panthera leo Felidae 177.3 Carnivore Terrestrial LCAR 1 -183.8 Panthera sp. Felidae Carnivore Terrestrial LCAR Panthera? Felidae Carnivore Terrestrial LCAR Panthera cf. pardus Felidae 28-90 Carnivore LCAR 1 Panthera pardus Felidae 28-90 Carnivore LCAR 1 Panthera pardus? Felidae Carnivore LCAR Papio sp./Parapapio Cerco. 22.8- Mixed Arboreal ARB 32 A/Parapapio C 37.7 Papionini indet. Cerco. 18.9 Mixed Terrestrial TERSM 7 Paracolobus mutiwa Cerco. 52.7 Browser Arboreal ARB 7 Paranthropus boisei Hominidae 34-49 Mixed Terrestrial MXM 28 Parapapio Cerco. 26.99 Mixed Arboreal/ ARB 8, 10 Terrestrial Parapapio B Cerco. 21.8 Mixed Arboreal/ ARB 8, 10 Terrestrial Parmularius Bovidae 88 Grazer Terrestrial GRM Parmularius altidens Bovidae Grazer Terrestrial GRM 3 Parmularius Bovidae 100- Grazer Terrestrial GRM 14, angusticornis 225 23 Parmularius cf. Bovidae 95.8 Grazer Terrestrial GRM 3 altidens Parmularius eppsi Bovidae Grazer Terrestrial GRM 3 Parmularius rugosus Bovidae Grazer Terrestrial GRM 14 Parmularius sp. new Bovidae Grazer Terrestrial GRM * Pelorovis Bovidae 281- Grazer Terrestrial GRL oldowayensis 641.3 Pelorovis sp. Bovidae Grazer Terrestrial GRL Pelorovis turkanensis Bovidae 302- Grazer Terrestrial GRL 1038. 8 Potamochoerus Suidae 46- Omnivore Terrestrial OMNL 1 porcus 130 Prototocyon recki Canidae Terrestrial INS 15 Rabaticeras sp. Bovidae 7-16 Browser Terrestrial TERSM 23 Redunca sp. Bovidae 19-95 Grazer Terrestrial GRM Rhinocolobus Cerco. 29.8- Frugivore/ Arboreal ARB

Species Family Body Diet Substrate Ecotype Refs. Mass (kg) turkanensis 41.5 folivore Pseudocivetta ingens Viverridae 35.8- Carnivore LCAR 42 Pseudocivetta sp. Viverridae Carnivore LCAR Pseudocivetta sp.? Viverridae Carnivore LCAR Sigmoceros sp. Bovidae 157- Grazer Terrestrial GRM 1 204 Sivatherium Giraffidae 1680- Browser Terrestrial MEGBR maurusium 5390 Syncerus acoelotus Bovidae 497.4 Grazer Terrestrial GRL 1 Syncerus caffer Bovidae 300- Grazer Terrestrial GRL 1 900 Syncerus sp. Bovidae Grazer Terrestrial GRL Taurotragus arkelli Bovidae 400- Browser Terrestrial BRL 1 1000 Theropithecus Cerco. 21.5- Grazer Terrestrial GRM 5, 6 oswaldi 80.8 Theropithecus sp. Cerco. 63.1 Grazer Terrestrial GRM 6 Cf. Torolutra Mustelidae 19.97 Omnivore Aquatic AQ Torolutra cf. T. Mustelidae 16 Carnivore Aquatic AQ ougandensis Torolutra Mustelidae 19.97 Omnivore Aquatic AQ Torolutra sp.? Mustelidae Omnivore Aquatic AQ Tragelaphini sp. Bovidae 71.9- Mixed Terrestrial MXM 578.4 Tragelaphus nakuae Bovidae 574.3- Browser Terrestrial BRL 598.3 Tragelaphus scriptus Bovidae 69.8- Browser Terrestrial BRM 111.6 Tragelaphus cf. Bovidae Browser Terrestrial BRM scriptus Tragelaphus aff. Bovidae Browser Terrestrial BRM scriptus Tragelaphus sp. Bovidae Mixed Terrestrial MXM 1 Tragelaphus Bovidae 168- Browser Terrestrial BRM strepsiceros 285.9 Ugandax sp. Bovidae 268.2 Grazer Terrestrial GRM Ursidae indet. Ursidae Omnivore Terrestrial OMNL 33 Vulpes cf. zerda Canidae Omnivore Terrestrial OMNSM 1

A matrix of the absolute number of species in each ecotype at each site was constructed.

The site matrix included modern sites and fossil sites. The Turkana fauna was divided by member. Several multivariate methods were used to analyze the data.

Multivariate Analyses

Correspondence analysis was used to explore patterns in the data between sites.

Analyses were performed using PAST (Paleontological Statistics, Hammer et al. 2001).

Non-metric multidimensional scaling (NMDS), in which dissimilarity is used to show differences among objects, was also used as an alternative technique (Kenkel and Orlóci

1985, Gotelli and Ellison 2004). NMDS is based on a distance matrix and preserves the relative rank of differences between variables (Hammer et al. 2001). For NMDS analyses, Euclidean distance was used as the measure of dissimilarity. Euclidean distance is a simple measurement of distance between two points, and does not introduce extra assumptions. In all of these analyses, sites with similar ecological properties were grouped closer together in multivariate space. The loadings were analyzed to determine which ecotypes caused separation between the sites. Modern sites were analyzed to see how ecotypes related to environmental differences. East Asian and East African sites were then compared to determine how they differed ecologically. Ecotypes were also compared within East Asia to look at differences between hominin and non-hominin sites.

Comparing Groups of Sites using Centroids:

The centroids for ancient sites from East Asia and East Africa, and for more

specific groupings, such as hominin sites within East Asia, were computed from the

factor loadings of the correspondence analysis. Centroids were used as a means of

comparing the mean position of groups of sites in multivariate space in a quantitative

manner. Computing an average representation of a group of sites allowed comparison of

90 each site to the group mean (Euclidean distance to the centroid, mean Euclidean distance to the centroid and maximum Euclidean distance to the centroid). These measures provided a quantitative way of comparing the amount of disparity within a group, to show whether it is relatively homogeneous or heterogeneous. Similar measurements were computed by Rodríguez (2004, 2006) and Rodríguez et al. (2006).

Within each group, the following were calculated:

• Euclidean distance to the centroid for each member of the group

• Mean Euclidean distance to the centroid and maximum distance to the centroid for each group, which measures whether the group is homogeneous or heterogeneous.

• MANOVA was used to test whether the centroids of each group (including groups of fossil sites) were different.

Comparisons were made between East Asian and East African sites to determine how each group of sites differed, and how as a group early Pleistocene East Asian hominin sites differed in terms of heterogeneity compared to East African sites.

MANOVA was also used to compare the centroid locations of ancient and modern sites to answer the question of whether ancient and modern environments were similar.

The heterogeneity or homogeneity of Plio-Pleistocene sites, as shown by mean

Euclidean distance to the centroid, shows to what degree sites within groups (such as all

Asian fossil sites) were similar or disparate in ecological structure. Disparities in ecological structure may indicate the sampling of different habitats within the ancient

East Asian subset.

Summary :

The focus of this chapter was to set out the standards used to distinguish ecological categories (ecotypes) and to assign species to an ecotype. The statistics used to compare sites and assemblages were also discussed.

Ecotypes were constructed to classify modern and ancient mammals ecologically.

All animals that fit within specific parameters of body mass, diet, and substrate were combined into a single ecotype regardless of taxonomic affiliation. Use of ecotypes permitted the comparison of the taxonomically distinct East Asian and East African faunas. Assignment to dietary categories in ungulates was based upon hypsodonty. When the hypsodonty index was not available, information from the literature, including isotopic studies and functional morphology, was used to classify species. Carnivores were classified into dietary categories using ecomorphological indices, such as relative blade length of the first lower molar, and relative grinding area of the molar dentition.

Body mass was estimated using regression equations based on dental dimensions. Body mass category cut-offs reflect predation patterns. Carnivores over approximately 20 kg hunt larger prey. Ungulates over 300 kg are usually only hunted by lions and tigers, while animals over 1000 kg are not regularly preyed upon by modern carnivores. The substrate categories of terrestrial, arboreal or aquatic, were assigned based on information from the literature.

Ecological structure was analyzed using a matrix consisting of the number of species falling into each ecotype at each site. Correspondence analysis (CA) and non- metric multidimensional scaling (NMDS) were used to analyze the matrix and explore

92 differences between sites. Three sets of analyses were performed. The ecological structure model was validated by examining modern sites alone. The modern site analysis shows the interaction of ecological structure, environment and biogeography. Modern and ancient sites were analyzed together to look at how ancient habitats compared to modern ones and to gauge the degree of difference between ancient sites as compared to differences between modern sites. Finally, ancient sites were considered alone to look at environmental and ecological patterns between the regions and within East Asia.

The results of the CA were analyzed using centroids, averages of the loading factors from each site within a grouping (such as one of Bailey’s divisions or all East

Asian fossil sites). MANOVA was used to determine whether the centroids were significantly different statistically. Mean and maximum Euclidean distance to the centroid were calculated within each centroid to show whether the sites within that centroid were similar to each other, with a small mean distance, or whether the centroid grouping contains very disparate sites with a larger mean distance.

Chapter 4: Results of the ecological structure analysis

This chapter discusses the results of the ecological structure analysis of East

Asian and East African faunas. The ecological structure method is applied to modern

faunas to show how structural differences relate to habitat and geographic position. Then

modern and ancient environments are compared using correspondence analysis and non-

metric multidimensional scaling analysis (NMDS). These analyses show how differences

between ancient faunal assemblages (or sites) relate to environment and geography, and

how differences between ancient sites compare with those between modern ones. Ancient faunal assemblages are compared using correspondence analysis and NMDS.

Modern Faunal Assemblages :

Modern faunal assemblages were analyzed to show how ecological structure

relates to different environments and geographic regions. Modern faunas from national

parks and nature preserves were categorized using ecotypes and analyzed in a

correspondence analysis, using an input matrix composed of the number of species of a

particular ecotype at each site (Figure 4.1). On the first three axes, the correspondence

analysis explains 28.8%, 18.7%, and 9.74% of the variance, respectively. Geography

plays a significant role in ecological structuring, probably due to historical differences.

The first axis shows a separation between many of the African and Asian sites.

African sites tend to cluster together, including the groups of African savannas, tropical-

subtropical steppes and two sites that are mixtures of tropical-subtropical steppe and

savanna. The continent supports a large number of ungulate, carnivore and primate

species. African savannas and tropical-subtropical steppes plot to one side of the axis

94 with browsing and grazing herbivores of all ecotypes (medium, large and mega-sized).

On the other end of the axis, Southeast Asian savannas, rainforests and subtropical lowlands plot near frugivores, large omnivores, arboreal vegetation feeders, mixed feeders of 20-300 kg, and small omnivores. This mix of species is consistent with forested vegetation. Savannas and rainforests are geographically clustered. African rainforests have a very large number of frugivorous species, while savannas from Africa,

Southeast Asia and India have different ecological structures, forming a geographic gradient.

The second axis separates Eurasian settings such as temperate desert and temperate steppe as well as subtropical lowlands from environments including Southeast

Asian savannas, African savannas and rainforests. Ecotypes that plot low on the second axis include large omnivores, mixed feeders of 20 to 300 kg, small carnivores and omnivores, and aquatic mammals. Frugivores, medium browsers, mega-sized browsers and grazers, and insectivores fall on the high end of the second axis with the rainforests and savannas. Frugivores and insectivores are indicative of year-round warm climates.

The environmental separation on the second axis is not perfect, as it groups prairie and temperate desert, both relatively dry divisions, with subtropical lowlands. Both prairies and temperate deserts have small numbers of species, and their ecological structure is simple, causing the multivariate analysis to place it near the subtropical forests, which also have relatively small faunas.

Altitude is a part of the classification of the Bailey ecoregions at the division level. Divisions are divided into lowland and mountainous categories. Habitat in mountainous divisions includes a succession of environments that appear with increasing

95 altitude. This series of habitats may lead to greater variety in ecological structure. The subtropical mountains division includes a variety of ecological structures, as shown by the sites that overlap with a number of other ecological divisions.

The centroids for each division, for subdivisions (such as a centroid for each geographic savanna grouping), and for super-categories that include both lowland and mountainous division subtypes (such as the category of all subtropical, which includes subtropical lowlands and subtropical mountains) were calculated. Mean distance to the centroid, maximum distance to the centroid and the most distant site are shown in Table

4.1. Based on the scatterplot of the correspondence analysis, it was expected that divisions containing sites from a wide geographic range would be more disparate, having a larger mean Euclidean distance to the centroid (MEDC) and more variation in ecological structure. For the most part, this expectation was met. Mean distances to the centroid were highest for all tropical-subtropical steppes. This category includes lowland and mountain sites with both African and Asian localities. They were also high for the super-category of all savannas, which includes sites from Southeast Asia, India and

Africa. MEDC, which measures disparity between the sites, is expected to be greater in sites from geographically distant localities due to the influence of geography on ecological structure. Sites from a common region may share species and aspects of structure. This correspondence analysis shows that ecological structure reflects both geography and environmental conditions.

Table 4.1 The MEDC (mean Euclidean distance to the centroid) for each modern division. The MAX EDC, the greatest distance to the centroid, and the site that is at that maximum distance is listed for each division. Savanna divisions are broken into geographic subunits. When divisions contained both mountainous and lowland terrain, any group with more than one site was analyzed separately, while the super-category including both altitudinal types is labeled the “all” category. Divisions MEDC MAX Max Distance Site EDC Subtropical Lowlands 0.362 0.581 Maolan Temperate Steppe 0.469 0.469 NA Indian Savanna 0.484 0.807 Great Himalayan Savanna/Tropical Subtropical Steppe 0.563 0.563 NA Prairie 0.572 0.690 Palava Tropical Subtropical Desert 0.582 0.582 NA African Savanna 0.604 0.990 Gombe Subtropical (All) 0.605 1.510 Jiuzhaigou SE Asian Savanna 0.614 0.811 Doi Chang Dao Subtropical Mountains 0.656 1.130 Jiuzhaigou Tropical Subtropical Steppe Mountains 0.722 0.722 NA Rainforest Lowlands 0.732 0.842 Alaungdaw Rainforest 0.776 0.845 Medog Savanna (All) 0.786 1.230 Moremi Temperate Desert 0.861 0.848 Repetek Tropical Subtropical Steppe Lowlands 0.915 1.450 Boucle Tropical Subtropical Steppe (All) 0.967 1.640 Boucle

Modern Environments NMDS Analysis :

Non-metric multidimensional scaling (NMDS) analysis with Euclidean distance as the similarity measure was also used to analyze the entire modern dataset (Figure 4.2).

NMDS preserves ranked differences between the datapoints (Hammer et al. 2009). The

NMDS scaling plot shows two African rainforests are clearly separate from the other types of sites. African rainforests have many species of frugivores and are very different in ecological structure compared with savannas and with temperate and dry domain environments. These results suggest the utility of comparing ecological structure without the divergent rainforest sites. For this reason, CA and NMDS analyses were performed

97 on modern sites excluding rainforests. Analyses combining ancient and modern data were also performed with and without rainforests.

African Subtropical Rainforests Mountains 0.8 FRUG BRM MEGGR 0.6 MEGBR African savannas INS

MXL 0.4 GRL

0.2 Axis 2 ARB GRM BRL 0 MEGMX LCAR Tropical- TERSM Subtropical -0.2 Steppe AQ Lowlands SCAROMNSM (African sites) MXM -0.4 SE Asian Savannas OMNL -0.6 Subtropical Lowlands -0.8 -1.8 -1.5 -1.2 -0.9 -0.6 -0.3 0 0.3 0.6 Axis 1 Figure 4.1 Scatterplot of the correspondence analysis of the modern faunal assemblages, and ecotypes. The two circled African rainforests are outliers. Abbreviations: SCAR: Small carnivore; LCAR: Large carnivore; INS: insectivore; FRUG: frugivore; AQ: Aquatic; ARB: Arboreal vegetation feeder; TERSM: small terrestrial vegetation feeder; BRM: Medium browser (20-300 kg); GRM: Medium grazer; MXM: Medium mixed feeder; BRL: Large browser (300- 1000 kg); GRL: Large grazer; MXL: Large mixed feeder; MEGBR: Mega-browser (>1000 kg); MEGGR: Mega-grazer; MEGMX: Mega mixed feeder; OMNSM: Small omnivore (1-20 kg); OMNL: Large omnivore (>20 kg). Key to symbols: Indian Savanna; Tropical Subtropical Steppe/Savanna; Hot continental; Prairie; African savanna; Tropical-subtropical steppe; Rainforest; Subtropical lowlands; Temperate desert; Subtropical mountains; ; East Asian savanna; Tropical-Subtropical Desert; Tropical-Subtropical Steppe Mountains; Temperate Steppe

0.06

0.03 African Rainforests

-0.03

Coordinate 2 Coordinate -0.06

-0.09

-0.12

-0.15

-0.18

-0.7 -0.6 -0.5 -0.4 -0.3 -0.2 -0.1 0 0.1 0.2 Coordinate 1

Figure 4.2 Non-metric multidimensional scaling analysis using Euclidean distance of modern divisions. African rainforests, which are circled, are clearly shown as outliers in this analysis. Abbreviations: SCAR: Small carnivore; LCAR: Large carnivore; INS: insectivore; FRUG: frugivore; AQ: Aquatic; ARB: Arboreal vegetation feeder; TERSM: small terrestrial vegetation feeder; BRM: Medium browser (20-300 kg); GRM: Medium grazer; MXM: Medium mixed feeder; BRL: Large browser (300-1000 kg); GRL: Large grazer; MXL: Large mixed feeder; MEGBR: Mega-browser (>1000 kg); MEGGR: Mega-grazer; MEGMX: Mega mixed feeder; OMNSM: Small omnivore (1-20 kg); OMNL: Large omnivore (>20 kg). Key to symbols: Indian Savanna; Tropical Subtropical Steppe/Savanna; Hot continental; Prairie; African savanna; Tropical-subtropical steppe; Rainforest; Subtropical lowlands; Temperate desert; Subtropical mountains; ; East Asian savanna; Tropical-Subtropical Desert; Tropical-Subtropical Steppe Mountains; Temperate Steppe

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Correspondence Analysis of Modern Sites Excluding Rainforests :

Further analysis was focused on savannas, and on environments from temperate and dry settings. Another correspondence analysis in which rainforests were excluded is shown in Figure 4.3. The first axis explains 30.6% of the variance while the second and third explain 15.2% and 10.7%, respectively. On the first axis, there was separation between African and Asian environmental groupings. All browsing and grazing ecotypes are placed near the African savannas and tropical-subtropical steppes. Frugivores, omnivores, arboreal vegetation feeders and medium mixed feeders are placed on the other end of the axis. Most temperate and dry sites are found low on the second axis, separated from tropical-subtropical steppes, savannas and subtropical lowlands. The temperate and dry site groups have fewer species. Mean Euclidean distances to the centroid are listed in Table 4.2. Mean distances to the centroid were highest for tropical- subtropical steppes. Savannas also had high MEDC values, showing more variation in structure.

Table 4.2 Modern site group centroids excluding rainforests, with the maximum Euclidean distance to the centroid and the locality with that distance are listed. Site Group MEDC MAX EDC Max Site Subtropical Lowlands 0.371 0.586 Maolan Temperate Steppe 0.458 0.458 NA Indian Savanna 0.490 0.810 Great Himalayan Savanna/Tropical Subtropical Steppe 0.554 0.554 NA Prairie 0.556 0.680 Palava Tropical Subtropical Desert 0.579 0.579 NA African Savanna 0.613 1.000 Gombe Subtropical (All) 0.618 1.523 Jiuzhaigou SE Asian Savanna 0.635 0.849 Doi Chiang Dao Subtropical Mountains 0.670 1.145 Jiuzhaigou Tropical Subtropical Steppe Mountains 0.725 0.725 NA Savanna (All) 0.804 1.238 Doi Chiang Dao Temperate Desert 0.850 1.154 Astrakhanskiy Tropical Subtropical Steppe Lowlands 0.893 1.408 Boucle Tropical Subtropical Steppe (All) 0.959 1.592 Boucle

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SE Asian Savannas FRUG 0.6 MEGGR ARB BRM INS 0.3 MEGBR MXL Axis 2 GRL 0 LCARTERSMMEGMX GRM OMNSMMXM

-0.3 SCAR OMNL African Savannas and Tropical- BRL -0.6 Subtropical Subtropical Steppes Lowlands AQ -0.9

-1.2 Prairie, temperate desert, temperate steppe, tropical-subtropical desert -1.5

-1.8

-1.8 -1.5 -1.2 -0.9 -0.6 -0.3 0 0.3 0.6 Axis 1 Figure 4.3 Correspondence analysis scatterplot of modern faunal sites excluding rainforests. Abbreviations: SCAR: Small carnivore; LCAR: Large carnivore; INS: insectivore; FRUG: frugivore; AQ: Aquatic; ARB: Arboreal vegetation feeder; TERSM: small terrestrial vegetation feeder; BRM: Medium browser (20-300 kg); GRM: Medium grazer; MXM: Medium mixed feeder; BRL: Large browser (300-1000 kg); GRL: Large grazer; MXL: Large mixed feeder; MEGBR: Mega-browser (>1000 kg); MEGGR: Mega-grazer; MEGMX: Mega mixed feeder; OMNSM: Small omnivore (1-20 kg); OMNL: Large omnivore (>20 kg). Key to symbols: Indian Savanna; Tropical Subtropical Steppe/Savanna; Hot continental; Prairie; African savanna; Tropical-subtropical steppe; Rainforest; Subtropical lowlands; Temperate desert; Subtropical mountains; ; East Asian savanna; Tropical-Subtropical Desert; Tropical-Subtropical Steppe Mountains; Temperate Steppe

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NMDS Modern Environmental Groups Excluding Rainforests

A NMDS analysis with modern sites excluding rainforests was used to study relationships between the other modern site groups (Figure 4.4). African savannas are spread over the first axis, with Mamili and Gombe particularly widely separated. Gombe has more frugivores and arboreal vegetation feeders, while Mamili has more species of browsers, grazers and mixed feeders. Gombe, an African savanna, is plotted with other divisions, such as Southeast Asian savannas and subtropical lowlands. These divisions on the high end of the first axis have more ecotypes of small-bodied animals and represent more forested environments. On the second axis, African savannas and an African tropical-subtropical steppe/savanna (Serengeti) are separated from a southeastern Asian savanna (Daweishan). Similar factors appear to separate the sites on the second axis with sites having more browsers, grazers and mixed feeders separated from the sites higher on the axis with fewer species in those ecotypes and more of the smaller ecotypes.

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0.24 Daweishan

0.18 Queen Elizabeth 0.12 Gombe

0.06 Coordinate 2

-0.06

-0.12 Mamili

-0.18 Moremi

-0.24 Serengeti

-0.3 -0.36 -0.3 -0.24 -0.18 -0.12 -0.06 0 0.06 0.12 Coordinate 1

Figure 4.4 NMDS of modern sites excluding rainforests. Symbols and abbreviations are as in Figure 4.1.

Summary of Ecotype Analyses of Modern Faunas :

Analysis of the modern faunas shows both geographic and environmental

influences on the placement of sites in multivariate space. NMDS analysis of all sites

underscores the differences between African rainforests and other modern sites, most

likely due to the large number of frugivores. Due to these differences, a correspondence

analysis without rainforests was undertaken. This CA showed similar patterns, with

African savannas and tropical-subtropical steppes plotting with browsers and grazers,

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while other divisions loaded with medium-sized mixed feeders, frugivores, and

omnivores. When the NMDS was repeated without rainforests, African savannas showed the most variation, with a contrast between sites with many large grazing and browsing ecotypes, and those with greater proportions of smaller ecotypes. From these analyses, it is evident that African rainforests are significantly different from the other sites, and that comparison using ecotypes shows differences in structure between different environments. Comparisons of modern faunas with those of Plio-Pleistocene sites will be

done using all modern sites as well as using a dataset of modern sites excluding

rainforests.

Plio-Pleistocene and Modern Sites Combined :

Plio-Pleistocene sites and modern localities grouped by division and geographic

location were analyzed together in a correspondence analysis. The modern and Plio-

Pleistocene sites were combined in order to facilitate environmental interpretation. These

comparisons show how the distribution of ecotypes in ancient sites can be interpreted

environmentally. The combination of ancient and modern faunas also shows whether

ancient faunas had analogs in modern environmental types. Were any of the ancient sites

like modern savannas and how were they were different? Also, were any of the ancient

sites from South China similar to modern subtropical forests? Finally, comparison of the

faunas of ancient and modern sites together is helpful in showing the magnitude of

difference between groups of sites. How does the amount of disparity in Asian or African

fossil sites compare with the amount of disparity in modern divisions? Greater amounts

of disparity imply ecological structures more diverse than those found in modern

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divisions, which are grouped by environments. This problem is addressed by looking at

mean Euclidean distance to the centroid (MEDC) and maximum distance to the centroid

(Max EDC), which measure how different each site is from the division average.

However, error may be introduced into this analysis because of the fact that the modern

and ancient faunal lists were compiled and sorted into ecotypes in different ways.

In the correspondence analysis of all modern sites and the Plio-Pleistocene sites,

the first axis explains 30.7% of the variation, the second 12.4% and the third 10.3%

(Figure 4.5). The first axis shows some separation between the ancient sites and the

modern groups. The ancient East Asian and African assemblages overlap in distribution, though they are mostly separate from modern sites. Sites such as Serengeti, Boucle,

Moremi, Tsavo and Jiuzhaigou had ecological structures most similar to those of the ancient sites. The Asian and African fossil assemblages are drawn together by browsers and grazers of all sizes, as well as by mega-sized mixed feeders. The fossil sites have many more species of these ungulates than any of the modern sites. At the other end of the first axis, the ecotypes of frugivores, small carnivores, small omnivores, and insectivores plot with site groups that include Southeast Asian savannas, rainforests and subtropical forests.

The second axis shows some separation based upon climate. Many of the sites from the drier environmental types, such as prairies, steppes and deserts plot low on the second axis. In Eurasia, these sites are relatively impoverished in terms of the number of faunal elements. High on the second axis are African rainforests, Southeast Asian savannas, as well as some African savannas, which plot with ecotypes such as frugivores, small arboreal vegetation feeders and insectivores. The third axis (Figure 4.6) separates

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Asian and African sites: the southern Asian sites were associated with browsers while the northern and African localities had the grazing ecotypes.

Table 4.3 shows the mean distance to the centroid for modern site groups and for ancient sites. A number of possible groupings were calculated for the ancient fossil assemblages based on geography, time period and whether hominin remains or archaeological evidence was found at the site. The correspondence analysis shows that the ancient fossil assemblages grouped closely together, though the Asian sites appear to occupy more space. This is confirmed by the MEDC, which shows that the African faunal assemblages had a smaller mean distance to the centroid than the group of all

Asian sites, and the group of Asian hominin sites. The larger centroid size of hominin sites in East Asia indicates that the places where hominins are found in East Asia were more disparate in ecological structure than the East African assemblages. However, the

East African sites used for the comparison come from a relatively small area of the

African continent where many species were shared.

The MEDC values show that the group of sites from northern China is more disparate than the south China assemblage. Ancient South China sites are thought to have been similar to subtropical forests, but the South China fossil sites do not group with the modern subtropical lowland and mountainous forests. The Nihewan sites have a high

MEDC despite their common location in the same basin and the fact that they contain many of the same species. The small sample sizes from Majuangou, Xiaochangliang and

Donggutuo contribute to the large MEDC. North China fossil sites may sample several habitat groups rather than a single habitat. The centroid for East Asian hominin sites has the largest mean distance to the centroid of any grouping. The Asian hominin sites as a

107 group are more disparate in ecological structure than any of the modern divisions including all savannas, indicating that the sites where hominins have been found sample multiple types, including diverse types of temperate or humid tropical habitats.

Table 4.3 Mean distance to the centroid and maximum distance to the centroid are shown for modern environmental groupings as well as for groupings of the ancient fossil assemblages. Environmental or Regional Group MEDC MAX Site with maximum EDC MEDC Olduvai 0.355 0.355 NA Subtropical Lowlands 0.394 0.648 Maolan Indian Savanna 0.456 0.811 Great Himalayan Prairie 0.477 0.552 Palava Temperate Steppe 0.503 0.503 NA Koobi Fora 0.527 0.976 Chari Savanna/Tropical Subtropical Steppe 0.537 0.537 NA Tropical Subtropical Desert 0.577 0.577 NA South China 0.581 0.830 Yuanmou West Turkana 0.582 0.701 Lokalalei African Savanna 0.591 1.014 Gombe Subtropical All 0.615 1.427 Jiuzhaigou Subtropical Mts. 0.640 1.064 Jiuzhaigou SE Asian Savanna 0.640 0.867 Doi Chang Dao Ancient African Sites 0.707 1.261 Chari Tropical Subtropical Steppe Mts. 0.767 0.767 NA Tropical Subtropical Steppe Lowlands 0.783 1.164 Boucle Savanna 0.784 1.209 Doi Chang Dao Rainforest Lowlands 0.806 0.914 Alaungdaw Temperate Desert 0.839 1.144 Astrakhanskiy Rainforest ALL 0.844 0.933 Tai Tropical Subtropical Steppe All 0.896 1.313 Baimaxueshan Pliocene 0.898 1.196 Nihewan Non-Hominin 1.003 1.251 Linyi Nihewan 1.064 1.596 Majuangou North China 1.242 1.601 Donggutuo Asia 1.271 2.007 Donggutuo Pleistocene 1.370 1.853 Donggutuo Hominins 1.460 1.767 Donggutuo

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Frugivores, small carnivores and omnivores, 1.25 insectivores Browsers, Grazers, FRUG Mega-Mixed Feeders Rainforests 1 SE Asian Savannas 0.75 African Savannas East Asian Fossil Sites 0.5 Donggutou Majuangou MEGMX Xiaochangliang ARB MEGGR 0.25 Nariokotome Oko ke INS

Kaitio GRL Chari Longdan Linyi KBS Lokalalei BRM Longgupo Axis 2 NatooNihewan GongMXL Mohu. OMNL 0 Kalochoro Yuanmou MXM UB Olduvai_I Olduvai_2 Jianshi MEGBR BRL LCAR OMNSM

GRM AQ -0.25 East African Fossil Subtropical

Sites Lowland

SCAR TERSM

-0.5 Tropical-Subtropical

Steppes

-0.75 Temperate Desert

-1 -1.5 -1.2 -0.9 -0.6 -0.3 0 0.3 0.6 0.9 Axis 1

Figure 4.5 Scatterplot of a correspondence analysis of the modern sites and ancient fossil assemblages. Key: Ancient East Asian faunal assemblages Ancient East African faunal assemblages. Indian Savanna; Tropical Subtropical Steppe/Savanna; Hot continental; Prairie; African savanna; Tropical-subtropical steppe; Rainforest; Subtropical lowlands; Temperate desert; Subtropical mountains; ; East Asian savanna; Tropical-Subtropical Desert; Tropical-Subtropical Steppe Mountains; Temperate Steppe

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Grazers 0.9 East African Chari Fossil sites Nariokotome 0.6 Kaitio MEGGR GRL GRM Natoo Okote 0.3 Olduvai_2 Olduvai_I. . AQ KBS Xiaochangliang Kalochoro Majuangou SCAR . Donggutou Lokalalei ARB. . MEGMX . 0 . FRUG. OMNL. INS OMNSM MXM . . Axis 3 TERSM. Longdan LCAR -0.3 Linyi Nihewan

Uppe r Burgi Jianshi -0.6 MXL BRL Gongwangling MEGBR -0.9 Longgupo BRM Browsers East Asian -1.2 Fossil sites Mohui Yuanmou -1.5

-1 -0.75 -0.5 -0.25 0 0.25 0.5 0.75 1 1.25 Axis 2

Figure 4.6 Scatterplot of a correspondence analysis of the modern sites and ancient fossil assemblages, axes 2 and 3. Key: Ancient East Asian faunal assemblages; Ancient East African faunal assemblages. Indian Savanna; Tropical Subtropical Steppe/Savanna; Hot continental; Prairie; African savanna; Tropical- subtropical steppe; Rainforest; Subtropical lowlands; Temperate desert; Subtropical mountains; ; East Asian savanna; Tropical-Subtropical Desert; Tropical-Subtropical Steppe Mountains; Temperate Steppe

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NMDS Analysis of Modern and Plio-Pleistocene Faunal Assemblages :

A non-metric multidimensional scaling analysis with all modern and ancient sites showed a very small overlap between the Asian and the African fossil sites (Figure 4.7).

The NMDS analysis separates sites that have the greatest differences. There is more difference between the African rainforests and the Plio-Pleistocene East African fossil sites than between the two sets of ancient sites. Modern site types overlap with each other on the first two axes, with the exception of the African rainforests and some African savannas. The first axis separates East African fossil sites from the African rainforests.

The sites differ in the number of browsers, grazers, and mixed feeders, which are more numerous in the East African fossil sites, such as the KBS member, and less common in the African rainforest. Frugivores and insectivores are found in the modern African rainforest, but are uncommon in the East African fossil sites. The second axis shows a

contrast between African savannas and East Asian fossil sites. Among the possible

factors contributing to the separation is the greater number of smaller ecotypes in the

modern savanna sites compared with the ancient sites.

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East Asian Fossil Sites 0.16 Longdan

0.12 Longgupo Yuanmou Nihewan Jianshi Mohui 0.08 Gongwangling Nariokotome Majuangou Linyi Donggutou Kaitio 0.04 Xiaochangliang. LokalaleiKalochoro . Natoo Coordinate 2 Chari. .. . Olduvai 2 0 . . Okote . .

-0.04 Olduvai I KBS African Rainforests -0.08 Upper Burgi

East African -0.12 Fossil Sites

-0.16 African savannas

-0.2 -0.24 -0.16 -0.08 0 0.08 0.16 0.24 0.32 Coordinate 1

Figure 4.7 NMDS analysis of ancient and modern sites. Key: Ancient East Asian faunal assemblages Ancient East African faunal assemblages. Indian Savanna; Tropical Subtropical Steppe/Savanna; Hot continental; Prairie; African savanna; Tropical-subtropical steppe; Rainforest; Subtropical lowlands; Temperate desert; Subtropical mountains; ; East Asian savanna; Tropical-Subtropical Desert; Tropical-Subtropical Steppe Mountains; Temperate Steppe

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Correspondence Analysis of Ancient and Modern Assemblages (Excluding Rainforests) :

Due to the evidence from the NMDS that the rainforests are one of the most

divergent groups, and from the correspondence analysis showing that African rainforests

seem to be significantly different from other humid tropical sites, as well as from other

temperate and dry environments, a CA of ancient sites with a group of modern

comparators excluding rainforests was performed. In this CA, the first axis explains

31.8% of the variation, with the second and third axes explaining 11.2% and 10.7% of the

variation, respectively.

The scatterplot (Figure 4.8) shows separation between ancient faunal assemblages

and most modern sites on the first axis. The second axis separates Plio-Pleistocene Asian

and African faunal assemblages. The ecotypes plotted with the ancient faunal

assemblages include all browsers and grazers, and large and mega-sized mixed feeders.

At the opposite end of the first axis modern site groups plot with ecotypes including

small carnivores and omnivores, frugivores, insectivores and small terrestrial vegetation

feeders. All of the modern sites group together, separate from the ancient sites. This is

due in part to the large number of species of ungulates in the categories of browsers,

grazers and mixed feeders in the ancient sites, compared with the smaller faunas in all of

the modern sites. Also, modern sites contain more small animals, which are often not

preserved in fossil assemblages. Though parts of the variation in ecological structure

reflect differences in habitat and vegetation, species richness and taphonomy also play

roles in the placement of sites.

On the second axis, the Asian fossil sites plot with browsing ungulates and large and mega-sized mixed feeders, while African fossil assemblages plot with a

113 concentration of grazers. Plio-Pleistocene Asian sites are spread over the third axis

(figure 4.9). Ancient southern China sites and Gongwangling are found lower on the axis with the browsing ecotypes, while Plio-Pleistocene northern Asian sites overlap with

Southeast Asian savannas.

The mean Euclidean distance to the centroid (MEDC) (Table 4.4) shows that hominin site faunas in Plio-Pleistocene East Asia have the largest MEDC, showing that they inhabited ecologically disparate sites. Northern East Asia also has a large MEDC compared with the modern divisions, which may imply that more than one habitat is represented in this sample of sites from northern Asia. The North China sites also span a long time range and vegetation is known to have fluctuated over time, with warm and humid periods, as well as cooler and drier ones (Wu et al. 2007). Within northern East

Asia, the Nihewan Basin has a large MEDC despite the sites’ similarity in species composition and geographic proximity. Southern East Asian fossil sites have less disparity in ecological structure than northern China and are similar to the values for

West Turkana and Koobi Fora. The large value for the East Asian hominin sites implies that hominins occurred in ecologically disparate places. Results of the MANOVA (Table

4.5) indicate that African fossil sites are statistically significantly different from many modern environments, including modern savannas. Asian fossil sites were significantly different from SE Asian savannas and from subtropical forests, showing that the fossil sites differed in structure from the modern environments in the region.

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Browsers and Mixed feeders East Asian Fossil Sites 1.25 Yuanmou Mohui

1 Frugivores, small BRM Longgupo omnivores, 0.75 Gongwangling insectivores, MEGBR MXL Jianshi small 0.5 BRL carnivores Axis 2 MEGMX Linyi 0.25 . . Donggutou NihewanLongdan FRUG OMNL MXMLCAR . . OMNSM 0 Xiaochangliang ARB Majuangou TERSM .

Lokalalei . INS Upper_Burgi. KBS -0.25 Kalochoro SCAR Okote Olduvai_I . Natoo Olduvai_2 AQ

Kaitio GRL GRM -0.5 Nariokotome MEGGR

Grazers Chari -0.75 East African Fossil sites

-1 -1.5 -1.2 -0.9 -0.6 -0.3 0 0.3 0.6 0.9 Axis 1

Figure 4.8 Correspondence analysis scatterplot of modern and ancient faunal assemblages, with modern rainforests excluded. Key: Ancient East Asian faunal assemblages; Ancient East African faunal assemblages. Indian Savanna; Tropical Subtropical Steppe/Savanna; Hot continental; Prairie; African savanna; Tropical-subtropical steppe; Subtropical lowlands; Temperate desert; Subtropical mountains; ; East Asian savanna; Tropical-Subtropical Desert; Tropical- Subtropical Steppe Mountains; Temperate Steppe

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1.5 MEGMX

1.2 SE Asian Savannas Donggutou East 0.9 African Xiaochangliang Fossil Sites FRUG 0.6 Nihewan Majuangou OMNL Nariokotome Axis 3 East Asian Natoo . ARB Linyi Fossil Sites 0.3 MEGGR OMNSM Okote MXM GRL KBS . ... Longdan Kaitio . . Lokalalei Gongwangling . . 0 . Upper. Burgi. .. Jianshi Kalochoro. . . Chari AQ Olduvai_I LCAR Longgupo Olduvai_2 . SCAR GRM. MXL -0.3 BRL Mohui MEGBR TERSM . BRM INS

-0.6 Browsers Yuanmou

-0.9 -1 -0.75 -0.5 -0.25 0 0.25 0.5 0.75 1 1.25 Axis 2 Figure 4.9 Correspondence analysis scatterplot of modern sites (excluding rainforests) and ancient fossil assemblages, axes 2 and 3. Key: Ancient East Asian faunal assemblages; Ancient East African faunal assemblages. Indian Savanna; Tropical Subtropical Steppe/Savanna; Hot continental; Prairie; African savanna; Tropical-subtropical steppe; Subtropical lowlands; Temperate desert; Subtropical mountains; ; East Asian savanna; Tropical-Subtropical Desert; Tropical- Subtropical Steppe Mountains; Temperate Steppe

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Table 4.4 MEDC and maximum distance to the centroid for modern and ancient sites excluding rainforests. Grouping MEDC MAX Max Site EDC Olduvai Gorge 0.349 0.349 NA Subtropical Lowlands 0.408 0.661 Maolan Indian Savanna 0.460 0.819 Great Himalayan Prairie 0.472 0.545 Palava Temperate Steppe 0.497 0.497 NA Koobi Fora 0.524 0.967 Chari Savanna/Tropical Subtropical Steppe 0.536 0.536 NA West Turkana 0.572 0.689 Lokalalei South China Fossil Sites 0.579 0.829 Yuanmou Tropical Subtropical Desert 0.581 0.581 NA African Savanna 0.609 1.042 Gombe Subtropical All 0.629 1.439 Jiuzhaigou Subtropical Mountains 0.656 1.082 Jiuzhaigou Southeast Asian Savanna 0.664 0.914 Doi Chang Dao African Fossil Sites 0.697 1.255 Chari Tropical Subtropical Steppe Lowlands 0.780 1.151 Boucle Tropical Subtropical Steppe Mts. 0.781 0.781 NA Savanna 0.808 1.296 Doi Chang Dao Temperate Desert 0.843 1.153 Astrakhanskiy Tropical Subtropical Steppe All 0.905 1.349 Baimaxueshan Non-Hominin Asian Fossil Sites 0.977 1.213 Linyi Nihewan Basin (all) 1.043 1.552 Majuangou North China Fossil Sites 1.212 1.564 Donggutuo Asia Fossil Sites 1.242 1.953 Donggutuo Hominins in East Asia Fossil Sites 1.426 1.718 Donggutuo

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Table 4.5 MANOVA results for the centroids of African and Asian fossil sites from the ancient and modern CA without rainforests. Values above the diagonal are p-values, values below are Bonferroni corrected. Some modern divisions contain too few sites to be tested. Abbreviations: N: NA because the test could not be performed. Trop. Subtrop. Steppe Trop. Subtrop. Steppe Mts. Temperate steppe Prairie Savanna/Trop. Subtrop. Steppe India Savanna SE Asia Savanna Africa Savanna Subtropical Mountains Subtropical Africa fossil sites Asia Hominins Asia Non-Hominins Trop. Subtrop. steppe -- N N N N 0.15 0.048 0.42 N 0.11 0.005 0.253 0.099 Trop. Subtrop. steppe Mts. N -- N N N N 0.357 0.035 N N 0.027 N N Temp. Steppe N N -- N N N 0.262 0.005 N N 0.001 N N Prairie N N N -- N 0.305 0.021 0.012 N 0.039 0.000 0.172 0.034 Savanna Trop.- Subtrop. steppe N N N N -- N 0.194 0.030 N N 0.093 N N India Savanna 1 N N 1 N -- 0.040 0.025 0.058 0.025 0.000 0.056 0.048 SE Asia Savanna 1 1 1 1 1 1 -- 0.021 0.283 0.231 0.000 0.000 0.000 Africa Savanna 1 1 0.55 1 1 1 1 -- 0.130 0.002 0.000 0.005 0.001 Subtrop. Mts. N N N N N 1 1 1 -- 0.056 0.003 0.206 0.011 Subtrop. Low. 1 N N 1 N 1 1 0.233 1 -- 0.000 0.017 0.004 African fossil 3.1 sites 0.57 1 0.16 0.028 1 0.012 E-05 0.006 0.280 0.005 -- 0.001 0.000 Asian Hom. Sites 1 N N 1 N 1 0.0004 0.505 1 1 0.153 -- 0.433 Asian Non-Hom Sites 1 N N 1 N 1 0.048 0.069 1 0.470 0.014 1 --

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NMDS of Plio-Pleistocene and Modern Faunas (excluding Rainforests) :

The non-metric multidimensional scaling analysis of Plio-Pleistocene faunal assemblages and modern sites excluding rainforests shows separation on the first axis between a Southeast Asian savanna locality and the KBS member fauna (Figure 4.10).

While the KBS fauna has many browsers, grazers and mixed feeders, Southeast Asian savannas have more small ecotypes, including small carnivores and omnivores, which are rarely preserved. There is separation on the first axis between the Asian and African fossil faunas. The second axis shows a progression of sites with a large group containing the modern sites, a group of East African fossil sites, and then the East Asian fossil faunas. The East Asian sites also show separation between many of the sites with smaller faunas lower on the second axis and larger faunas on the positive end.

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Longdan East Asian Fossil Sites

0.16 Longgupo Yuanmou 0.12 Mohui Nihewan Gongwangling 0.08 Jianshi Nariokotome Majuangou Kaitio Xiaochangliang Linyi Kalochoro Donggutou Lokalalei Natoo 0.04 Chari Olduvai 2 . Okote 0

Olduvai I KBS -0.04

Upper Burgi -0.08

Coordinate 2 East African -0.12 Fossil Sites African -0.16 Savannas . -0.2 -0.18 -0.12 -0.06 0 0.06 0.12 0.18 0.24 0.3 0.36 Coordinate 1

Figure 4.10 NMDS plot of Plio-Pleistocene assemblages with modern sites (excluding rainforests). Key: Ancient East Asian faunal assemblages; Ancient East African faunal assemblages. Indian Savanna; Tropical Subtropical Steppe/Savanna; Hot continental; Prairie; African savanna; Tropical-subtropical steppe; Subtropical lowlands; Temperate desert; Subtropical mountains; ; East Asian savanna; Tropical-Subtropical Desert; Tropical-Subtropical Steppe Mountains; Temperate Steppe

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Correspondence Analysis with Plio-Pleistocene Faunal Assemblages :

Analysis of the Plio-Pleistocene faunal assemblages alone was performed in order

to examine patterns between the ancient sites. Unlike other comparisons, this analysis

uses only ancient assemblages. The data were derived from faunal site lists, and

adaptations were assigned using dental measurements. The ancient sites were studied

alone in this section to examine the effects of geography and environment, particularly for the East Asia localities.

A correspondence analysis was also done with only the Plio-Pleistocene faunal assemblages. The correspondence analysis explained 31.2% of the variance on the first axis, 16.8% on the second and 14.8% on the third axis (Figure 4.11). The first axis separates most East Asian and East African sites. East African sites form a much tighter cluster than the East Asian set, indicating that they have more similarity in their ecological structures than East Asian sites do. East African sites plot with ecotypes such as grazers, insectivores, arboreal vegetation feeders and aquatic mammals. The Asian sites are associated with a variety of ecotypes including small omnivores, browsers and mixed feeders and small terrestrial mammals. The fact that the East African sites form a tight cluster in the multivariate analysis may be due to their location in a single geographic region, leading to greater ecological similarity compared with the East Asian sites sampled here. A future sample with African fossil sites from different regions could be used to investigate ecological structure differences in Plio-Pleistocene Africa as well.

The second axis shows some geographic structure in the group of East Asian sites.

The south China sites, as well as the sites of Gongwangling and Longdan, are found higher on the axis, at the same level as the East African faunas, such as Olduvai I, Okote

121 and Upper Burgi. These sites have higher amounts of arboreal vegetation feeders, frugivores, and browsers, which is consistent with the forested environments. Like the south China sites, Longdan also has a frugivorous species, and a small terrestrial mammal, while Gongwangling has arboreal vegetation feeders. The lower end of the second axis includes the other north China sites, and is associated with ecotypes such as mega-sized mixed feeders and browsers. The placement of Longdan and Gongwangling also reflects their geographic position on the southern edge of north China. The position of the Longdan fauna in multivariate space could reflect a relatively southern location or a warmer time period during the Pliocene. Likewise, the position of the Gongwangling fauna in the CA could reflect its geographic position, or it could indicate that some faunal elements from southern China were able to disperse through the Qingling Mountains during a warmer period.

Comparison of the mean distance to the centroid (Table 4.6) shows that the average distance to the centroid is greater in Asia than in Africa, which is logical because the African sites are concentrated in a small geographic region, whereas the Asian sites are spread over a larger area. Approximately 800 km separates Olduvai Gorge from sites in Lake Turkana, whereas the East Asian sites span approximately 1,160 km from north to south and 800 km from east to west. The mean distance to the centroid for the group of

Asian hominin sites was also greater than that of the non-hominin sites, and greater than

African sites. Of the hominin sites, Yuanmou has the greatest distance to the centroid.

Yuanmou has many browsers and mixed feeders, possibly contributing to its distance from the centroid. Gongwangling is similar to Yuanmou in the types of ecotypes represented, though it contains fewer species. Xiaochangliang, Majuangou and

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Donggutuo are small assemblages with few species. The small number of species makes their ecological patterns somewhat difficult to interpret, but they are similar to each other because they share ecotypes. A MANOVA of the centroid values (Table 4.7) shows that

African fossil sites are significantly different in ecological structure from both the Asian hominin and non-hominin sites. However, the Asian hominin and non-hominin sites are not significantly different from each other. This agrees with the placement of sites in the

CA, which is primarily influenced by geography and environment. Hominins did not occur in a single type of ecological structure. The non-hominin East Asian sites sampled here do not belong to a single type of ecological structure in which hominins did not occur. Though there were likely to have been environments in which hominins were not found, there is no evidence of a systematic difference between the hominin and non- hominin sites sampled here. Rather, hominins occurred in a variety of ecological structures. Based on the sites sampled, there is no evidence that hominins were selecting certain habitats within East Asia.

Table 4.6 Mean and maximum distance to the centroid for Plio-Pleistocene sites.

Grouping MEDC MAX Max Site EDC Olduvai Gorge 0.442 0.442 NA West Turkana 0.454 0.538 Lokalalei Koobi Fora 0.511 0.905 Chari South China 0.597 0.744 Yuanmou Ancient African Sites 0.601 1.143 Chari Nihewan sites 0.818 1.075 Majuangou Non-Hominin East Asian sites 0.827 1.013 Longdan North China sites 0.947 1.235 Longdan Ancient Asian sites 0.987 1.189 Longdan Hominin sites in East Asia 1.047 1.175 Yuanmou

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Table 4.7 MANOVA results for the centroids of African and Asian fossil sites (ancient CA only), for sites grouped by hominin presence compared with Africa. Values above the diagonal are p-values, values below are Bonferroni corrected. Africa Asian Asian Non-hominins Hominin Africa 0 0.000132755 0.0008368 Asian Non- 0.000398 0 0.513968 hominins Asian Hominins 0.00251 1 0

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Browsers and mixed feeders, small INS 1.2 omnivores, small terrestrial mammals East African Fossil Sites 0.8 East Asian Fossil Sites FRUG Olduvai_I SCAR ARB Chari 0.4 Okote TERSM Longdan Longgupo UB KBS Mohui Jianshi BRL Kalochoro GRM MEGBR AQ 0 Yuanmou BRM GongwanglingLCAR Olduvai_2 Axis 2 MXM Lokalalei GRL Natoo Kaitio -0.4 OMNSM MXL OMNL Nariokotome South China Majuangou MEGGR Fossil Sites Linyi -0.8 Nihewan Xiaochangliang Grazers, insectivores, -1.2 Most North arboreal vegetation China Sites Donggutou feeders, aquatic -1.6 mammals

MEGMX -1 -0.75 -0.5 -0.25 0 0.25 0.5 0.75 1 Axis 1

Figure 4.11 Correspondence analysis scatterplot of Plio-Pleistocene sites. The north China sites are the Nihewan, Donggutuo, Xiaochangliang, Majuangou, Linyi, Longdan and Gongwangling. They are split up in the CA scatterplot, while the south China sites (circled) are found together. The hominin sites are Donggutuo, Xiaochangliang, Majuangou, Yuanmou and Gongwangling. They are found in a variety of ecological structures. Key: Ancient East Asian faunal assemblages; Ancient East African faunal assemblages

Summary :

The ecological structure of ancient sites in East Asia and East Africa, along with a

set of modern sites for comparison, was analyzed to explore the question of whether East

African and East Asian faunas were ecologically similar to different. A related question

looks at whether the East Asian hominin sites were ecologically different from the Asian

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non-hominin sites. Ecological structure analyses were done using modern faunas, modern

and ancient faunas and ancient faunas alone.

The correspondence analysis of all modern faunas shows the influence of

geography, with divisions from Africa, such as African savannas and tropical-subtropical

steppes, separated from Asian divisions such SE Asian savannas and subtropical forests.

Environmentally and ecologically, there is a gradient between drier divisions and those

with more humid climates. African rainforests are an outlier in terms of ecological

structure due to the large number of frugivorous species. Due to the effect of the African

rainforests on the placement of the rest of the fauna, subsequent analyses were performed

with and without this group.

Ancient and modern sites were analyzed together to look at whether ancient sites had modern analogues and to put differences between ancient sites in context. Plio-

Pleistocene sites from both Asia and Africa are separated from modern sites of all divisions on the first axis of the correspondence analysis, indicating that African sites were not similar to modern savannas, and Asian fossil sites did not have the same structure as modern subtropical forests. All browsers, grazers, as well as large and mega- sized mixed feeders plot with the Plio-Pleistocene assemblages, while modern sites group with small carnivores and omnivores, frugivores, insectivores and small terrestrial vegetation feeders. Asian and African fossil sites are separated from each other on the second axis. On the second axis, the Asian fossil sites plot with browsing ungulates and large and mega-sized mixed feeders, while African fossil assemblages plot with a concentration of grazers. Measurements of centroid statistics for the ancient and modern groups shows that the set containing East Asian hominin sites has the largest mean

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distance to the centroid, indicating that this group contains the most ecologically

disparate sites.

The Plio-Pleistocene site analysis showed the East Asian and East African sites

are separated on the first axis. East African sites plot with ecotypes such as grazers,

insectivores, arboreal vegetation feeders and aquatic mammals. The Asian sites are

associated with a variety of ecotypes including small omnivores, browsers and mixed

feeders and small terrestrial mammals. The East African sites form a much tighter cluster

than the East Asian set, indicating that the African sites are more similar in ecological

structure compared with East Asia.

Within East Asia, the correspondence analysis shows ecological and geographic

patterning. South China sites as well as the northern sites of Gongwangling and Longdan, have a similar ecological structure, with the ecotypes of arboreal vegetation feeders, frugivores, and browsers, which is which is consistent with more closed habitats. These sites are distinct from a second set of sites, all located in northern China. Examination of the centroids of the Plio-Pleistocene sites shows that the average distance to the centroid is greater in Plio-Pleistocene Asian groupings than in African groupings. African sites are concentrated in a smaller geographic area compared with the Asian sites. The mean distance to the centroid for the group of Asian hominin sites was also greater than that of

the Asian non-hominin sites, and greater than that of African sites.

This chapter shows that hominins did not occur in a single type of ecological

structure. While the ecological structures of Plio-Pleistocene East Asian sites differ due

to geography and environment, the sites do not separate into hominin and non-hominin

groups. Within the Asian Plio-Pleistocene sites sampled here, hominins do not appear to

127 have been excluded from any particular ecological structure. Although hominins would not have lived in all possible environments (rainforests, deserts and tundra would have been unlikely during the early Pleistocene), there is no evidence for systemic differences between the temperate hominin and non-hominin sites sampled here. Rather, hominins occurred in diverse ecological structures.

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Chapter 5: Background to Carnivore Ecomorphology

Ecomorphological Analysis of Carnivore Guilds :

Ecomorphological analysis is based on functional morphological characteristics that relate to aspects of ecology. Using a system of measurements related to ecology, carnivores from diverse species can be compared in terms of traits such as feeding adaptations, locomotion, prey capture and body mass. This project focuses on feeding adaptations and body mass, as derived from dental measurements. Feeding adaptations are used to classify carnivores into dietary classes such as highly carnivorous, omnivorous, or bone-crackers. These traits are examined for all species and tested for the groups of carnivores in East Asian and East African ancient faunal assemblages.

Groups of carnivores found at a particular site, region or time period are known as guilds. A guild is defined as a group of species that uses a resource in a similar way (Root

1967). For carnivores, the resources used are meat and bone from hunted or scavenged prey. The aim of comparing carnivore guilds from different times and places is to use ecomorphological traits to show which species are ecological avatars, that is, different species that have similar adaptations and fill similar roles in the community.

Fossil carnivore guilds have been analyzed ecomorphologically in several previous studies (Lewis 1995, 1997; Van Valkenburgh 1985, 1988, 1989; Viranta and

Andrews 1995; Wesley-Hunt 2005). Modern carnivore species from sites located in diverse environments were classified into body size and feeding categories to serve as references for dental variation in fossil materials. The combination of species present at each location was used to study ecological variation between carnivores from different habitats (Van Valkenburgh 1988). These modern carnivore guilds differed in composition

129 between various habitats, especially in terms of the number of species in each dietary class (Van Valkenburgh 1988). In particular, the number of species in dietary categories such as flesh-specialists and bone-crackers varied in different regions. Temperate environments lack insectivores. Omnivorous species are most prevalent in thickly vegetated forests with multiple canopy layers, and are also found in seasonal environments (Viranta and Andrews 1995).

During the Cenozoic, different species have occupied the ecological roles of hypercarnivore and omnivore, and at times, some of the roles were unfilled (Wesley-Hunt

2005). Analysis of prey capture and locomotion showed that carcass availability was likely to have differed in South and East Africa during the Plio-Pleistocene due to the presence of different morphotypes of carnivores in the two regions (Lewis 1995, 1997).

The extinction of carnivores as the African guild transitioned to its modern form may have provided opportunities for hominins (Lewis 1997). Analysis of character traits within a set of co-occurring carnivores may also show character displacement, in which an increase in dissimilarity helps to minimize overlap in resource use and competition

(Mooney and Cleland 2001). Character displacement has been found in the European

Pleistocene in the traits of felids and canids (García and Virgós 2007), and in Miocene hyenas (Werdelin 1996). Character displacement may also occur as a result of immigration of a new species into a guild (Ricklefs and Schluter 1993).

Dental morphology and Carnivore Feeding Behaviors :

Carnivore dental morphology relates to diet, phylogeny and to prey capture style.

Carnivores consume animal parts including skin, meat and bone, as well as plants. Blade-

130 like structures are used to cut through skin and bone, while large, blunt-topped cones are used to break bone (Lucas and Luke 1984). Wide, flat teeth are used for grinding.

Canines :

The canines are used for stabbing and slicing. Their shape relates in part to prey- killing behavior as well as to activities such as bone-cracking. Canines may be used to stab and hold prey, as in modern felids and mustelids, or to open slash wounds, as in canids (Biknevicius and Van Valkenburgh 1996). With a base that is approximately equal in length and width, the canines of modern felids are able to resist strain from all directions caused by struggling prey (Biknevicius and Van Valkenburgh 1996). Felid canines are stronger than those of canids, especially on the anteroposterior axis (Van

Valkenburgh and Ruff 1987). Strength on the mediolateral axis is related to bite force

(Van Valkenburgh and Ruff 1987). Modern hyaenid canines are strong in the anteroposterior and mediolateral axes (Van Valkenburgh and Ruff 1987). The round shape of the canine base helps resist the stress of bone-cracking (Van Valkenburgh and

Ruff 1987, Biknevicius and Van Valkenburgh 1996). Canines shaped like those of modern canids encounter stress in a single direction when used for slashing prey

(Biknevicius and Van Valkenburgh 1996). Saber-toothed felids used their upper canines to slash flesh, while the incisors may have been used to grip the flesh and prevent damage to the canines (Biknevicius et al. 1996).

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Premolars:

Modern hyenas use their premolars to crush bone. The rounded shape helps in withstanding stress (Lucas 1979). Premolars that are relatively large compared to body mass indicate a bone-crushing adaptation (Van Valkenburgh 1989). Felids may also use their premolars to break very small bones (Biknevicius and Van Valkenburgh 1996,

Schaller 1972, Blumenschine 1987). Other carnivores that consume bone, such as canids and mustelids, use postcanine teeth (Biknevicius and Van Valkenburgh 1996). Rounder premolars are often associated with diets that include large portions of non-vertebrate foods, as well as bone crushing (Van Valkenburgh 1989). Ursids, in contrast to the general pattern, have very small premolars.

Carnassials:

4 The carnassials, P and M 1, are used to slice flesh. Cusps are modified on both carnassial teeth into slicing blades that shear past each other when the jaw is closed.

Cutting efficiency is increased by reduction of the protocone and enlargement of the metastyle in the upper fourth premolar (Biknevicius and Van Valkenburgh 1996). The metaconid and talonid of the first lower first molar are reduced to increase cutting function (Biknevicius and Van Valkenburgh 1996). Relatively longer trigonid blades compared with the length of the first lower molar are associated with greater proportions of meat in the diet.

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Post-carnassial Molars:

Canids and mustelids use post-carnassial molars to crush hard and tough substances such as bones (Ewer 1973, Riley 1985, Biknevicius and Van Valkenburgh

1996). The protocone is also involved in crushing. Crushing specializations include blunt cusps able to withstand stress and a basin into which the tooth occludes (Lucas 1994).

The upper M 1 is expanded in mustelids as a specialization for crushing (Biknevicius and

Van Valkenburgh 1996). Modern hyaenids have reduced post-carnassial dentitions, using their premolars for bone crushing. Omnivores and species with significant non-vertebrate components in their diet use the post-carnassial area and lower carnassial talonid as a crushing and grinding area (Van Valkenburgh 1989).

Body Size:

Body size is an important determinant of prey size and competitive interactions

(Carbone et al. 1999, Van Valkenburgh 2001). Body size is also an important factor in carnivore competition. It is particularly important when determining whether an animal is able to retain control of a kill (Van Valkenburgh 2001). Larger carnivores are able to retain control over or steal carcasses from smaller animals, though packs of smaller carnivores are sometimes able to keep control (Eaton 1979, Lamprecht 1978, Van

Valkenburgh 2001). Smaller, solitary carnivores may lose carcasses (Van Valkenburgh

2001, Eaton 1979). Carnivores that weigh less than 10 kg in body mass are more likely than larger species to be killed by other carnivores (Caro and Stoner 1993). Among modern species, large hyenas such as Crocuta crocuta may steal kills from Lycaon pictus

or Acinonyx (Creel and Creel 1996, Fanshawe and Fitzgibbon 1993, Kruuk 1972, Mills

133 and Biggs 1993, Schaller 1972). Panthera leo and Crocuta crocuta may steal each others’ kills, with the outcome dependent on the number of individuals present (Cooper

1991).

Prey size follows similar patterns. Large carnivores and group hunters are able to kill larger prey as well as small animals, while small carnivores concentrate on smaller animals (Ewer 1973, Gittleman 1985, Mills 1990, Caro 1994, Lewis 1997). For extant carnivores, large prey over 300 kg is occasionally captured by lions and tigers (Schaller

1972; Seidensticker and McDougal 1993, Pienaar 1969). These species also take smaller prey. Many species, including hyenas, wild dogs, cheetah, leopard, lions and tigers, hunt prey from a middle range of body mass (20-300 kg) (Kruuk and Turner 1967; Lewis

1997; Schaller 1973). Carnivores over 21.5 kg in body mass tend to prey upon animals that weigh more than 45% of their own body mass (Carbone et al. 1999). Small prey under 20 kg may be hunted by wolves, jackals and hunting dogs, while some the diets of some jackals and striped and brown hyenas include substantial amounts of plant materials

(Lewis 1997).

Species within a guild that have the same feeding adaptations and the approximately the same body size are likely to have been competitors. For instance, the hypercarnivorous felids Acinonyx pardinensis , Panthera gombaszoegensis , and Panthera pardus are thought to have been in the same size range, and to have been competitors during the European early Pleistocene (García and Virgós 2007). Similar principles would have applied to the East Asian and East African carnivore guilds. Carnivores of similar size and feeding adaptations would have played similar roles in the carnivore guilds of East Asia and East Africa (though there may have been differences in their

134 locomotor adaptations). Hominins, as a new member of the carnivore guild in East Asia, would have also been competitors for prey. Interspecific competition for prey and theft of carcasses affects the distribution of modern carnivores. In places where there are large populations of spotted hyenas and lions, populations wild logs ( Lycaon pictus ) are smaller compared with places where lions and hyenas are less common (Creel and Creel

1996). There is more competition between hyenas and wild dogs in open habitat where carcasses are easily located (Creel and Creel 1998). The effects of interspecific competition may be particularly important for medium-sized carnivores (Creel and Creel

1998). The addition of hominin scavengers to the carnivore guild would have affected the distribution of other carnivores, particularly those from which they stole carcasses.

Hominin Meat-Eating:

Hominin diets shifted during the late Pliocene to include more animal products.

Evidence for this dietary shift includes archaeological assemblages with cut-and percussion marked bones, which show that hominins used tools to access animal meat and marrow (Bunn and Kroll 1986; de Heinzelin 1999; Semaw 2000; Semaw et al. 2003).

The co-occurrence of carnivore and hominin damage on the same bones shows that these resources were contested with other members of the carnivore guild (Blumenschine 1995;

Dominguez-Rodrigo et al. 2005; Potts, 1988; Shipman 1986). However, while previously, it was thought that sites in Olduvai Gorge represented assemblages collected by hominins (e.g., Leakey 1971), many of the assemblages in Bed I have been identified as produced by carnivore activities (Domínguez-Rodrigo 2007a, b, Egeland 2007, Faith et al. 2009). An important exception is FLK Zinjanthropus , which shows signs of being a

135 processing location (Domínguez-Rodrigo and Barba 2006, Faith et al. 2009, Bunn and

Kroll 1986). Formerly, marks interpreted as toothmarks were used to support the idea that

FLK Zinj represented a site with substantial carnivore involvement in a sequence of carnivore-hominin-carnivore (Blumenschine 1995). Although the matter is contested, reanalysis shows that biochemical marks were mistaken for toothmarks, supporting the idea that hominins acquired and butchered the carcasses at this site (Domínguez-Rodrigo and Barba 2006).

Factors Affecting Carcass Availability :

Feeding adaptations of carnivores and carcasses

The relative availability of scavengeable carcasses can be inferred from carnivore hunting and feeding adaptations (Marean 1989). Hypotheses have focused on the roles of flesh-slicing and bone-crushing carnivores in particular. Low numbers of bone-crackers in a carnivore guild would lead to greater carcass availability, especially of parts such as heads and marrow (Blumenschine 1987, Blumenschine et al. 1994). Modern spotted hyenas ( Crocuta crocuta ) can consume an entire carcass. Large numbers of bone- cracking hyaenids may have been serious competition for scavenging hominins (Turner

1992). Competition from bone-cracking hyaenids could have been significant whether the hyaenids came from a single species or from multiple bone-cracking species, provided the animals were abundant. Researchers have speculated that saber-toothed felids would have left useable meat and bone on their kills that hominins could have exploited (Ewer

1954, Blumenschine 1987, Marean 1989, Arribas and Palmqvist 1999). However,

Marean and Ehrhardt (1995) found that Homotherium from North America defleshed

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long bones and transported skeletal elements away from the kill site to be gnawed. Based

on this evidence, they believe that Homotherium would have provided only “moderate” amounts of scavengeable meat for hominins (Marean and Ehrhardt 1995).

Carnivore body size

Predator body size would have impacted the likelihood of successful scavenging

(Van Valkenburgh 2001). It would have been more difficult for hominins to compete against large predators directly. Larger predators also tend to take larger prey. However, a large carcass is also more likely to be stolen and is more difficult to defend

(Biknevicius and Van Valkenburgh 1996). Carnivore behaviors such as pack-hunting, tree-caching and habitat preference would also affect the likelihood of hominin scavenging success. Pack-hunting carnivores are able to hunt larger prey than their body size would normally permit (Van Valkenburgh 2001). Prey cached in trees could have been raided by hominins able to climb trees (Cavallo and Blumenschine 1989).

Habitat and carnivore competition

Carcass availability on different areas of the landscape may have varied due to the habitat preferences of the local carnivore populations. Riparian woodlands are thought to be low-competition settings because there are few closed-habitat large African carnivores, while open settings are thought to be higher risk settings, with high competition from bone-crushing competitors (Blumenschine et al. 1994). Behavioral studies on modern African carnivores indicated that competition among carnivores is higher in open habitats (Kruuk 1972, Schaller 1972, Sinclair 1979). Carcasses were

137 available for longer periods of time in riparian woodlands in the Serengeti compared with open grasslands (Blumenschine 1986). Kills are also more easily spotted in open area, where the presence of vultures may alert carnivores to the carcass (Cavallo 1997). Study of carcasses at the Maasai Mara and Tsavo shows that carcasses were consumed within hours in open country, often with many carnivores present (Domínguez-Rodrigo 2001).

In woodlands, carcasses remained on the landscape longer because it took hyenas, which consume a carcass completely, longer to find them (Domínguez-Rodrigo 2001).

Behavioral differences in lions were observed in different habitats outside the Serengeti, with lions remaining longer and consuming more of the carcass when it was located in riparian woodlands (Domínguez-Rodrigo 2001). Lions may transport carcasses to shrublands to feed on it over several days (Rudnai 1973).

However, much research on carcass availability has been based upon East African savannas, which tend to be drier than savannas in other regions. Virunga, in central

Africa, is less seasonal, has more tall grass and has more resident (as opposed to seasonally migrating) ungulates (Tappen 1995). Due to the environmental differences, scavenging opportunities, as inferred from bone deposition, differ in Virunga. There, more bone deposition occurs in open grassland compared with woodland riverine areas

(Tappen 1995). This may be ultimately traced to differences in habitat use by lions, which hunt in the tall grass at Virunga, but hunt in more wooded areas in the Serengeti

(Tappen 1995). More bones tend to be found where lions are the dominant predator, and where there were few hyenas (Tappen 1992, 1995). Seasonality also affected carcass availability. Whereas the Serengeti has a seasonal abundance of carcasses, scavengeable remains are a more constant, low-level food source at Virunga (Tappen 1995).

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Dominguez-Rodrigo (2001) theorized that competition is greater in the more arid

savannas and that resources there are used more completely. Ancient ecosystems in

which the dominant predators did not crush bone and in which substantial prey biomass

occurred would have provided the most scavenging opportunities (Tappen 1995).

Carcass availability and persistence on the landscape is also affected by the

number of carnivores present. When more carnivores are present in the fauna, carcasses are more likely to be consumed completely (Ayeni 1975, Blumenschine 1986, Kruuk

1972, Schaller 1972, Van Valkenburgh 2001). Where there are fewer carnivore species present, or where their territories do not overlap, competition levels are expected to be low and carcasses may persist longer or may be consumed only by a single species.

Bone assemblages accumulated by hyaenids occurring together with stone tools provide evidence that hominins and carnivores foraged in the same locations, though some of these assemblages were primarily accumulated by carnivores. Olduvai Bed I contained a variety of landscapes that included marshland (Hay 1976, Bonnefille 1984,

Peters and Blumenschine 1995, Domínguez-Rodrigo 2001) and riverine forest (Sikes

1994). Paleoenvironmental data from carcass butchery sites from the Okote member show hominin activity in a variety of different habitats including swamps, open grassland, wooded areas, and gallery forests (Pobiner 2008).

Habitat differences, specifically the prevalence of open or closed areas, may contribute to differences in competition for carcasses in Africa and East Asia. Of the East

Asian large carnivores, Panthera pardus (leopard) and Panthera tigris (tiger) are both

closed habitat animals. Modern Cuon alpinus inhabits forests, alpine areas, and scrub

jungle (Nowak 1999) while Ursus thibetanus lives in deciduous forests and brush areas

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(Nowak 1999). There is little evidence on competitive patterns of the carnivores within

closed habitats, but competition does influence carnivore behavior. Tree-caching in East

Asian leopards does not occur in the absence of other predators, such as P. tigris

(Muckenhirn and Eisenberg 1973). In an Indian tropical forest, tigers, leopards and dholes coexist as predators by hunting different species and sizes of prey (Karanth and

Sunquist 2000). These predators used the same forest habitats, and no predator excluded another from the habitat (Karanth and Sunquist 2000).

Prey Size

The ideal prey size for hominins has been disputed. Based on the size differences

between hominin-marked and carnivore-marked bones, Monahan (1996) believed that

hominins at Olduvai Bed II primarily foraged on animals of Bunn’s size classes 3 and 4

while the carnivore-marked bones were from animals of sizes 1 and 2. Okote faunas

contained similar numbers of cut marked bones of mammals of size classes 1-2 and 3-4,

while sizes 5-6 had few marks (Pobiner 2008). However, these findings are difficult to

interpret because the probability of cut marks being preserved may depend on the size

class of the mammal, but investigators do not agree on whether sizes 1 and 2 or sizes 3

and 4 should display more cut marks. The number of cutmarks on mammals sized 1-4

may signal either preference or greater availability as scavengeable material relative to

sizes 5-6 (Pobiner 2008).

Based on the number of potential predators on hominins and the difficulty of

retaining carcasses, preference for small prey would have allowed hominins to butcher

rapidly for transport or consumption (Brantingham 1998). Small carcasses could also

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have been obtained from leopard tree-caches (Cavallo and Blumenschine 1989). Data

from Olduvai Bed II also suggest that hominins were able to obtain relatively complete

carcasses (Monahan 1996). Studies of hominin-transported assemblages from Koobi Fora

and Olduvai Bed I suggest that hominins were intermediate between top predators and

confrontational scavengers in their bone transport (Brantingham 1998). Later analysis of

Olduvai Bed I, however, showed no evidence that hominins selectively transported small

or large sized animals (Faith et al. 2009).

Scavenging, Carnivores and Dispersal:

Use of animal tissues would have been advantageous for a dispersing hominin, since it would have allowed occupation of temperate regions with fewer year-round plant

resources, and would have mitigated the difficulties of using unfamiliar and possibly

toxic plant materials. Percussion damage on Nihewan bones dated to 1.66 Ma shows that

hominins extracted marrow from them as in Africa (Zhu et al. 2004). Arribas and

Palmqvist (1999) argued that the unconsumed flesh and bone left behind by saber-toothed

felids would have been an important resource for dispersing hominins in unfamiliar

territory.

Carnivore guild of East Africa :

African Carnivores :

The species diversity of African carnivores was reduced after 3.3 Ma, with an extinction rate peak at 3.0 and a steady rate of extinctions after 1.8 Ma (Werdelin and

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Lewis 2005). The species that went extinct may have included a large proportion of habitat specialists (Werdelin and Lewis 2005, Peters et al. 2008).

African Canidae :

Canids are relatively rare in the East African fossil record, a fact which Werdelin and Lewis (2005) attribute to their preference for open habitats, rather than the more vegetated lakeside habitats of Lake Turkana. Specimens attributed to Canis mesomelas from Koobi Fora and C. mesomelas from Olduvai are dentally distinct and probably came from different species (Werdelin and Lewis 2005). Canis lycaonoides , found at Olduvai

Bed II, is thought to be part of the Lycaon lineage (Martínez-Navarro and Rook 2003).

Prototocyon recki is found at Olduvai I and is a possible ancestor of Otocyon (Werdelin and Lewis 2005).

African Ursidae :

Ursid specimens have been found at Hadar and in the Tulu Bor Member. Ursids were present in Africa from the Late Miocene to the Late Pliocene (Werdelin and Lewis

2005).

African Hyaenidae :

Hyaena makapani is the ancestor of Hyaena hyaena . H. makapani has fewer bone cracking adaptations and is smaller than H. hyaena . Other bone-cracking hyaenids include Crocuta ultra , Crocuta dietrichi , and Crocuta crocuta . Chasmaporthetes , a hyaena with a more sectorial and less bone-cracking dentition than Crocuta and Hyena ,

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has been found at Olduvai. Pachycrocuta has not been found in East Africa during the

time period considered (Werdelin and Lewis 2005).

African Felidae :

Large felids in Africa include Dinofelis , Homotherium , and Megantereon, as well

as Panthera pardus, P. leo and Acinonyx . Megantereon and Homotherium may have become extinct after the Okote Member, while Dinofelis might have survived longer

(Werdelin and Lewis 2005). Modern pantherines Panthera leo and Panthera pardus appear in Olduvai Bed I (Werdelin and Lewis 2005). Acinonyx fossils are larger than the

modern species (Werdelin and Lewis 2005). Werdelin and Lewis (2005) note that there is

no simultaneous extinction of sabertooth felids at 1.5 Ma.

The species identification of many specimens of Megantereon is uncertain

because isolated teeth do not distinguish between M. ekidoit and M. whitei (Werdelin and

Lewis 2002). Megantereon is inferred to have been a closed habitat animal based upon its

short distal elements implying a lesser degree of cursoriality compared with modern open

habitat carnivores (Marean 1989, Lewis 1997, Werdelin and Lewis 2001).

Homotherium was a very large sabertoothed felid. Its large body size would have

enabled it to hunt the largest prey animals. Based on post-cranial remains, it was

relatively cursorial (and thus may have lived in more open habitats) and had slightly

reduced prey grappling capabilities (Lewis 1997). Homotherium latidens postcranial

material from an early Pleistocene site in Spain is consistent with a relatively cursorial,

cantering gait, while claw reduction was interpreted as an adaptation to improve traction

while running (Antón et al. 2005). It may have hunted in packs (Lewis 1997, Marean and

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Ehrhardt 1995, Antón et al. 2005). Pack-hunting behavior has been disputed on the

grounds that Homotherium does not display healed fractures which would indicate group

provisioning (Rawn-Schatzinger 1992). Although it has been suggested that carcasses left

by this animal would have provided substantial scavengeable meat (Ewer 1954,

Blumenschine 1987, Marean 1989, Arribas and Palmqvist 1999), a Homotherium den in

North America shows that the animal consumed most of the flesh from the bones

(Marean and Ehrhardt 1995). Based on the relatively gracile postcranial skeleton of

Homotherium latidens , Antón et al. (2005), conclude that though the canines would have made Homotherium a very efficient hunter of large prey, it would not have taken prey significantly larger than a lion. The reconstruction of Homotherium latidens as relatively gracile is in contrast to reconstructions of Homotherium body mass based on dental

estimators (Antón et al. 2005).

Dinofelis probably frequented more closed habitats, having short distal elements

(Marean 1989, Lewis 1997, Werdelin and Lewis 2001). Dinofelis was able to rotate its

forelimb, and was found to be similar in other post-cranial characteristics to modern prey-

grappling felids (Lewis 1997). Dinofelis , which was larger than modern leopards, may

have been too heavy to regularly cache food in trees (Lewis 1997). It was probably able

to hunt larger prey than modern felids. The upper canines of Dinofelis were similar to

those of P. leo and P. tigris , probably due to convergence (Werdelin and Lewis 2001).

However, D. piveteaui and D. aronoki , which are found in this dissertation sample, have

features typical of machairodonts (Werdelin and Lewis 2001).

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Panthera pardus has been identified as a potential source of size 1 and 2 carcasses

for hominin scavengers living in more wooded settings (Cavallo and Blumenschine

1989).

African Mustelidae :

The mustelids include species of Enhydriodon . Some members of this lineage have lion-sized postcrania (Werdelin and Lewis 2005, Lewis 2008). Torolutra is found at

Koobi Fora and West Turkana, Aonyx is found at Olduvai Bed I and in the KBS member, and Mellivora is found at Koobi Fora (Werdelin and Lewis 2005).

African Viverridae :

Several large species of viverrid were present in East Africa, including

Pseudocivetta ingens found at Koobi Fora and Olduvai Bed II (Werdelin and Lewis

2005). Werdelin and Lewis (2005) found a shift from smaller more arboreal forms to larger, terrestrial species in later assemblages.

African Herpestidae :

Many of the species present are too small to be included in this study. Werdelin and Lewis (2005) note that terrestrial mongooses are more common when there are other paleoclimatic indicators of the spread of savannas.

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Carnivore Guild of East Asia :

Asian Carnivores:

The patterns of the East Asian Plio-Pleistocene carnivore guild as a whole are not

well known. Carnivore families and species are described below. Some of the Chinese

species described here represent endemic species, such as the giant panda, while others

are similar morphologically to species found in Europe and may represent geographic

variants. Many of the species are poorly known compared with African carnivores, due in part to fewer comparative studies and less fossil material. The carnivore guild as a whole shows turnover during the Pleistocene (Qiu 2006), though it is not clear that there was ecological change along with the taxonomic change. In North China, many new taxa appeared at 2.6 Ma. Some were immigrants from North America, while others evolved in situ. The new species and the faunal makeup of the North China faunas persisted until about 1.3 Ma (Qiu 2006). At 1.3 Ma, faunal turnover occurred again (Qiu 2006).

Chasmaporthetes and Eirictis went extinct, while more derived species evolved from earlier variants (such as species of canids and saber-toothed felids). Current evidence places the arrival of hominins between the two turnover events. The causal connection between the carnivore guild turnover and the arrival of hominins may be due to hominin competition with carnivores, or to hominin opportunism in colonizing an area with a guild in transition due to other factors.

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Asian Canidae :

Several genera of canids were present in East Asian Plio-Pleistocene. Among

these are Canis chihliensis , which Teilhard and Piveteau (1930) characterized as similar to the European Canis etruscus . Canis variabilis is reported from Gongwangling, and is described as a relatively small species (Qiu et al. 2004). Canis teilhardi, Canis longdanensis and Canis brevicephalus are large-sized canids from Longdan. Sinicuon

(formerly Cuon ) cf. dubius has also been reported from Longdan. Vulpes chikushanensis is thought to be ancestral to V. corsac (Qiu et al. 2004). Nyctereutes sinensis shared many features with the European species N. megamastoides (Tedford and Qiu 1991).

Asian Hyaenidae :

Most of the hyena species found in East Asia during this time period are synonymous with Pachycrocuta brevirostris (Werdelin and Solounias 1991).

Pachycrocuta licenti was a very large hyena and would have been able to consume carcasses more completely than extant C. crocuta (Turner and Antón 1996). Its size would have facilitated the hunting of large prey. Turner and Antón (1996) suggest that its size would have helped in preying upon medium sized ungulates and their young, but that group action would have been important, particularly since running would have been energetically costly. It has relatively short distal limbs compared to modern hyenas, indicating less cursoriality or an adaptation for carcass transport (Turner and Antón

1996). Pachycrocuta is commonly found in Asian sites. It was the main bone accumulator at Riwat (Dennell et al. 2008), Longgupo (Wood and Turner 1995), and

Zhoukoudian (Boaz et al. 2000). Other hyaenids present include Chasmaporthetes , which

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is thought to have an ecological role similar to a large wild dog (Galiano and Frailey

1977). Chasmaporthetes is not a bone cracking hyena, and has a cursorial skeleton

(Kurtén and Werdelin 1988). Crocuta honanensis is also present.

Asian Felidae :

Megantereon has been recorded in many sites. It was an ambush predator.

Isotopic studies of specimens from Venta Micena showed that it focused on forest browsers and that it was ecologically separated from Homotherium , which hunted grazing animals presumably found in open habitats (Palmqvist et al. 2003). Palmqvist et al.

(2007) found that Chinese early Pleistocene Megantereon specimens were similar to M. cultridens in dental dimensions. Other species present in East Asia include Sivapanthera , an extinct cheetah larger than modern Acinonyx (Qiu et al. 2004), Panthera palaeosinensis and Panthera tigris . Smaller felids include Felis teilhardi , which is similar to Lynx in morphology, though much smaller. Lynx shansius is also present, but larger than living Lynx lynx and similar to Lynx issiodorensis (Qiu et al. 2004). The other small felids grouped in Felis sp. are poorly understood (Qiu 2006).

Asian Ursidae :

Many of the Ursus specimens in the East Asian Plio-Pleistocene are allocated to

Ursus sp. or to Ursus cf. etruscus due to lack of definitive materials and taxonomic study, though they are similar to the extant Ursus thibetanus (Zheng 2004). However, Ursus cf. etruscus from the Nihewan was smaller and more primitive than the European form, while the specimen from Gongwangling was probably part of Ursus thibetanus (Qiu

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2006). There are also species of Ailuropoda in China, including A. microta , A. wulingshanensis and A. melanoleuca . Ailuropoda specimens are common in southern

China and are only found in the northern region at Gongwangling (Qiu 2006).

Asian Mustelidae :

Most species from this family are poorly understood and have few fossil remains.

There are many fossils of meline species, including M. teilhardi and M. chiai in north

China (Qiu 2006). Lutrines were represented by Lutra licenti .

Summary :

This section discusses carnivore ecomorphology, interactions between hominins and carnivores and characteristics of the Plio-Pleistocene carnivores of East Asia and

East Africa. Ecomorphological analysis is based on morphological characteristics that are related to aspects of ecology, such as feeding adaptations, body mass, locomotion and prey capture. This project focuses on body mass and feeding adaptations, as derived from dental measurements. The groups of carnivores found in the same time period or region are referred to as guilds, which are groups of species that use resources in a similar way

(Root 1967). Ecological avatars are species that have similar adaptations and fill similar roles in the community.

Feeding adaptations are inferred by the morphology of the dentition. Canine shape is related to prey capture (stabbing or slashing), as well as to withstanding strain from other activities, such as bone-crushing. Premolars are used by hyaenids to crush bone. The

4 carnassials, M 1 and P , have blades that are used to slice flesh. Longer trigonid blades in

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the first lower molars are associated with increased proportions of meat in the diet. Post-

carnassial molars are used to grind hard and tough material.

Body size is an important determinant of prey size and competitive interactions

(Carbone et al. 1999). Larger carnivores can take larger prey, and retain or gain

possession of prey more often. Grouping behavior can facilitate the hunting of larger prey and retention of carcasses against larger but solitary competitors.

Hominin and carnivore interactions during the Plio-Pleistocene would have involved both predation and competition for carcasses. Competition occurred as hominin diets came to include more animal products during the late Pliocene. Bone-cracking hyaenids would have competed with hominins for carcasses, while sabertoothed felids may have produced carcasses with scavengeable material. Carcasses availability varies with the adaptations of the predators present. More carcasses would be found in areas with fewer bone- cracking predators. Animal products would have been helpful for dispersing hominins colonizing temperate regions with plants that would have had different characteristics than those in East Africa.

Species diversity among East African carnivores was reduced after 3.3 Ma (Werdelin and

Lewis 2005). Among the types of carnivores that went extinction were Ursids, which are not found during the focal time period. Also, the large, bone-cracking hyaenid Pachycrocuta was

extinct by this time. In East Asia, taxonomic turnover among North China carnivores has been

found at 2.6 Ma and 1.3 Ma (Qiu 2006). In contrast to Africa during this period, Pachycrocuta is

commonly found in East Asia during the Plio-Pleistocene. Ursus is also present throughout the

period. Differences between the carnivore species in Plio-Pleistocene East Asia and East Africa

and the implications for hominins will be explored in further chapters.

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Chapter 6: Carnivore Ecomorphology Methods

Overview :

The methods used to evaluate carnivore ecomorphology in East Africa and East

Asia are described here. Samples, measurements, composite animals, index choice, disparity categories and the analytical methods used are described. The analysis is described for 1) index values and then 2) category values.

Samples :

All available adult specimens from the order Carnivora from the focal sites in

East Asia and East Africa were measured (appendix tables 6.1-6.12). Species attributions

were based upon species lists for each site, and upon taxonomic revisions when available

(Deng et al. 2008; Dong et al. 2000; Flynn et al. 1991; Hu and Qi 1978; Huang and Fang

1991; Lin et al. 1978; Pu and Qian 1977; Qian and Zhou 1991; Qiu et al. 2004; Teilhard

and Piveteau 1930; Turner et al. 1999; Wang et al. 2007; Werdelin and Lewis 2005;

Werdelin and Solounias 1991; Werdelin personal communication; Zheng 2004; Zhu et al.

2008). When specimens were not available, information from the literature was used as a

supplement. Specimens labeled aff. or cf. were treated as full members of the species in

order to avoid generating too many possible species categories. Canis cf. mesomelas and

Canis mesomelas from the African fossil sample are exceptions. The specimens from

Lake Turkana (Canis cf. mesomelas ) are not conspecific with those from Olduvai ( Canis

mesomelas ), and neither is conspecific with the modern species (Werdelin, pers. comm.).

Samples were also measured from modern carnivores of the families Felidae,

Canidae, Ursidae, Hyaenidae, Herpestidae, Mustelidae and Viverridae (Tables 3.6-3.8,

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Tables 6.1-6.2). Dimensions measured on the modern taxa were used to show how each ecomorphological index related to differences in feeding adaptations. Modern carnivore specimens were measured at the National Museum of Natural History, Smithsonian

Institution, in Washington DC. Modern sample species were chosen to include dietary variants, with a focus on 1) larger species, and 2) genera that were represented in the fossil record. Carnivores had to be over 1 kg in body mass to be included. Information from a greater variety of taxa was drawn from the literature (e.g., Friscia et al. 2007; Van

Valkenburgh and Koepfli 1993) when necessary.

Sample sizes of some carnivore species are small. This is typical of carnivores, which are usually less numerous in faunas than herbivore species. All available fossil specimens were measured. Due to the small sample sizes, it is difficult to get a sense of the amount of variation in each index of the carnivore species. It is possible that some of the specimens measured are not representative of the values of the fossil species, or that the composite indices created from certain specimens do not correspond to the mean values for the species. Some species of carnivores may not be represented in the fossil record of either sites or in regions altogether. Small carnivores are less likely to be preserved in the faunas. Carnivores that were originally rare are less likely to have been preserved also.

There are very few carnivore species and specimens from the sites of

Xiaochangliang, Donggutuo, Majuangou, and Linyi. For the first three sites, the possible carnivore faunas are represented by proxy with the carnivores from the Nihewan Basin sensu stricto . This fauna dates from approximately the same time period and includes many species of carnivores. However, it is possible that the Nihewan fauna s.s . does not

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accurately represent the carnivore faunas from the time period of Majuangou,

Donggutuo, and Xiaochangliang.

Table 6.1 Modern Felidae samples. References: 1. Van Valkenburgh 1988 2. Nowak 1999 Species Dietary Class Specimens Sex Acinonyx jubatus Carnivore (1) USNM 361490 Female USNM 540003 Unknown Lynx lynx Carnivore (2) USNM 198468 Female Panthera leo Carnivore (1) USNM 182309 Female USNM 184816 Male Panthera pardus Carnivore (1) USNM 156284 Male USNM 162147 Male Panthera tigris Carnivore (1) USNM 253285 Unknown USNM 278470 Male Uncia uncial Carnivore (1) USNM 084091 Female USNM 176048 Female

Table 6.2 Modern Hyaenidae samples. References: 1. Van Valkenburgh 1988

Species Dietary Class Specimens Sex Crocuta crocuta Meat/Bone (1) USNM 163103 Female USNM 163104 Male USNM 367384 Male USNM 367385 Female Hyaena hyaena Meat/Bone (1) USNM 183040 Female USNM 182045 Male USNM 253282 Unknown USNM 318112 Female Parahyaena brunnea Meat/Bone (1) USNM 221088 Male USNM 267891 Female USNM 296135 Female USNM 429177 Male

Carnivore adaptations: Diet and Prey Properties :

Modern carnivore dietary classifications are necessary to determine how each index relates to ecological categories. Modern carnivores that represent certain dietary types were used to determine cut-offs for disparity categories and to assign ecotypes

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(chapter 3). Dietary classifications were based on previous work. The possible categories

include carnivore, meat/bone feeder, omnivore, insectivore and herbivore. For Canidae,

species were divided into highly carnivorous, which take prey larger than themselves, or

which have high proportions of meat in their diets, moderately carnivorous, and

omnivorous (Van Valkenburgh and Koepfli 1993). Moderately carnivorous canids

consume vegetation as well as vertebrate prey (Van Valkenburgh and Koepfli 1993).

Modern Ailuropoda is an herbivore. Information from the literature as well as from

previous ecomorphological studies was used in categorizing modern species.

Indices:

Ecomorphological indices were chosen to contain information about feeding adaptations, hunting behavior and body mass (tables 6.3 and 6.4). These indices are based on craniodental materials. The indices pertain to specifics such as the amount of meat in the diet, the relative occlusal area devoted to grinding or slicing, the degree of adaptation for hard object feeding or the style of prey capture. The utility of each index varies for each family. Ecological interpretation was done within families. A measure of third premolar size was not used in this project because of the small amount of available data. However, as the main bone-crushing tooth, its relative size is important for comparing hyaenid adaptations. Due to the importance of premolars, measurements of the fourth lower premolar (RPS and P4WL) were included in the indices.

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Table 6.3 Ecomorphological measurement descriptions. The measurements used are listed with a description of the calculation method. Areas were calculated as the maximum length multiplied by the maximum width of the tooth unless otherwise indicated. Indices in this table were derived from the work of Van Valkenburgh 1988, Van Valkenburgh and Koepfli 1993, Friscia et al. 2007 and Sacco and Van Valkenburgh 2004. Index Definition Measurement RBL Relative blade length of Length of the trigonid divided by the the first molar molar anteroposterior length of the tooth RGA Relative grinding area of Summed areas of all grinding areas present, the mandibular dentition including the M1 talonid, M2, and M3, divided by the length of the blade RUMGA Relative grinding area of Summed areas of all grinding areas present, the maxillary dentition including M 1 and M 2, divided by the length of P 4 PROTO Size of the protocone Maximum width of P 4 measured at the protocone relative to the length of P 4 divided by the maximum anteroposterior length of the P 4 P4WL Lower P4 shape Width of p 4 divided by length

CWL Relative elongation of the Upper canine width divided by length upper canine UM21 Relative sizes of the Square root of the area of the M 2 divided by the upper molars square root of the area of the M 1 LBM Log Body Mass Body mass estimated using regression of the first lower molar; Log 10 of the predicted body mass. RPS Relative size of the lower Lower P4 width divided by the cube root of body P4 mass

Table 6.4 Ecological traits and ecomorphological indices. The traits associated with each index are listed. References: 1. Van Valkenburgh 1988; 2. Van Valkenburgh and Koepfli 1993; 3. Friscia et al. 2007; 4. Van Valkenburgh 1989; 5. Biknevicius and Van Valkenburgh 1996, 6. Carbone et al. 1999. Index Indications RBL Amount of the dentition devoted to slicing; Highly carnivorous species have greater relative blade lengths, compared to omnivorous or less carnivorous species (1) RGA and Relative amount of slicing compared with the amount of grinding area in the RUMGA molars. Highly carnivorous taxa have smaller relative grinding areas compared with omnivores (1, 2) PROTO The protocone tends to be large in insectivores and omnivores. Reduced protocone size increases slicing efficiency while a large-sized protocone aids bone cracking (1, 2, 3, 4). P4WL This index measures the relative shape of the premolar. Rounder premolars tend to be better for breaking material such as bone (1, 5) CWL A round shape indicates that prey is killed with a single strong bite. Narrow

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Index Indications canines are used to slash prey (1). UM21 This is used to assess the amount of dentition devoted to slicing compared with grinding. Taxa that are more omnivorous have greater amounts of the dentition devoted to grinding (1). LBM Larger carnivores are able to hunt a wider range of prey animals (6) RPS Premolars are relatively larger in bone-crushing and in more carnivorous species (4).

Measurements and Calculations :

Adult specimens were measured using digital calipers. Modern adult samples were selected based upon completeness of dentition. The modern measurements were averaged to create a composite representation. Fossil taxon measurements were also combined to create composite animals.

Composites :

A composite is an index value with input from more than one specimen for a species. Composites were necessary because many fossils are fragmentary or consist of isolated teeth. Specimen averages have been used to create composites in previous studies (Van Valkenburgh 1988; Lewis 1995, 1997, Wesley-Hunt 2005). Composites were created when single specimens did not have all the dimensions necessary for an index. For example, a composite may be made for the index C1m1 with the upper canine area taken from one specimen, and the m 1 area taken from another specimen of the same species, or indices such as RUMGA might be constructed from isolated teeth of the same species at a site. Composites as average representations of a species were also made. An example of this type might include an RBL ratio value constructed from the average of all available blade lengths from a species divided by all available m 1 lengths to create an

156 average representation of that species’ RBL value. Composites were checked against the index values for individual specimens. When possible, composites were made from specimens from the same species at the same site. When that was not possible, composites were made within the same species within East Asia or East Africa. The modern composites were made to increase the ease of comparison of indices across all modern species. In the future, additional fossil materials may permit the study of species from specific localities which were made into composites for this study. More complete specimens may be compared individually to look at trends within species, as well as whether composites here accurately reflect the species’ adaptations.

Analysis of the Index Values :

Values for the indices were calculated for each available species at a site (tables

6.5 and 6.6). Analysis of the indices focused on determining whether the species were avatars, that is, whether taxa from different species had comparable craniodental adaptations. The indices were also used to characterize differences between species ecologically. The carnivore guilds from East Asian and East African were compared in terms of their ecological composition. The East Asian guild was also compared over time to look at whether there was ecological turnover.

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Table 6.5 Index values for fossil carnivore species. Canis cf. mesomelas from the Upper Burgi Formation is not conspecific with Canis mesomelas from Olduvai. Neither form is conspecific with the modern C. mesomelas . The original labels are retained, however, pending taxonomic revision. Species Family Site RBL RGA RUMGA RPS Canis brevicephalus Canidae Longdan 0.731 0.732 0.866 2.975 Canis cf. mesomelas Canidae Kalochoro 0.682 ? ? 1.733 Canis cf. mesomelas Canidae KBS ? ? ? 2.036 Canis cf. mesomelas Canidae Natoo ? ? ? ? Canis cf. mesomelas Canidae Upper Burgi 0.667 ? 0.917 ? Canis chihliensis Canidae Nihewan 0.691 0.787 0.865 2.522 Canis longdanensis Canidae Longdan 0.684 ? 0.831 2.569 Canis lupus Canidae Nihewan 0.703 0.763 ? 2.545 Canis lycaonoides Canidae Olduvai II 0.720 0.751 0.783 2.709 Canis mesomelas Canidae Olduvai I 0.648 0.892 0.861 1.968 Canis palmidens Canidae Nihewan 0.565 ? 0.845 2.198 Canis sp. Canidae Donggutuo ? ? ? ? Canis sp. Canidae Linyi ? ? ? ? Canis sp. Canidae Upper Burgi ? ? ? ? Canis teilhardi Canidae Longdan 0.693 0.731 0.874 2.575 Canis variabilis Canidae Gongwangling 0.663 0.767 ? 2.174 Canis yuanmouensis Canidae Yuanmou ? ? 0.878 ? Cuon dubius Canidae Jianshi ? 0.587 0.833 2.105 Cuon dubius Canidae Longgupo 0.695 0.587 0.835 2.105 Cuon dubius Canidae Mohui ? ? 0.835 ? Nyctereutes cf. sinensis Canidae Longgupo ? ? ? ? Nyctereutes cf. sinensis Canidae Nihewan ? ? 0.976 ? Prototocyon recki Canidae Olduvai I 0.605 1.516 1.607 2.086 Sinicuon cf. dubius Canidae Longdan ? ? 0.835 ? Vulpes cf. zerda Canidae KBS ? ? ? ? Vulpes cf. zerda Canidae Upper Burgi ? ? ? ? Vulpes chikushanensis Canidae Longdan 0.630 0.950 0.991 1.706 Vulpes sp. Canidae Nihewan ? ? ? ? Acinonyx jubatus Felidae Olduvai II 1 1 ? ? Acinonyx sp Felidae Okote 1 1 ? ? Acinonyx sp Felidae Upper Burgi 1 1 ? ? Acinonyx sp. Felidae KBS 1 1 ? ? Caracal sp. Felidae Upper Burgi 1 1 ? ? Dinofelis aronoki Felidae Upper Burgi 1 1 ? 1.752 Dinofelis piveteaui Felidae Okote 1 1 0.153 1.733 Dinofelis sp. D Felidae Olduvai I 1 1 ? ? Dinofelis sp. D Felidae Olduvai II 1 1 ? ? Felis microta Felidae Longgupo 1 1 ? 1.609 Felis sp. Felidae Longgupo 1 1 ? ? Felis teilhardi Felidae Jianshi 1 1 ? 1.834 Felis teilhardi Felidae Longdan 1 1 ? 2.019 Felis teilhardi Felidae Longgupo 1 1 ? 1.834 Felis teilhardi Felidae Mohui 1 1 ? 1.834 Homotherium cf. crenatidens Felidae Longgupo 1 1 ? ? Homotherium cf. crenatidens Felidae Nihewan 1 1 0.112 ?

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Species Family Site RBL RGA RUMGA RPS Homotherium crendatidens Felidae Longdan 1 1 0.104 ? Homotherium crendatidens Felidae Jianshi 1 1 0.104 ? Homotherium davitasvili Felidae Haiyan 1 1 ? 1.805 Homotherium sp. Felidae Kaitio 1 1 ? ? Homotherium sp. Felidae Kalochoro 1 1 ? ? Homotherium sp. Felidae KBS 1 1 ? 1.216 Homotherium sp. Felidae Okote 1 1 ? ? Homotherium sp. Felidae Upper Burgi 1 1 ? ? Lynx shansius Felidae Longdan 1 1 0.206 2.016 Lynx shansius Felidae Nihewan 1 1 ? 2.033 Machairodus sp. Felidae Olduvai II 1 1 ? ? Megantereon nihowanensis Felidae Gongwangling 1 1 ? 1.719 Megantereon nihowanensis Felidae Longdan 1 1 0.216 1.998 Megantereon nihowanensis Felidae Nihewan 1 1 ? 1.910 Megantereon nihowanensis Felidae Yuanmou 1 1 ? ? Megantereon nihowanensis Felidae Haiyan 1 1 ? ? Megantereon sp. Felidae Chari 1 1 ? ? Megantereon sp. Felidae Jianshi 1 1 ? ? Megantereon sp. Felidae Kaitio 1 1 ? ? Megantereon sp. Felidae KBS 1 1 ? ? Megantereon sp. Felidae Linyi 1 1 ? ? Megantereon sp. Felidae Upper Burgi 1 1 ? ? Megantereon whitei Felidae Okote 1 1 0.196 1.737 Metailurus sp. Felidae Olduvai II 1 1 ? ? Panthera cf. leo Felidae KBS 1 1 ? ? Panthera cf. leo Felidae Upper Burgi 1 1 ? ? Panthera cf. palaeosinensis Felidae Longgupo 1 1 ? ? Panthera cf. pardus Felidae Longgupo 1 1 ? ? Panthera cf. tigris Felidae Gongwangling 1 1 ? ? Panthera leo Felidae Okote 1 1 ? ? Panthera leo Felidae Olduvai I 1 1 ? ? Panthera leo Felidae Olduvai II 1 1 ? 2.399 Panthera palaeosinensis Felidae Longdan 1 1 ? ? Panthera pardus Felidae Gongwangling 1 1 ? 2.495 Panthera pardus Felidae Jianshi 1 1 ? 2.432 Panthera pardus Felidae Olduvai I 1 1 ? ? Panthera pardus Felidae Olduvai II 1 1 ? ? Panthera pardus Felidae Upper Burgi 1 1 ? 2.279 Panthera pardus ? Felidae KBS 1 1 ? ? Panthera sp. Felidae Mohui 1 1 ? ? Panthera sp. Felidae KBS 1 1 ? ? Panthera sp. Felidae Upper Burgi 1 1 ? ? Panthera tigris Felidae Yuanmou 1 1 ? ? Sivapanthera linxiaensis Felidae Longdan 1 1 0.223 2.055 Sivapanthera pleistocaenicus Felidae Gongwangling 1 1 ? 2.035 Sivapanthera pleistocaenicus Felidae Jianshi 1 1 0.224 ? Sivapanthera pleistocaenicus Felidae Longgupo 1 1 ? ? Sivapanthera pleistocaenicus Felidae Nihewan 1 1 ? ? Atilax paludinosus Herpestidae Olduvai II 0.653 ? ? ?

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Species Family Site RBL RGA RUMGA RPS Herpestes primitivus Herpestidae Olduvai I 0.750 0.767 ? 2.397 Mungos dietrichi Herpestidae Olduvai I 0.550 1.750 ? 3.468 Chasmaporthetes cf Hyaenidae Nihewan ossifragus ? ? ? ? Chasmaporthetes nitidula Hyaenidae Olduvai II 0.827 0.238 ? 2.148 Chasmaporthetes progressus Hyaenidae Longdan ? ? 0.246 ? Crocuta cf. ultra Hyaenidae Chari ? ? ? ? Crocuta crocuta Hyaenidae Kalochoro ? ? ? ? Crocuta crocuta Hyaenidae Nariokotome ? ? ? ? Crocuta crocuta Hyaenidae Olduvai I 0.866 0.217 ? 2.802 Crocuta crocuta Hyaenidae Olduvai II ? ? ? ? Crocuta dietrichi Hyaenidae KBS ? ? ? ? Crocuta dietrichi Hyaenidae Upper Burgi 0.886 0.183 ? 2.558 Crocuta honanensis Hyaenidae Longdan 0.820 0.283 0.210 3.137 Crocuta honanensis Hyaenidae Haiyan 0.854 0.246 ? 3.360 Crocuta sp. Hyaenidae KBS 0.828 0.252 ? 3.270 Crocuta sp. Hyaenidae Okote ? ? ? ? Crocuta sp. Hyaenidae Upper Burgi ? ? ? ? Crocuta ultra Hyaenidae KBS 0.875 0.200 ? 2.795 Crocuta ultra Hyaenidae Lokalalei 0.859 0.217 ? 2.826 Crocuta ultra Hyaenidae Okote 0.844 0.197 ? 2.650 Crocuta ultra Hyaenidae Upper Burgi ? ? ? ? Hyaena cf. makapani Hyaenidae KBS ? ? ? ? Hyaena hyaena Hyaenidae Olduvai I ? ? 0.190 ? Hyaena hyaena Hyaenidae Olduvai II ? ? ? ? Hyaena makapani Hyaenidae Upper Burgi 0.797 0.329 0.346 2.866 Hyaena sp. Hyaenidae Kaitio 0.779 0.377 ? 2.778 Hyaena sp. Hyaenidae Kalochoro 0.783 0.369 ? 2.781 Hyaena sp. Hyaenidae KBS ? ? ? ? Hyaena sp. Hyaenidae Linyi ? ? ? ? Hyaena sp. Hyaenidae Okote ? ? ? ? Hyaena sp. Hyaenidae Upper Burgi 0.768 0.337 0.275 2.709 Pachycrocuta brevirostris Hyaenidae Gongwangling ? ? 0.182 ? Pachycrocuta brevirostris Hyaenidae Jianshi 0.827 0.273 0.238 3.460 Pachycrocuta brevirostris Hyaenidae Longdan 0.837 0.266 0.219 3.572 Pachycrocuta brevirostris Hyaenidae Longgupo 0.808 0.301 ? 3.252 Pachycrocuta brevirostris Hyaenidae Mohui ? ? ? ? Pachycrocuta brevirostris Hyaenidae Nihewan 0.840 0.196 0.218 3.397 Pachycrocuta brevirostris Hyaenidae Xiaochangliang ? ? ? ? Pachycrocuta brevirostris Hyaenidae Yuanmou 0.857 0.240 0.242 3.325 Pachycrocuta brevirostris Hyaenidae Haiyan ? ? ? ? Pachycrocuta sp. Hyaenidae Majuangou ? ? ? ? Pliocrocuta perrieri Hyaenidae Haiyan 0.804 0.306 ? 3.480 Aonyxini Mustelidae KBS ? ? ? ? Arctonyx cf. minor Mustelidae Longgupo 0.493 1.173 ? 1.676 Arctonyx collaris Mustelidae Mohui ? ? 1.020 ? Arctonyx collaris Mustelidae Jianshi ? ? 1.020 ? cf. Torolutra Mustelidae Okote ? ? ? ? Eirictis robusta Mustelidae Longdan 0.645 0.816 0.709 1.943

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Species Family Site RBL RGA RUMGA RPS Enhydriodon sp. Mustelidae Upper Burgi ? ? ? ? Lutra licenti Mustelidae Nihewan ? ? ? ? Lutra maculicollis Mustelidae Olduvai I ? ? ? ? Lutra maculicollis Mustelidae Olduvai II ? ? ? ? Lutra sp. Mustelidae Jianshi ? ? 0.628 ? Lutrinae Mustelidae Upper Burgi 0.680 ? ? 2.262 Martes sp. Mustelidae Mohui 0.643 ? ? ? Martes sp. Mustelidae Xiaochangliang ? ? ? ? Martes sp. 1 Mustelidae Jianshi 0.706 ? ? 2.086 Martes sp. 2 Mustelidae Jianshi 0.630 ? ? ? Meles cf. chiai Mustelidae Longgupo 0.454 ? ? 1.650 Meles cf. leucurus Mustelidae Gongwangling 0.410 ? ? 1.462 Meles chiai Mustelidae Nihewan 0.573 ? ? 1.503 Meles teilhardi Mustelidae Longdan 0.374 ? 1.335 1.591 Mellivora benfieldi Mustelidae Upper Burgi 0.760 ? ? ? Mellivora sp. Mustelidae Okote ? ? ? ? Torolutra ? Mustelidae Okote ? ? ? ? Torolutra ? Mustelidae Upper Burgi ? ? ? ? Torolutra cf. ougandensis Mustelidae KBS ? ? ? ? Torolutra cf. ougandensis Mustelidae Upper Burgi 0.646 0.803 ? ? Torolutra sp. Mustelidae KBS 0.685 0.738 ? 2.153 Prionodon sp. Prionodontidae Jianshi 0.779 0.483 ? 1.034 Ailuropoda melanoleuca Ursidae Gongwangling 0.635 ? ? ? Ailuropoda microta Ursidae Longgupo ? ? 1.432 ? Ailuropoda microta Ursidae Mohui ? ? 1.443 ? Ailuropoda wulingshanensis Ursidae Jianshi ? ? 1.130 ? Ursus aff. thibetanus Ursidae Longgupo 0.529 2.024 1.915 ? Ursus cf. etruscus Ursidae Gongwangling ? ? 1.901 ? Ursus cf. etruscus Ursidae Nihewan ? ? ? ? Ursus sp. Ursidae Jianshi 0.562 1.872 1.890 ? Ursus sp. Ursidae Linyi ? ? ? ? Ursus thibetanus Ursidae Mohui ? ? ? ? Megaviverra pleistocaenica Viverridae Longgupo ? ? 0.791 2.304 Paguma Viverridae Mohui ? ? ? ? Pseudocivetta ingens Viverridae KBS 0.589 1.245 ? ? Pseudocivetta ingens Viverridae Olduvai I 0.598 1.245 ? ? Pseudocivetta ingens Viverridae Olduvai II 0.598 1.245 ? ? Pseudocivetta ingens Viverridae Upper Burgi ? ? ? ? Pseudocivetta sp Viverridae Upper Burgi ? ? ? ? Viverra sp. Viverridae Jianshi 0.607 ? ? 2.254 Viverricula malaccensis Viverridae Yuanmou ? ? ? 3.230 Viverridae (KBS) Viverridae KBS ? ? ? ? Viverridae (UB) Viverridae Upper Burgi ? ? ? 1.890

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Table 6.6 Index values for fossil carnivore species, continued. Species Family Site P4WL PROTO CWL UM21 LBM Canis brevicephalus Canidae Longdan 0.541 0.509 0.680 0.568 1.471 Canis cf. mesomelas Canidae Kalochoro 0.428 ? ? ? 1.023 Canis cf. mesomelas Canidae KBS 0.439 ? ? ? ? Canis cf. mesomelas Canidae Natoo ? ? ? ? ? Canis cf. mesomelas Canidae Upper Burgi ? 0.455 0.678 0.571 ? Canis chihliensis Canidae Nihewan 0.471 0.445 0.620 0.557 1.353 Canis longdanensis Canidae Longdan 0.478 0.520 0.648 0.543 1.334 Canis lupus Canidae Nihewan 0.440 ? ? ? 1.236 Canis lycaonoides Canidae Olduvai II 0.457 0.716 0.675 0.518 1.368 Canis mesomelas Canidae Olduvai I 0.430 0.456 ? 0.607 1.102 Canis palmidens Canidae Nihewan 0.423 0.533 0.569 0.554 1.357 Canis sp. Canidae Donggutuo ? ? ? ? ? Canis sp. Canidae Linyi ? ? ? ? ? Canis sp. Canidae Upper Burgi ? ? ? ? ? Canis teilhardi Canidae Longdan 0.477 0.535 0.636 0.573 1.384 Canis variabilis Canidae Gongwangling 0.495 ? 0.698 ? 1.265 Canis yuanmouensis Canidae Yuanmou ? 0.502 ? 0.580 ? Cuon dubius Canidae Jianshi 0.449 0.548 0.668 0.475 ? Cuon dubius Canidae Longgupo ? 0.534 ? 0.457 1.451 Cuon dubius Canidae Mohui ? ? ? 0.457 ? Nyctereutes cf. sinensis Canidae Longgupo ? ? ? ? ? Nyctereutes cf. sinensis Canidae Nihewan 0.455 0.517 0.733 0.651 ? Prototocyon recki Canidae Olduvai I 0.505 0.711 0.738 0.953 0.332 Sinicuon cf. dubius Canidae Longdan ? ? ? 0.457 ? Vulpes cf. zerda Canidae KBS ? ? ? ? ? Vulpes cf. zerda Canidae Upper Burgi ? ? ? ? ? Vulpes chikushanensis Canidae Longdan 0.392 0.495 0.606 0.668 0.856 Vulpes sp. Canidae Nihewan ? ? ? ? ? Acinonyx jubatus Felidae Olduvai II ? ? ? 0 ? Acinonyx sp Felidae Okote ? ? ? 0 ? Acinonyx sp Felidae Upper Burgi ? ? ? 0 ? Acinonyx sp. Felidae KBS ? ? ? 0 ? Caracal sp. Felidae Upper Burgi ? ? ? 0 1.150 Dinofelis aronoki Felidae Upper Burgi 0.486 ? ? 0 2.283 Dinofelis piveteaui Felidae Okote 0.462 0.328 0.577 0 2.115 Dinofelis sp. D Felidae Olduvai I ? ? ? 0 ? Dinofelis sp. D Felidae Olduvai II ? ? ? 0 ? Felis microta Felidae Longgupo 0.481 ? ? 0 0.854 Felis sp. Felidae Longgupo ? ? ? 0 ? Felis teilhardi Felidae Jianshi ? 0.491 ? 0 1.296 Felis teilhardi Felidae Longdan 0.491 ? ? 0 1.258 Felis teilhardi Felidae Longgupo ? ? ? 0 1.335 Felis teilhardi Felidae Mohui ? ? ? 0 1.540 Homotherium cf. Felidae Longgupo crenatidens ? 0.509 ? 0 ? Homotherium cf. Felidae Nihewan crenatidens ? ? 0.409 0 ? Homotherium crendatidens Felidae Longdan ? 0.272 0.434 0 ? Homotherium crendatidens Felidae Jianshi ? 0.303 0.452 0 2.397 162

Species Family Site P4WL PROTO CWL UM21 LBM Homotherium davitasvili Felidae Haiyan 0.494 ? ? 0 2.275 Homotherium sp. Felidae Kaitio ? ? ? 0 ? Homotherium sp. Felidae Kalochoro ? ? ? 0 ? Homotherium sp. Felidae KBS 0.396 ? ? 0 2.589 Homotherium sp. Felidae Okote ? ? ? 0 ? Homotherium sp. Felidae Upper Burgi ? ? ? 0 ? Lynx shansius Felidae Longdan 0.476 0.494 0.762 0 1.457 Lynx shansius Felidae Nihewan ? ? ? 0 1.392 Machairodus sp. Felidae Olduvai II ? ? 0.548 0 ? Megantereon nihowanensis Felidae Gongwangling 0.444 ? 0.268 0 2.003 Megantereon nihowanensis Felidae Longdan 0.449 0.437 0.530 0 1.990 Megantereon nihowanensis Felidae Nihewan 0.451 ? ? 0 2.035 Megantereon nihowanensis Felidae Yuanmou ? ? ? 0 ? Megantereon nihowanensis Felidae Haiyan ? ? 0.525 0 ? Megantereon sp. Felidae Chari ? ? ? 0 ? Megantereon sp. Felidae Jianshi ? ? 0.470 0 ? Megantereon sp. Felidae Kaitio ? ? ? 0 ? Megantereon sp. Felidae KBS ? ? ? 0 ? Megantereon sp. Felidae Linyi ? ? ? 0 ? Megantereon sp. Felidae Upper Burgi ? ? ? 0 ? Megantereon whitei Felidae Okote 0.391 0.436 0.604 0 1.575 Metailurus sp. Felidae Olduvai II ? ? 0.712 0 ? Panthera cf. leo Felidae KBS ? ? ? 0 ? Panthera cf. leo Felidae Upper Burgi ? ? ? 0 ? Panthera cf. palaeosinensis Felidae Longgupo ? ? ? 0 ? Panthera cf. pardus Felidae Longgupo ? ? ? 0 1.914 Panthera cf. tigris Felidae Gongwangling ? ? ? 0 ? Panthera leo Felidae Okote 0.478 ? ? 0 ? Panthera leo Felidae Olduvai I ? ? ? 0 ? Panthera leo Felidae Olduvai II 0.502 ? 0.793 0 2.256 Panthera palaeosinensis Felidae Longdan ? ? 0.906 0 ? Panthera pardus Felidae Gongwangling 0.533 ? 0.780 0 1.956 Panthera pardus Felidae Jianshi 0.503 0.524 0.893 0 1.918 Panthera pardus Felidae Olduvai I ? ? ? 0 ? Panthera pardus Felidae Olduvai II ? ? ? 0 ? Panthera pardus Felidae Upper Burgi 0.461 ? ? 0 1.567 Panthera pardus ? Felidae KBS ? ? ? 0 ? Panthera sp. Felidae Mohui ? ? ? 0 ? Panthera sp. Felidae KBS ? 0.468 ? 0 ? Panthera sp. Felidae Upper Burgi ? ? ? 0 1.192 Panthera tigris Felidae Yuanmou ? ? ? 0 ? Sivapanthera linxiaensis Felidae Longdan 0.491 0.479 0.786 0 1.995 Sivapanthera Felidae Gongwangling pleistocaenicus 0.495 ? ? 0 2.026 Sivapanthera Felidae Jianshi pleistocaenicus ? 0.442 ? 0 ? Sivapanthera Felidae Longgupo pleistocaenicus ? ? 0.761 0 ? Sivapanthera Felidae Nihewan pleistocaenicus ? ? ? 0 ?

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Species Family Site P4WL PROTO CWL UM21 LBM Atilax paludinosus Herpestidae Olduvai II ? ? ? ? 0.636 Herpestes primitivus Herpestidae Olduvai I 0.437 0.576 ? ? 0.150 Mungos dietrichi Herpestidae Olduvai I 0.570 1.066 ? ? 0.030 Chasmaporthetes cf. Hyaenidae Nihewan ossifragus ? ? ? 0 ? Chasmaporthetes nitidula Hyaenidae Olduvai II 0.411 ? ? 0 1.936 Chasmaporthetes Hyaenidae Longdan progressus ? 0.554 0.723 0 ? Crocuta cf. ultra Hyaenidae Chari ? ? ? 0 ? Crocuta crocuta Hyaenidae Kalochoro 0.634 ? ? 0 ? Crocuta crocuta Hyaenidae Nariokotome ? ? ? 0 ? Crocuta crocuta Hyaenidae Olduvai I 0.630 ? ? 0 1.962 Crocuta crocuta Hyaenidae Olduvai II 0.611 ? ? 0 ? Crocuta dietrichi Hyaenidae KBS ? ? ? 0 ? Crocuta dietrichi Hyaenidae Upper Burgi 0.609 ? ? 0 2.001 Crocuta honanensis Hyaenidae Longdan 0.645 0.552 0.755 0 1.883 Crocuta honanensis Hyaenidae Haiyan 0.648 ? ? 0 1.839 Crocuta sp. Hyaenidae KBS 0.694 ? ? 0 1.895 Crocuta sp. Hyaenidae Okote ? ? ? 0 ? Crocuta sp. Hyaenidae Upper Burgi ? ? ? 0 ? Crocuta ultra Hyaenidae KBS 0.606 0.522 0.779 0 2.009 Crocuta ultra Hyaenidae Lokalalei 0.650 0.544 ? 0 2.034 Crocuta ultra Hyaenidae Okote 0.549 0.518 ? 0 1.943 Crocuta ultra Hyaenidae Upper Burgi ? ? ? 0 ? Hyaena cf. makapani Hyaenidae KBS ? ? ? 0 ? Hyaena hyaena Hyaenidae Olduvai I ? 0.558 ? 0 ? Hyaena hyaena Hyaenidae Olduvai II ? ? ? 0 ? Hyaena makapani Hyaenidae Upper Burgi 0.531 0.609 0.683 0 1.550 Hyaena sp. Hyaenidae Kaitio 0.510 ? ? 0 1.812 Hyaena sp. Hyaenidae Kalochoro 0.560 ? ? 0 1.649 Hyaena sp. Hyaenidae KBS ? ? ? 0 ? Hyaena sp. Hyaenidae Linyi ? ? ? 0 ? Hyaena sp. Hyaenidae Okote ? ? ? 0 ? Hyaena sp. Hyaenidae Upper Burgi 0.525 0.569 0.631 0 1.662 Pachycrocuta brevirostris Hyaenidae Gongwangling ? 0.617 ? 0 ? Pachycrocuta brevirostris Hyaenidae Jianshi 0.697 0.587 0.776 0 2.049 Pachycrocuta brevirostris Hyaenidae Longdan 0.718 0.603 0.806 0 1.950 Pachycrocuta brevirostris Hyaenidae Longgupo 0.619 0.594 ? 0 1.997 Pachycrocuta brevirostris Hyaenidae Mohui ? ? ? 0 ? Pachycrocuta brevirostris Hyaenidae Nihewan 0.634 0.606 0.743 0 2.021 Pachycrocuta brevirostris Hyaenidae Xiaochangliang ? ? ? 0 ? Pachycrocuta brevirostris Hyaenidae Yuanmou 0.639 0.658 0.900 0 2.082 Pachycrocuta brevirostris Hyaenidae Haiyan 0.636 ? 0.734 0 1.914 Pachycrocuta sp. Hyaenidae Majuangou ? ? ? 0 ? Pliocrocuta perrieri Hyaenidae Haiyan 0.689 ? ? 0 ? Aonyxini Mustelidae KBS ? ? ? 0 ? Arctonyx cf. minor Mustelidae Longgupo 0.600 ? ? 0 0.929 Arctonyx collaris Mustelidae Mohui ? 0.857 ? 0 ? Arctonyx collaris Mustelidae Jianshi ? 0.857 ? 0 ? cf. Torolutra Mustelidae Okote ? ? ? 0 ? 164

Species Family Site P4WL PROTO CWL UM21 LBM Eirictis robusta Mustelidae Longdan 0.557 0.621 0.809 0 1.114 Enhydriodon sp. Mustelidae Upper Burgi ? ? ? 0 ? Lutra licenti Mustelidae Nihewan ? ? ? 0 ? Lutra maculicollis Mustelidae Olduvai I ? ? ? 0 ? Lutra maculicollis Mustelidae Olduvai II ? ? ? 0 ? Lutra sp. Mustelidae Jianshi ? 0.893 ? 0 ? Lutrinae Mustelidae Upper Burgi 0.511 ? ? 0 1.337 Martes sp. Mustelidae Mohui ? ? ? 0 ? Martes sp. Mustelidae Xiaochangliang ? ? ? 0 ? Martes sp. 1 Mustelidae Jianshi 0.481 ? ? 0 0.133 Martes sp. 2 Mustelidae Jianshi ? 0.550 ? 0 1.101 Meles cf. chiai Mustelidae Longgupo 0.517 ? ? 0 1.148 Meles cf. leucurus Mustelidae Gongwangling 0.564 ? ? 0 1.442 Meles chiai Mustelidae Nihewan 0.580 ? ? 0 1.145 Meles teilhardi Mustelidae Longdan 0.595 0.808 0.733 0 1.220 Mellivora benfieldi Mustelidae Upper Burgi ? ? ? 0 0.787 Mellivora sp. Mustelidae Okote ? ? ? 0 ? Torolutra ? Mustelidae Okote ? ? ? 0 ? Torolutra ? Mustelidae Upper Burgi ? ? ? ? 1.212 Torolutra cf. ougandensis Mustelidae KBS ? ? ? 0 ? Torolutra cf. ougandensis Mustelidae Upper Burgi ? ? ? 0 1.096 Torolutra sp. Mustelidae KBS 0.544 ? ? 0 1.300 Prionodon sp. Prionodontidae Jianshi 0.403 ? ? ? 0.043 Ailuropoda melanoleuca Ursidae Gongwangling ? ? ? ? 1.973 Ailuropoda microta Ursidae Longgupo ? 0.556 ? 1.118 1.939 Ailuropoda microta Ursidae Mohui ? 0.579 ? ? ? Ailuropoda wulingshanensis Ursidae Jianshi ? 0.424 ? 1.136 1.980 Ursus aff. thibetanus Ursidae Longgupo ? 0.700 0.721 1.240 1.895 Ursus cf. etruscus Ursidae Gongwangling ? 0.731 ? 1.372 ? Ursus cf. etruscus Ursidae Nihewan ? ? ? ? ? Ursus sp. Ursidae Jianshi ? 0.726 0.759 ? 1.890 Ursus sp. Ursidae Linyi ? ? ? ? ? Ursus thibetanus Ursidae Mohui ? ? ? 1.221 ? Megaviverra pleistocaenica Viverridae Longgupo 0.478 0.728 ? 0.628 1.571 Paguma Viverridae Mohui ? 0.706 ? ? ? Pseudocivetta ingens Viverridae KBS ? ? ? ? 1.558 Pseudocivetta ingens Viverridae Olduvai I ? ? ? ? 1.625 Pseudocivetta ingens Viverridae Olduvai II ? ? ? ? 1.625 Pseudocivetta ingens Viverridae Upper Burgi ? ? ? ? ? Pseudocivetta sp Viverridae Upper Burgi ? ? 0.791 ? ? Viverra sp. Viverridae Jianshi ? 0.542 ? 1.234 1.059 Viverricula malaccensis Viverridae Yuanmou 0.469 0.620 ? ? 0.535 Viverridae (KBS) Viverridae KBS ? 0.843 0.778 ? ? Viverridae (UB) Viverridae Upper Burgi 0.428 0.709 ? ? 1.406

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Principal Components Analysis :

Principal components analysis (PCA) was used to explore differences between

species in families using index values as input. This method of analysis was chosen as a

way to look directly at the index values of the various species. It also serves as a test of

the category system, described below. Similar results from the PCA and the

correspondence analysis of the categories would indicate that the categories reflect the

actual ecological adaptations of the animals. Only indices that came directly from a

species at a site were used in this analysis.

Index Categories and Correspondence Analysis:

Further ecomorphological analyses split the index values into categories. These categories were analyzed using correspondence analysis (CA) and disparity analysis.

Disparity was measured as mean pairwise dissimilarity. In this analysis, each index was divided into character state categories (described below) to reflect ecological differences.

These categories make large and ecologically important differences between species readily apparent. Correspondence analysis was used because it plots species and indices together for easier interpretation. However, the disadvantage of categories is that they may sometimes separate species that are close in value but happen to fall on opposite sides of the category cut-offs.

Like the PCA analysis, the category analysis looks at the questions of whether the carnivore guilds of East Asia and East Africa were similar in the types of species present.

Did they contain avatars or all species, and if not, which species and adaptations were different? This analysis also looks at whether there were changes in the types of

166 adaptations present in East Asia from the Pliocene to the Pleistocene. Similarity in many categories was used to show which species may have been avatars in the traits examined here.

Creating Categories :

Categories reflecting both the ecological adaptations of modern species and the distribution patterns of fossil species were created (table 6.7). Values for each index for modern and ancient species were plotted. Composite values of fossil species from each available site were used in the plots. Natural breaks between the fossil taxa were used in combination with ecological data from modern taxa to determine breaks between categories.

Table 6.7 Category cut-off values. Category values for each index are listed with the trend in values. Index Categories Trend RBL 1. 0.9-1 Carnivorous taxa have relatively larger blade lengths 2. 0.9-0.76 compared with more omnivorous species. 3. 0.76-0.69 4. 0.69-0.66 5. <0.66 RGA 1. 0 Carnivorous taxa have smaller relative grinding area 2. >0 – 0.38 compared with more omnivorous taxa. 3. 0.38-.75 4. 0.75-1 5. 1-1.5 6. >1.5 RUMGA 1. 0-0.047 Carnivores and bone-crushers have relatively small 2. 0.47-0.92 values, while omnivorous and moderately 3. 0.93-1.3 carnivorous species have larger values. 4. 1.3-1.8 5. >1.8 PROTO 1. <0.47 Carnivorous taxa tend to have smaller ratios 2. 0.47-0.7 compared with omnivores. 3. >0.7

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Index Categories Trend P4WL 1. >0.6 Bone crushers and omnivorous viverrids have larger 2. 0.6-0.49 ratios, while carnivores and omnivores tend to have 3. <0.49 smaller ratios. CWL 1. <0.63 Modern felids and hyaenids tend to have high ratios, 2. 0.63-0.7 while moderately carnivorous and omnivorous 3. 0.7-0.8 species have species have smaller values. Saber- 4. >0.8 toothed felids have smaller values. UM21 1. 0 Omnivores tend to have larger values of this index. 2. <0.61 Carnivores have less grinding area. 3. 0.61-0.66 4. 0.66-1 5. 1-1.14 6. >1.14 LBM 1. 0.1 Carnivores of increasing size according to natural 2. 1-1.332438 groupings of the body masses of extant carnivores 3. 1.332438-2 (Lewis and Werdelin 2007). 4. >2 RPS 1. >3 Bone-crushers and omnivorous viverrids and 2. 3-2.51 herpestids have large values. More carnivorous and 3. <2.51 omnivorous taxa have smaller values.

Category Scores and Missing Information :

Categories were assigned based on the index values of each species at a site.

Species that were present at different sites sometimes had differing amounts of information for each of the index categories. In that case, category scores (Table 6.8) from species at sites with more information were used to fill in missing information. For this analysis, taxa classified as “aff.” or “cf.” were treated as members of the species. The species pools of East Africa and East Asia were kept separate. When two or more different category classifications were found for an index within a species, classifications were only applied to the missing data if 1) a majority of classifications were in one particular category or 2) if information existed to support a certain classification. When no pattern or information existed, the category scores for the index were left as question

168

marks, signifying missing data. Some specimens had no information available and

missing information could not be filled in. When sufficient information was not available, the species was excluded from the correspondence analysis of the categories. This category assignment procedure assumed that the taxonomic attribution of each of the species at the sites was correct, and that the category score obtained for individuals at one site was a valid reflection of the species’ properties at other sites. Categories were necessarily broad and so much variation between individual specimens is contained with

one category.

Analyzing the category scores :

The number of differences in the categories between species reflects the degree of

difference between species. The number of differences between species is quantified

using the Hamming distance, which is the number of character state differences divided

by the total number of characters analyzed for that pair of taxa (Hamming 1950).

Category scores were analyzed using correspondence analysis (CA) of the categories and

Non-metric Multi-Dimensional Scaling analysis (NMDS) of the Hamming distance

matrix. The Hamming distances between pairs of species may be calculated regardless of

the number of traits for which there is information, although they are a more reliable

source of information about the overall amount of disparity between species when more indices are available. NDMS of the Hamming distance was used to look at overall differences between the species.

The results of the NMDS and CA were used to look at carnivores within families to characterize taxa by their feeding adaptations and to differentiate taxa from each other.

169

Comparisons of the trends within a family over time were done in order to examine

whether species with similar feeding adaptations were always present, or whether the

feeding and body mass compositions of a guild differed over time.

Using the factor loadings of the CA, centroids were calculated for the carnivore families in East Asia and East Africa, as well as for the carnivore guilds in the ancient sites. Both sets of centroid positions were tested for significant differences using

MANOVA. These tests help examine whether the families and sites occupy statistically different locations in multivariate space, and thus whether they are composed of different adaptations. Mean distance to the centroid was calculated for the families as well as for the sites. The mean distance to the centroid (MEDC) shows the amount of disparity between groups of sites and African and Asian representatives of the carnivore families.

The MEDC comparisons show which families or sites had a greater diversity of adaptations, as indicated by the distribution of sites and the distance to the centroid.

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Table 6.8 Category scores for all carnivore taxa used in the CA analysis. RPS RPS RBL RBL RGA RGA LBM LBM CWL CWL UM21 UM21 P4WL P4WL PROTO PROTO RUMGA RUMGA

Canis brevicephalus Longdan 3 3 2 2 2 2 2 2 3 Canis cf. mesomelas Kalochoro 4 ? ? 3 3 ? ? ? 2 Canis cf. mesomelas Natoo 4 ? ? 3 3 ? ? ? 2 Canis cf. mesomelas KBS ? ? 2 3 3 1 2 2 ? Canis cf. mesomelas Upper Burgi ? ? 2 3 3 1 2 2 ? Canis mesomelas Olduvai I 5 4 2 3 3 1 2 2 2 Canis chihliensis Nihewan 3 4 2 2 3 1 1 2 3 Canis longdanensis Longdan 4 ? 2 2 3 2 2 2 3 Canis lupus Nihewan 3 4 ? 2 3 ? ? ? 2 Canis lycaonoides Olduvai II 3 4 2 2 3 3 2 2 3 Canis palmidens Nihewan 5 ? 2 3 3 2 1 2 3 Canis teilhardi Longdan 3 3 2 2 3 2 2 2 3 Canis variabilis Gongwangling 4 3 ? 3 2 ? 2 ? 2 Cuon dubius Jianshi 3 3 2 3 3 2 2 2 3 Cuon dubius Longgupo 3 3 2 3 3 2 2 2 3 Cuon dubius Mohui 3 3 2 3 3 2 2 2 3 Sinicuon cf. dubius Longdan 3 3 2 3 3 2 2 2 3 Nyctereutes cf. sinensis Longgupo ? ? 3 ? 3 2 3 3 ? Nyctereutes cf. sinensis Nihewan ? ? 3 ? 3 2 3 3 ? Prototocyon recki Olduvai I 5 6 4 3 2 3 3 4 1 Vulpes chikushanensis Longdan 5 4 3 3 3 2 1 3 1 Acinonyx jubatus Olduvai II 1 1 ? ? ? ? ? 1 ? Acinonyx sp. KBS 1 1 ? ? ? ? ? 1 ? Acinonyx sp. Okote 1 1 ? ? ? ? ? 1 ? Acinonyx sp. Upper Burgi 1 1 ? ? ? ? ? 1 ? Caracal sp. Upper Burgi 1 1 ? ? ? ? ? 1 2 Dinofelis aronoki Upper Burgi 1 1 ? 3 3 ? ? 1 4 Dinofelis piveteaui Okote 1 1 1 3 3 1 1 1 4 Dinofelis sp. D Olduvai I 1 1 ? ? ? ? ? 1 ? Dinofelis sp. D Olduvai II 1 1 ? ? ? ? ? 1 ? Felis microta Longgupo 1 1 ? 3 3 ? ? 1 1 Felis sp. Longgupo 1 1 ? ? ? ? ? 1 ? Felis teilhardi Jianshi 1 1 ? 3 2 2 ? 1 2 Felis teilhardi Longdan 1 1 ? 3 2 2 ? 1 2 Felis teilhardi Longgupo 1 1 ? 3 2 2 ? 1 3 Felis teilhardi Mohui 1 1 ? 3 2 2 ? 1 3 Homotherium cf. crenatidens Longgupo 1 1 1 ? ? 2 1 1 4 Homotherium cf. crenatidens Nihewan 1 1 1 ? ? ? 1 1 4 Homotherium crendatidens Longdan 1 1 1 ? ? 1 1 1 4 Homotherium Jianshi 1 1 1 ? ? 1 1 1 4 171

RPS RPS RBL RBL RGA RGA LBM LBM CWL CWL UM21 UM21 P4WL P4WL PROTO PROTO RUMGA RUMGA

crendatidens Homotherium davitasvili Haiyan 1 1 ? 3 2 ? ? 1 4 Homotherium sp. Kaitio 1 1 ? ? ? ? ? 1 ? Homotherium sp. Kalochoro 1 1 ? ? ? ? ? 1 ? Homotherium sp. KBS 1 1 ? 3 3 ? ? 1 4 Homotherium sp. Okote 1 1 ? ? ? ? ? 1 ? Homotherium sp. Upper Burgi 1 1 ? ? ? ? ? 1 ? Lynx shansius Longdan 1 1 1 3 3 2 3 1 3 Lynx shansius Nihewan 1 1 1 3 3 2 3 1 3 Machairodus Olduvai II 1 1 ? ? ? ? 1 1 ? Megantereon nihowanensis Gongwangling 1 1 1 3 3 1 1 1 4 Megantereon nihowanensis Longdan 1 1 1 3 3 1 1 1 3 Megantereon nihowanensis Nihewan 1 1 1 3 3 1 1 1 4 Megantereon nihowanensis Yuanmou 1 1 1 3 3 1 1 1 4 Megantereon nihowanensis Haiyan 1 1 1 3 3 1 1 1 4 Megantereon sp . Chari 1 1 ? ? ? ? ? 1 ? Megantereon sp . Jianshi 1 1 ? ? ? ? 1 1 ? Megantereon sp . Kaitio 1 1 ? ? ? ? ? 1 ? Megantereon sp . KBS 1 1 ? ? ? ? ? 1 ? Megantereon sp . Linyi 1 1 ? ? ? ? ? 1 ? Megantereon sp . Upper Burgi 1 1 ? ? ? ? ? 1 ? Megantereon whitei Okote 1 1 1 3 3 1 1 1 3 Metailurus Olduvai II 1 1 ? ? ? ? 3 1 ? Panthera cf. leo KBS 1 1 ? 3 3 ? 3 1 4 Panthera cf. leo UB 1 1 ? 3 3 ? 3 1 4 Panthera leo Okote 1 1 ? 3 3 ? 3 1 4 Panthera leo Olduvai I 1 1 ? 3 2 ? 3 1 4 Panthera leo Olduvai II 1 1 ? 3 2 ? 3 1 4 Panthera cf. tigris Gongwangling 1 1 ? ? ? ? ? 1 ? Panthera tigris Yuanmou 1 1 ? ? ? ? ? 1 ? Panthera palaeosinensis Longdan 1 1 ? ? ? ? 4 1 ? Panthera cf. palaeosinensis Longgupo 1 1 ? ? ? ? 4 1 ? Panthera cf . pardus Longgupo 1 1 ? 3 2 2 4 1 3 Panthera pardus Gongwangling 1 1 ? 3 2 2 3 1 3 Panthera pardus Jianshi 1 1 ? 3 2 2 4 1 3 Panthera pardus Olduvai I 1 1 ? 3 3 ? ? 1 3 Panthera pardus Olduvai II 1 1 ? 3 3 ? ? 1 3 Panthera pardus Upper Burgi 1 1 ? 3 3 ? ? 1 3 Panthera pardus ? KBS 1 1 ? 3 3 ? ? 1 3 Panthera sp. Mohui 1 1 ? ? ? ? ? 1 ?

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RPS RPS RBL RBL RGA RGA LBM LBM CWL CWL UM21 UM21 P4WL P4WL PROTO PROTO RUMGA RUMGA

Panthera sp. KBS 1 1 ? ? ? 1 ? 1 ? Sivapanthera linxiaensis Longdan 1 1 1 3 2 2 3 1 3 Sivapanthera pleistocaenicus Gongwangling 1 1 1 3 2 1 3 1 4 Sivapanthera pleistocaenicus Jianshi 1 1 1 3 2 1 3 1 4 Sivapanthera pleistocaenicus Longgupo 1 1 1 3 2 1 3 1 4 Sivapanthera pleistocaenicus Nihewan 1 1 1 3 2 1 3 1 4 Herpestes primitivus Olduvai I 3 4 ? 3 3 2 ? ? 1 Mungos dietrichi Olduvai I 5 6 ? 1 2 3 ? ? 1 Chasmaporthetes nitidula Olduvai II 2 2 ? 3 3 ? ? 1 3 Crocuta cf. ultra Chari 2 2 ? 2 1 2 3 1 3 Crocuta ultra KBS 2 2 ? 2 1 2 3 1 4 Crocuta ultra Lokalalei 2 2 ? 2 1 2 3 1 4 Crocuta ultra Okote 2 2 ? 2 2 2 3 1 3 Crocuta ultra Upper Burgi 2 2 ? 2 2 2 3 1 3 Crocuta crocuta Kalochoro 2 2 ? 2 1 ? ? 1 3 Crocuta crocuta Nariokotome 2 2 ? 2 1 ? ? 1 3 Crocuta crocuta Olduvai I 2 2 ? 2 1 ? ? 1 3 Crocuta crocuta Olduvai II 2 2 ? 2 1 ? ? 1 3 Crocuta dietrichi KBS 2 2 ? 2 1 ? ? 1 4 Crocuta dietrichi Upper Burgi 2 2 ? 2 1 ? ? 1 4 Crocuta honanensis Longdan 2 2 1 1 1 2 3 1 3 Crocuta honanensis Haiyan 2 2 1 1 1 2 3 1 3 Crocuta sp. KBS 2 2 ? 1 1 ? ? 1 3 Hyaena makapani Upper Burgi 2 2 1 2 2 2 2 1 3 Hyaena cf. makapani KBS 2 2 1 2 2 2 2 1 3 Hyaena sp. Kaitio 2 2 ? 2 2 ? ? 1 3 Hyaena sp. Kalochoro 2 2 ? 2 2 ? ? 1 3 Hyaena sp. Upper Burgi 2 2 1 2 2 2 2 1 3 Pachycrocuta brevirostris Gongwangling 2 2 1 1 1 2 ? 1 4 Pachycrocuta brevirostris Jianshi 2 2 1 1 1 2 3 1 4 Pachycrocuta brevirostris Longdan 2 2 1 1 1 2 4 1 3 Pachycrocuta brevirostris Longgupo 2 2 1 1 1 2 ? 1 3 Pachycrocuta brevirostris Mohui 2 2 1 1 1 2 ? 1 4 Pachycrocuta brevirostris Nihewan 2 2 1 1 1 2 3 1 4 Pachycrocuta brevirostris Xiaochangliang 2 2 1 1 1 2 3 1 4 Pachycrocuta brevirostris Yuanmou 2 2 1 1 1 2 4 1 4 Pachycrocuta brevirostris Haiyan 2 2 1 1 1 2 3 1 3 Pliocrocuta perrieri Haiyan 2 2 ? 1 1 ? ? 1 3 Arctonyx cf. minor Longgupo 5 5 ? 3 1 ? ? 1 1 Eirictis robusta Longdan 5 4 2 3 2 2 4 1 2 Lutrinae Upper Burgi 4 ? ? 3 2 ? ? 1 3 Martes sp. Mohui 5 ? ? ? ? ? ? 1 ?

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RPS RPS RBL RBL RGA RGA LBM LBM CWL CWL UM21 UM21 P4WL P4WL PROTO PROTO RUMGA RUMGA

Martes sp. 1 Jianshi 3 ? ? 3 3 ? ? 1 1 Martes sp. 2 Jianshi 5 ? ? ? ? 2 ? 1 2 Meles cf. chiai Longgupo 5 ? ? 3 2 ? ? 1 2 Meles chiai Nihewan 5 ? ? 3 2 ? ? 1 2 Meles cf. leucurus Gongwangling 5 ? ? 3 2 ? ? 1 3 Meles teilhardi Longdan 5 ? 4 3 2 3 3 1 2 Mellivora benfieldi Upper Burgi 3 ? ? ? ? ? ? 1 1 Torolutra cf. Ougandensis KBS 5 4 ? ? ? ? ? 1 2 Torolutra cf. Ougandensis Upper Burgi 5 4 ? ? ? ? ? 1 2 Torolutra sp. KBS 4 3 ? 3 2 ? ? 1 2 Ailuropoda microta Longgupo ? ? 4 ? ? 2 ? 5 3 Ailuropoda microta Mohui ? ? 4 ? ? 2 ? 5 3 Ailuropoda wulingshanensis Jianshi ? ? 3 ? ? 1 ? 5 3 Ursus aff. thibetanus Longgupo 5 6 5 ? ? 3 3 6 3 Ursus thibetanus Mohui 5 6 5 ? ? 3 3 6 3 Ursus sp. Jianshi 5 6 5 ? ? 3 3 6 3 Megaviverra pleistocaenica Longgupo ? ? 2 3 3 3 ? 3 3 Prionodon sp. Jianshi 2 3 ? 3 3 ? ? ? 1 Viverra sp. Jianshi 5 ? ? 3 ? 2 ? ? 2

Summary:

This chapter describes the methods used to characterize fossil carnivore species ecomorphologically as well as the statistics used to analyze them. The purpose of the ecomorphological analysis was to compare the feeding adaptations of large carnivores from Plio-Pleistocene East Asia and East Africa. The ecomorphology of the carnivore guild in East Asia was also compared over time from the Late Pliocene to the Early

Pleistocene.

All available adult fossil carnivore samples were measured for all possible ecomorphological measures. Modern carnivores were also measured. Ecomorphological indices, which were based on dental measurements, relate to feeding adaptations, hunting

174 behavior and body mass. The indices measure the relative size of the blade on the M 1

(RBL) and the relative amount of grinding area in the molars (RGA, RUMGA and

UM21). Longer blade lengths and smaller amounts of grinding area are traits correlated with increased carnivory, whereas greater amounts of grinding area with relatively small blades are related to increased omnivory. PROTO measures the relative size of the protocone in P 4, which is relatively large in bone-crackers compared with flesh-specialist carnivores. P4WL measures the shape of the P 4 and RPS its relative size. Rounded and relatively large lower fourth premolars are correlated with bone-cracking. CWL measures the shape of the upper canine, which is narrow in species that slash their prey and rounded in stabbing species. Body size is measured by LBM, log body size. Larger animals are capable of hunting larger prey, and are typically dominant in interspecific interactions.

Due to the fact that many specimens were incomplete, composites of index values were made using input from more than one specimen. The values of the indices from ancient and modern species were separated into categories that reflected differences in ecology and natural breaks in the data. The data from the indices was analyzed in several ways. The values of the indices were analyzed using PCA, while the categorical data was analyzed using correspondence analysis. Centroids were calculated for carnivore families from Asia and Africa (e.g., the group of all felids from Africa and a group of all felids from Asia). They were also calculated for sites with large carnivore faunas (e.g., all carnivores from Gongwangling and Olduvai Bed I). Mean Euclidean distance to the centroid was calculated for all centroids. MANOVA was used to examine whether these centroids were significantly different statistically.

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Chapter 7: Carnivore Ecomorphology Results:

The objectives of this chapter are to compare carnivores from East Asia and East

Africa ecomorphologically, to characterize carnivores that were found in East Asian hominin sites and to look at East Asian carnivore adaptations and guild occupation patterns through time. Body mass and feeding adaptations were compared in each family and overall using PCA and CA of categories.

This chapter will look at how the carnivores from East Asia and East Africa compare in terms of feeding adaptations and body mass to determine whether the two guilds had sets of ecologically similar carnivores or whether there were differences that dispersing hominins would have encountered. The question of change over time from the

Pliocene to the Pleistocene in East Asia was also addressed by considering whether there was change in the carnivore guild, as shown by the appearance or disappearance of certain carnivore types. Turnover in the carnivore guild could have had an important effect upon the opportunities for hominins, in terms of niches that may have been available. Turnover later in the early Pleistocene could also signal the effect of hominins upon the carnivore guild.

Canidae

East Africa versus East Asia :

Many more species of canids are found in the East Asian fossil record than in East

African sites examined here, using CA, PCA and NMDS analysis.

On the first axis of the correspondence analysis (Figure 7.1), there is separation between canids with larger grinding areas and small bodies, and those with larger bodies

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and adaptations for more carnivorous diets. Increased grinding area is present in taxa

such as Prototocyon recki , which uses its supernumerary dentition to consume insects,

and in Vulpes chikushanensis from Longdan. Canids such as Cuon dubius /Sinicuon dubius , Canis teilhardi, Canis brevicephalus , Canis longdanensis, Canis chihliensis and

Canis lycaonoides show more adaptations toward meat-consumption and large body size.

They have less grinding area (RGA, RUMGA, UM21), and more relative blade length

(RBL). Canids such as C. variabilis , the C. cf. mesomelas from Koobi Fora and West

Turkana, and Canis mesomelas from Olduvai (which are not conspecific), C. lupus

(Nihewan) and Nyctereutes sinensis have more intermediate adaptations. On the second

axis, a group of species including Vulpes chikushanensis , C. mesomelas from Olduvai, C.

cf. mesomelas from Koobi Fora, C. palmidens and C. chihliensis plot with RBL, RPS and

P4WL. Characteristics of some of these species are shorter relative blade lengths, smaller

values of PROTO, and smaller ratios of P4WL, which may indicate omnivorous

tendencies. PROTO and CWL plot low on the second axis. Both Prototocyon recki and

Canis lycaonoides have large values for the PROTO index.

Non-metric multidimensional scaling analysis of the Hamming distance (Figure

7.2) emphasizes how different Prototocyon recki is from the other canids. Canis brevicephalus , Canis teilhardi , Canis longdanensis, Canis palmidens , Canis chihliensis and Canis lycaonoides plot within a smaller distance, having similar amounts of difference between them.

On the PCA (Figure 7.3), traits such as RGA, RUMGA and UM21, which are indicative of post-carnassial dentition devoted to grinding, load positively on the first axis, while LBM and RPS load negatively (Figures 7.4 and 7.5). On the second axis,

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RPS, RGA, RUMGA and to a lesser extent UM21 load positively, while LBM loads

negatively. The first axis divides species that are more omnivorous, or that have more

molar grinding area, such as Prototocyon recki , Nyctereutes , Vulpes chikushanensis ,

Canis mesomelas and Canis cf. mesomelas from species that more carnivorous. On the second axis, larger species such as C. teilhardi , C. longdanensis , C. brevicephalus , “ C. lupus ,” Canis chihliensis from East Asia and Canis lycaonoides from East Africa group together as larger, carnivorous species.

Avatars :

Species such as Canis brevirostris , Canis teilhardi , Canis longdanensis from the

Pliocene site of Longdan, Cuon dubius and Canis lycaonoides from Olduvai Bed II share

ecological features. Compared with other canids, they have relatively low values for

RGA, RUMGA and UM21, indicating more carnivorous adaptations. They also tend to

have high body masses. Species including Canis variabilis, C. chihliensis and Canis

palmidens and C. mesomelas from Olduvai, West Turkana and Koobi Fora also have

intermediate RGA, RUMGA and UM21 values. However, C. mesomelas and C. cf.

mesomelas were smaller in size than the others. Vulpes chikushanensis (Longdan),

Nyctereutes sinensis (Longgupo and Nihewan) and Prototocyon recki (Olduvai I) also

have large grinding areas and small body masses. However, none of these smaller species

were analogous to each other. Information for the African species V. cf. zerda is not

available and its comparability to V. chikushanensis is unknown. Many smaller species of

this type may be missing from the fossil record. Prototocyon recki has unique adaptations

for insectivory and is not comparable to the other ancient taxa sampled here.

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East Asia Pliocene to early Pleistocene :

Highly carnivorous canids :

Many large and highly carnivorous species are found at Longdan. Canis chihliensis from the Nihewan has carnivorous tendencies. It is uncertain whether the

Canis sp. found at the hominin site of Donggutuo was similar to this species. Cuon

dubius and Sinicuon dubius also represent a carnivorous lineage that is present during the

Pliocene and the Pleistocene.

Moderately carnivorous canids :

The moderately carnivorous canid C. palmidens is present in the Haiyan

Formation (Yushe Basin) and in the Nihewan Basin. “ Canis lupus,” which can be interpreted as moderately or highly carnivorous, was present at the Nihewan. Canis variabilis is also present at Gongwangling. The canid at Donggutuo could also belong to any of these lineages. The presence of this ecological type of canid through

Gongwangling appears to indicate some continuity.

Omnivorous canids :

Omnivorous, small canids are not well sampled in the fossil record, probably due to their small body mass. It is not known how V. chikushanensis and Vulpes sp. at the

Nihewan were related. Species of Vulpes were probably present at other East Asian sites,

but would not have had a substantial impact on hominin foraging activities due to its

small size.

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More insectivorous or More carnivorous Meso_Old1 0.16 omnivorous Palmidens

0.12 Vulpes_Long RBL Chih_Nih Meso_KBS_UB 0.08 RPS P4wl 0.04 Axis 2 Meso_Kalo_Natoo RGA 0 Variabilis Long_Long LBM Cuon/Sinicuon -0.04 UM21RUMGA Lupus_Nih -0.08 Teilhardi Nyctereutes Brevi_Long -0.12 Prototo_Old1 Lycaon -0.16 CWL

Proto -0.2 -0.4 -0.32 -0.24 -0.16 -0.08 0 0.08 0.16 0.24 Axis 1

Figure 7.1 Scatterplot of the CA for Canidae category scores plotted with indices. Refer to table 6.4 for names and meanings of indices. Key: Red: Canids from Pliocene sites of Longdan and Longgupo. Purple circles: Canids from the Nihewan s.s . (representing canids that may have been present in Nihewan hominin sites), Yuanmou, and Gongwangling. Blue: East African canids. Green: Canids from Jianshi and Mohui. Abbreviations: Prototo_Old1: Prototocyon recki, Olduvai I; Vulpes_Long: Vulpes chikushanensis , Longdan; Meso_Old1: Canis mesomelas , Olduvai I; Meso_Kalo_Natoo: Canis mesomelas Kalochoro and Natoo members; Variabilis: Canis variabilis , Gongwangling; Lupus_Nih: Canis “lupus” from Nihewan; Nyctereutes: Nyctereutes sinensis , Longdan and Nihewan; Palmidens: Canis palmidens , Nihewan; Chih_Nih: Canis chihliensis , Nihewan; Long_Long: Canis longdanensis , Longdan; Cuon/Sinicuon: Cuon dubius/Sinicuon dubius (same species), Longdan, Longgupo, Jianshi, Mohui; Teilhardi: Canis teilhardi , Longdan; Brevi_Long: Canis brevicephalus , Longdan; Lycaon: Canis lycaonoides , Olduvai II.

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0.25

0.2 Variabilis

0.15 Meso_KBS_UB 0.1 Prototo_Old1 Coordinate 2 Meso_Kalo_Natoo Meso_Old1 0.05 Cuon/Sinicuon 0 Palmidens Long_Long -0.05 Teilhardi Vulpes_Long -0.1 Brevi_Long Lycaon Chih_Nih -0.15 Lupus_Nih Nyctereutes -0.2 -0.24 -0.16 -0.08 0 0.08 0.16 0.24 0.32 0.4 Coordinate 1

Figure 7.2: Scatterplot of NMDS analysis of Hamming distances for Canidae. Key: Red: Canids from Pliocene sites of Longdan and Longgupo. Purple circles: Canids from the Nihewan s.s . (representing canids that may have been present in Nihewan hominin sites), Yuanmou, and Gongwangling. Blue: East African canids. Green: Canids from Jianshi and Mohui. Abbreviations: Prototo_Old1: Prototocyon recki, Olduvai I; Vulpes_Long: Vulpes chikushanensis , Longdan; Meso_Old1: Canis mesomelas , Olduvai I; Meso_Kalo_Natoo: Canis mesomelas Kalochoro and Natoo members; Variabilis: Canis variabilis , Gongwangling; Lupus_Nih: Canis “lupus” from Nihewan; Nyctereutes: Nyctereutes sinensis , Longdan and Nihewan; Palmidens: Canis palmidens , Nihewan; Chih_Nih: Canis chihliensis , Nihewan; Long_Long: Canis longdanensis , Longdan; Cuon/Sinicuon: Cuon dubius/Sinicuon dubius (same species), Longdan, Longgupo, Jianshi, Mohui; Teilhardi: Canis teilhardi , Longdan; Brevi_Long: Canis brevicephalus , Longdan; Lycaon: Canis lycaonoides , Olduvai II.

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Component 2 Prototocyon_recki

0.6

More carnivorous More omnivorous 0.48 or increased Canis_brevicephalus grinding area 0.36

Canis_lycaonoides 0.24 Canis_lupus Canis_longdanensis Canis_teilhardi 0.12 Canis_chihliensis Nyct_Nih -0.9 -0.6 -0.3 0.3 0.6 0.9 1.2 1.5 1.8 Component 1 Canis_variabilis -0.12 Canis_palmidens Moderately meso_Old Carnivorous -0.24 Meso_KBS-UB Canids Cuon_Jian Vulpes_chikushanensis

Cuon_Wush-0.36 meso_Kalo

Figure 7.3 PCA Scatterplot of index values for Canidae from East Africa and East Asia. Key: Red: Canids from Pliocene sites of Longdan and Longgupo. Purple circles: Canids from the Nihewan s.s . (representing canids that may have been present in Nihewan hominin sites), Yuanmou, and Gongwangling. Blue: East African canids. Green: Canids from Jianshi and Mohui. Abbreviations: Prototo_Old1: Prototocyon recki, Olduvai I; Vulpes_Long: Vulpes chikushanensis , Longdan; Meso_Old1: Canis mesomelas , Olduvai I; Meso_Kalo_Natoo: Canis mesomelas Kalochoro and Natoo members; Variabilis: Canis variabilis , Gongwangling; Lupus_Nih: Canis “lupus” from Nihewan; Nyctereutes: Nyctereutes sinensis , Longdan and Nihewan; Palmidens: Canis palmidens , Nihewan; Chih_Nih: Canis chihliensis , Nihewan; Long_Long: Canis longdanensis , Longdan; Cuon/Sinicuon: Cuon dubius/Sinicuon dubius (same species), Longdan, Longgupo, Jianshi, Mohui; Teilhardi: Canis teilhardi, Longdan; Brevi_Long: Canis brevicephalus , Longdan; Lycaon: Canis lycaonoides , Olduvai II.

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0.8

0.6

0.4 0.3647 0.335

0.2 0.1931 0.03914 Loading 0 -0.02316 -0.06083 0.02466

-0.2

-0.4

-0.6 -0.6012 -0.5913

-0.8

-1 RBL RGA RUMGA RPS P4WL PROTO CWL UM21 LBM

Figure 7.4 Loadings for component 1 of the PCA of Canidae fossils

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0.8 0.7613

0.6

0.4 0.391 0.3544

0.2 0.2149 0.0856 0.05832 0.1396 0 0.0169

-0.2 -0.2552

-0.4

-0.6

-0.8

-1 RBL RGA RUMGA RPS P4WL PROTO CWL UM21 LBM

Figure 7.5 Loadings for component 2 of the PCA for Canidae fossils.

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Hyaenidae

East Africa versus East Asia:

The East African and East Asian Hyaenidae show substantial differences. The

East Asian Plio-Pleistocene contained a variety of hyenas, including Crocuta honanensis,

Chasmaporthetes progressus and Pachycrocuta brevirostris . East Africa contains species

belonging to the genera Crocuta , Chasmaporthetes and Hyaena . Pachycrocuta became extinct in East Africa before 2.5 Ma (Werdelin and Lewis 2005).

The CA analysis shows that the Asian and African hyenas are substantially different (Figure 7.6). On the first axis, there is a separation between Pachycrocuta brevirostris and Chasmaporthetes nitidula . Pachycrocuta, Pliocrocuta, Crocuta honanensis and Crocuta sp. from KBS load with LBM and CWL, while

Chasmaporthetes nitidula loads with P4WL and RPS. Chasmaporthetes nitidula has small ratios for P4WL and RPS, which are indicative of carnivory. Chasmaporthetes progressus from Longdan did not have sufficient information to be included in analysis, but it would have lessened the difference between the hyenas in East Asia and East

Africa. On the negative end of the first axis, there are many large bodied taxa as well as higher ratios for CWL, indicating greater canine strength. On the second axis, P4WL loads on the negative end, while RPS loads on the positive end. African taxa including some Crocuta ultra , Crocuta dietrichi , and Crocuta crocuta , have lower values of RPS than Pachycrocuta and Crocuta honanensis , indicating less bone crushing robusticity.

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African Crocuta species are similar to Pachycrocuta in their size, but differ in the relative size of their premolars, which would have aided in bone cracking. Pachycrocuta

and Hyaena differ in terms of body mass. Hyaena specimens have slightly lower RBL

ratios, indicating that their relative blade lengths are somewhat smaller. Hyaena

specimens also tend to have higher RGA values than Pachycrocuta . Crocuta sp. from the

KBS member is similar to Asian Pachycrocuta .

NMDS scaling analysis of the Hamming distances shows the differences between the East Asian and East African Hyaenidae, as well as differences between the East

African species (Figure 7.7). The analysis shows differences between a set including older species such as Pachycrocuta brevirostris from Longgupo, Longdan and Haiyan, and Crocuta honanensis that are distinct from Pachycrocuta brevirostris from sites that include the early Pleistocene sites, as well as the Nihewan, Jianshi and Mohui. The differences between Chasmaporthetes nitidula and the other species are emphasized by the distance between that species and others.

The PCA scatterplot of Hyaenidae indices separates most ancient East Asian and

East African taxa from each other on the first axis (Figure 7.8). The exception is Crocuta sp. from KBS, which plots near Crocuta honanensis from Longdan. The index RPS

(relative premolar size) loads positively on the first axis (Figure 7.9). High RPS values,

such as those for Pachycrocuta , indicate a relatively larger premolar, which is correlated

with greater bone cracking ability, though the p 4 is only slightly involved in bone

cracking itself. Pachycrocuta , Crocuta honanensis , and Pliocrocuta are all located at the

positive end of the first axis, while the African species Crocuta , Crocuta ultra, Crocuta

dietrichi , Hyaena , Hyaena makapani and Chasmaporthetes nitidula all plot low on the

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first axis. RUMGA loads on the negative side of the first axis. RUMGA values were not

available for many species. RUMGA tends to be a larger index value in Hyaena , which

was found in East Africa. LBM loads negatively on the second component (Figure 7.10).

The larger taxa include African Crocuta species, as well as some of the Asian

Pachycrocuta . RGA, RPS and RUMGA load on the positively on the second axis. These species tend to have greater grinding area and relatively larger premolars.

East Asia Pliocene to Early Pleistocene:

The PCA scatterplot shows some changes over time in the adaptations of the

Hyaenidae. Species from the Haiyan Formation, including Crocuta honanensis ,

Pachycrocuta brevirostris and Pliocrocuta perrieri are plotted with species from

Longdan ( Pachycrocuta brevirostris and Crocuta honanensis ). They have similar

characteristics in terms of body mass and RPS values. Other representatives of

Pachycrocuta from later sites, such as Longgupo, Jianshi, the Nihewan and the

Pleistocene site of Yuanmou have slightly different values of RPS and LBM.

The CA analysis may also show differences over time. Pachycrocuta brevirostris

from older sites including Haiyan, Longdan and Longgupo shows similarities with other

older taxa, such as C. honanensis . This group of species also tends to be smaller in body

mass than the other Asian hyenas. Though Pachycrocuta and Crocuta honanensis were

similar in the characteristics examined here, they must have had other traits that separated

them ecologically. The role of Chasmaporthetes in the Hyaenidae in East Asia is not

clear, although it may have functioned in a role similar to a hypercarnivorous canid.

Chasmaporthetes is not recorded after the Nihewanian. The two species of

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Chasmaporthetes recorded during this time period, Ch. cf. ossifragus and Ch. progressus ,

are most likely representatives of Ch. lunensis (Kurtén & Werdelin, 1988; Galiano &

Frailey, 1977). Overall, these results show the loss of diversity within the Asian

Hyaenidae over this time period, with fewer species found in the early Pleistocene sites than in the earlier sites.

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0.16 RPS

0.12 Dietrichi_KBS_UB Ultra_KBS_Loka Ultra_Chari 0.08 Crocuta

Chasmaporthetes is present in both regions, but data were only 0.04 available for the African specimen. Chasmaporthetes is LBM not found in East Asia after the Axis 2 0 Nihewan.

CWL UM21RBLRGARUMGAProto Maka_Hyaena_UB Pachy_Gong, Nih, Xiao, Jianshi, Mohui -0.04 Pachy_Yuan Pliocrocuta Ultra_UB_OK Chasma Hyaena_Kalo_Kait Pachy_Hai, Wushan, Crocuta sp. KBS Honanensis Pachy_Long -0.08

Asian Pachycrocuta, Crocuta and Pliocrocuta differed from -0.12 African Crocuta and Hyaena. Pliocrocuta and Crocuta honanensis were similar in the characteristics examined here to Pachycrocuta. They are not found in the sites examined after the Nihewan. -0.16 P4WL

-0.18 -0.12 -0.06 0 0.06 0.12 0.18 0.24 0.3 0.36 Axis 1

Figure 7.6 CA Scatterplot of Hyaenidae category scores. Key: Red: Hyaenids from Pliocene sites of Longdan and Longgupo. Purple circles: hyaenids from the Nihewan s.s . (representing canids that may have been present in Nihewan hominin sites), Yuanmou, and Gongwangling. Blue: East African hyaenids. Green: hyaenids from Jianshi and Mohui. Abbreviations: Dietrichi: Crocuta dietrichi (KBS, UB); Ultra: Crocuta ultra (KBS, Lokalalei, Chari, UB, Okote) Pachy: Pachycrocuta brevirostris (Gongwangling, Nihewan, Xiaochangliang, Jianshi, Mohui, Yuanmou, Haiyan, Longgupo (Wushan) and Longdan). Honanensis: Crocuta honanensis (Longdan, Haiyan); Pliocrocuta: Pliocrocuta perrieri (Haiyan); Hyaena: Hyaena sp. (Kalochoro, Kaitio, UB); Maka: Hyaena makapani (UB); Chasma: Chasmaporthetes nitidula (Olduvai II).

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0.15

Diet_KBS_UB

0.1 Ultra_KBS_Loka

0.05 Hyaena_Kalo_Kait Pachy_Jian_Mohui Ultra_UB_OK Ultra_Chari Pachy_Nih_Xiao Crocuta 0 Maka_Hy_UB Pachy_Gong Pachy_Yuan

Coordinate 2

-0.05 Pachy_Hai Pliocrocuta Honan_Long_Hai

Crocuta_KBS Pachy_Wush -0.1 Pachy_long

-0.15

Chasmaporthetes is very -0.2 different from the other species; Chasma African Hyaena and Crocuta

-0.25 differ from the Asian Pachycrocuta

-0.3 -0.24 -0.18 -0.12 -0.06 0 0.06 0.12 0.18 0.24 Coordinate 1 Figure 7.7 NMDS scatterplot of Hyaenidae Hamming distances. Key: Red: Hyaenids from Pliocene sites of Longdan and Longgupo. Purple circles: hyaenids from the Nihewan s.s . (representing canids that may have been present in Nihewan hominin sites), Yuanmou, and Gongwangling. Blue: East African hyaenids. Green: hyaenids from Jianshi and Mohui. Figure Abbreviations: Dietrichi: Crocuta dietrichi (KBS, UB); Ultra: Crocuta ultra (KBS, Lokalalei, Chari, UB, Okote) Pachy: Pachycrocuta brevirostris (Gongwangling, Nihewan, Xiaochangliang, Jianshi, Mohui, Yuanmou, Haiyan, Longgupo (Wushan) and Longdan). Honanensis: Crocuta honanensis (Longdan, Haiyan); Pliocrocuta: Pliocrocuta perrieri (Haiyan); Hyaena: Hyaena sp. (Kalochoro, Kaitio, UB); Maka: Hyaena makapani (UB); Chasma: Chasmaporthetes nitidula (Olduvai II).

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Component 2 0.36 Maka_UB Large 0.3Species Hy_UB 0.24 Hy_Kalo Relatively Large 0.18 Premolars

0.12 Hy_Kait Honanensis_Haiyan Plio_Hai 0.06 Crocuta_KBS Honanensis_Long Pachy_Long -0.8 -0.64 -0.48 -0.32 -0.16 0.16 0.32 0.48 Component 1 Pachy_Wush -0.06 Ultra_OK Crocuta_Old1 Pachy_NIh Pachy_Jian -0.12 Chasma_Old2 Ultra_KBS Pachy_Yuan Ultra_Loka -0.18 Dietrichi_UB

Figure 7.8 PCA Scatterplot of Hyaenidae index values. Key: Red: Hyaenids from Pliocene sites of Longdan and Longgupo. Purple circles: hyaenids from the Nihewan s.s . (representing canids that may have been present in Nihewan hominin sites), Yuanmou, and Gongwangling. Blue: East African hyaenids. Green: hyaenids from Jianshi and Mohui. Abbreviations: Dietrichi: Crocuta dietrichi (KBS, UB); Ultra: Crocuta ultra (KBS, Lokalalei, Chari, UB, Okote) Pachy: Pachycrocuta brevirostris (Gongwangling, Nihewan, Xiaochangliang, Jianshi, Mohui, Yuanmou, Haiyan, Longgupo (Wushan) and Longdan). Honanensis: Crocuta honanensis (Longdan, Haiyan); Pliocrocuta: Pliocrocuta perrieri (Haiyan); Hyaena: Hyaena sp. (Kalochoro, Kaitio, UB); Maka: Hyaena makapani (UB); Chasma: Chasmaporthetes nitidula (Olduvai II).

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0.9775

0.8

0.6

0.4

0.2 0.1693 Loading 0.003543 0.0335 0.1088 0 0.003482 -0.02552 0.04694 0

-0.2

-0.4

-0.6

-0.8

-1 RBL RGA RUMGA RPS P4WL PROTO CWL LBM UM21

Figure 7.9 Component 1 PCA loadings for Hyaenidae

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0.8

0.6

0.4 0.3108 0.2 0.1297 0.02739 Loading 0.1008 0 0 -0.1153 -0.2 -0.172 -0.1938

-0.4

-0.6

-0.8 -0.8917 -1 RBL RGA RUMGA RPS P4WL PROTO CWL LBM UM21

Figure 7.10 Component 2 PCA loadings for Hyaenidae

Felidae

East Africa versus East Asia:

The correspondence analysis of the Felidae excluded species that only have

information for RBL, RGA and UM21, as these characters do not vary within the family.

The scatterplot (Figure 7.11) shows separation between many of the saber-toothed felids

and the rest of the Felidae. Homotherium and Megantereon fall on the negative end of the first axis, with the indices LBM and P4WL. Panthera pardus and Panthera

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palaeosinensis are plotted on the other end of the first axis with CWL. Modern

pantherines have more conically shaped canines, which are able to resist stress in

multiple directions. On the second axis, PROTO plots on the negative end, while LBM

plots on the positive side with many of the larger species.

The NMDS scatterplot of Hamming distances (Figure 7.12) highlights differences on the negative end of the first axis between species such as Panthera pardus ,

Sivapanthera linxiaensis , and Felis teilhardi, while Homotherium and Megantereon are on the positive end. On the second axis, “ Machairodus ” (probably Homotherium ) from

Olduvai II, Megantereon (from Jianshi) and Panthera pardus is differentiated from

Sivapanthera pleistocaenicus , Homotherium davitashvili and Panthera leo . For the most part, the sabertooth species are more similar to each other than to the other felids.

The PCA scatterplot of Felidae (Figure 7.13) shows a division of the family along the first axis that is primarily due to differences in LBM (Figure 7.14). Most large bodied felids load on the negative side of the first axis. RPS loads slightly positively on the first axis. RPS loads negatively on the second axis, along with CWL (Figure 7.15). Felids with high RPS and canine WL values included Panthera pardus from the Upper Burgi member, Jianshi, and Gongwangling, Sivapanthera from Gongwangling and Longdan,

Panthera leo from Olduvai II, and Lynx shansius from Longdan. Smaller relative premolar size and ratios of canine width to length were found in Homotherium from the

KBS member, Dinofelis and Megantereon .

Panthera pardus :

Panthera pardus is found in both Asia and Africa. Relatively few indices are available to compare the specimens available here. The Upper Burgi specimen is smaller

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than the two from Jianshi and Gongwangling, though it is in the same category. It is not

possible to determine from these data whether African P. pardus is smaller in general.

The CA analysis shows that small differences between the African and Asian specimens are related to the value of P4WL, which is somewhat smaller in the African specimen, indicating that the Asian leopards may have had more robust premolars. The NMDS plot of the Hamming distances shows the distance between the Panthera pardus specimens from East Asia and East Africa.

Panthera leo , Panthera tigris, and Panthera palaeosinensis :

Panthera leo fossil specimens are found in East Africa while Panthera tigris and

Panthera palaeosinensis are found in East Asia. Data from these species are limited.

They share values of RBL and RGA, as do all felids. Both have a relatively high canine

WL index, as would be expected for pantherines. Correspondence analysis showed

differences between Panthera leo from Olduvai, which had a larger ratio of P4WL compared with P. leo from Koobi Fora, which indicates greater robusticity. Panthera palaeosinensis has a larger CWL index. The NMDS analysis of the Hamming distances showed considerable differences between Panthera palaeosinensis and Panthera leo .

Sivapanthera and Acinonyx :

Acinonyx and Sivapanthera are genera of cheetahs. Values are not available to differentiate fossils of African Acinonyx from East Asian Sivapanthera . Sivapanthera linxiaensis is slightly smaller than S. pleistocaenicus . S. pleistocaenicus is in a higher category for PROTO showing more carnivorous adaptations. These differences lead to substantial distance between S. linxiaensis and S. pleistocaenicus in the NMDS Hamming

distance plot. Although S. linxianensis and P. pardus from Gongwangling plot together in

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the CA, other P. pardus are in larger CWL categories and other Sivapanthera species are larger in body size.

Megantereon :

Megantereon is present in both Africa and East Asia. Megantereon sp. specimens from the Kaitio, Upper Burgi, KBS, and Chari are too incomplete to include the sort of ecomorphological information required for these analyses. This discussion is based on M.

whitei from the Okote member. This specimen of Megantereon whitei is smaller in estimated body mass than those of Megantereon nihowanensis from Gongwangling and the Nihewan. Megantereon nihowanensis has larger values of RUMGA, and slightly larger values of P4WL compared with M. whitei , implying greater robusticity in the dentition.

The correspondence analysis shows that Megantereon whitei and Megantereon nihowanensis from Longdan have no differences in the categories for which information is available. The NDMS plot of Hamming distances shows differences in the

Megantereon species that are probably related to the number of available indices.

Megantereon nihowanensis from other sites differs in categories such as body mass, which may be due to intraspecific variation. Megantereon sp. from Jianshi plots with species of Homotherium because of lack of information for many indices. Although M.

whitei falls into the same categories as Megantereon nihowanensis , it differs in other characteristics, such as the relative size of the dentition compared with skull size

(Werdelin and Lewis 2002, Werdelin, pers. comm.) as well as the relative reduction of the anterior cheek teeth (Palmqvist et al. 2007).

Homotherium :

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Homotherium and Homotherium cf. crenatidens fossils are found in East Asia,

while Homotherium sp. fossils are found in East Africa. For the most part, the placement

of Homotherium species on the CA plot reflects typical sabertooth adaptations.

Differences, such as between Homotherium from Nihewan compared with Longdan and

Jianshi reflect missing data. The specimen from Longgupo has an anomalous value for

PROTO, placing it in a different category. Homotherium davitashvili (Haiyan) and

Homotherium sp. from the KBS have additional information available, such as the index

RPS, for which they have values that would be expected for a highly carnivorous animal,

and P4WL, for which H. davitashvili is only slightly different in values, though in a

different category.

Homotherium sp. from KBS is larger than Asian Homotherium . Homotherium sp.

from the KBS member also differs from H. davitashvili from the Haiyan in the RPS index

with H. davitashvili having a larger relative premolar size. Homotherium sp. from KBS

also has a smaller value of P4WL than H. davitashvili. Due to these differences,

Homotherium davitashvili and Homotherium sp. from the KBS member are placed

relatively far apart on the PCA scatterplot.

Dinofelis :

Dinofelis fossils are only found in the East African sample, though the genus is

known from Asian localities in China and the Siwaliks. There is no information available

about Dinofelis sp. D from Olduvai beyond general felid characteristics. There are no

categorical differences between Dinofelis aronoki and Dinofelis piveteaui for the indices

available. Differences in the placement of these species on the scatterplot reflect the

additional information available for D. piveteaui . D. piveteaui is in a low CWL category,

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which is typical of sabertoothed felids, though its value is higher than those of

Homotherium . The category assignments for RUMGA and PROTO point to highly

carnivorous adaptations. Thus, Dinofelis piveteaui plots near Megantereon and

Homotherium , indicating similarity in the ecomorphological characteristics measured

here. Dinofelis species are also closer to other sabertooth species in the NMDS Hamming

distance plot. However, in other craniodental traits, Dinofelis shares characteristics with

Panthera .

In the PCA scatterplot, Dinofelis plots with other sabertoothed felids. Dinofelis

piveteaui has a slightly greater RUMGA value than Homotherium and a greater RUMGA

than Megantereon . The value for PROTO is smaller than many of the PROTO values for

other felids. CWL is similar to the value for Megantereon , and not as large as that for

Panthera .

Lynx shansius , Felis teilhardi , Felis microta , Felis sp. and Caracal :

In the NMDS scatterplot, these species are separated from each other, indicating differences in feeding adaptations and body mass. In the CA scatterplot, Lynx shansius does not appear to have counterparts in the East African Felidae. It differs in size from many of the other African felids. Lynx tends to have bigger RPS values than Asian Felis and is larger than Felis teilhardi and Caracal . In the CA analysis, these felids are located

low on the second axis. They share features of an intermediate PROTO value with

Panthera pardus and Sivapanthera linxiaensis . Lynx shansius also shares features of

CWL with many of the non-sabertooth taxa. Felis teilhardi differs from Lynx shansius

and Felis microta in the index P4WL, in which F. teilhardi has a value that might

indicate more robusticity. This is shared with P. leo , P. pardus and Sivapanthera . Lynx

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shansius and F. microta have a value that indicates more flesh consumption. In the PCA

scatterplot, F. microta differs in its small body mass, which drives its position on the first axis. Differences in the body mass of these smaller felids would have had a significant effect upon their competitive interactions (Dayan et al. 1990).

East Asia Pliocene to Early Pleistocene:

Pliocene and Pleistocene species of Homotherium appear to be similar in craniodental adaptations, such as RUMGA and CWL, and in the values of LBM. This niche for a hypercarnivorous hunter of large prey appears to have been continuously filled. Species of Megantereon , which are present at many sites, are similar in the indices of LBM, CWL, and P4WL. The RPS value of the Gongwangling Megantereon is smaller, but it is not out of the range for the genus. The Sivapanthera and Lynx lineages are present from the Pliocene into the Early Pleistocene. These lineages are found at Longdan and at later Pleistocene sites with little change in the available measurements of the properties examined here. Differences between S. linxiaensis and S. pleistocaenicus are due to body mass. Panthera pardus also shows little change between older sites such as

Longgupo and younger ones. The correspondence analysis shows that there is no size trend through time for F. teilhardi body mass differences. The fate of lineages such as

Felis sp. at Longgupo is not known, but these felids are too small to have had a significant effect upon the niche of hominins. Panthera palaeosinensis is found at the older sites of Longdan and Longgupo. It is not known whether this form is ancestral to P. tigris , which is found at Gongwangling and Yuanmou and did not have sufficient

199

information to be included in the analysis. If P. tigris at those sites had similar body mass, it would likely represent a similar species.

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Large Species

0.15 Sivapanthera_pleistocaenicus LBM

0.1 Sabertoothed Felids P_Leo_Old Homotherium_davitasvili Homotherium_crendatidens Panthera_leo P_paleosinensis CWL Panthera_KBS 0.05 Homotherium_KBS D_piveteaui_OK D_aronoki_UB Metailurus Megantereon_nihowanensis P_pardus_Jian 0 Homotherium_cf._crenatidens Panthera_cf._pardus Axis 2 Machairodus RBLRGARUMGARPSUM21 Sivapanthera_linxiaensis -0.05 Megantereon_Jian Panthera_pardus P_Pardus_Gong Homotherium_cf._crenatidens P4wl Lynx_shansius M_niho_Long Felis_teilhardi -0.1 Megantereon_whitei

-0.15 Caracal_UB Proto Felis_teilhardi -0.2

-0.25

F_microta_Wush -0.24 -0.16 -0.08 0 0.08 0.16 0.24 0.32 Axis 1 Figure 7.11 CA Scatterplot of Felidae category scores. Key: Red: Felids from Pliocene sites of Longdan and Longgupo. Purple circles: Felids from the Nihewan s.s . (representing felids that may have been present in Nihewan hominin sites), Yuanmou, and Gongwangling. Blue: East African felids. Green: felids from Jianshi and Mohui. Abbreviations: Sivapanthera_pleistocaenicus: Sivapanthera pleistocaenicus (Jianshi, Gongwangling, Longgupo, Nihewan); P_Leo_Old: Panthera leo (Olduvai); Homotherium_davistashvili: Homotherium davitashvili (Haiyan); P_palaeosinensis: Panthera palaeosinensis (Longdan); Panthera_leo (Okote); Homotherium_crenatidens: H. crenatidens (Longgupo, Nihewan, Longdan, Jianshi); D_piveteaui: Dinofelis piveteaui (Okote); D_aronoki_UB: Dinofelis aronoki (UB); Metailurus (Olduvai II); Megantereon_nihowanensis: M. nihowanensis (Gongwangling, Longdan, Nihewan, Yuanmou, Haiyan); Homotherium cf. crenatidens (Gongwangling, Nihewan, Longgupo); Megantereon _Jian: Megantereon sp. (Jianshi); Machairodus (Olduvai II); M_niho_Long: Megantereon nihowanensis (Longdan); Megantereon whitei (Okote); Homotherium (KBS); P_pardus_Jian: Panthera pardus (Jianshi); Panthera cf. pardus (Longgupo); P_pardus_Gong: P. pardus (Gongwangling); Panthera pardus (Olduvai I, II, UB, KBS); Felis teilhardi (Jianshi, Longdan, Longgupo, Mohui); Caracal_sp_UB: Caracal sp. (UB); Lynx_shansius (Longdan, Nihewan); F_microta_Wush: Felis microta (Longgupo)

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0.16 Metailurus Homotherium_davitasvili 0.12 Panthera_leo Sivapanthera_pleistocaenicus

0.08 Panthera_sp . Felis_teilhardi Panthera_leo 0.04 Homotherium_crendatidens Coordinate 2 Homotherium_ Sivapanthera_linxiaensis Dinofelis_aronoki 0 Panthera_pardus Caracal_sp cf._crenatidens Homotherium_sp . Felis_teilhardi Dinofelis_piveteaui -0.04 Panthera_pardus Megantereon_nihowanensis

Lynx_shansius Felis_microta -0.08 Megantereon_nihowanensis

Panthera_palaeosinensis Megantereon_whitei -0.12 Panthera_pardus Machairodus Megantereon_sp. -0.16

-0.2 -0.25 -0.2 -0.15 -0.1 -0.05 0 0.05 0.1 0.15 Coordinate 1 Figure 7.12 NMDS scatterplot of Hamming distances for Felidae. Key: Red: Felids from Pliocene sites of Longdan and Longgupo. Purple circles: Felids from the Nihewan s.s . (representing felids that may have been present in Nihewan hominin sites), Yuanmou, and Gongwangling. Blue: East African felids. Green: felids from Jianshi and Mohui. Abbreviations: Sivapanthera_pleistocaenicus: Sivapanthera pleistocaenicus (Jianshi, Gongwangling, Longgupo, Nihewan); P_Leo_Old: Panthera leo (Olduvai); Homotherium_davistashvili: Homotherium davitashvili (Haiyan); P_palaeosinensis: Panthera palaeosinensis (Longdan); Panthera_leo (Okote); Homotherium_crenatidens: H. crenatidens (Longgupo, Nihewan, Longdan, Jianshi); D_piveteaui: Dinofelis piveteaui (Okote); D_aronoki_UB: Dinofelis aronoki (UB); Metailurus (Olduvai II); Megantereon_nihowanensis: M. nihowanensis (Gongwangling, Longdan, Nihewan, Yuanmou, Haiyan); Homotherium cf. crenatidens (Gongwangling, Nihewan, Longgupo); Megantereon _Jian: Megantereon sp. (Jianshi); Machairodus (Olduvai II); M_niho_Long: Megantereon nihowanensis (Longdan); Megantereon whitei (Okote); Homotherium (KBS); P_pardus_Jian: Panthera pardus (Jianshi); Panthera cf. pardus (Longgupo); P_pardus_Gong: P. pardus (Gongwangling); Panthera pardus (Olduvai I, II, UB, KBS); Felis teilhardi (Jianshi, Longdan, Longgupo, Mohui); Caracal_sp_UB: Caracal sp. (UB); Lynx_shansius (Longdan, Nihewan); F_microta_Wush: Felis microta (Longgupo)

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Component 2

Homo_KBS 0.64 Many forms are present throughout the time studied including Homotherium, Larger species 0.48 Megantereon , Sivapanthera , Panthera pardus and Lynx.

Meg_Gong 0.32 Microta_Wush

Meg_whitei_Ok Di no_aron_UB Dino_pivet_OK 0.16 Teilhardi_Moh Teilhardi_Jian Homo_Dav_Hai Teilhardi_Wush Meg_Nih Homo_Nih Homo_Jian Homo_Long -0.8 -0.6 -0.4 -0.2 Siva_Jian 0.2 0.4 0.6 0.8 Meg_Long Teilhardi_Long Component 1 Lynx_Nih Siva_Gong Lynx_Long -0.16 Siva_Long

-0.32 Panthera_pardus_UB

-0.48 Panthera_leo_Old2 Panthera_pardus_Jian Panthera_pardus_Gong Figure 7.13 PCA Scatterplot of Felidae index values. Key: Red: Felids from Pliocene sites of Longdan and Longgupo. Purple circles: Felids from the Nihewan s.s . (representing felids that may have been present in Nihewan hominin sites), Yuanmou, and Gongwangling. Blue: East African felids. Green: felids from Jianshi and Mohui. Abbreviations: Meg_Gong: Megantereon nihowanensis (Gongwangling); Meg_whitei: Megantereon whitei (Okote); Microta_Wush: Felis microta (Longgupo); Dino_pivet_Ok: Dinofelis piveteaui (Okote); D_aron_UB: Dinofelis aronoki (UB); Homo_Dav_Hai: Homotherium davitashvili (Haiyan); Meg_Nih: Megantereon nihowanensis (Nihewan); Meg_Long: Megantereon nihowanensis (Longdan); Siva_Gong: Sivapanthera pleistocaenicus (Gongwangling); Siva_Long: Sivapanthera linxiaensis (Longdan); Homo_Nih: Homotherium crenatidens (Nihewan); Homo_Long: Homotherium crenatidens (Longdan); Siva_Jian: Sivapanthera pleistocaenicus (Jianshi); Panthera_pardus_UB: Panthera pardus (UB); Teilhardi_Wush: Felis teilhardi (Longgupo); Teilhardi_Moh: Felis teilhardi (Mohui); Teilhardi_Jian: Felis teilhardi (Jianshi); Teilhardi_Long: Felis teilhardi (Longdan); Lynx_Nih: Lynx shansius (Nihewan); Lynx_Long: Lynx shansius (Longdan); Panthera_pardus_Jian: Panthera pardus (Jianshi); Panthera_pardus_Gong: Panthera pardus (Gongwangling)

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0.8

0.6

0.4

0.2 0.06208 0.03622 Loading 6.366E-17 -1.628E-42 0 0 0.01186 0.007829 0.02637

-0.2

-0.4

-0.6

-0.8 -0.997

-1 RBL RGA RUMGA RPS P4WL PROTO CWL UM21 LBM

Figure 7.14 PCA loadings for component 1 for Felidae.

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0.8

0.6

0.4

0.2

Loading -8.39E-17 -0.04741 -2.773E-40 0 0 -0.01065 -0.07438 -0.06919 -0.2 -0.2783 -0.4

-0.6

-0.8 -0.9539

-1 RBL RGA RUMGA RPS P4WL PROTO CWL UM21 LBM

Figure 7.15 PCA loadings for component 2 for Felidae.

Ursidae

East Africa versus East Asia:

Ursidae became extinct in East Africa before the focal time period of this study

(Werdelin and Lewis 2005). Therefore, they are only a factor for hominins in East Asia,

though it is possible that other East African taxa played the roles of the ursids. East Asian

ursids include Ailuropoda and Ursus .

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The CA scatterplot for Ursidae shows separation between two of the Ailuropoda species and Ursus on the first axis (Figure 7.16). RPS and P4WL were dropped from this analysis because no values were available for these indices for ursids. Ailuropoda melanoleuca from Gongwangling could not be included due to lack of information.

RUMGA and PROTO load on the positive end of the first axis with Ursus , which has a larger upper molar grinding area and is in a higher category for PROTO. These characteristics are indicative of omnivory. RBL, RGA, CWL and LBM load on the negative side of the first axis. All ursids in this set are in the same category for body mass, and the others are not available for both Ursus and Ailuropoda .

UM21 loads high on the second axis of the CA. UM21 is greater in Ursus than in

Ailuropoda , indicating a greater grinding capacity. Ailuropoda has less grinding area than

Ursus , which is characteristic of that herbivorous genus (Sacco and Van Valkenburgh

2004). Both the CA and the NMDS Hamming distance analysis shows that the Ursus species do not differ from each other in category scores or in disparity (Figure 7.17).

Ailuropoda wulingshanensis differs from A. microta in its smaller RUMGA value, which implies less upper molar grinding area, and in its smaller value of PROTO. Given the small number of categories available, Hamming distances between A. microta and A. wulingshanensis are fairly great.

In the PCA (Figure 7.18), Ailuropoda species are clearly distinguished from the

Ursus species. RUMGA loads positively on the first axis (Figure 7.19), with the Ursus species, which is an expected pattern for ursids. LBM loads slightly negatively. UM21 loads on the positively on the second axis, while RUMGA loads negatively (Figure 7.20).

206

The species within each genus differ from each other in small ways. Ursus cf. etruscus

from Gongwangling has a greater UM21 than Ursus aff. thibetanus (Longgupo) and U. thibetanus (Mohui), implying greater molar grinding area. Also, Ursus aff. thibetanus from Longgupo had a greater value of RGA compared with Ursus sp. from Jianshi.

East Asia Pliocene to Early Pleistocene:

Ailuropoda microta , found in the Pliocene site of Longgupo and at Mohui, shows

no differences in category assignment over time. Ailuropoda wulingshanensis differed in

some categories, but does not appear to be fundamentally different. Zheng et al. (2004)

believe A. wulingshanensis to be an intermediate form between A. microta and A.

melanoleuca . A. wulingshanensis from Jianshi is described as intermediate in body mass

between A. microta and A. melanoleuca , with A. melanoleuca as the larger species

(Zheng et al. 2004). On the PCA scatterplot, A. melanoleuca is similar to A. microta from

Mohui and A. wulingshanensis from Jianshi. The evidence seems to indicate that the niche was filled continuously.

Ursus is recorded from the Pliocene site of Longgupo. There is also an Ursus

specimen from Longdan that was not yet available for study. There is no evidence here to

show consistent differences over time between the Ursus specimens from East Asia.

These analyses do not support the idea of a Pliocene to Pleistocene change in the ecomorphology of Ursus feeding adaptations. Ursus cf . etruscus from Gongwangling

differs from the other Ursus in the ratio value of UM21, which is larger in that species.

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0.04

0.032 UM21

0.024 Large Molar Grinding Areas Ailuropoda_wulingshanensis Omnivory 0.016

0.008 Ursus_thibetanus Ursus_aff._thibetanus Axis 2 Ursus_sp. 0

-0.008 LBMRBLRGACWL RUMGA Proto -0.016

-0.024 Ailuropoda_microta

-0.2 -0.15 -0.1 -0.05 0 0.05 0.1 0.15 0.2 0.25 Axis 1

Figure 7.16 CA scatterplot of Ursidae category scores. Key: Red: Ursids from Pliocene sites of Longdan and Longgupo. Green: ursids from Jianshi and Mohui. The species present are: Ailuropoda wulingshanensis (Jianshi); Ailuropoda microta (Longgupo and Mohui); Ursus thibetanus (Mohui); Ursus aff. thibetanus (Longgupo); Ursus sp. (Jianshi).

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0.25

Ailuropoda_wulingshanensis 0.2

0.15

0.1

Coordinate 2 0.05 Ursus_thibetanus Ursus_sp. 0 Ursus_aff._thibetanus

-0.05

-0.1

-0.15 Ailuropoda_microta

-0.2 -0.5 -0.4 -0.3 -0.2 -0.1 0 0.1 0.2 0.3 Coordinate 1

Figure 7.17 NMDS Scatterplot of Hamming distance values for Ursidae. Key: Red: Ursids from Pliocene sites of Longdan and Longgupo. Green: ursids from Jianshi and Mohui. The species present are: Ailuropoda wulingshanensis (Jianshi); Ailuropoda microta (Longgupo and Mohui); Ursus thibetanus (Mohui); Ursus aff. thibetanus (Longgupo); Ursus sp. (Jianshi).

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Very large relative grinding Etruscus_Gong Ursus specimens are similar area through time and Ailuropoda is 0.08 continuously present as an herbivore 0.06

0.04

Microta_Mohui Melano_Gong 0.02 Wuling_Jian Large relative grinding area

-0.5 -0.4 -0.3 -0.2 -0.1 0.1 0.2 0.3 Component 1 -0.02

-0.04 Thibet_Wush

Microta_Wush -0.06

Component 2 Ursus_Jian

Figure 7.18 PCA Scatterplot of Ursidae index values. Key: Red: Ursids from Pliocene sites of Longdan and Longgupo. Purple: Gongwangling; Green: ursids from Jianshi and Mohui. The species present are: Wuling_Jian: Ailuropoda wulingshanensis (Jianshi); Microta_Mohui/Microta_Wush: Ailuropoda microta (Mohui and Longgupo); Thibet_Wush: Ursus aff. thibetanus (Longgupo); Ursus_Jian: Ursus sp. (Jianshi); Etruscus_Gong: Ursus etruscus (Gongwangling).

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0.9189 0.8

0.6

0.4 0.3358

0.2 0.1893 Loading 0 -0.0286 0.002262 -0.0005676 -0.07813 -0.2

-0.4

-0.6

-0.8

-1 RBL RGA RUMGA PROTO CWL UM21 LBM

Figure 7.19 Loadings for component 1 of PCA scatterplot for Ursidae.

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0.9175 0.8

0.6

0.4

0.2844 0.2 0.1892 0.1196 Loading 0 0.007498 -0.03001

-0.2 -0.162

-0.4

-0.6

-0.8

-1 RBL RGA RUMGA PROTO CWL UM21 LBM

Figure 7.20 Loadings for component 2 of PCA scatterplot for Ursidae.

Mustelidae :

East Africa versus East Asia:

Many different kinds of mustelids are present in the East African and East Asian

carnivore guilds. Species are distinguished by differences in LBM and in RBL. Though

the East Asian guild preserves a number of species, there is limited information available

from the East African Plio-Pleistocene mustelids. The East African species that are

preserved tend to have high LBM, indicating that the larger species are more likely to be

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preserved. Also, many of the East African mustelids are aquatic (Werdelin and Lewis

2005).

On the CA for Mustelidae (Figure 7.21), P4WL and RGA plot on the negative end

of the axis with taxa such as Martes sp. 1 and Mellivora sp. while LBM and RBL plot on the positive end with taxa such as Meles cf. leucurus . Martes sp. 1 and Mellivora sp. have small body masses and relatively large blade lengths (RBL). Meles cf. leucurus has high body mass, and a relatively small relative blade length. The first axis shows a general contrast between more carnivorous and omnivorous mustelids. The NMDS scatterplot of

Hamming distances (Figure 7.22) shows the similarities between the species of Martes sp. 1 (Jianshi) and Mellivora benfieldi , as well as the similarities between a cluster of a cluster of species with small relative blade lengths that includes Meles , Martes sp. 2,

Eirictis robusta and Torolutra ougandensis .

The PCA scatterplot shows very little overlap between the mustelids of East

Africa and East Asia (Figure 7.23). LBM loads high on the first axis, while RPS and RBL load negatively (Figure 7.24). On the second axis, RPS loads positively, while RGA and

RUMGA load negatively. The East Asian sample includes omnivorous mustelids such as

Meles, Arctonyx and Martes sp. 2. Mustelids from East Africa of similar size, including

Torolutra from the KBS member, have high RPS values, which may relate to feeding on hard objects such as mollusks. Eirictis robusta from Longdan combines traits of relatively high RPS with large ratios for RGA and RUMGA. Another type of mustelid from East Africa is Mellivora , which differs from Asian species in this study in body size and high RBL value. The genus is not represented in the Asian localities sampled.

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Axis 2 0.25

0.2 Arctonyx_cf._minor

0.15 Relatively Carnivorous Large body mass 0.1 High Relative Blade Length RBL RGA 0.05 Eirictis_robusta Martes_sp. Meles_chiai/cf. chiai Meles_teilhardi Martes_sp._2 0 CWL Torolutra_cf._Ougandensis RUMGAProtoUM21RPS Mellivora_benfieldi -0.05 Meles_cf._leucurus

Torolutra_sp. -0.1 Lutrinae Martes_sp._1 LBM -0.15 P4wl -0.2 -0.3 -0.24 -0.18 -0.12 -0.06 0 0.06 0.12 0.18 0.24 Axis 1

Figure 7.21 CA Scatterplot of Mustelidae category scores. Red: mustelids from Pliocene sites of Longdan and Longgupo. Purple circles: Mustelids from the Nihewan s.s . (representing mustelids that may have been present in Nihewan hominin sites), Yuanmou, and Gongwangling. Blue: East African mustelids. Green: mustelids from Jianshi and Mohui. Species: Martes sp. 1 (Jianshi); Mellivora benfieldi (UB); Arctonyx cf. minor (Longgupo); Meles teilhardi (Longdan); Martes sp. (Mohui); Torolutra sp. (KBS); Eirictis robusta (Longdan); Meles chiai /Meles cf. chiai (Nihewan and Longgupo); Martes sp. 2 (Jianshi); Torolutra cf. ougandensis (UB); Lutrinae (UB); Meles cf. leucurus (Gongwangling)

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0.25 Torolutra sp. Lutrinae 0.2

0.15

0.1 Coordinate 2 0.05 Meles_teilhardi Meles_cf._leucurus 0 Meles_chiai Eirictis_robusta

Torolutra_cf._Ougandensis Martes sp 2 -0.05 Mellivora_benfield Martes_sp._1 Martes sp. -0.1

-0.15 Arctonyx_cf._minor

- -0.48 -0.4 -0.32 -0.24 -0.16 -0.08 0 0.08 0.16 Coordinate 1

Figure 7.22 NMDS Scatterplot of Hamming distances for Mustelidae. Red: mustelids from Pliocene sites of Longdan and Longgupo. Purple circles: Mustelids from the Nihewan s.s . (representing mustelids that may have been present in Nihewan hominin sites), Yuanmou, and Gongwangling. Blue: East African mustelids. Green: mustelids from Jianshi and Mohui. Species: Martes sp. 1 (Jianshi); Mellivora benfieldi (UB); Arctonyx cf. minor (Longgupo); Meles teilhardi (Longdan); Martes sp. (Mohui); Torolutra sp. (KBS); Eirictis robusta (Longdan); Meles chiai /Meles cf. chiai (Nihewan and Longgupo); Martes sp. 2 (Jianshi); Torolutra cf. ougandensis (UB); Lutrinae (UB); Meles cf. leucurus (Gongwangling)

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0.5 Lutrinae_UB

Toro_KBS 0.4

0.3 Eirictis 0.2 High body mass Omnivorous 0.1 Carnivorous Toro_Oug_UB Martes2_Jian -0.96 -0.8 -0.64 -0.48 -0.32 -0.16 0.16 0.32 0.48 Component 1 Mellivora_UB -0.1 Chiai_Wush Martes1_Jian -0.2 Leucurus_Gong Chiai_Nih Arctonyx_Wush MelTeil_Long -0.3 Component 2

Figure 7.23 PCA Scatterplot for index values for Mustelidae. Red: mustelids from Pliocene sites of Longdan and Longgupo. Purple circles: Mustelids from the Nihewan s.s . (representing mustelids that may have been present in Nihewan hominin sites), Yuanmou, and Gongwangling. Blue: East African mustelids. Green: mustelids from Jianshi and Mohui. Species: Martes1_Jian: Martes sp. 1 (Jianshi); Mellivora_UB: Mellivora benfieldi (UB); Arctonyx_Wush: Arctonyx cf. minor (Longgupo); MelTeil_Long: Meles teilhardi (Longdan); Toro_KBS: Torolutra sp. (KBS); Eirictis: Eirictis robusta (Longdan); Chiai_Nih and Chiai_Wush: Meles chiai /Meles cf. chiai (Nihewan and Longgupo); Martes2_Jian: Martes sp. 2 (Jianshi); Toro_Oug_UB: Torolutra cf. ougandensis (UB); Lutrinae_UB: Lutrinae (UB); Leucurus_Gong: Meles cf. leucurus (Gongwangling)

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0.8865 0.8

0.6

0.4

0.2 0.05779 Loading 0.07574 -0.009194 0 9.836E-41 -0.01382 0.03118

-0.2 -0.2299

-0.4 -0.3885

-0.6

-0.8

-1 RBL RGA RUMGA RPS P4WL PROTO CWL UM21 LBM

Figure 7.24 PCA of Mustelidae loadings for component 1

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0.8 0.8025

0.6

0.4 0.4408

0.2694 0.2 0.02354 Loading 0 5.902E-39 -0.05266 -0.06482

-0.2 -0.2095 -0.1939

-0.4

-0.6

-0.8

-1 RBL RGA RUMGA RPS P4WL PROTO CWL UM21 LBM

Figure 7.25 PCA of Mustelidae loadings for component 2

East Asia Pliocene to Early Pleistocene:

Meles :

Species of Meles persist through the Pliocene and into the early Pleistocene.

Though the species show some differences in RBL and RPS, there are no consistent

trends, and there is nothing to indicate that the niche changed substantially. In the

correspondence analysis (Figure 7.21), the amount of information available for the Meles

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species differed, but with the exception of M. leucurus , which differs in body mass, they

are largely similar in categories.

Arctonyx :

Species of Arctonyx are present from sites such as Longgupo, Jianshi and Mohui

(A. collaris from these two latter sites did not have sufficient information to be included

in the CA), indicating presence of the genus from the late Pliocene to early Pleistocene.

The indices available do not show substantial differences and none of the categories

differ.

Herpestidae, Prionodontidae and Viverridae :

East Africa versus East Asia

Herpestidae are found only in the East African carnivore guild, while Viverridae

are found in both. The specimen of Prionodon from the Prionodontidae was found at

Jianshi. Many of the species in these families are small and less likely to be preserved.

The CA (Figure 7.26) of these animals shows a separation between more

carnivorous animals, such as Prionodon , and more omnivorous taxa, such as Mungos dietrichi . Mungos dietrichi loads with RBL and RGA. It has a small relative blade length and a relatively large grinding area. LBM loads high on the second axis, with the species that are larger in body mass, including Megaviverra pleistocaenica and Viverra sp.

(Jianshi). RUMGA, UM21 and P4WL plot on the negative side of the second axis. In this group, information is only available for M. pleistocaenica for RUMGA and UM21. That species is in categories indicative of more carnivorous tendencies. For P4WL, only M. dietrichi has a value that tends toward omnivory. The NMDS scatterplot (Figure 7.27)

219 emphasizes the differences between the more omnivorous Mungos dietrichi and the carnivorous Megaviverra , Prionodon and Herpestes .

On the PCA scatterplot (Figure 7.28), RPS loads positively and LBM loads negatively on the first component (Figure 7.29). For the second component, RPS and

LBM load negatively and RGA and RBL load positively (Figure 7.30). Megaviverra and

Pseudocivetta are similar in body mass, but cannot be compared in other indices.

Pseudocivetta has omnivorous features including small relative blade length, and a large amount of lower molar grinding area. A specimen of Viverridae sp. from the Upper Burgi member is similar to Megaviverra though it has a lower body mass and RPS score.

Prionodon sp. from Jianshi is a small creature with carnivorous dietary adaptations.

Viverra sp. from Jianshi might have some omnivorous adaptations, with a relatively high value for UM21 and a lower RBL value. However, Viverra sp. and Viverricula malaccensis from Yuanmou have lower values of PROTO, indicating more carnivory.

East Asia Pliocene to Early Pleistocene:

Viverrids are not sufficiently sampled in East Asia to determine whether there was continuity of adaptations from the Pliocene through the early Pleistocene. Species are not shared between sites and many species are missing from the record. These species are unlikely to have affected hominins in a substantial way.

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LBM 0.36

0.3 Larger Body Mass 0.24

0.18 Megaviverra pleistocaenica

0.12 Carnivorous Omnivorous Axis 2 Viverra sp. 0.06 RBL

0 RPS

Proto RGA -0.06 P4WL UM21RUMGA Mungos dietrichi -0.12 Herpestes primitivus Prionodon sp.

-0.18 -0.4 -0.32 -0.24 -0.16 -0.08 0 0.08 0.16 0.24 0.32 Axis 1

Figure 7.26 CA Scatterplot of category scores for Herpestidae, Prionodontidae and Viverridae. Key: Red: Viverrids from Longgupo. Blue: Herpestids from East Africa; Green: Viverridae and Prionodontidae from Jianshi. Species: Megaviverra pleistocaenica (Longgupo); Viverra sp. (Jianshi); Prionodon sp. (Jianshi); Herpestes primitivus (Olduvai I); Mungos dietrichi (Olduvai I)

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Viverra p. 0.4

0.32

0.24 Coordinate 2

0.16 Herpestes primitivus 0.08

-0.08

Mungos dietrichi -0.16 Prionodon sp.

-0.24 Megaviverra pleistocaenica

-0.36 -0.24 -0.12 0 0.12 0.24 0.36 0.48 0.6 Coordinate 1

Figure 7.27 NMDS Scatterplot of Hamming distances for Viverridae, Herpestidae and Prionodontidae species. Key: Red: Viverrids from Longgupo. Blue: Herpestids from East Africa; Green: Viverridae and Prionodontidae from Jianshi. Species: Megaviverra pleistocaenica (Longgupo); Viverra sp. (Jianshi); Prionodon sp. (Jianshi); Herpestes primitivus (Olduvai I); Mungos dietrichi (Olduvai I)

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1.8 Component 2 Prionodon 1.5

1.2

0.9

Herp_Prim 0.6

0.3 Mungos_diet Viverricula -0.9 -0.6 -0.3 Viverra_Jian 0.3 0.6 0.9 1.2 1.5 1.8 Viverridae_UB Component 1 -0.3 Megaviv_Wush Pseudocivetta_KBS -0.6 Pseudocivetta_Old Larger Body Mass

-0.9

Figure 7.28 PCA scatterplot of index values for Viverridae, Herpestidae and Prionodontidae. Key: Red: Viverrids from Longgupo. Blue: Herpestids from East Africa; Green: Viverridae and Prionodontidae from Jianshi. Species: Megaviv_Wush: Megaviverra pleistocaenica (Longgupo); Viverra_Jian: Viverra sp. (Jianshi); Prionodon: Prionodon sp. (Jianshi); Herp_Prim: Herpestes primitivus (Olduvai I); Mungos_diet: Mungos dietrichi (Olduvai I); Viverridae_UB: Viverridae (UB); Pseudocivetta_Old: Pseudocivetta ingens (Olduvai I and II); Pseudocivetta_KBS: Pseudocivetta ingens (KBS); Viverricula: Viverricula malaccensis (Yuanmou)

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0.8 0.8055

0.6

0.4

0.2592 0.2 0.0137 Loading 0.09527 0 3.536E-18 -0.03454 0.0468

-0.2

-0.4

-0.5208 -0.6

-0.8

-1 RBL RGA RUMGA RPS P4WL PROTO UM21 LBM

Figure 7.29 Component 1 loadings for PCA of Viverridae, Herpestidae and Prionodontidae.

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0.8

0.6

0.4

0.2 0.03267 Loading 0.08626 1.679E-18 0 -0.02501 -0.01835

-0.2

-0.3306 -0.4 -0.4273

-0.6

-0.8 -0.8359

-1 RBL RGA RUMGA RPS P4WL PROTO UM21 LBM

Figure 7.30 Component 2 loadings for PCA of Viverridae, Herpestidae and Prionodontidae.

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Table 7.1 P-values (above) and Bonferroni values (below) for the MANOVA of carnivore family centroids in East Asia and East Africa. Less than 0.0001 is indicated by a *.

Felidae Felidae Asia Felidae Africa Hyaenidae Asia Hyaenidae Africa Canidae Asia Canidae Africa Ursidae Mustelidae Asia Mustelidae Africa Felidae NA 0.0009 * * * * * * * Asia Felidae 0.032 NA * * * * * * * Africa Hyaen. * * NA * * * * * * Asia Hyaen. * * * NA * * * * * Africa Canid. * * * * NA 0.599 * * 0.0001 Asia Canid. * * * * 1 NA 0.022 0.033 0.010 Africa Ursid. * * * * 0.0002 0.796 NA * 0.014 Asia Mustel. * * * * * 1 * NA 0.404 Asia Mustel. * * 0.0005 * 0.003 0.372 0.497 1 NA Africa

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Table 7.2 Mean distance to the centroid, maximum distance to the centroid, species and site of maximum distance for carnivore families from Africa and Asia using CA analysis loadings. Family Continent MDC Max Species Site MDC Hyaenidae Asia 0.106 0.191 Pliocrocuta perrieri Haiyan Hyaenidae Africa 0.133 0.256 Chasmaporthetes nitidula Olduvai II Felidae Africa 0.140 0.321 Dinofelis piveteaui Okote Mustelidae Africa 0.144 0.194 Mellivora benfieldi Upper Burgi Canidae Asia 0.187 0.278 Canis palmidens Nihewan Ursidae Asia 0.189 0.230 Ailuropoda wulingshanensis Jianshi Mustelidae Asia 0.195 0.390 Arctonyx cf. minor Longgupo Canidae Africa 0.243 0.386 Prototocyon recki Olduvai I Felidae Asia 0.248 0.405 Felis microta Longgupo

Table 7.3 Mean distance to the centroid, maximum distance to the centroid, species and site of maximum distance for carnivore guilds at various sites using CA analysis loadings. Locality MDC Max Species MDC Olduvai II 0.243 0.378 Canis lycaonoides Upper Burgi 0.268 0.374 Megantereon whitei Okote 0.290 0.387 Torolutra sp . KBS 0.292 0.486 Torolutra cf . ougandensis Haiyan 0.331 0.502 Megantereon nihowanensis Longgupo 0.374 0.638 Arctonyx cf . minor Yuanmou 0.378 NA NA Olduvai I 0.383 0.577 Mungos dietrichi Mohui 0.387 0.514 Arctonyx cf . minor (representing A. collaris ) Jianshi 0.392 0.579 Arctonyx cf . minor (representing A. collaris ) Nihewan 0.396 0.502 Ursus sp . Gongwangling 0.410 0.505 Ursus sp . Longdan 0.609 1.292 Vulpes chikushanensis

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Table 7.4 MANOVA of carnivore guilds (using CA) at selected sites in East Africa and East Asia. Overall, the sites were not different (p=0.4349). Bonferroni post-hoc pairwise comparisons (below the diagonal) are not significant.

Olduvai I Olduvai II Upper Burgi Nihewan Longgupo Longdan KBS Okote Gongwangling Haiyan Jianshi Mohui Old. I --- 0.83 0.48 0.43 0.34 0.43 0.71 0.81 0.60 0.45 0.77 0.64 Old. II 1 --- 0.73 0.14 0.19 0.11 0.54 0.55 0.42 0.34 0.48 0.32 UB 1 1 --- 0.15 0.04 0.48 0.93 0.97 0.19 0.23 0.37 0.31 Nihewan 1 1 1 --- 0.33 0.9 0.11 0.32 0.81 0.83 0.73 0.70 Longgupo 1 1 1 1 --- 0.72 0.02 0.19 0.83 0.24 0.98 0.88 Longdan 1 1 1 1 1 --- 0.22 0.27 0.22 0.75 0.88 0.44 KBS 1 1 1 1 1 1 --- 0.88 0.26 0.49 0.22 0.31 Okote 1 1 1 1 1 1 1 --- 0.13 0.13 0.65 0.29 Gong- 1 1 1 1 1 1 1 1 --- 0.47 0.98 0.96 wangling Haiyan 1 1 1 1 1 1 1 1 1 --- 0.6 0.79 Jianshi 1 1 1 1 1 1 1 1 1 1 --- 0.92 Mohui 1 1 1 1 1 1 1 1 1 1 1 ---

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Correspondence Analysis of all Families:

Correspondence analysis of the categories of all families together was untaken to

look at large scale pattern and overlap between East Africa and East Asia. The CA

scatterplot (Figure 7.31) shows a division on the first axis between Felidae and species

such as Mungos dietrichi , Arctonyx minor , and Prototocyon recki . Indices such as LBM and RPS load low on the first axis. Many of the felids have high body mass and relatively small premolar size. The indices RBL and RGA are plotted on the high end of the axis with species that have relatively small blade lengths and relatively large grinding areas.

Species within families tended to plot together on the first and second axes, with members more similar to each other than to members of other families. The first axis separates hypercarnivorous flesh-eating species from omnivorous species with large molars. It also separates large species on the negative end of the axis from the smaller species on the positive end.

On the second axis, UM21, P4WL and RUMGA plot on the negative end, while

CWL and PROTO load on the positive end. Ailuropoda , which has a large amount of upper molar grinding area loads low on the second axis. CWL and PROTO tend to be high in pantherine felids and hyaenids. Taxa that have larger premolars plot higher on the axis, while those with larger molars are found in lower positions.

In this overall comparison, the most noticeable differences between the Asian and

African carnivore guilds are in the Hyaenidae. Asian Pachycrocuta differs from African

Crocuta and Hyaena . Much of this difference is due to relative size of the premolars, which is very large in the Asian species. Chasmaporthetes nitidula from Olduvai plots

229

lower on the second axis, near the felids, which is expected given that species’ felid-like

adaptations. Chasmaporthetes was also present in East Asia during this time, but insufficient information was available and it could not be included in this analysis.

Among the felids, Panthera pardus from the Upper Burgi plots lower on the second axis than P. pardus specimens from Asia. This could represent a difference between two populations of the same species. Alternatively, it could be normal variation between specimens. Due to limited availability of P. pardus craniodental specimens from East

Africa, both alternatives remain possible. Overall, felids occupy approximately the same multivariate space in the axes shown in both Africa and Asia. Ursus and Ailuropoda do not have counterparts among the East African fossil faunas sampled here. Ursus is a large omnivore. There are no large omnivores among the Carnivora in Africa during the late

Pliocene and early Pleistocene. The extinction of bears in Africa may be due to the presence of hominin species as competing large omnivores. Among the Canidae, there are differences between Canis mesomelas from Olduvai, which seems to have more grinding adaptations, Canis cf. mesomelas from Kalochoro and Natoo, which are similar to Canis teilhardi and Canis longdanensis , and Canis cf. mesomelas from the Upper

Burgi, which is similar to Nyctereutes and Megaviverra . Canis lycaonoides , from

Olduvai II, is similar to Asian species that also have highly carnivorous adaptations, including Canis brevicephalus , C. teilhardi and C. longdanensis .

On the third axis (Figure 7.32) UM21 plots high, with RUMGA, CWL and LBM.

This axis distinguishes Ailuropoda and Ursus from the other taxa. Pachycrocuta also loads high on the third axis. On the negative end of the third axis, the indices PROTO,

RPS and RBL are plotted. Taxa that are found low on the third axis include mustelids,

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herpestids, Viverra and Prionodon , Canis mesomelas from Olduvai, Chasmaporthetes and Megantereon . Many of these taxa have small relative blade lengths and small RPS values, indicating omnivory or carnivory.

Despite the overlap on the first two axes as shown in the scatterplot, the families

Felidae and Hyaenidae, divided into African and Asian groups, had statistically different centroid locations on the CA plot from all other groups (Table 7.1). For the Canidae and

Mustelidae, African and Asian guilds were not statistically different from each other. The

MDC (Mean Distance to the Centroid) shows how much difference there is between members of the guild. It was calculated for both African and Asian representatives of families (Table 7.2) and for sites (Table 7.3). Hyaenidae were the least disparate, with the smallest mean distances to the centroid, indicating that most members had similar adaptations. Chasmaporthetes specimens from East Asia did not have enough information to be included or this genus would have increased the amount of disparity in the Asian carnivore guild. Felids in East Asia had the largest MDC, probably because they are well-sampled, with Panthera, Homotherium , Megantereon, and Sivapanthera as well as smaller felids present in the sample. Canids in Africa also had a high MDC due to the presence of Prototocyon recki , which is very different from the other fossil canids.

For sites (Table 7.4), Longdan had the largest MDC, with Vulpes chikushanensis as the most distant species. Longdan has an usually large sample of carnivores. Torolutra is most the most distant taxa from the centroid at the KBS and Okote members, highlighting the differences between this type of otter and many other taxa that are preserved in the carnivore guild.

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Hypercarnivores: Large size, Pachy_Yuan relatively large Pachy_Long Large Premolars blade lengths, Pachy_Nih 0.32 small grinding Pachy_Jian Pachy_Hai areas Pachy_Xiao Honan_Long_Hai Pachy_Gong Pachy_Moh 0.24 Pachy_Wush CWL Pliocroc_Hai Eirictis_robusta Mungos_dietrichi Ultra_KBS_Loka Croc_KBS Ultra_Chari 0.16 Dietrichi_KBS_UB Arctonyx_cf._minor Croc_Kalo_Nario_Old Martes_sp. Torolutra_cf._Ougandensis Panthera_cf._pardus Ultra_OK_UB P_paleosinensis Proto Meles_leuc. Martes_sp._2 P_pardus_Jian Hyaena_UB Lutrinae Meles_chiai RBL Omnivores/ 0.08 LBM Maka_UB_KBS Torolutra_sp. Insectivores: Siva_linx. Meles_teilhardi Hyaena_Kait_Kalo RGA Large grinding P_Leo_Old Viverra_sp. Siva_pleisto P_Pardus_Gong areas, small Metailurus Mellivora_benfieldi Variabilis 0 Lupus_Nih body mass, Lynx_shansiusHomotherium_davitasvili Lycaonoides Panthera_leo Felis_teilhardi Chasma Brevi_Long small relative Homo_cren._KBS Meso_Kalo_Natoo Felis_teilhardi Martes_sp._1 Long_Long Herpestes_primitivus D_aronoki_UB Caracal_UB -0.08 C. Teilhardi Meso_Old1 Panthera_KBS Cuon/Sinicuon Panthera_pardus RPS Prionodon_sp. Prototo_Old1 Homotherium_crendatidens RUMGA Palmidens -0.16 Megantereon_Jian Megaviverra_pleistocaenicaChih_Nih Machairodus F_microta_Wush Nyctereutes Ursus_thibetanus D_piveteaui_OK P4wl Meso_KBS_UB Ursus_sp. Megantereon_nihowanensis -0.24 M_niho_Long Vulpes_Long Axis 2 Megantereon_whitei

-0.32 Large UM21 Molars Ailuropoda_microta Ailuropoda_wulingshanensis -0.4 -0.4 -0.3 -0.2 -0.1 0 0.1 0.2 0.3 0.4 0.5 Axis 1

Figure 7.31 Scatterplot of CA of carnivores from East Asia and East Africa. Note the separation between Asian and African Hyaenidae and the presence of Ursidae in East Asia, which are not present in East Africa during the period studied.

Key to symbols: Canidae from Africa: Green diamonds; Canidae from Asia: Purple circles; Felidae from Africa: Black dot; Felidae from Asia red cross; Hyaenidae from Africa: blue squares; Hyaenidae from Asia: pink squares; Ursids: blue circles; Mustelids from Africa: blue triangles; Mustelids from Asia: yellow triangles; Herpestids from Africa: blue stars; Viverrids and Prionodon from Asia: green x.

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Ailuropoda_microta

0.32 Ailuropoda_wulingshanensis

UM21 Ursus_sp. Large Premolars Ursus_thibetanus 0.24 Large molars Pachy_Yuan Pachy_Xiao_Nih Pachy_Long Pliocroc_Hai Pachy_Hai Pachy_Jian Croc_KBS Pachy_Gong 0.16 Nyctereutes Pachy_Moh Honan_Long_Hai Pachy_Wush Prototo_Old1 Dietrichi_KBS_UB Ultra_KBS_Loka P_paleosinensis Ultra_Chari RUMGA CWL 0.08 P_Leo_Old Croc_Kalo_Nario_Old Axis 3 Megaviverra_pleistocaenica LBMP_pardus_Jian Brevi_Long Metailurus Panthera_cf._pardus Homotherium_davitasviliProto Ultra_OK_UB Panthera_KBS Hyaena_Kait_Kalo Caracal_UB Felis_teilhardi 0 Lycaon RGA Sivapanthera_pleistocaenicus Teilhardi Panthera_leo Sivapanthera_linxiaensis Megantereon_Jian Felis_teilhardi Maka_UB_KBS Machairodus D_aronoki_UB Homotherium_crendatidens Hyaena_UB Meso_KBS_UB Chih_Nih Cuon Long_Long Lynx_shansius Mungos_dietrichi -0.08 Panthera_pardus Lupus_Nih Vulpes_Long Chasma Meles_teilhardi Prionodon_sp. Eirictis_robusta F_microta_Wush Variabilis Lutrinae -0.16 D_piveteaui_OK Herpestes_primitivus Arctonyx_cf._minor Megantereon_nihowanensis RPS Mellivora_benfieldiRBLMartes_sp. P4WL Meso_Old1 Torolutra_sp.Torolutra_cf._Ougandensis M_niho_Long Palmidens Meso_Kalo_Natoo Meles_cf._leucurus Megantereon_whitei Viverra_sp. Martes_sp._2 -0.24 Martes_sp._1 Meles_chiai

-0.4 -0.32 -0.24 -0.16 -0.08 0 0.08 0.16 0.24 0.32 Axis 2

Figure 7.32 Scatterplot of CA of carnivores from East Asia and East Africa, second and third axes. Key to symbols: Canidae from Africa: Green diamonds; Canidae from Asia: Purple circles; Felidae from Africa: Black dot; Felidae from Asia red cross; Hyaenidae from Africa: blue squares; Hyaenidae from Asia: pink squares; Ursids: blue circles; Mustelids from Africa: blue triangles; Mustelids from Asia: yellow triangles; Herpestids from Africa: blue stars; Viverrids and Prionodon from Asia: green x.

Summary :

Results of the carnivore ecomorphology analysis are presented here. The aims of this analysis were to answer the questions of whether carnivore guilds in East Asia and

East Africa contained animals with similar sets of adaptations and whether the ecomorphological composition of the carnivore guild changed from the Late Pliocene to

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the Early Pleistocene in East Asia. Analyses were done for each family and for carnivores

as a whole.

There are several species of large and highly carnivorous Canidae in East Asia

and one species of that type in East Africa ( Canis lycaonoides ). There appears to be a

reduction in the number of species of highly carnivorous Canis in East Asia over time.

Cuon , another highly carnivorous species, persists in East Asia. Moderately carnivorous

species are found in both Africa and East Asia, and persist in East Asia over time.

The East Asian hyaenid species Pachycrocuta brevirostris , Pliocrocuta perrieri

and Crocuta honanensis have large body masses and very large fourth premolars. They

are very different from East African Hyaena and Crocuta . After the Pliocene, the only

hyaenid of this type sampled in East Asia is Pachycrocuta. Chasmaporthetes , which is found in both Africa and Asia, could not be compared directly here because of lack of information about the Asian species. Chasmaporthetes in general has a sectorial dentition, suited to carnivory rather than bone-crushing.

Felidae differ primarily in body size. Felids have very similar dental characteristics related to hypercarnivory and ecomorphological distinctions lie in body mass differences, and in postcranial adaptations for capturing prey that were not part of this study. In East Asia, Homotherium and Megantereon , which are hypercarnivorous large

prey hunters, are present throughout the Late Pliocene to Early Pleistocene time interval.

Sivapanthera , Panthera pardus and Lynx were also consistently present and remained similar

over time in their feeding adaptations.

Ursids were extinct in Africa during the late Pliocene to Early Pleistocene. Ursus species

were omnivorous and do not appear to show significant changes in feeding adaptations or body

mass in East Asian sites over time.

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Mustelids, viverrids, herpestids and prionodontids are difficult to compare between Africa and Asia because of few specimens are available. More species were likely to have been present during the Plio-Pleistocene in both Africa and Asia that are not preserved. Many of these species would have been too small to have had a significant impact upon hominins.

The correspondence analysis of all families together shows broad separation between large hypercarnivorous animals and those that are smaller and have adaptations for omnivory or insectivory. Species with large premolars (Hyaenidae) and those with large molars (Ursidae) are also separated. Hyaenidae from East Asia and East Africa are separated in the CA scatterplot and have statistically significantly different centroids. The presence of the Ursidae in East Asia but not in Africa also represents a substantial difference in the carnivore guilds. East Africa does not have any large Carnivora filling the role of omnivore during the late Pliocene and early Pleistocene.

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Chapter 8: Discussion

Ecological Structure of Plio-Pleistocene East Africa and East Asia:

This section will present an overview of results from the ecological structure

section of the dissertation, review confounding factors and information from other studies

and discuss how the data relate to the initial questions concerning differences in

ecological structure in East Asia and East Africa and differences in hominin and non-

hominin environments within East Asia. Climatic and environmental indicators in East

Asia will be discussed. The roles of habitat similarity and hominin habitat uniformity,

particularly with regard to savannas, will be considered. Environmental reconstructions

from other initial dispersal regions will also be reviewed. The section will finish with the

implications of these data for the conditions of initial dispersal and the potential for future

projects.

Ecological Structure of Modern Sites :

Modern sites were analyzed using ecotypes in order to validate the ecological

structure model. Specifically, these analyses showed that both geography and

environmental conditions influence the placement of sites in correspondence analysis and

NMDS scatterplots. African rainforests were particularly different in comparison to the other environments sampled. Due to these differences, modern and Plio-Pleistocene localities were analyzed both with and without rainforest localities.

African modern sites tend to cluster on the scatterplot away from the Asian localities. Part of this geographic difference in multivariate placement may be due to the larger numbers of species of browsers, grazers and mixed feeders on the African

236 continent compared with other modern localities on other continents. Also, the barriers between divisions in Africa do not appear to be sufficient to prevent species from one area from colonizing another area, indicating that the species have ecological flexibility.

Modern and Ancient Sites Combined :

The faunas of modern and ancient sites were analyzed together to show the magnitude of difference between ancient sites and to aid in environmental interpretation.

The combined analysis is also useful in showing the extent to which ancient site environments can be interpreted in modern terms. Did ancient environments resemble modern savannas or subtropical forests?

Correspondence analysis showed separation between ancient and modern faunas in multivariate space, indicating ecological structure differences. The ancient sites plot with browsers, grazers and large and mega-sized mixed feeders. The modern sites plot with ecotypes such as small carnivores and omnivores, frugivores, insectivores and small terrestrial vegetation feeders. This combined ecological structure analysis indicates that the Plio-Pleistocene sites studied here were not equivalent to modern savannas or to subtropical forests, though there were similarities. Ancient faunas had many species of ungulate browsers, grazers and mixed feeders. African savannas are the nearest analogue to this. Other habitat types may have been analogous prior to human-mediated faunal extinctions. Comparisons between modern subtropical divisions and south China fossil faunas are more difficult. Modern subtropical areas are forested and south China sites have ecological indicators of closed-habitat conditions. However, they do not overlap in multivariate space. Based on the data in this project, neither savannas nor subtropical

237 forests can be said to be precise analogues to the ancient habitats in which hominins were found.

Ecological Structure in Plio-Pleistocene Africa and East Asia :

Correspondence analysis of the ecotypes of Plio-Pleistocene sites shows separation between East Asian and East African sites. Grazers of all sizes, as well as the insectivore and aquatic ecotypes plot with the East African sites, while mixed feeders and browsers, as well as small terrestrial mammals and omnivores grouped with the East

Asian sites. The correspondence analysis shows that East Asian and East African environments had different ecological structures. It also shows a large amount of ecological diversity within East Asia. In particular, most of the northern Asian sites are separated from the south China sites. The exceptions are the north China sites of

Longdan, which has a small non-carnivore fauna, and Gongwangling. The fauna of

Gongwangling has been noted for its southern elements. The southern sites plot with ecotypes such as browsers, medium and large size mixed feeders, small terrestrial vegetation feeders, small carnivore and omnivores.

There are several possible confounding factors in an analysis comparing modern and ancient sites together, and in comparing faunal assemblages deposited and collected in different conditions and locations. Potentially significant factors, including taphonomy, time averaging, and difficulties in the comparability of modern and ancient localities, are discussed below.

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Taphonomy :

Taphonomic effects may have influenced the patterns seen here. Small animals are less likely to be preserved than larger ones (Behrensmeyer 1991). Ungulates may be more likely to be preserved compared with small carnivores or omnivores, or arboreal or small terrestrial mammals. Smaller species, especially small arboreal taxa, are important for the environmental interpretation of sites. This lack of preservation may increase apparent differences between modern and ancient faunas. For example, modern subtropical forests have a diversity of smaller taxa while many of the larger species of ungulates are extinct or are very rare. In fossil faunas, the larger taxa are more likely to be preserved while the smaller species are not. Many small carnivores, omnivores, terrestrial vegetation feeders (TERSM) and arboreal vegetation feeders may have been present at the sites studied here, but not found in the fossil record.

Time-averaging :

The faunas at these sites have been time-averaged. The amount of time represented by the faunas of each site varies. Some of the species from the faunas might not have actually coexisted. Some of the herbivore species, which are very abundant in the ancient faunas, may have existed at different times. Also, the faunal assemblages may represent slightly different habitats that existed in sequentially over time. Species indicative of different habitats may indicate fluctuations in vegetation over short periods of time, as indicated by pollen evidence (Bonnefille 1995). They may also indicate habitat mosaics. In the future, additional data may allow faunal lists to be broken into smaller slices of time, but time-averaging at some temporal length will not be eliminated.

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Comparability of Modern Faunas:

There are several problems with using modern faunas to compare with the fossil faunas. These problems include differences in the amount of information available for modern species, as well as the means of obtaining that information, potential problems with the classification of modern localities, and impoverishment of some of the modern faunas. Modern species are well-known and surveys have established the large mammalian inhabitants of modern nature reserves to a high degree of accuracy. In comparison, some specimens in fossil faunas cannot be identified. Some fossils of non- comparable elements (such as antlers or horn cores and teeth) may be labeled as separate species when incomplete specimens are present in the collection. Chinese species in particular may be taxonomically oversplit, leading to false perceptions of taxonomic and ecological diversity. Moreover, a fossil fauna may not preserve all the species that were present in a paleocommunity. The level of information about modern faunas also differs from that about fossil faunas. Ecological information about body mass, diet and substrate is well-known for most modern species and was obtained from the literature. For some species, observations on the range of body masses and diet percentages are available.

Information on fossil species was obtained from calculations based upon dental materials.

Biases may exist because the information was collected in different ways.

Bailey’s ecoregion system was used to classify modern localities from Eurasia and Africa. This system identified analogous regions from both continents, such as rainforests and savannas. However, the ecological structures of the rainforests and savannas from Eurasia and Africa differ. There is also evidence of geographic

240 substructure in the overall modern fauna analysis in the way that the tropical-subtropical steppes (all found in Africa) plot with the African savannas, while Asian localities tend to group together. This geographic patterning indicates that the savannas and rainforests from Asia and Africa that were used here do not have similar ecological structures. This could indicate that Eurasian and African savannas and rainforests do not belong in the same environmental divisions despite environmental similarities in temperature and precipitation. It would also be consistent with the idea that multiple ecological structures are possible in similar environments (Lewontin 1969).

These different ecological structures may relate to differences in the history of the regions. Using systems of biomes (which grouped sites based on landforms, vegetation and climate) and Bailey’s divisions, Rodríguez (2004, 2006) found that recent communities grouped into the same environmental classification did not always have similar ecological structures. In seven out of eleven divisions, the modern faunas grouped into the same ecological division were not significantly similar in ecological structure

(Rodríguez 2006). In a wider test of the ecological structures of a variety of Holarctic communities, Rodríguez et al. (2006) found that broad scale environmental differences

(such as between moist and very arid localities) were reflected in the placement of communities in multivariate space. Statistically significant ecological structure differences were not found between many of the sites in different divisions (Rodríguez et al. 2006). Nevertheless, the ecological structure reflected general environmental trends.

Biogeography also has an effect upon ecological structure as shown in multivariate analyses (Rodríguez et al. 2006). Faunas from different biogeographic regions but from the same divisions were often separated in multivariate space on at least

241 one axis (Rodríguez et al. 2006). The content of the species pools, which vary in the amount of species diversity that they host, and the topography and landscape heterogeneity of the localities themselves may cause the biogeographic differences.

Historical differences may ultimately underlie the pattern, creating differences in the regional species pools (Ricklefs 1987, 1989, Ricklefs and Schluter 1993). Absences of taxa or ecological types, which may be due to extinction or taxa that never dispersed into certain areas, have the strongest effect upon the composition of the regional species pool

(Rodríguez et al. 2006, Nieto et al. 2005). Modern African localities, on average, have greater species richness compared with other locations, with the exception of the Oriental realm (Nieto et al. 2005). Modern African localities have greater large mammal diversity because fewer extinctions have occurred in the past compared with other regions, and because of the diversification of large mammals with body masses between 200-1000 kg in body mass (Nieto et al. 2005). Also, some modern faunas, particularly in Eurasia, may not be representative due to human hunting and population growth eradicating many native mammalian species and altering the ecological structure.

Ancient Sites and Region Size :

The two Plio-Pleistocene regions vary in size. The distance between Olduvai

Gorge and the northern side of Lake Turkana is approximately 800 km and encompasses nearly seven degrees of latitude. The East Asian sites span over 1,160 km and 17° from north to south, and about 800 km and 13° from east to west. The East Asian faunas come from areas that are presently differentiated into the Palearctic realm in the north and the

Oriental realm in the south. Coming from a smaller geographic area and within a single

242 biogeographic region, the East African Plio-Pleistocene faunas would be expected to be more similar to each other compared with the East Asian faunas that come from a larger region.

These patterns may affect the placement of modern and ancient localities in multivariate space and affect the possible conclusions. The greater diversity of African localities in the past contributed to differences in ecological structure compared with East

Asia. That same species diversity also separates modern African savannas and tropical- subtropical steppes from vegetationally and climatically comparable regions. Patterns within East Asia are consistent with closed and open habitat differences. The ecological diversity within East Asia compared with East Africa was expected given the larger size of the East Asian region. Future analyses comparing African sites from a larger geographic region would help show the amount of ecological structure variation throughout the range of hominins.

East Asia and East African Ecological Structures and Initial Hypotheses:

Hominins occurred in sites with faunas that had diverse ecological structures. The ecology of the East African hominin sites at Lake Turkana and Olduvai Gorge differs from that of the East Asian sites as a whole, and from the East Asian hominin sites in particular. This answers the question of whether hominin initial dispersal sites in East

Asia were ecologically similar to a selection of comparative sites in East Africa. They were not. The differences may relate to habitat differences between Asia and East Africa, as well as to historical differences between the regions in terms of species diversity.

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Furthermore, there is evidence that hominins were associated with several different habitats within East Asia. The ecological structures of the initial dispersal sites vary, with indications that some, such as Gongwangling and Yuanmou, occurred in more closed habitats, while Majuangou, Donggutuo and Xiaochangliang in the Nihewan were associated with a different ecological structure. Hominins occurred in ecologically diverse settings outside of Africa, rather than a single type of structure. The Asian structures are more diverse than the ones from Africa, though this is probably due to the smaller area of the African sample compared with the large area covered by the Asian sites. Differences between the Asian sites reflect environmental differences between northern and southern sites.

Environmental Data and Ecological Structure :

Environmental data from East Africa indicates climatic deterioration. Pedogenic carbonates are interpreted to show woodland to open savanna transition between 3 to 1

Ma (Cerling 1992, Cerling et al. 1988). Arid-adapted mammals appear between 2.5 and

1.8 Ma (Behrensmeyer et al. 1997). Hypsodont bovids and suids appear, indicating the expansion of open habitats, while forest taxa went extinct (Bobe and Behrensmeyer

2004). Reed (1997), using ecological structure methods, found a transition between open woodland in the Upper Burgi to scrub woodland in the KBS with more grazers, to edaphic grassland in the Okote member. However, Bonnefille (1995) recorded climatic fluctuations between cold and dry and humid during the time of the Upper Burgi and the

Okote members based on pollen evidence. The association between the African Plio-

Pleistocene habitats and presence of numerous grazing species is consistent with the

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expansion of open habitats after 2.5 Ma. East African ancient assemblages were plotted

with ecotypes including grazers of all sizes, as well as insectivores, arboreal vegetation

feeders and small terrestrial mammals. However, species from many other groups were

present, indicating that while this was a trend, other habitats existed. Humid periods

recorded in the pollen data may have contributed to the survival of closed-habitat

animals.

A similar climatic trend of aridification and cooling occurred in East Asia in the

time period studied. Aridification was due to increased northern hemisphere glaciation

(Tian et al. 2002, Ding et al. 2005) and uplift of the Tibetan plateau (Guo et al. 2002).

More detailed pollen data from loess plateau locality shows fluctuations over time,

despite an overall trend of cooling and the spread of open habitats. At approximately 2.5

Ma, the climate shifted from warm and humid to arid with an open habitat. That in turn

shifted to a forest-steppe after 1.85 Ma (Wu et al. 2007). Open steppes were not detected at this location until after 0.95 Ma. Faunal taxonomic data suggests that northern and southern China had differing regional climates during the late Pliocene, with southern

China having a more subtropical climate (Jablonski et al. 2000).

Ecological structure data supports the idea of regional climatic differentiation between north and south China after 2.5 Ma. Southern China sites (as well as

Gongwangling and Longdan) plot with browsing ecotypes of all sizes, as well as medium and large sized mixed feeders. Many of these animals are cervids, which is consistent with a more closed habitat. The southern China sites do not plot with modern subtropical forests, however. While these sites seem to have had a warm and moist climate conducive to the growth of forest and woody vegetation, they are not precise analogues for modern

245 subtropical forests in the area. These southern sites include the hominin site of Yuanmou.

Pollen from the site of Yuanmou is consistent with an interpretation of a mosaic of habitats that included forested areas as well as open habitat. Trees, including pine, and herbaceous vegetation were reported (Qian and Zhou 1991, Zhu et al. 2008).

The northern sites (with the exceptions of Gongwangling and Longdan, which has a sparse non-carnivore fauna), plot with mega-sized mixed feeders, megafauna found in steppe environments. On the second axis, they are also found with mega-sized grazers.

The faunas of the hominin sites of Xiaochangliang, Donggutuo and Majuangou are difficult to interpret because they have few species. Pollen from Majuangou suggests changes in vegetation over time. The presence of grazers and relative lack of browsing species is consistent with more open habitats in northern China. However, true open steppes may not have arisen until later in the Pleistocene. Sites in the Nihewan have many species of mixed feeders and grazers, while the species composition at Yuanmou and Gongwangling tends toward mixed feeders and browsers. While the Nihewan seems to have fewer closed-habitat indicators, it is difficult to make interpretations of habitat from dietary classifications. Mixed feeding species may live in open, closed or intermediate habitats. This is consistent with environmental data on glaciation and aridification, although more information from large faunas is needed to determine if this is a general pattern. The fact that the northern sites of Gongwangling and Longdan do not conform to this pattern could relate to either changes over time or to regional differences.

The Longdan fauna dates to the late Pliocene and was deposited before the other sites in the overall ecological structure comparison. Regional climate differences and temporal fluctuations may also be reflected in these sites’ placement. Longdan and Gongwangling

246 may represent warmer and more humid periods. Also, Gongwangling may be part of a regional fauna that included southern migrants.

Ecological diversity in initial dispersal sites :

From the ecological data examined here, Yuanmou and Gongwangling do not show the same patterns as Xiaochangliang, Donggutuo and Majuangou. Gongwangling and Yuanmou are also distinct from the fauna of the Nihewan Basin in terms of ecological structure. The Nihewan fauna sensu stricto may be a better representative of the faunas in the Nihewan region than the hominin sites located in the Nihewan Basin of

Xiaochangliang, Donggutuo and Majuangou, because it includes many more species. The distinctions between these hominin sites are shown in the correspondence analysis.

Hominins were found in a variety of ecological settings in East Asia. Environmental and ecological data from other initial dispersal sites in Asia are discussed below.

Other initial dispersal sites suggest evidence of ecological variation compared with Africa as well. The hominin site of Dmanisi in western Asia has a fauna indicating the presence of several different types of habitat in the local area. Pollen analysis

(Klopotovskaja et al., 1989; Kvavadze, 1997) shows evidence of shrubs and herbs. The large number of cervid species from Dmanisi reflects input from multiple habitat types

(Gabunia et al. 2000b). While many of the cervids in the fauna are thought to have been forest-dwellers, some species are from open area and ecotones (Gabunia et al. 2000b).

Different altitude zones from the mountains are also represented (Gabunia et al. 2000b).

Overall, the environment at Dmanisi is reconstructed as an area of shrubs and grass, with forests near the rivers and in the mountains. An ecological analysis of the Dmanisi fauna

247 based on the relationship of diet and substrate to environment suggested a grassland environment (Palmqvist 2002).

The Sangiran paleosols show an open environment (Bettis et al. 2009). The site contained a lake-margin with marshes and plants such as sedges, ferns, grass and a few trees. Sedges, grass, ferns and some trees from the nearby higher ground were interpreted as savanna-like open woodland (Bettis et al. 2009). Open woodland settings expanded as glacial periods caused aridification. The mix of different habitats would have hosted different types of mammals, which may have offered a variety of resources (Bettis et al.

2009).

Comparing these data with the ecological structure analysis and with each other is difficult, which is the reason all faunas in this project were evaluated in the same way in an explicitly comparative framework. At Dmanisi, open grassland was surrounded by forested mountains. Grazers would have inhabited the grasslands, while browsers would have lived in the forests. The fauna of Dmanisi may have a slightly different type of ecological structure compared with the northern China sites at the same latitude, depending on factors such as the dietary adaptations of the cervids and the proportions of grazers. However, Dmanisi’s high latitude and geographic position would have affected the ecological structure, differentiating it from closed habitats in southern China by lack of ecotypes such as frugivores and insectivores. In this way, Dmanisi was probably similar to Nihewan sites based on signs of open habitat, but somewhat different due to a large cervid fauna. Some of these cervids might have mixed feeding or browsing adaptations. Analysis of more sites in North China with larger faunas, would clarify the range of variation in northern sites. The open habitat savannas described from Java would

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probably have a combination of tropical ecotypes and grazers in open habitat, with some

browsing species. There would also have been input from other areas. Periodic isolation

and extinction may also have reduced the number of species present. The site in Java

would probably have shared some features with the East Asian fossil sites sampled here,

but would have probably had differences in ecological structure due to the size of the

fauna and the combination of open habitat and lower latitude.

Overall, the evidence from Dmanisi and Java points to sites in which a mix of

habitat types were present. At Dmanisi, there were both grasslands and forests, while

marsh, grassland and open woodland were present at Sangiran. Ecotype proportions in

East Asian sites show that many sites had animals that presumably inhabited open and

closed habitats, indicating that they contained mixtures of various habitats as well. The

diversity of ecological structures found in the East Asian fossil sites indicates that

hominins were able to live in a variety of ecological structures. The potential differences

between these sites and the environments described for Dmanisi and Java would be an

interesting extension of this project in the future.

Habitat similarity :

Habitat similarity has been considered to be a very important factor in determining the course of hominin dispersals. Similar habitats could relate to a variety of aspects of environment, including climate, seasonality, faunal similarity and vegetation

(open grasslands versus more closed woodland or forest).

Occupation of northern habitats in Asia (40°N at Dmanisi and the Nihewan

Basin) suggests that the level of seasonality experienced there was tolerable to hominins.

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Hominins also occupied a significant latitudinal range in Africa. The spread of savanna- like habitats may have facilitated hominin dispersal into Asia (Dennell 2004). The spread of savanna-like vegetation is thought to have signaled the availability of similar resources, such as carcasses. African species at ‘Ubeidiya may be indicative of a general opportunity for dispersal for hominins and other mammals (Tchernov 1987, 1992a, b).

However, the African species found in the Levant did not disperse into East Asia (Potts and Teague, in press). Taxonomic faunal similarity does not appear to have been important to hominins in dispersing into East Asia. Faunal similarity as measured in terms of ecological similarity in this project shows that hominins were found in various types of environments.

The distribution of hominin and non-hominin ape fossils in East Asia during the early Pleistocene has been used to argue for an environmental divide in China between densely forested sites, and those that were open. Ciochon (2009) argues that the open vegetation sites and the resources they supported were the hominin habitat, while sites in other habitats supported an ecologically different type of ape. South China has been considered an ecological and geographic barrier to dispersal of hominins into the East

Asian region (Luchterhand 1984). However, the Yuanmou hominin fossils show that hominins occurred in South China. The Qingling Mountains, which divide north China with its Palearctic species from the South China Oriental realm, were a filter than a biogeographic barrier (Rolland 2001). The fact that the ranges of northern and southern species shift around the Qingling Mountains during cold or warm climatic periods (Xu

1988) supports the idea that these mountains were not an insurmountable barrier.

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South China has been described in the anthropological literature as a subtropical

forest, which is the dominant type of vegetation in the region currently. Subtropical

forests in south China have been diagnosed on the basis of the presence of the Stegodon -

Ailuropoda fauna, a complex of co-occurring taxa that may include Ailuropoda,

Stegodon, Gigantopithecus and Pongo. The South China sites group together in the ecological structure analysis, but there is some variation in their faunal structures. Also,

some sites that do not possess the Stegodon -Ailuropoda fauna (i.e., Gongwangling and

Longdan) are placed near the Stegodon -Ailuropoda sites in multivariate space due to

ecostructural similarities. The placement probably reflects the sites’ relatively southerly

position on the continent. The Longdan fauna may reflect a warmer period when closed

habitats were more common. Gongwangling may also contain a fauna reflecting moister

and warmer time when southern elements were able to cross the Qingling Mountains. The

Stegodon -Ailuropoda faunas in the Plio-Pleistocene sites are not unique in structure

compared with all other sites in East Asia. The group of relatively closed habitat

ecostructures includes the hominin sites of Gongwangling and Yuanmou. Based on these

data, hominins in East Asia were not so habitat-specific that they were not able to exist in

any closed-habitat communities.

The faunas described as Stegodon -Ailuropoda may vary as well. For instance,

except for Mohui, all the south China faunas studied here include a species of Equus ,

showing that while many faunal elements are consistent with closed habitats, at least

some open habitat was also present. Many of these sites may have thus been mosaics.

Other Stegodon -Ailuropoda faunas (such as those that also possess Pongo and

Gigantopithecus ) may be more densely vegetated examples. Furthermore, regions such as

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South China may have varied in their habitability over time, with some climatic

fluctuations bringing more favorable conditions. Further research on the development of

faunas in South China may show other gradients in space or time in this area.

Circumstances of initial dispersal :

Dispersal ability of hominins has been tied to the presence or absence of physical

barriers to dispersal, climate or habitat occupation barriers, competition (Turner 1992),

parasite load (Bar-Yosef and Belfer-Cohen 2001) and changes within Homo resulting in new ecological roles or capabilities. Among the changes are the use of stone tool technology to access animal food, which is visible in the archaeological record around approximately 2.6 Ma (de Heinzelin et al. 1999, Domínguez-Rodrigo et al. 2005), but possibly was present earlier. Carnivory and larger body size are traits correlated to range expansion (Antón et al. 2002). Increased carnivory, facilitated by the use of tools, would have opened up additional niche space for hominins. As a social omnivore able to use tools to access flesh and marrow, hominins would have used resources in a different way compared with local species, thus increasing the chances of successful dispersal.

Competition and the hominin niche will be discussed further in the carnivore section.

The timing and route of dispersal from Africa into Asia are related and have been debated. The current consensus on the timing of initial dispersal to around 1.8 Ma reflects the convergence of recent finds and dates of sites such as Dmanisi, Java, Yuanmou and the Nihewan. However, there is insufficient evidence to rule out an earlier hominin presence. The data presented here show hominins occurred in a variety of ecological structures, even in their (currently known) initial dispersal sites. There is nothing to

252 suggest that hominin dispersal could not have occurred earlier. A test of the ecological structure of African localities from a range of geographic locations and a longer time period prior to dispersal would establish the range of ecological structure conditions to which hominins had adapted in the past. Given the range of conditions that hominins were able to tolerate, as shown by the ecological structures of the initial dispersal sites studied here, there is no evidence that hominin dispersal was dependent upon any specific environment or event, such as the spread of savannas. Though hominins may have favored open habitats compared with tropical rainforests, they also occurred in sites with closed habitats. It is more likely that the timing of hominin dispersal was tied to geological events that opened dispersal pathways out of Africa and into other regions of

Eurasia.

Future Work :

Certain areas touched upon in this work would benefit from further research. The

East Asian ecological dataset should be expanded to include more sites. Samples of older faunas would be useful to test patterns of ecological turnover over time, particularly after the onset of glaciation and to look at the effects of regionalization with more Early

Pleistocene faunas from different areas. An expanded dataset may also include new ecotype information from faunal elements for which data were limited or unavailable.

Additional sites, particularly those with larger faunas, would be helpful in understanding the ecological structure of northern China during the Plio-Pleistocene.

Ecological structure data from other initial dispersal sites, such as Dmanisi and

Java, would be used to build a picture of the ecological variation in Eurasian initial

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dispersal sites. Although environmental data from these other initial sites is available, it is

difficult to compare between sites. This project was explicitly designed to provide a

means of comparing taxonomically distinct faunas. The comparative ecotype method

could usefully be extended to other regions. Additional African fossil faunal samples will

also be needed to assess the degree of variation in hominin environments. These faunas

should come from diverse geographic regions to match Asian data, which come from

different regions. The ecological structure could then be used to look at wider trends of

environmental change as it related to hominin dispersal.

Ecomorphology of East Asian and East African Carnivores :

This section will review the main conclusions from the carnivore ecomorphology

analysis pertaining to both the question of overall similarity or differences between the

East Asian and East African carnivore guilds and the question of changes in the East

Asian guild over time. Discussion of complications resulting from small sample size and missing data follows. The implications of the results with regard to the ecomorphological estimation variables, Asian and Asian comparisons, particularly concerning

Pachycrocuta , Canis and Ursidae, habitat and scavenging, the biogeographical implications of the invasion of the carnivore guild, the effect of hominin immigration on carnivores and finally the differences between carnivore- or hominin-centered approaches will be discussed. Future work and a summary of the main conclusions will follow.

Ecomorphological Differences between Asian and African Plio-Pleistocene Carnivores :

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Although there are specific small differences between East Asian and East

African carnivores, the biggest differences are within the family Hyaenidae.

Pachycrocuta brevirostris and Crocuta honanensis differ from African Hyaena and

Crocuta . Many of the differences between them relate to premolar size, which is larger in the Asian species. This may indicate differences in bone cracking abilities, though this information was based on the P4, which is not greatly involved in bone cracking.

Chasmaporthetes progressus , for which insufficient information was available for inclusion in the correspondence analysis, probably occupied a similar ecological niche to

Chasmaporthetes nitidula in East Africa. The Felidae were most likely similar in Africa and Asia, if one accounts for the fact that some genera are present but do not have sufficient information available about them to include in the CA. Some continental differences, such as between Panthera pardus in Asia and Africa may be of significance but require more data to confirm the pattern. The felid dentition tends to be very similar in all species. Differences in ecological niche may be related to body size, habitat and locomotion. Locomotion and prey capture information are not available for Asian species at this time due to lack of postcranial specimens. Differences between canid species are unclear, although most forms have similar avatars present on the other continent. The

African species Prototocyon recki , which is insectivorous, is an exception to this. Ursids demonstrate another major difference between the carnivores of East Asia and East

Africa. Ursids are present in East Asia, but not in East Africa during the time period focused on in this study. While Ailuropoda was essentially an herbivore (despite arguments that some species may have been more omnivorous, a conclusion that could

255 not be verified here), Ursus was a large omnivore. The East African sites studied here had no large omnivores among the Carnivora.

Changes in the East Asian Carnivore Guild over Time :

Assessment of changes to the East Asian carnivore guild was focused on large carnivores from the families Felidae, Hyaenidae, Canidae and Ursidae. In the Felidae,

Canide and Hyaenide, there is some evidence of a loss of diversity between the Pliocene sites and the later Pleistocene sites. In Longdan, there are three species of large, highly carnivorous canids. In the Nihewan, only Canis chihliensis could be interpreted in this way. Other sites do not have a highly carnivorous Canis , although Cuon persists in south

China. This may signal a loss of diversity in northern China, if the Longdan canids coexisted, and if highly carnivorous forms are not found later in China. Within the Felidae, most of the larger species persist without ecomorphological changes as signaled in their dentition. Dinofelis cristata , which was present in East Asia, though not in the sites sampled, is found at sites that are thought to be from the Early Pleistocene (Werdelin and

Lewis 2001). Dinofelis cristata was probably more pantherine in its dental properties

(Werdelin and Lewis 2001). Panthera palaeosinensis is recorded from Longdan and possibly from Longgupo but is not found in the early Pleistocene, while Panthera tigris appears in the early Pleistocene. Many differences between these two large pantherines may have been related to locomotion and postcranial features. The Hyaenidae also show species loss, with neither Pliocrocuta perrieri nor Crocuta honanensis found in any of the later hominin sites. Combined with the loss of Chasmaporthetes after the Nihewan,

256 this shows that the Asian Hyaenidae in the sites examined here were dominated solely by

Pachycrocuta brevirostris , rather than a group of hyena species.

Confounding Factors :

The results may be influenced by the treatment of missing information due to incomplete specimens, missing species known to be present and missing species not preserved in the fossil record at all. To maximize the number of comparisons available, data from different specimens of the same species at the same site, and sometimes at different sites, were combined to make composite animals, which were then used in the assignment of categories and as representative values for the PCA. The values used for categorical classification and in the PCA are averages. Specimens from the same species at different sites may be used to create a composite, which has the effect of drawing those sites together artificially. Due to this procedure, temporal and geographic variation may be lost. Variation within a species at a site may also have been lost by using only the mean values. Small sample sizes may also have affected the index values. The small samples for some carnivore species may not have yielded representative index values for their species. Small sample size also made it difficult to look at the range of values possible for each species for the indices. In future work, the question of variation within the data, especially with regard to temporal or geographic trends, may be dealt with in a carnivore-guild centric project by comparing the trends of important indices.

Taxonomic uncertainty may also have affected the results. When taxonomic reviews were available, they were used to consolidate species that had been split into

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many invalid categories. However, many of the species have never been studied

comparatively. Therefore, in some cases different species names may refer to the same

species.

Some species are known to have been present at certain sites, but they could not

be analyzed due to insufficient information. Examples include Chasmaporthetes

progressus and Panthera tigris from East Asia, and Acinonyx from East Africa. In these cases, information from other sites was not available or was also insufficient to fill in the data needed for a comparison using correspondence analysis. Due to these limitations, the carnivore guilds as analyzed are incomplete, and information about trends and the roles these unrepresented species played in the carnivore guilds at each site and region must be approximated. In future work, fossil specimens of these missing species could be measured from other sites to make a more complete picture of the regional carnivore guilds.

Finally, some species were most likely present, but are not found in the fossil carnivore guilds. Species that are unlikely to be preserved or to be collected from fossil sites include those that were small in body size, species that were rare in the original fauna and species that lived in environments in which fossilization was unlikely. In this analysis, species from Viverridae, Herpestidae, Mustelidae as well as small Canidae are most likely to have been deleted from the record due to lack of preservation. These families were likely to have been more diverse during the Plio-Pleistocene. However, while these forms were important in order to understand the overall structure and function of the carnivore guild, they were unlikely to have interacted substantially with hominins.

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Interpretation :

Importance of body mass for distinguishing carnivores :

The most important indices for distinguishing carnivores were body mass (LBM) and relative premolar size (RPS), which has body mass as one of its components. Some of the particular importance of body mass may derive from the fact that certain families, like felids, have very similar dental specializations. All felids have a value of 1 for relative blade length (RBL), and a value of 0 for relative grinding area for lower molars

(RGA). Thus, sabertoothed felids, which are extremely hypercarnivorous, are not distinguished by many indices from other felids. Felids that possess similar dental adaptations are separated ecologically by body size and by postcranial adaptations for prey capture and locomotion (García and Virgos 2007, Lewis 1997). Additional data from postcrania on other traits would help distinguish species and compare African and

Eurasian specimens. Postcranial measurements would also be helpful in estimating body mass more accurately. In this project, body mass estimates were based upon dental measurements. Teeth are not weight bearing elements and the M 1 is under selection as a cutting blade and may not be the best estimator of body mass for some families, though it was the only means available of making estimates for this project. Postcranial elements typically yield better estimates of body mass for comparison of carnivore adaptations.

East Asia and East Africa compared

Pachycrocuta brevirostris , with its large body and dental size, has emerged as one of the most different aspects of the East Asian carnivore guild compared with East

Africa. Hominins would have had to compete with this animal. It would have been

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capable of destroying carcasses, and if it traveled in packs, confrontational scavenging

would have been difficult due to Pachycrocuta ’s large size. Pachycrocuta is present at

nearly every East Asian site (except for Majuangou and Donggutuo, which have very

small faunas). Though hominins would have encountered only one species acting as a

bone-cracking competitor, that species, Pachycrocuta , may have been very abundant. In

the case of high abundance, the absence of multiple species may have made little

difference for hominins in terms of the amount of scavengeable matter available.

Differences may have been evident in the lack of specialized predators on smaller

carcasses. However, the number of specimens and the widespread distribution of

Pachycrocuta may imply that scavengeable carcasses were readily available. Changes in

the East Asian carnivore guild through time complicate the picture, with Crocuta

honanensis and Pliocrocuta perrieri , both similar to Pachycrocuta in craniodental adaptations studied here, disappearing after the Pliocene. The number of niches for bone- cracking hyaenids may have been reduced, or the niches of these species may have been filled by Pachycrocuta .

East African and East Asian habitats might also differ in their canids. In ancient

East African sample, there are several species labeled Canis mesomelas (not conspecific) that appear to have been moderately carnivorous smaller canids, while there was only a single species of larger, highly carnivorous canid, Canis lycaonoides . In contrast, the East

Asian fossil record from the sample sites includes multiple species of highly carnivorous canids, such as Canis brevicephalus , Canis longdanensis , Canis teilhardi and Canis chihliensis . Three of these species are found at Longdan, which spans a long time period.

These canid species may not have been contemporaneous. It is also possible that the

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reduction of highly carnivorous Canis from the three found at Longdan to the single highly carnivorous species C. chihliensis from the Nihewan, represents a substantial

change in the East Asian carnivore guild. Several different species of highly carnivorous

Canis yielded to the single species found during the Nihewan.

Could hominins have been competitors with those species of highly carnivorous

canids in East Asia, leaving niche space for only one species of this type, similar to the

situation in East Africa? In Africa, other open country carnivores, as well as hominins,

which are also social animals, may have been sufficient competition that Canis

lycaonoides was the only species of large carnivorous canid present. In East Asia, the

extinction of large, highly carnivorous canids may have provided an opportunity for

hominins. Alternatively, pressure from hominins may have contributed to the extinction

of these forms. It is also possible that highly carnivorous species of Canis inhabited East

Asia into the early Pleistocene and that they are not recorded in the sites sampled here or in the East Asian fossil record as it is currently known. In order to better determine how and when the carnivore guild of East Asia changed with respect to Canis (and how that contrasts with Africa), more information on the first and last appearances of these species is needed.

Another important difference may relate to the role of ursids as omnivores. East

Africa did not have ursids during the focal time period. The large omnivore role was filled by some suid species, and by hominin species. Hominin species may have been so effective at filling of the large omnivore niche that ursids were not able to occupy African habitats after approximately 2.6 Ma. This again may relate to hominin tool use. Suids were also present in East Asia, though they are not well-studied. In East Asia, Homo was

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unique in the fauna as a tool-using omnivore and facultative carnivore. Still, the ursid

characteristics measured here do not show changes, and Ursus thibetanus is still present in the region, possibly indicating that the impact of hominins on that omnivore was minimal.

Pliocene to Early Pleistocene carnivore guilds

Species loss between the Pliocene and early Pleistocene sites implies a carnivore guild in transition that could have facilitated hominin colonization. Invasions of a new area are more likely when the fauna is unstable, leading to opportunities for an invader

(Brown 1989, Vermeij 1991). The colonization of East Asia by Homo may have been facilitated by the fact that the carnivore guild may have been shrinking, with fewer species of hyaenids and canids present. This would imply that fewer bone cracking species were present during the early Pleistocene. A reduction in the number of bone cracking species may have opened opportunities for hominins (Blumenschine 1987,

Blumenschine et al. 1994, Turner 1992). The loss of highly carnivorous species, such as

Chasmaporthetes and several of the large canids, would imply that fewer large carnivores filled the niches in the middle of the carnivore hierarchy. These middle-hierarchy carnivores may have been killers or scavengers of smaller prey. The large sabertoothed felids Homotherium and Megantereon would continue to be top predators, and

Pachycrocuta was most likely the top scavenger, able to displace other species from carcasses, and able to hunt large animals based on its large size. Hominin opportunities may have consisted of scavenging from carcasses that were not already taken by

Pachycrocuta , and stealing from groups of smaller carnivores.

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Habitat and scavenging opportunity :

Research on relative availability of carcasses by habitat and the amount of competition between carnivores (and thus the relative availability of carcasses for hominins) is focused on modern African carnivores and savannas. Range overlap between two or more larger species, such as lions or hyenas, resulted in full utilization of carcasses (Ayeni 1975, Blumenschine 1986, Kruuk 1972, Schaller 1972, Van

Valkenburgh 2001). Carcasses are more easily spotted in open habitat (Cavallo 1997). In

East Africa, some studies have indicated that carcasses located in open habitat were consumed quickly, leaving carcasses from the woodland areas on the landscape longer, though findings in areas other than the Serengeti conflict with these conclusions

(Blumenschine 1986, Domínguez-Rodrigo 2001). Behavioral studies also indicate that the levels of competition between carnivore species are higher in open habitat (Kruuk

1972, Schaller 1972, Sinclair 1979). However, in savannas in other African regions, such as Central Africa, which have resident ungulates and a different vegetation structure due to tall grass, carcass availability patterns are different, with more carcasses available in open areas (Tappen 1995). Increased carcass availability in modern African habitats overall may be related to locations with many lions (which leave behind utilizable carcasses) but without substantial numbers of hyenas. However, none of the ancient habitats was directly analogous ecologically to the sites where carnivore competition has been studied.

All of the East Asian sites sampled here have Pachycrocuta present, and large felids are present at many of the sites, particularly in south China. The presence (and

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possible abundance) of a very large scavenger in all of the hominin sites, may have meant

that no particular habitat had an exceptionally low number of competitors. Asia contains

fossil sites with faunas that have ecological structures that can be interpreted as primarily

open or closed habitats. In the more closed vegetation habitats where hominin remains

have been found, such as Gongwangling and Yuanmou, closed habitat predators, such as

P. tigris and Megantereon are also present. South China closed habitats may have been high-competition settings, and with Pachycrocuta present, there may not have been large amounts of scavengeable material available. More data on the locomotion and habitat preferences of the predators would be needed to look at relative amounts of competition in various parts of the sites.

Hominin versus carnivore guild centric

The ecomorphological research here was approached with the idea of looking at the differences in the carnivore guild and how that would have affected opportunities for hominins. Hominins and carnivores are linked. The adaptations and capabilities of hominins may have constrained the carnivore guilds, leading to extinctions, making hominins relevant. Research focused on the carnivore guild alone would have handled the data differently, possibly making more use of the values and ranges. Information may have included other ecological variables, as well as taxonomically relevant variables.

Analysis would have considered the functions of carnivore species within their guild in relation to each other, over time and space. Here, the focus has been on testable variation between East Africa and East Asia and on variation within East Asia over time. This comparative focus does not require complete reconstructions of the carnivore guilds.

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The effect of hominins on the carnivore guild :

Pachycrocuta brevirostris became extinct in East Africa by approximately 2.5

Ma, but persisted in South Africa until as late as 1.5 Ma (Werdelin 1999). In contrast, in

East Asia, this species existed until the Middle Pleistocene (Qiu 2006). The East African extinction occurred at approximately the same time as the first known stone tools.

Although the correspondence between the extinction of the top predator, Pachycrocuta ,

and the visibility of tools on the landscape is striking, it is not necessarily a causal

relationship. The carnivore guild was already declining in species richness after 3.3 Ma

and the extinction of Pachycrocuta may have been part of this general trend (Werdelin

and Lewis 2005). However, competitive hominins may have contributed to the decline in

carnivore species richness. Species such as Pachycrocuta persist for much longer in East

Asia. This raises the question of the effect of hominins on the carnivore guild. If

hominins were partially responsible for the extinctions in the African carnivore guild,

with which they coexisted for a long period of time, how would the East Asian carnivores

have been affected by the intrusion of hominins into the carnivore guild? Some effects

may have been delayed as many carnivore species in East Asia, such as Homotherium

and Pachycrocuta , persist until the Middle Pleistocene when hominins in East Asia may

have been more numerous. Further work including a longer time span would be necessary

to look at long-term interactions of hominins and carnivores in this fauna.

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Future Work :

Future work on the ecomorphological traits of carnivores would involve adding information to the current database of carnivore traits. Information will be obtained by measuring specimens of carnivores from species for which there are currently no data.

Sites from earlier in the Pliocene will be added in order to look at ecological change in the carnivore guild. Glaciation began around 2.8 to 2.5 Ma. Data from the time prior to the glacial cycles will be required to examine ecological turnovers that may have occurred. With the additional data, patterns of ecological loss could further tested. The ecological patterns shown in this project may be only part of a larger event related to climate change. Additional data from a longer time span may clarify patterns. Ecological structure and carnivore ecomorphology data combined from a Pliocene to Pleistocene time span would show how changes in the carnivore guild related to changes in large mammals as a whole, and how that may have correlated with environmental shifts.

Another avenue of comparison between East Asian and East African carnivores would be the inclusion of postcranial data, thus including inferences about locomotion and prey capture. Although there are no postcranial remains in China for the carnivore species,

European specimens from the same species may be used as proxies. This information may provide differentiation among the members of the families of Felidae and Canidae, for example.

Summary :

The main finding of this project with regard to overall ecological structure of mammalian faunas is that Plio-Pleistocene East Africa and East Asia were ecologically

266 different. The numbers of grazing species in East Africa compared with the number of browsers and mixed feeders in East Asia drove the differences. Neither the fossil sites in

East Asia nor those in East Africa were similar in ecological structure to modern environments, including habitats such as savannas and subtropical forests. From the diversity of ecological structures found in East Asia, and found in Asian hominin sites, it is evident that hominins occurred in a variety of ecological settings during their first known dispersal.

A number of factors may have confounded the results of this analysis. These factors include taphonomic bias, which may have resulted in fewer small bodied species being preserved, time-averaging of faunas, differences in the region size included by the

Plio-Pleistocene East African and East Asian sites.

East Asian ecological structure is consistent with environmental data that suggests differentiation between north and south China during the early Pleistocene.

Concentration of browsers in South China and lack of many browsing species in many of the North China sites, such as in the Nihewan, is consistent with northern hemisphere glaciation and aridification.

As with the initial dispersal sites within East Asia that include a variety of ecological structures, the initial dispersal sites outside of East Asia, Dmanisi and

Sangiran, also appear to have different environments compared with East Africa, showing that Homo was capable of colonizing disparate ecological settings.

The results bear on two theories of hominin dispersal in East Asia. Ciochon

(2009) hypothesized that hominins were excluded from forested southern China habitats with Stegodon-Ailuropoda faunas. This analysis shows that the sites with Stegodon-

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Ailuropoda faunas are part of a larger subset of sites that plot with a concentration of browsing ecotypes and are consistent with closed habitat. Two of those closed habitat

sites have hominin remains, indicating that hominins were not excluded from that general

habitat. Some of the sites may also have had habitat mosaics rather than a single type of

vegetation structure. The general differences in ecological structure in hominin sites

make it unlikely that hominins were strictly dependent upon savanna habitats, though

savannas may have facilitated their dispersal. Hominin dispersal does not appear to have

been dependent upon the spread of particular habitats.

Future work on this part of the project will involve expanding the ecological

structure dataset. In particular, the East Asian data will be expanded to include more

sites, particularly from the Pliocene and from north China, to look at the possibility of

structural turnover between the late Pliocene and early Pleistocene and to better

understand the ecological structures in North China, respectively. Ecological data from

other Asian initial sites would enable better comparison of the circumstances of hominin

dispersal. Finally, expansion of the African data would show the variation in African

settings.

The analysis of carnivore ecomorphology shows that while many of the

carnivores were largely similar ecomorphologically between Asia and Africa, there were

several important differences. Pachycrocuta brevirostris differed from East African

Crocuta and Hyaena in indices related to premolar size. The presence of the family

Ursidae in the East Asian carnivore guild also distinguishes the two regions. East Africa

had no large omnivores from the Carnivora during this period. The East Asian carnivore

guild shows some change over time, with the number of species of large, bone-cracking

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hyaenids decreasing to include only Pachycrocuta . There was also a reduction in the number of hypercarnivorous canid species from the Late Pliocene to the Early

Pleistocene.

Despite the reduction of species diversity in the Hyaenidae, Pachycrocuta is frequently found in East Asian early Pleistocene sites. Due to its large size and ability to consume a carcass completely, Pachycrocuta brevirostris would have been a formidable competitor for hominins. The reduction of the number of canid species may also have resulted in ecological opportunities for hominins. With the widespread Pachycrocuta

present, there may not have been low-competition settings for carnivores in East Asia.

Hominins coexisted with the East African carnivore guild for a long time.

However, it is possible that interactions between carnivores and hominins led to

extinctions in the African carnivore guild after 3.3 Ma. Hominin dispersal to East Asia

led to contact with carnivores that had never interacted with hominins before. These

interactions may have affected the Asian carnivore guild. Some species that went extinct

in East Africa, such as Pachycrocuta brevirostris , persisted for a much longer time in

East Asia. The overall effect of hominins on the carnivore guild is unknown.

In the future, data on carnivores over a longer time period will be collected. The

database will also aim to fill in data from missing species and to add data from more

traits, including possibly locomotor and prey capture characteristics.

In summary, the study of the ecological settings of hominin dispersal shows that

Homo occurred in diverse ecological structures. Hominins in East Asia interacted with a

carnivore guild that differed in several ways from that in East Africa, possibly including

more competition from bone-cracking hyaenids.

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Appendices

Table A6.1 Measureable specimens of Canidae from East Asian sites are listed with museum accession number and available teeth. Only adult specimens were measured. Species Number Site Mandibular Maxillary Dentition Dentition Canis variabilis V2936a Gongwangling P2, M 1-M2 Canis variabilis V2936b Gongwangling M1-M3 Canis variabilis V2936c Gongwangling C-P4 Canis variabilis V2936d Gongwangling P3-M2 Canis variabilis V2936e Gongwangling M1-M2 Canis variabilis V2936f Gongwangling M2 Canis variabilis V2936g Gongwangling M1 Canis variabilis V2936h Gongwangling M1 Canis variabilis V2936i Gongwangling P1-P2 Canis variabilis V2936j Gongwangling M1 Canis variabilis V2936k Gongwangling I1-I3 Cuon dubius V13398.1 Jianshi P2 Cuon dubius V13398.2 Jianshi M1 Cuon dubius V13398.3 Jianshi M1 Cuon dubius V13398.4 Jianshi M2 Cuon dubius V13399 Jianshi P2 Cuon dubius V13402.1 Jianshi P3 Cuon dubius V13402.2 Jianshi P4 Cuon dubius V13402.3 Jianshi M1 Cuon dubius V13402.4 Jianshi M1 Cuon dubius V13403.1 Jianshi C Cuon dubius V13403.3 Jianshi P3 Cuon dubius V13403.4 Jianshi P3 Cuon dubius V13403.5 Jianshi P4 Cuon dubius V13404.1 Jianshi C Cuon dubius V13404.10 Jianshi P4 Cuon dubius V13404.11 Jianshi M2 Cuon dubius V13404.2 Jianshi C Cuon dubius V13404.2 Jianshi P2 Cuon dubius V13404.3 Jianshi P3 Cuon dubius V13404.4 Jianshi P4 Cuon dubius V13404.5 Jianshi P4 Cuon dubius V13404.6 Jianshi P4 Cuon dubius V13404.7 Jianshi M1 Cuon dubius V13404.8 Jianshi M1 Cuon dubius V13404.9 Jianshi P3 Canis sp . V6751.1 Linyi M1 (BROKEN) Canis sp . V6751.2 Linyi M1 Canis sp . V6751.3 Linyi C Canis brevicephalus HMV1172 Longdan I3- M2 Canis teilhardi HMV 101 Longdan I1- M2 Canis teilhardi HMV 1145 Longdan I3-C; P 2- M2 Canis teilhardi HMV 1147 Longdan I1-M2 2 2 Canis teilhardi HMV 1149 Longdan I2-M2 I - M Canis teilhardi HMV 1150 Longdan I1- M2 Canis teilhardi HMV 1151 Longdan I1-I3; P 1- M2 Canis teilhardi HMV 1153 Longdan I3- M2 Canis teilhardi HMV 1154 Longdan I1-I3; P1- M2 Canis teilhardi HMV 1155 Longdan P1; P 3- M2

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Species Number Site Mandibular Maxillary Dentition Dentition Canis teilhardi HMV 1156 Longdan I1- M2 Canis teilhardi HMV 1157 Longdan I2-C; P 2-P3; M 1 Canis teilhardi HMV 1162 Longdan P2-M1 Canis teilhardi HMV 1164 Longdan P1-M3 Canis teilhardi HMV 1165 Longdan I2-M1 Canis teilhardi HMV 1167 Longdan I1-M2 1 2 Canis teilhardi HMV 1168 Longdan I1-M3 I - M 1 2 3 2 Canis teilhardi HMV 1169 Longdan I1-C; P 2-M2 I -I ; C; P - M Canis teilhardi HMV 1173 Longdan C- M2 Canis teilhardi HMV 1174 Longdan I1- M2 Canis teilhardi HMV 1176 Longdan P2-M2 Canis teilhardi HMV 1177 Longdan I1-M2 Canis teilhardi HMV 1178 Longdan I2; C-M2 Canis teilhardi HMV 1179 Longdan I1-M2 Canis teilhardi HMV 1180 Longdan I2-P3; M 2 Canis teilhardi HMV 1181 Longdan C-P2; P 4-M2 Canis teilhardi HMV 1182 Longdan I2-C; P 2-M2 Canis teilhardi HMV 1183 Longdan I1-C; P 2-M2 Canis teilhardi HMV 1185 Longdan C-M3 Sinicuon dubius V13543 Longdan M1 3 2 Vulpes chikushanensis V13533 Longdan I1-M3 I -M Cuon dubius CV 892 Longgupo P4- M1 Cuon dubius CV 893 Longgupo P4 Cuon dubius CV 895 Longgupo M1 Cuon dubius CV 896 Longgupo M1 Cuon dubius CV 896.1 Longgupo M2 Cuon dubius MH 269 Mohui M1 Canis chihliensis TNP 161 Nihewan I2-I3; P 2-M3 Canis chihliensis TNP 162 Nihewan I1-M2 Canis chihliensis TNP 163 Nihewan I3-P1; P 3-M3 Canis lupus TNP 18752 Nihewan P3-M3 Canis lupus TNP 18753 Nihewan M2 Canis lupus TNP 18798 Nihewan C-M2 Canis palmidens TNP 18754 Nihewan C-M1 Canis palmidens TNP 190 Nihewan I1-C; P 2-M2 Nyctereutes sinensis TNP 167 Nihewan I1-I3; P 1-M2 Nyctereutes sinensis TNP 171 Nihewan C-P4 Nyctereutes sinensis TNP 197 Nihewan I1-M2

Table A6.2 Measureable specimens of Felidae from East Asian sites are listed with museum accession number and available teeth. Only adult specimens were measured.

Species Museum Site Mandibular Maxillary Number Dentition Dentition Megantereon nihowanensis V2979.1 Gongwangling C Panthera cf. tigris V5421 Gongwangling P4 Megantereon nihowanensis RV 45019 Haiyan I1-C; P 3 Homotherium davitasvili RV 45016 Haiyan? I1-C; P 4- M1 Homotherium davitasvili RV 45017 Haiyan? I1-I3; P 3- M1 Homotherium davitasvili RV 45018 Haiyan? I1-C Felis teilhardi V13443.1 Jianshi M1 Felis teilhardi V13443.2 Jianshi M1 Felis teilhardi V13443.3 Jianshi P4

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Species Museum Site Mandibular Maxillary Number Dentition Dentition Homotherium sp. V13438.1 Jianshi C Homotherium sp. V13438.2 Jianshi C Homotherium sp. V13438.3 Jianshi P4 Homotherium sp. V13439 Jianshi M1 Megantereon sp. V13440.1 Jianshi C Panthera pardus V13444.1 Jianshi P3 Panthera pardus V13444.2 Jianshi P4 Panthera pardus V13444.3 Jianshi P4 Panthera pardus V13444.4 Jianshi P4 Panthera pardus V13444.5 Jianshi M1 Panthera pardus V13444.6 Jianshi M1 Panthera pardus V13445 Jianshi P3- P4 Panthera pardus V13446.1 Jianshi P4 Panthera pardus V13446.2 Jianshi P4 Panthera pardus V13446.3 Jianshi P4 Panthera pardus V13446.4 Jianshi P4 Panthera pardus V13446.5 Jianshi M1 Panthera pardus V13446.6 Jianshi M1 Panthera pardus V13446.8 Jianshi M1 Panthera pardus V13446.9 Jianshi C Panthera pardus V13447.1 Jianshi P4 Panthera pardus V13447.2 Jianshi M1 Panthera pardus V13448.2 Jianshi P4 Panthera pardus V13448.3 Jianshi M1 Panthera pardus V13448.4 Jianshi M1 Panthera pardus V13448.4 Jianshi M1 Panthera pardus V13449.1 Jianshi P3 Panthera pardus V13449.3 Jianshi M1 Panthera pardus V13450.1 Jianshi P4 Panthera pardus V13450.2 Jianshi P4 Panthera pardus V13450.3 Jianshi P4 Panthera pardus V13450.5 Jianshi P4 Panthera pardus V13450.6 Jianshi M1 Panthera pardus V13450.7 Jianshi M1 Panthera pardus V13450.X Jianshi P4 Sivapanthera pleistocaenicus V13451 Jianshi P4 Sivapanthera pleistocaenicus V13452 Jianshi P4 Sivapanthera pleistocaenicus V13453.1 Jianshi P3 Sivapanthera pleistocaenicus V13453.2 Jianshi P3 Sivapanthera pleistocaenicus V13453.3 Jianshi P3 Homotherium crenatidens HMV 1213 Longdan I1-C; P 4 Lynx shansius HMV 1226 Longdan I1-M1 1 3 1 Lynx shansius HMV 1228 Longdan I1-C; P 3- M1 I -C; P -M Lynx shansius HMV 1230 Longdan C- P4 Lynx shansius HMV 1232 Longdan I1-C; P 3- M1 Lynx shansius HMV 1233 Longdan I1- P4 Lynx shansius HMV 1234 Longdan I1- M1 Lynx shansius HMV 1236 Longdan C- M1 Lynx shansius M290 Longdan I1- P4 Lynx shansius V 13539 Longdan P3 Lynx shansius V13540 Longdan I1-C; P 3- P4 Megantereon nihowanensis HMV 1215 Longdan P3-M1 Megantereon nihowanensis HMV 1220 Longdan I1-M1 Megantereon sp. MB001 Longdan I3-C; P 3-M1

294

Species Museum Site Mandibular Maxillary Number Dentition Dentition Panthera palaeosinensis V13538 Longdan I1-C; P 2- P3 Sivapanthera linxiaensis V13536 Longdan C; P 2- P4 Sivapanthera linxiaensis V13537 Longdan I1-C; P 4 Felis microta CV908 Longgupo I3- M1 Felis teilhardi CV909 Longgupo M1 Homotherium crenatidens CV809 Longgupo P4 Homotherium crenatidens CV894 Longgupo P4 Homotherium crenatidens CV894B Longgupo P4 Panthera pardus CV904.2 Longgupo M1 Panthera pardus CV904.4 Longgupo M1 Panthera pardus CV904.5 Longgupo M1 Sivapanthera pleistocaenicus CV903.1 Longgupo C Sivapanthera pleistocaenicus CV903.2 Longgupo C Sivapanthera pleistocaenicus CV903.3 Longgupo C Sivapanthera pleistocaenicus CV904.1 Longgupo C Sivapanthera pleistocaenicus CV904.2 Longgupo C Sivapanthera pleistocaenicus CV904.3 Longgupo C Sivapanthera pleistocaenicus CV905 Longgupo C Felis teilhardi MH156 Mohui M1 Felis teilhardi MH381 Mohui M1 Homotherium crenatidens T32067 Nihewan C-M1 Megantereon nihowanensis TNP 175 Nihewan P3- M1

Table A6.3 Measureable specimens of Hyaenidae from East Asian sites are listed with museum accession number and available teeth. Only adult specimens were measured.

Species Number Site Mandibular Maxillary Dentition Dentition Pachycrocuta brevirostris V2939.1 Gongwangling P1-M1 Pachycrocuta brevirostris V7293 Haiyan P2-P4 Pliocrocuta perrieri V7290 Haiyan I2-C; P 2-M1 Pliocrocuta perrieri V7291 Haiyan I3; P 3-M1 Pachycrocuta brevirostris V13423.1B Jianshi P2-P4 Pachycrocuta brevirostris V13423.2 Jianshi P1, P 3 Pachycrocuta brevirostris V13423.3 Jianshi M1 Pachycrocuta brevirostris V13425.10 Jianshi P4 Pachycrocuta brevirostris V13425.13 Jianshi P3 Pachycrocuta brevirostris V13425.14 Jianshi P3 Pachycrocuta brevirostris V13425.15 Jianshi P4 Pachycrocuta brevirostris V13425.16 Jianshi P4 Pachycrocuta brevirostris V13425.17 Jianshi M1 Pachycrocuta brevirostris V13425.18 Jianshi M1 Pachycrocuta brevirostris V13425.19 Jianshi M1 Pachycrocuta brevirostris V13425.4 Jianshi P2 Pachycrocuta brevirostris V13425.4 Jianshi M1 Pachycrocuta brevirostris V13425.5 Jianshi C, P 2 Pachycrocuta brevirostris V13425.6 Jianshi C Pachycrocuta brevirostris V13425.8 Jianshi P3 Pachycrocuta brevirostris V13426 Jianshi P4 Pachycrocuta brevirostris V13427.1 Jianshi C, P 2-M1 Pachycrocuta brevirostris V13427.15 Jianshi C Pachycrocuta brevirostris V13427.16 Jianshi C Pachycrocuta brevirostris V13427.18 Jianshi C

295

Species Number Site Mandibular Maxillary Dentition Dentition Pachycrocuta brevirostris V13427.19 Jianshi C Pachycrocuta brevirostris V13427.2 Jianshi P3-M1 Pachycrocuta brevirostris V13427.20 Jianshi C Pachycrocuta brevirostris V13427.21 Jianshi C Pachycrocuta brevirostris V13427.24 Jianshi P2 Pachycrocuta brevirostris V13427.25 Jianshi P2 Pachycrocuta brevirostris V13427.26 Jianshi P2 Pachycrocuta brevirostris V13427.27 Jianshi P2 Pachycrocuta brevirostris V13427.28 Jianshi P2 Pachycrocuta brevirostris V13427.29 Jianshi P2 Pachycrocuta brevirostris V13427.3 Jianshi C, P 2, P 4 Pachycrocuta brevirostris V13427.30 Jianshi P3 Pachycrocuta brevirostris V13427.31 Jianshi P3 Pachycrocuta brevirostris V13427.33 Jianshi P3 Pachycrocuta brevirostris V13427.34 Jianshi P3 Pachycrocuta brevirostris V13427.35 Jianshi P3 Pachycrocuta brevirostris V13427.36 Jianshi P3 Pachycrocuta brevirostris V13427.37 Jianshi P4 Pachycrocuta brevirostris V13427.38 Jianshi P4 Pachycrocuta brevirostris V13427.4 Jianshi P4-M1 Pachycrocuta brevirostris V13427.40 Jianshi P4 Pachycrocuta brevirostris V13427.41 Jianshi P4 Pachycrocuta brevirostris V13427.42 Jianshi P4 Pachycrocuta brevirostris V13427.43 Jianshi P4 Pachycrocuta brevirostris V13427.45 Jianshi C Pachycrocuta brevirostris V13427.46 Jianshi C Pachycrocuta brevirostris V13427.47 Jianshi C Pachycrocuta brevirostris V13427.48 Jianshi P3 Pachycrocuta brevirostris V13427.49 Jianshi P3 Pachycrocuta brevirostris V13427.50 Jianshi P3 Pachycrocuta brevirostris V13427.51 Jianshi P3 Pachycrocuta brevirostris V13427.52 Jianshi P3 Pachycrocuta brevirostris V13427.53 Jianshi P3 Pachycrocuta brevirostris V13427.54 Jianshi P4 Pachycrocuta brevirostris V13427.55 Jianshi P4 Pachycrocuta brevirostris V13427.56 Jianshi P4 Pachycrocuta brevirostris V13427.57 Jianshi P4 Pachycrocuta brevirostris V13427.58 Jianshi P4 Pachycrocuta brevirostris V13427.59 Jianshi P4 Pachycrocuta brevirostris V13427.60 Jianshi P4 Pachycrocuta brevirostris V13427.62 Jianshi M1 Pachycrocuta brevirostris V13427.63 Jianshi M1 Pachycrocuta brevirostris V13427.64 Jianshi M1 Pachycrocuta brevirostris V13427.65 Jianshi M1 Pachycrocuta brevirostris V13427.66 Jianshi M1 Pachycrocuta brevirostris V13428 Jianshi P4 Pachycrocuta brevirostris V13429.1 Jianshi C Pachycrocuta brevirostris V13429.2 Jianshi P3 Pachycrocuta brevirostris V13430.10 Jianshi P3-P4 Pachycrocuta brevirostris V13430.11 Jianshi P2 Pachycrocuta brevirostris V13430.12 Jianshi M1 Pachycrocuta brevirostris V13430.4 Jianshi P2 Pachycrocuta brevirostris V13430.5 Jianshi P3 Pachycrocuta brevirostris V13430.7 Jianshi P3

296

Species Number Site Mandibular Maxillary Dentition Dentition Pachycrocuta brevirostris V13431.1 Jianshi P2-M1 Pachycrocuta brevirostris V13431.2 Jianshi P3-P4 Pachycrocuta brevirostris V13431.4 Jianshi C Pachycrocuta brevirostris V13431.5 Jianshi P4 Pachycrocuta brevirostris V13431.6 Jianshi M1 Pachycrocuta brevirostris V13431.7 Jianshi M1 Pachycrocuta brevirostris V13432.1 Jianshi P3 Pachycrocuta brevirostris V13432.10 Jianshi P3 Pachycrocuta brevirostris V13432.11 Jianshi P4 Pachycrocuta brevirostris V13432.13 Jianshi P4 Pachycrocuta brevirostris V13432.18 Jianshi P2 Pachycrocuta brevirostris V13432.19 Jianshi P2 Pachycrocuta brevirostris V13432.2 Jianshi P4-M1 Pachycrocuta brevirostris V13432.20 Jianshi P2 Pachycrocuta brevirostris V13432.21 Jianshi P2 Pachycrocuta brevirostris V13432.22 Jianshi P3 Pachycrocuta brevirostris V13432.23 Jianshi P3 Pachycrocuta brevirostris V13432.24 Jianshi M1 Pachycrocuta brevirostris V13432.25 Jianshi M1 Pachycrocuta brevirostris V13432.26 Jianshi M1 Pachycrocuta brevirostris V13432.4 Jianshi C Pachycrocuta brevirostris V13432.5 Jianshi C Pachycrocuta brevirostris V13432.6 Jianshi C Pachycrocuta brevirostris V13432.7 Jianshi C Pachycrocuta brevirostris V13432.8 Jianshi C Pachycrocuta brevirostris V13432.9 Jianshi P2 Pachycrocuta brevirostris V13433.1 Jianshi P2 Pachycrocuta brevirostris V13433.2 Jianshi P3 Pachycrocuta brevirostris V13433.3 Jianshi P3 Pachycrocuta brevirostris V13433.4 Jianshi P3 Pachycrocuta brevirostris V13433.4 Jianshi P4 Pachycrocuta brevirostris V13433.5 Jianshi P4 Pachycrocuta brevirostris V13433.7 Jianshi M1 Pachycrocuta brevirostris V13434 Jianshi P4 Pachycrocuta brevirostris V13436.1 Jianshi C Pachycrocuta brevirostris V13436.5 Jianshi P3 Pachycrocuta brevirostris V13436.7 Jianshi P4 Pachycrocuta brevirostris V13437.1 Jianshi P2 Pachycrocuta brevirostris V13437.2 Jianshi M1 Pachycrocuta brevirostris V134XX.1 Jianshi P4 Chasmaporthetes progressus HMV 1197 Longdan I1-P4 Chasmaporthetes progressus HMV 1198 Longdan I1-C; P 2-P4 Crocuta honanensis HMV 1205 Longdan I1-M1 Crocuta honanensis HMV 1207 Longdan C; P 2-M1 4 Crocuta honanensis HMV 236 Longdan C; P 2-M1 C-P 1 1 Crocuta honanensis V13535 Longdan P3 I -M Pachycrocuta brevirostris HMV 1200 Longdan I1-I3; P 1-M1 Pachycrocuta brevirostris HMV 1201 Longdan C; P 2-M1 Pachycrocuta brevirostris HMV 1203 Longdan I1-M1 Pachycrocuta brevirostris V13551 Longdan I1-M1 Pachycrocuta brevirostris CV789.4 Longgupo M1 Pachycrocuta brevirostris CV876.1 Longgupo P4 Pachycrocuta brevirostris CV876.10 Longgupo P4 Pachycrocuta brevirostris CV876.11 Longgupo P4

297

Species Number Site Mandibular Maxillary Dentition Dentition Pachycrocuta brevirostris CV876.12 Longgupo P4 Pachycrocuta brevirostris CV876.3 Longgupo P4 Pachycrocuta brevirostris CV876.4 Longgupo P4 Pachycrocuta brevirostris CV876.6 Longgupo P4 Pachycrocuta brevirostris CV876.8 Longgupo P4 Pachycrocuta brevirostris CV876.9 Longgupo P4 Pachycrocuta brevirostris CV877.11 Longgupo P3 Pachycrocuta brevirostris CV877.12 Longgupo P3 Pachycrocuta brevirostris CV877.13 Longgupo P3 Pachycrocuta brevirostris CV877.2 Longgupo P3 Pachycrocuta brevirostris CV877.3 Longgupo P3 Pachycrocuta brevirostris CV877.4 Longgupo P3 Pachycrocuta brevirostris CV877.5 Longgupo P3 Pachycrocuta brevirostris CV877.6 Longgupo P3 Pachycrocuta brevirostris CV877.7 Longgupo P3 Pachycrocuta brevirostris CV877.8 Longgupo P3 Pachycrocuta brevirostris CV878.1 Longgupo P4 Pachycrocuta brevirostris CV878.2 Longgupo P4 Pachycrocuta brevirostris CV878.3 Longgupo P4 Pachycrocuta brevirostris CV878.4 Longgupo P4 Pachycrocuta brevirostris CV878.5 Longgupo P4 Pachycrocuta brevirostris CV878.6 Longgupo P4 Pachycrocuta brevirostris CV878.7 Longgupo P4 Pachycrocuta brevirostris CV878.8 Longgupo P4 Pachycrocuta brevirostris CV878.9 Longgupo P4 Pachycrocuta brevirostris CV879.16 Longgupo M1 Pachycrocuta brevirostris CV879.18 Longgupo M1 Pachycrocuta brevirostris CV879.3 Longgupo M1 Pachycrocuta brevirostris CV879.5 Longgupo M1 Pachycrocuta brevirostris CV879.6 Longgupo M1 Pachycrocuta brevirostris CV879.7 Longgupo M1 Pachycrocuta brevirostris CV879.8 Longgupo M1 Pachycrocuta brevirostris CV879.9 Longgupo M1 Pachycrocuta brevirostris CV881 Longgupo C Pachycrocuta brevirostris CV882.1 Longgupo P2 Pachycrocuta brevirostris CV882.10 Longgupo P2 Pachycrocuta brevirostris CV882.11 Longgupo P2 Pachycrocuta brevirostris CV882.12 Longgupo P2 Pachycrocuta brevirostris CV882.2 Longgupo P2 Pachycrocuta brevirostris CV882.3 Longgupo P2 Pachycrocuta brevirostris CV882.4 Longgupo P2 Pachycrocuta brevirostris CV882.5 Longgupo P2 Pachycrocuta brevirostris CV882.6 Longgupo P2 Pachycrocuta brevirostris CV882.7 Longgupo P2 Pachycrocuta brevirostris CV882.9 Longgupo P2 Pachycrocuta brevirostris CV883.1 Longgupo P2 Pachycrocuta brevirostris CV883.2 Longgupo P2 Pachycrocuta brevirostris CV883.3 Longgupo P2 Pachycrocuta brevirostris CV883.4 Longgupo P2 Pachycrocuta brevirostris CV883.5 Longgupo P2 Pachycrocuta brevirostris CV883.6 Longgupo P2 Pachycrocuta brevirostris CV884.1 Longgupo P3 Pachycrocuta brevirostris CV884.10 Longgupo P3 Pachycrocuta brevirostris CV884.11 Longgupo P3

298

Species Number Site Mandibular Maxillary Dentition Dentition Pachycrocuta brevirostris CV884.12 Longgupo P3 Pachycrocuta brevirostris CV884.13 Longgupo P3 Pachycrocuta brevirostris CV884.14 Longgupo P3 Pachycrocuta brevirostris CV884.15 Longgupo P3 Pachycrocuta brevirostris CV884.16 Longgupo P3 Pachycrocuta brevirostris CV884.2 Longgupo P3 Pachycrocuta brevirostris CV884.3 Longgupo P3 Pachycrocuta brevirostris CV884.4 Longgupo P3 Pachycrocuta brevirostris CV884.5 Longgupo P3 Pachycrocuta brevirostris CV884.6 Longgupo P3 Pachycrocuta brevirostris CV884.7 Longgupo P3 Pachycrocuta brevirostris CV884.8 Longgupo P3 Pachycrocuta brevirostris CV884.9 Longgupo P3 Pachycrocuta brevirostris CV903.2 Longgupo P4 Pachycrocuta brevirostris TNP 229 Nihewan P3-M1 Pachycrocuta brevirostris TNP 276 Nihewan P2-M1 Pachycrocuta brevirostris TNP 277 Nihewan C; P 3; M 1 Pachycrocuta brevirostris TNP 278 Nihewan I1-I3; P 1-P4 Pachycrocuta brevirostris TNP 68 Nihewan C; P 3-P4 Pachycrocuta brevirostris V4033 Yuanmou C-M1 Pachycrocuta brevirostris V5315 Yuanmou C; P 2-M1

Table A6.4 Measureable specimens of Ursidae from East Asian sites are listed with museum accession number and available teeth. Only adult specimens were measured. Species Number Site Mandibular Maxillary Dentition Dentition Ailuropoda melanoleuca V5412 Gongwangling M1 Ailuropoda wulingshanensis V13457 Jianshi P2-M2 Ursus sp. V13405.1 Jianshi P4 Ursus sp. V13405.2 Jianshi M1 Ursus sp. V13405.3 Jianshi M2 Ursus sp. V13405.4 Jianshi M1 Ursus sp. V13405.5 Jianshi M2 Ursus sp. V13405.6 Jianshi M2 Ursus sp. V13405.7 Jianshi M3 Ursus sp. V13405.8 Jianshi M3 Ursus sp. V13406.1 Jianshi M2 Ursus sp. V13406.2 Jianshi M3 Ursus sp. V13407.1 Jianshi M1 Ursus sp. V13408.1 Jianshi P4 Ursus sp. V13408.2 Jianshi M1 Ursus sp. V13408.3 Jianshi M1 Ursus sp. V13408.4 Jianshi M1 Ursus sp. V13408.5 Jianshi M2 Ursus sp. V13409.2 Jianshi M2 Ursus sp. V13410.1 Jianshi M1 Ursus sp. V13410.2 Jianshi M3 Ursus sp. V13411.1 Jianshi C; M 2 Ursus sp. V13411.10 Jianshi P4 Ursus sp. V13411.11 Jianshi M1 Ursus sp. V13411.12 Jianshi M1 Ursus sp. V13411.13 Jianshi M1 Ursus sp. V13411.13 Jianshi M2

299

Species Number Site Mandibular Maxillary Dentition Dentition Ursus sp. V13411.14 Jianshi M2 Ursus sp. V13411.15 Jianshi M2 Ursus sp. V13411.16 Jianshi C Ursus sp. V13411.17 Jianshi C Ursus sp. V13411.18 Jianshi C Ursus sp. V13411.19 Jianshi C Ursus sp. V13411.20 Jianshi C Ursus sp. V13411.21 Jianshi C Ursus sp. V13411.22 Jianshi C Ursus sp. V13411.23 Jianshi M1 Ursus sp. V13411.24 Jianshi M1 Ursus sp. V13411.25 Jianshi M1 Ursus sp. V13411.27 Jianshi M2 Ursus sp. V13411.28 Jianshi M3 Ursus sp. V13411.29 Jianshi M3 Ursus sp. V13411.3 Jianshi C; P 3 Ursus sp. V13411.4 Jianshi C Ursus sp. V13411.5 Jianshi C Ursus sp. V13411.6 Jianshi C Ursus sp. V13411.7 Jianshi C Ursus sp. V13411.8 Jianshi C Ursus sp. V13411.9 Jianshi P4 Ursus sp. V13411.X Jianshi C Ursus sp. V13412.1 Jianshi M1 Ursus sp. V13412.2 Jianshi M1 Ursus sp. V13412.3 Jianshi M1 Ursus sp. V13412.4 Jianshi M2 Ursus sp. V13413.1 Jianshi P4 Ursus sp. V13413.2 Jianshi M1 Ursus sp. V13413.3 Jianshi M1 Ursus sp. V13413.4 Jianshi M1 Ursus sp. V13414.1 Jianshi C Ursus sp. V13414.10 Jianshi M2 Ursus sp. V13414.11 Jianshi M2 Ursus sp. V13414.12 Jianshi M2 Ursus sp. V13414.13 Jianshi C Ursus sp. V13414.14 Jianshi C Ursus sp. V13414.16 Jianshi C Ursus sp. V13414.17 Jianshi C Ursus sp. V13414.18 Jianshi M1 Ursus sp. V13414.2 Jianshi C Ursus sp. V13414.20 Jianshi M2 Ursus sp. V13414.21 Jianshi M2 Ursus sp. V13414.22 Jianshi M2 Ursus sp. V13414.23 Jianshi M3 Ursus sp. V13414.4 Jianshi M1 Ursus sp. V13414.5 Jianshi M1 Ursus sp. V13414.6 Jianshi M2 Ursus sp. V13414.8 Jianshi M2 Ursus sp. V13414.9 Jianshi M2 Ursus sp. V13415.1 Jianshi M1 Ursus sp. V13415.2 Jianshi M2 Ursus sp. V13416.1 Jianshi M1-M2 Ursus sp. V13416.2 Jianshi M2

300

Species Number Site Mandibular Maxillary Dentition Dentition Ursus sp. V13416.3 Jianshi M2 Ursus sp. V134XX.2 Jianshi P4 Ursus sp. V6752 Linyi C Ailuropoda microta CV921.1 Longgupo M1 Ailuropoda microta CV921.3 Longgupo M1 Ailuropoda microta CV921.4 Longgupo M1 Ailuropoda microta CV921.5 Longgupo M1 Ailuropoda microta CV921.6 Longgupo M1 Ailuropoda microta CV922.1 Longgupo M2 Ailuropoda microta CV922.2 Longgupo M2 Ailuropoda microta CV922.3 Longgupo M2 Ailuropoda microta CV922.4 Longgupo M2 Ailuropoda microta CV922.6 Longgupo M2 Ailuropoda microta CV922.7 Longgupo M2 Ailuropoda microta CV922.8 Longgupo M2 Ailuropoda microta CV923.1 Longgupo P4 Ailuropoda microta CV923.2 Longgupo P4 Ailuropoda microta CV923.3 Longgupo P4 Ailuropoda microta CV923.4 Longgupo P4 Ailuropoda microta CV923.5 Longgupo P4 Ailuropoda microta CV923.6 Longgupo P4 Ailuropoda microta CV924.1 Longgupo P3 Ailuropoda microta CV924.2 Longgupo P3 Ailuropoda microta CV924.3 Longgupo P3 Ailuropoda microta CV924.4 Longgupo P3 Ailuropoda microta CV925.1 Longgupo P2 Ursus aff. thibetanus CV885.2 Longgupo P4 Ursus aff. thibetanus CV885.3 Longgupo P4 Ursus aff. thibetanus CV885.4 Longgupo P4 Ursus aff. thibetanus CV885.5 Longgupo P4 Ursus aff. thibetanus CV885.6 Longgupo P4 Ursus aff. thibetanus CV885.7 Longgupo P4 Ursus aff. thibetanus CV885.8 Longgupo P4 Ursus aff. thibetanus CV886.1 Longgupo M1 Ursus aff. thibetanus CV886.10 Longgupo M1 Ursus aff. thibetanus CV886.11 Longgupo M1 Ursus aff. thibetanus CV886.12 Longgupo M1 Ursus aff. thibetanus CV886.13 Longgupo M1 Ursus aff. thibetanus CV886.2 Longgupo M1 Ursus aff. thibetanus CV886.3 Longgupo M1 Ursus aff. thibetanus CV886.4 Longgupo M1 Ursus aff. thibetanus CV886.5 Longgupo M1 Ursus aff. thibetanus CV886.6 Longgupo M1 Ursus aff. thibetanus CV886.7 Longgupo M1 Ursus aff. thibetanus CV886.8 Longgupo M1 Ursus aff. thibetanus CV886.9 Longgupo M1 Ursus aff. thibetanus CV887.1 Longgupo M2 Ursus aff. thibetanus CV887.2 Longgupo M2 Ursus aff. thibetanus CV887.4 Longgupo M2 Ursus aff. thibetanus CV887.5 Longgupo M2 Ursus aff. thibetanus CV887.6 Longgupo M2 Ursus aff. thibetanus CV888 Longgupo C Ursus aff. thibetanus CV888.1 Longgupo C Ursus aff. thibetanus CV889.1 Longgupo M2

301

Species Number Site Mandibular Maxillary Dentition Dentition Ursus aff. thibetanus CV889.10 Longgupo M2 Ursus aff. thibetanus CV889.13 Longgupo M2 Ursus aff. thibetanus CV889.2 Longgupo M2 Ursus aff. thibetanus CV889.3 Longgupo M2 Ursus aff. thibetanus CV889.4 Longgupo M2 Ursus aff. thibetanus CV889.5 Longgupo M2 Ursus aff. thibetanus CV889.6 Longgupo M2 Ursus aff. thibetanus CV889.7 Longgupo M2 Ursus aff. thibetanus CV889.9 Longgupo M2 Ursus aff. thibetanus CV890.1 Longgupo M3 Ursus aff. thibetanus CV890.10 Longgupo M3 Ursus aff. thibetanus CV890.11 Longgupo M3 Ursus aff. thibetanus CV890.2 Longgupo M3 Ursus aff. thibetanus CV890.3 Longgupo M3 Ursus aff. thibetanus CV890.4 Longgupo M3 Ursus aff. thibetanus CV890.5 Longgupo M3 Ursus aff. thibetanus CV890.6 Longgupo M3 Ursus aff. thibetanus CV890.7 Longgupo M3 Ursus aff. thibetanus CV890.8 Longgupo M3 Ursus aff. thibetanus CV890.9 Longgupo M3 4 2 Ursus aff. thibetanus CV891 Longgupo M3 P -M Ursus aff. thibetanus CV897 Longgupo M1 Ursus aff. thibetanus CV898.1 Longgupo M1 Ursus aff. thibetanus CV898.2 Longgupo M1 Ailuropoda microta MH0021 Mohui M3 Ailuropoda microta MH0023 Mohui M3 Ailuropoda microta MH051 Mohui P4 Ursus sp. MH580 Mohui M1 Ursus sp. MH607 Mohui M1

Table A6.5 Measureable specimens of Mustelidae from East Asian sites are listed with museum accession number and available teeth. Only adult specimens were measured.

Species Number Site Mandibular Maxillary Dentition Dentition Meles leucurus V5411 Gongwangling C; P 2-M1 Arctonyx collaris V13421.1 Jianshi M1 Arctonyx collaris V13421.1 Jianshi M1 Lutra sp. V13422.1 Jianshi P4 Lutra sp. V13422.2 Jianshi M1 Martes sp. 1 V13417 Jianshi P2-M1 Martes sp. 2 V13418 Jianshi P4 Martes sp. 2 V13419 Jianshi M1 Martes sp. 2 V13420 Jianshi M1 Eirictis robusta HMV 1191 Longdan I1-M1 Eirictis robusta HMV 1192 Longdan I1-M2 Meles teilhardi HMV 1193 Longdan I1-M1 1 1 Meles teilhardi HMV 1194 Longdan I1- M1 I -M Meles teilhardi HMV 1195 Longdan I1-M1 Meles teilhardi MB005 Longdan I1-P4 Meles teilhardi V13534 Longdan I1-M1 Arctonyx cf. minor CV900 Longgupo P3-P4 Arctonyx cf. minor CV902 Longgupo C-M2

302

Species Number Site Mandibular Maxillary Dentition Dentition Meles cf. chiai CV899 Longgupo P4-M1 Meles cf. chiai CV901 Longgupo M1 Meles cf. chiai CV901.1 Longgupo M1 Meles cf. chiai CV901B Longgupo M1 Arctonyx collaris MH418 Mohui M1 Arctonyx collaris MH550 Mohui M1 Martes sp. MH 218 Mohui M1 Martes sp. MH 3000B Mohui M1 Meles chiai TNP 165 Nihewan C; M 1 Meles chiai TNP 321 Nihewan C-M1

Table A6.6 Measureable specimens of Viverridae and Prionodontidae from East Asian sites are listed with museum accession number and available teeth. Only adult specimens were measured.

Species Number Site Mandibular Maxillary Dentition Dentition Prionodon sp. V13456 Jianshi P4-M2 Viverra sp. V13454 Jianshi M1 Viverra sp. V13455 Jianshi P4 Megaviverra pleistocaenica CV913.2 Longgupo P4 Megaviverra pleistocaenica CV914.2 Longgupo M1 Megaviverra pleistocaenica CV914.3 Longgupo M1 Megaviverra pleistocaenica CV914.4 Longgupo M1 Megaviverra pleistocaenica CV915.3 Longgupo M2 Megaviverra pleistocaenica CV915.4 Longgupo M2 Megaviverra pleistocaenica CV916.1 Longgupo P4 Megaviverra pleistocaenica CV916.2 Longgupo P4 Paguma sp. MH366 Mohui P4 Paguma sp. MH427 Mohui M1 Paguma sp. MH591 Mohui P4 Viverricula malaccensis V5313A Yuanmou P1-P3 Viverricula malaccensis V5313B Yuanmou P3-P4 Viverricula malaccensis V5313C Yuanmou P4-M1 Viverricula malaccensis V5313D Yuanmou M1

Table A6.7 Measureable specimens of Canidae from East African sites (divided by Turkana Basin member or Olduvai Bed) are listed with museum accession number and available teeth. Only adult specimens were measured.

Species Number Member Mandibular Maxillary or Bed Dentition Dentition Canis cf. C. mesomelas WT 14664 Kalochoro C, P 1-P4, M 1-M2 Canis cf. C. mesomelas ER 668 KBS C, P 3 Canis cf. C. mesomelas ER 895 KBS P4-M2 Canis cf. C. mesomelas ER 332 KF P3, P 4, M 2 Canis mesomelas OLD 018 Olduvai I M2 Canis mesomelas OLD 068/6520 Olduvai I M1 Canis mesomelas OLD 102 Olduvai I P4 Canis mesomelas OLD 2671 Olduvai I M2 Canis mesomelas OLD 2902 Olduvai I P4 1 2 Canis mesomelas OLD 358 Olduvai I C-M3 P -M 303

Species Number Member Mandibular Maxillary or Bed Dentition Dentition Canis mesomelas OLD 60 Olduvai I P4-M2 Canis mesomelas OLD 6234 Olduvai I P4 Canis mesomelas OLD 6237 Olduvai I C-M1 Canis mesomelas OLD 6252 Olduvai I P4-M1 Canis mesomelas OLD 7304 Olduvai I C-M3 Canis mesomelas OLD b99a Olduvai I P3 Canis mesomelas OLD b99b Olduvai I M1 Canis mesomelas OLD b99c Olduvai I M2 Canis mesomelas OLD b99d Olduvai I P3 Canis mesomelas OLD b99e Olduvai I P1 Canis mesomelas OLD c999 Olduvai I P4-M2 Canis mesomelas OLD K14 Olduvai I P4-M2 Otocyon OLD 1314 Olduvai I P2 Otocyon OLD 1314b Olduvai I M2-M3 Otocyon OLD 960 Olduvai I P2-P3 Protocyon recki OLD 308 Olduvai I C-M2 Protocyon recki OLD 9317 Olduvai I C-M1 Prototocyon recki OLD 6275 Olduvai I C-M3 Prototocyon recki OLD 761 Olduvai I C-M3 Canis lycaonoides BM.M 15017 Olduvai II M1-M3 Canis lycaonoides BM.M 15018 Olduvai II M2-M3 Canis lycaonoides OLD 1C Olduvai II C-P4 2 Canis lycaonoides OLD 74 Olduvai II C-M3 C-M Canis cf. C. mesomelas ER 3767 UB C, P 1-M2 Canis sp. ER 3755 UB M2-M3 Canis sp. ER 44951 UB P2 Vulpes cf. V. zerda ER 3121 UB-KBS P1-P2

Table A6.8 Measureable specimens of Felidae from East African sites (divided by Turkana Basin member or Olduvai Bed) are listed with museum accession number and available teeth. Only adult specimens were measured.

Species Number Member or Bed Mandibular Maxillary Dentition Dentition Homotherium WT 17436 Kaitio P3-P4 Homotherium ER 44059 KBS I1 Homotherium ER 931 KBS P3-M1 Homotherium ? ER 44607 KBS P4 Panthera ? (not P. leo ) ER 44496 KBS P4 Dinofelis piveteaui ER 40482A Okote P3-M1 Dinofelis piveteaui ER 40482B Okote P3-P4 Dinofelis piveteaui ER 40482C Okote P4-M1 Dinofelis piveteaui ER 40482F Okote I1-I3 Dinofelis piveteaui ER 40482G Okote C Dinofelis piveteaui ER 666 Okote C, P 3 Felidae ER 44535 Okote P3 1 3 3 1 Megantereon whitei ER 793 Okote I2-I3, C, P 3-M1 I -I , C, P -M Panthera leo ER 874 Okote P4 Felidae (small) OLD 1d Olduvai 1 P3 Dinofelis D OLD 2642 Olduvai I I3 “Machairodus” OLD 6158 Olduvai II C (Homotherium ?) Metailurus OLD 2482 Olduvai II C 304

Species Number Member or Bed Mandibular Maxillary Dentition Dentition Panthera cf. leo OLD 1273A Olduvai II C, P 3-M1 Panthera cf. leo OLD 1273B Olduvai II P3-M1 Panthera leo OLD 1776 Olduvai II C Panthera leo OLD 301 Olduvai II C Panthera leo OLD 302 Olduvai II C Panthera leo OLD 303 Olduvai II I3 Acinonyx ER 3740 UB P3 cf. Caracal ER 3758 UB C, M 1 Dinofelis aronoki ER 1549 UB I1-I3, C, P 3-M1 Homotherium ER 1540 UB I3 Homotherium WT 37401 UB P3 Panthera pardus ER 3848 UB C, P 3-M1 Panthera sp. ER 44652 UB M1

Table A6.9 Measureable specimens of Hyaenidae from East African sites (divided by Turkana Basin member or Olduvai Bed) are listed with museum accession number and available teeth. Only adult specimens were measured.

Species Number Member Mandibular Maxillary or Bed Dentition Dentiton Hyaena sp. WT 17434 Kaitio P4-M1 Crocuta crocuta WT 14995 Kalachoro P3-P4 Hyaena sp. WT 14991 Kalachoro P2-M1 Crocuta sp. ER 28520 KBS P2-P3, M 1 Crocuta sp. ER 44346 KBS C Crocuta sp. ER 44346a KBS C Crocuta sp. ER 44433 KBS P3 1 4 Crocuta ultra ER 10078 KBS C, P 2-M1 C, P - P Crocuta ultra ER 358 KBS C, P 2-P4 Crocuta ultra ER 4387 KBS P2-P4 Crocuta ultra ER 4967 KBS P3-P4 Crocuta ultra ER 940 KBS P3-P4 Hyaena sp.? ER 1666 KBS P3, M 1 Crocuta ultra ER 1659 KBS-OK P2 Hyaena sp. ER 23094 KF C, P 2-M1 Hyaena sp. ER 44713 KF P4 4 Crocuta ultra WT 39209 Lokalalei P2-P4 P Crocuta crocuta WT 14989 Nariokotome C Crocuta sp. ER 40417 Okote P3 Crocuta ultra ER 360 Okote C, P 2-P3 Crocuta ultra ER 367 Okote P2-P3 Crocuta ultra ER 4421 Okote P3-P4 Crocuta ultra ER 667 Okote P3 Crocuta ultra ER 694 Okote P3 Crocuta ultra ER 723 Okote P2-M1 Hyaena sp. ER 40415 Okote P4 Crocuta crocuta OLD 263 Olduvai I P2-M1 Crocuta crocuta OLD 3131 Olduvai I P 3 Crocuta crocuta OLD 5349 Olduvai I P4 Crocuta crocuta OLD 5396 Olduvai I M1 Crocuta crocuta OLD 7577 Olduvai I C, P 3-M1 Crocuta crocuta OLD 7642 Olduvai I C, P 2-M1 Chasmaporthetes nitidula OLD a900 Olduvai II C; P 2-M1 305

Species Number Member Mandibular Maxillary or Bed Dentition Dentiton Crocuta crocuta OLD 1510 Olduvai II P3-P4 Hyaena hyaena OLD 2609 Olduvai II P2-M1 Crocuta cf. dietrichi ER 3745 UB C, P 2-P4 Crocuta dietrichi ER 3109 UB P2, P 4, M 1 Crocuta dietrichi ER 3753 UB P2 Crocuta dietrichi ER 721 UB C, P 2-M1 Crocuta dietrichi ? ER 1541 UB P3-P4 Crocuta dietrichi ? ER 3765 UB P2-P4 1 1 Hyaena makapani ER 1548A,B UB C, P 2-M1 C, P -M 1 1 Hyaena sp. ER 3766 UB P1-M1 C, P -M Hyaena sp. ER 669 UB P3 Crocuta ultra ER 896 UB-KBS P3-P4

Table A6.10 Measureable specimens of Mustelidae from East African sites (divided by Turkana Basin member or Olduvai Bed) are listed with museum accession number and available teeth. Only adult specimens were measured.

Species Number Site Mandibular Maxillary Dentition Dentition Torolutra ER 44462 KBS P3-M1 Enhydriodon ER 44722 Upper Burgi M1 Lutrinae ER 4568 Upper Burgi P2-M1 Mellivora benfieldi ER 3760 Upper Burgi M1 Torolutra cf. ougandensis ER 1486 Upper Burgi M1-M2 Torolutra cf. ougandensis ER 1486b Upper Burgi M1 Torolutra ? ER 5895 Upper Burgi P3-M1

Table A6.11 Measureable specimens of Herpestidae from East African sites (divided by Turkana Basin member or Olduvai Bed) are listed with museum accession number and available teeth. Only adult specimens were measured.

Species Number Member Lower Parts Upper Parts or Bed Galerella delibis OLD 2635 Olduvai I M1 Galerella primitivus OLD 764 Olduvai I P4 Galerella primitivus OLD 764b Olduvai I P4-M2 Mungos dietrichi OLD 018 Olduvai I C, M 2 Mungos dietrichi OLD 1A Olduvai I P3, P 4 Mungos dietrichi OLD 6128 Olduvai I C, P 2-M2 Mungos minutus OLD 762 Olduvai I P2, M 1 Mungos minutus OLD 763 Olduvai I M1 Atilax OLD 1349 Olduvai II M1 Atilax OLD 1b Olduvai II M1

306

Table A6.12 Measureable specimens of Viverridae from East African sites (divided by Turkana Basin member or Olduvai Bed) are listed with museum accession number and available teeth. Only adult specimens were measured.

Species Number Member Lower Parts Upper Parts or Bed Pseudocivetta ingens ER 2134 KBS M1 Pseudocivetta ingens ER 44505 KBS M1 Viverridae ER 5339 KBS I1- P4 Pseudocivetta ingens OLD 1F Olduvai M1 Viverridae OLD 1294 Olduvai P3-P4 Pseudocivetta ingens OLD 1J Olduvai I I3 2 Pseudocivetta ingens OLD 840 Olduvai II M2 M Pseudocivetta sp. ER 3105 Upper Burgi I3, C Pseudocivetta sp. ER 3757 Upper Burgi M1 Viverridae ER 3749 Upper Burgi P1-M1 Viverridae ER 878 Upper Burgi P4-M1 Viverridae WT 39333 Upper Burgi C, P 2-P3, M 1

307